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Along with sugar accumulation and colour development, softening is an important physiological change during the onset of ripening in fruits. In this work, we investigated the relationships among major events during softening in grape (Vitis vinifera L.) by quantifying elasticity in individual berries. In addition, we delayed softening and inhibited sugar accumulation using a mechanical growth-preventing treatment in order to identify processes that are sugar and/or growth dependent. Ripening processes commenced on various days after anthesis, but always at similarly low elasticity and turgor. Much of the softening occurred in the absence of other changes in berry physiology investigated here. Several genes encoding key cell wall-modifying enzymes were not up-regulated until softening was largely completed, suggesting softening may result primarily from decreases in turgor. Similarly, there was no decrease in solute potential, increase in sugar concentration, or colour development until elasticity and turgor were near minimum values, and these processes were inhibited when berry growth was prevented. Increases in abscisic acid occurred early during softening and in the absence of significant expression of the V. vinifera 9-cis-epoxycarotenoid dioxygenases. However, these increases were coincident with decreases in the abscisic acid catabolite diphasic acid, indicating that initial increases in abscisic acid may result from decreases in catabolism and/or exogenous import. These data suggest that softening, decreases in turgor, and increases in abscisic acid represent some of the earliest events during the onset of ripening. Later, physical growth, further increases in abscisic acid, and the accumulation of sugar are integral for colour development.

Premise of the Study The stem of Vitis vinifera, a climbing vine of global economic importance, is characterized by both wide and narrow vessels and high specific hydraulic conductivity. While the effect of drought stress has been studied in 1‐ and 2‐yr‐old stems, there are few data documenting effects of drought stress on the anatomical structure of the mature, woody stem near the base of the vine. Here we describe mature wood anatomical responses to two irrigation regimes on wood anatomy and specific hydraulic conductivity in Vitis vinifera Merlot vines. Methods For 4 years, irrigation was applied constantly at low, medium, or high levels, or at alternating levels at two different periods during the growing season, either early spring or late summer, resulting in late season or early spring deficits, respectively. The following variables were measured: trunk diameter, annual ring width and area, vessel diameter, specific hydraulic conductivity and stem water potential. Key Results High water availability early in the season (late deficit) resulted in vigorous vegetative growth (greater trunk diameter, ring width and area), wider vessels and increased specific hydraulic conductivity. High water availability early in the season caused a shift of the vessel population towards the wider frequency classes. These late deficit vines showed more negative water potential values late in the season than vines that received low but relatively constant irrigation. Conclusions We concluded that high water availability during vegetative growth period of Vitis increases vessels diameter and hydraulic conductivity and causes the vines to be more vulnerable to drought stress late in the season.

Across plants, leaves exhibit profound diversity in shape. As a single leaf expands, its shape is in constant flux. Plants may also produce leaves with different shapes at successive nodes. In addition, leaf shape varies among individuals, populations and species as a result of evolutionary processes and environmental influences. Because leaf shape can vary in many different ways, theoretically, the effects of distinct developmental and evolutionary processes are separable, even within the shape of a single leaf. Here, we measured the shapes of > 3200 leaves representing > 270 vines from wild relatives of domesticated grape (Vitis spp.) to determine whether leaf shapes attributable to genetics and development are separable from each other. We isolated latent shapes (multivariate signatures that vary independently from each other) embedded within the overall shape of leaves. These latent shapes can predict developmental stages independent from species identity and vice versa. Shapes predictive of development were then used to stage leaves from 1200 varieties of domesticated grape (Vitis vinifera), revealing that changes in timing underlie leaf shape diversity. Our results indicate that distinct latent shapes combine to produce a composite morphology in leaves, and that developmental and evolutionary contributions to shape vary independently from each other.

Background The mountainous region between the Caucasus and China is considered to be the center of domestication for grapevine. Despite the importance of Central Asia in the history of grape growing, information about the extent and distribution of grape genetic variation in this region is limited in comparison to wild and cultivated grapevines from around the Mediterranean basin. The principal goal of this work was to survey the genetic diversity and relationships among wild and cultivated grape germplasm from the Caucasus, Central Asia, and the Mediterranean basin collectively to understand gene flow, possible domestication events and adaptive introgression. Results A total of 1378 wild and cultivated grapevines collected around the Mediterranean basin and from Central Asia were tested with a set of 20 nuclear SSR markers. Genetic data were analyzed (Cluster analysis, Principal Coordinate Analysis and STRUCTURE) to identify groups, and the results were validated by Nei’s genetic distance, pairwise F ST analysis and assignment tests. All of these analyses identified three genetic groups: G1, wild accessions from Croatia, France, Italy and Spain; G2, wild accessions from Armenia, Azerbaijan and Georgia; and G3, cultivars from Spain, France, Italy, Georgia, Iran, Pakistan and Turkmenistan, which included a small group of wild accessions from Georgia and Croatia. Wild accessions from Georgia clustered with cultivated grape from the same area ( proles pontica ), but also with Western Europe ( proles occidentalis ), supporting Georgia as the ancient center of grapevine domestication. In addition, cluster analysis indicated that Western European wild grapes grouped with cultivated grapes from the same area, suggesting that the cultivated proles occidentalis contributed more to the early development of wine grapes than the wild vines from Eastern Europe. Conclusions The analysis of genetic relationships among the tested genotypes provided evidence of genetic relationships between wild and cultivated accessions in the Mediterranean basin and Central Asia. The genetic structure indicated a considerable amount of gene flow, which limited the differentiation between the two subspecies. The results also indicated that grapes with mixed ancestry occur in the regions where wild grapevines were domesticated.

Water deficits are known to alter fine root structure and function, but little is known about how these responses contribute to differences in drought resistance across grapevine rootstocks. The ways in which water deficit affects root anatomical and physiological characteristics were studied in two grapevine rootstocks considered as low–medium (101-14Mgt) and highly (110R) drought resistant. Rootstocks were grown under prolonged and repeated drying cycles or frequent watering (‘dry’ and ‘wet’ treatments, respectively), and the following parameters were evaluated: root osmotic and hydrostatic hydraulic conductivity (Lp os and Lp hyd, respectively), suberization, steady-state root pressure (P rs), sap exudation rates, sap osmotic potential, and exosmotic relaxation curves. For both rootstocks, the ‘dry’ treatment reduced fine root Lp, elicited earlier root suberization and higher sap osmotic potential, and generated greater P rs after rewatering, but the rootstocks responded differently under these conditions. Lp os, Lp hyd, and sap exudation rates were significantly higher in 110R than in 101-14Mgt, regardless of moisture treatment. Under ‘dry’ conditions, 110R maintained a similar Lp os and decreased the Lp hyd by 36% compared with ‘wet’ conditions, while both parameters were decreased by at least 50% for 101-14Mgt under ‘dry’ conditions. Interestingly, build-up of P rs in 110R was 34% lower on average than in 101-14Mgt, suggesting differences in the development of suberized apoplastic barriers between the rootstocks as visualized by analysis of suberization from fluorescence microscopy. Consistent with this pattern, 110R exhibited the greatest exosmotic Lp os (i.e. Lp os of water flowing from roots to the soil) as determined from relaxation curves under wet conditions, where back-flow may have limited its capacity to generate positive xylem pressure. The traits studied here can be used in combination to provide new insights needed for screening drought resistance across grapevine rootstocks.

Terroir, the unique interaction between genotype, environment, and culture, is highly refined in domesticated grape (Vitis vinifera). Toward cultivating terroir, the science of ampelography tried to distinguish thousands of grape cultivars without the aid of genetics. This led to sophisticated phenotypic analyses of natural variation in grape leaves, which within a palmate-lobed framework exhibit diverse patterns of blade outgrowth, hirsuteness, and venation patterning. Here, we provide a morphometric analysis of more than 1,200 grape accessions. Elliptical Fourier descriptors provide a global analysis of leaf outlines and lobe positioning, while a Procrustes analysis quantitatively describes venation patterning. Correlation with previous ampelography suggests an important genetic component, which we confirm with estimates of heritability. We further use RNA-Seq of mutant varieties and perform a genome-wide association study to explore the genetic basis of leaf shape. Meta-analysis reveals a relationship between leaf morphology and hirsuteness, traits known to correlate with climate in the fossil record and extant species. Together, our data demonstrate a genetic basis for the intricate diversity present in grape leaves. We discuss the possibility of using grape leaves as a breeding target to preserve terroir in the face of anticipated climate change, a major problem facing viticulture.

Background and Aims Predicting both plant water status and leaf gas exchange under various environmental conditions is essential for anticipating the effects of climate change on plant growth and productivity. This study developed a functional-structural grapevine model which combines a mechanistic understanding of stomatal function and photosynthesis at the leaf level (i.e. extended Farqhuhar-von Caemmerer-Berry model) and the dynamics of water transport from soil to individual leaves (i.e. Tardieu-Davies model). [br/] Methods The model included novel features that account for the effects of xylem embolism (f(PLC)) on leaf hydraulic conductance and residual stomatal conductance (g(0)), variable root and leaf hydraulic conductance, and the microclimate of individual organs. The model was calibrated with detailed datasets of leaf photosynthesis, leaf water potential, xylem sap abscisic acid (ABA) concentration and hourly whole-plant transpiration observed within a soil drying period, and validated with independent datasets of whole-plant transpiration under both well-watered and water-stressed conditions. [br/] Key Results The model well captured the effects of radiation, temperature, CO2 and vapour pressure deficit on leaf photosynthesis, transpiration, stomatal conductance and leaf water potential, and correctly reproduced the diurnal pattern and decline of water flux within the soil drying period. In silico analyses revealed that decreases in g(0) with increasing f(PLC) were essential to avoid unrealistic drops in leaf water potential under severe water stress. Additionally, by varying the hydraulic conductance along the pathway (e.g. root and leaves) and changing the sensitivity of stomatal conductance to ABA and leaf water potential, the model can produce different water use behaviours (i.e. iso- and anisohydric). [br/] Conclusions The robust performance of this model allows for modelling climate effects from individual plants to fields, and for modelling plants with complex, non-homogenous canopies. In addition, the model provides a basis for future modelling efforts aimed at describing the physiology and growth of individual organs in relation to water status.

To better understand water uptake patterns in root systems of woody perennial crops, we detailed the developmental anatomy and hydraulic physiology along the length of grapevine (Vitis berlandieri × Vitis rupestris) fine roots from the tip to secondary growth zones. Our characterization included the localization of suberized structures and aquaporin gene expression and the determination of hydraulic conductivity (Lp r ) and aquaporin protein activity (via chemical inhibition) in different root zones under both osmotic and hydrostatic pressure gradients. Tissue-specific messenger RNA levels of the plasma membrane aquaporin isogenes (VvPIPs) were quantified using laser-capture microdissection and quantitative polymerase chain reaction. Our results highlight dramatic changes in structure and function along the length of grapevine fine roots. Although the root tip lacked suberization altogether, a suberized exodermis and endodermis developed in the maturation zone, which gave way to the secondary growth zone containing a multilayer suberized periderm. Longitudinally, VvPIP isogenes exhibited strong peaks of expression in the root tip that decreased precipitously along the root length in a pattern similar to Arabidopsis (Arabidopsis thaliana) roots. In the radial orientation, expression was always greatest in interior tissues (i.e. stele, endodermis, and/or vascular tissues) for all root zones. High Lp r and aquaporin protein activity were associated with peak VvPIP expression levels in the root tip. This suggests that aquaporins play a limited role in controlling water uptake in secondary growth zones, which contradicts existing theoretical predictions. Despite having significantly lower Lp r , woody roots can constitute the vast majority of the root system surface area in mature vines and thus provide for significant water uptake potential.

The cultivation of grapevines has spanned millennia, leading to thousands of varieties through exchanges, mutations, and crosses between genotypes, as well probably as gene flow from wild populations. These varieties are typically categorized by regional origin and primary use, either for wine production or fruit consumption. France, within the Western European group, hosts many of the world's renowned wine grape varieties. However, the historical development of cultivated grapevines in France and in the world remains poorly understood. This study applies morphometry on 19,377 charred and waterlogged archaeological grape pips to investigate the evolutionary history of grapevine in France over the last 10,000 years. The study compares seed outlines and lengths, corrected for taphonomic distortions, with a reference collection of 80 wild and 466 modern domestic grapevine accessions. Findings reveal a shift from wild grapevine exploitation to the expansion of domestic varieties around 600-500 BCE, coinciding with Mediterranean cultural influences and the introduction of eastern grape types. The identification of the East-Table group, a group of varieties of eastern origin for fruit consumption, indicates that grapes were also grown for food, especially in Mediterranean regions and near urban areas, alongside wine production. Early French viticulture featured a notable presence of Western European wine-type grapevines. The abundance of pips with wild-like morphology suggests early cultivation involved plants at an initial domestication stage and gene flow between introduced and wild grapevines. As viticulture spread northward, wild and Eastern morphotypes declined, leading to the dominance of Western European wine types in inner France during the Middle Ages.

This study presents an extensive analysis of 1,768 well-preserved waterlogged archaeological grape pips covering approximately 7000 years of history. These samples originate from 25 Italian archaeological sites spanning from the Early Neolithic (6 th millemmium BC) to the Medieval period (8 th -14 th centuries AD). Employing geometric morphometrics and linear discriminant analyses, we compared these archaeological grape pips with modern reference collections to differentiate between wild and domestic grape types. Additionally, we analysed phenotypic changes in grape pip length and shape over the studied period to the present day to highlight traits associated with domestication syndrome. During the Early Neolithic, no evidence of morphologically domesticated grapes was observed. Data from Early Bronze Age sites (ca. 2050–1850 BC) display the same trend observed for the Early Neolithic period. The Middle Bronze Age sites (ca. 1600–1300 BC) continue to exhibit a predominance of wild grape pips. However, a notable transition occurs at the end of the Late Bronze Age (ca. 1300–1100 BC), with the majority of grape pips classified as domestic, indicating the definitive establishment of cultivation practices and selection of domestic grape by these communities. In the Iron Age, grape pips from Etruscan sites dating to the 4 th century BC are predominantly domestic, suggesting an advanced viticulture for this period. During the Roman period (1 st -6 th centuries AD), some sites exhibited a high presence of domestic grape pips and intermediate forms between wild and domestic morphotypes, suggesting introgression between local wild and domestic grape allowing the formation of new varieties. Finally, the Medieval period (8 th -14 th centuries AD) sites demonstrate a widespread prevalence of domestic grape pips across archaeological sites, indicating a reduction of intermediate forms between wild and domestic morphotypes and displaying morphometric characteristics entirely similar to modern domestic grape references. Overall, our study provides valuable insights into the evolution of grapevine cultivation in Italy, highlighting the gradual transition from wild to domesticated types over millennia.