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Understanding the physiological basis underlying the water stress responses in grapevine is becoming increasingly topical owing to the challenges that climate change will impose to grapevine agriculture. Here we used cv. Pinot gris (clone H1), grafted on a series of tolerant (1103Paulsen, P), sensitive (SO4) and recently selected (Georgikon28, G28, Georgikon121, G121, Zamor17, Z17) rootstocks. Plants were either subjected to reduced water availability (WS) or maintained at pot capacity (WW). Photosynthetic (light response curves), stomatal and in vivo gas exchange analysis were carried out as well as dynamics of daily water use (WU), leaf area accumulation with affordable RGB imaging pipelines and leaf water potential. Significant genotypic variation was recorded between rootstocks for most of the traits analyzed under optimal conditions with P and SO4 showing a more vigorous growth, higher CO2 assimilation rate, stomatal conductance and stomatal density per unit of leaf area than G28, G121, Z17 (p &lt, 0.001). Under WS, rootstocks induced different water stress response in Pinot gris, with G28 and G121 showing a higher sensitivity of water use to reduced water availability (WS) (p = 0.021) and no variation for midday leaf water potential until severe WS. P, Z17 and to some extent SO4 induced a pronounced near-anisohydric response with a general WU maintenance followed by reduction in leaf water potential even at high levels of soil water content. In addition, G28 and G121 showed a less marked slope in the linear relationship between daily water use and VPD (p = 0.008) suggesting elevated sensitivity of transpiration to evaporative demand. This led to an insensitivity for total dry weight biomass of G28 and G121 under WS conditions (p &lt, 0.001). This work provides: (i) an in-depth analysis for a series of preferable traits under WS in Pinot gris, (ii) a characterization of Pinot gris &times, rootstock interaction and a series of desirable traits under WS induced by several rootstocks, (iii) the potential benefit for the use a series of affordable methods (e.g., RGB imaging) to easily detect dynamic changes in biomass in grapevine and quickly phenotype genotypes with superior responses under WS. In conclusion, the near-isohydric and conservative behavior observed for G28 and G121 coupled with their low vigor suggest them as potential Pinot gris rootstock candidates for sustaining grapevine productivity in shallow soils likely to develop terminal stress conditions.

Under conditions of high transpiration and low soil water availability, the demand for water can exceed supply causing a reduction in water potential and a loss of cell turgor (wilting). Regulation of stomatal aperture mediates the loss of water vapour (gs), which in turn is dependent in part on the anatomical characteristics of stomatal density (SD) and stomatal size (SS). Anisohydric sugar beet (Beta vulgaris) is atypical, exhibiting wilting under high soil water availability. Spinach (Spinacia oleracea) belongs to the same family Chenopodiaceae s.s., but demonstrates a more typical wilting response. To investigate the role of stomatal dynamics in such behaviours, sugar beet and spinach leaves were exposed to step-changes in photosynthetic photon flux density (PPFD) from 250 to 2500 µmol m−2 s−1. Using a four log-logistic function, the maximum rate of stomatal opening was estimated. Concurrent measurements of SD and SS were taken for both species. While sugar beet coupled faster opening with smaller, more numerous stomata, spinach showed the converse. After exposure to drought, maximum gs was reduced in sugar beet but still achieved a similar speed of opening. It is concluded that sugar beet stomata respond rapidly to changes in PPFD with a high rate and magnitude of opening under both non-droughted and droughted conditions. Such a response may contribute to wilting, even under high soil water availability, but enables photosynthesis to be better coupled with increasing PPFD., Sugar beet is an important crop that is surprisingly prone to short-term reversible wilting of leaves. This wilting is driven by pores on the leaf surface (stomata) remaining open for photosynthesis when they would normally close in response to lowered water potential. This is termed anisohydric behaviour. The benefits to the plant are still unclear but here we show that sugar beet have an adaptation which helps improve function under these conditions. They possess many smaller stomata which rapidly respond to changes in light intensity, improving leaf productivity but also causing the high leaf evaporation which is associated with wilting.

Lysimeters are the reference method for determining ETc, but they are expensive and complex, which limits their use. The first objective of this work was to adjust and evaluate the robustness of sap flow sensors in order to determine the transpiration of a vineyard and, together with an evaporation model, to calculate the ETc of the vineyard. For this purpose, we compared water consumption data obtained from a vineyard weighing lysimeter (ETcLys) with the sum of transpiration obtained from sap flow sensors (TSF) and evaporation estimated empirically over four years (2012, 2013, 2014 and 2015). The second objective was to obtain the relationship between the vegetative growth and transpiration of the vines with different water availability (irrigation and rainfed treatments), as an alternative method for estimating vine water needs adjusted to their real development. The third and last objective was to evaluate the transpiration response of the vines when subjected to water stress. We carried out the work in an experimental vineyard which has a well-established weighing lysimeter. As a result, a good match was obtained between vine sap flow and transpiration (R2 = 0.85) as well as a good relationship between vegetative growth and vine transpiration (FiPAR: R2Irrigation = 0.34. R2Rainfed = 0.54, LAI: R2Irrigation = 0.68. R2Rainfed = 0.53).

As a major driving element of the structure and function of arid and semiarid ecosystems, rainfall is the essential factor limiting plant biological processes. To clarify the characteristics of transpiration and responses to summer rainfall, sap flow density (Fd) of Pinus tabulaeformis and Hippophae rhamnoides was monitored using thermal dissipation probes. In addition, midday leaf water potential (ψm) and leaf stomatal conductance (Gs) were also analyzed to determine water use strategies. The results indicated that the diurnal variation in the normalized Fd values exhibited a single‐peak curve for P. tabulaeformis, while H. rhamnoides showed multiple peaks. The normalized Fd for P. tabulaeformis remained relatively stable regardless of rainfall events. However, there was also a significant increase in the normalized Fd for H. rhamnoides in response to rainfall in June and August (p

We report physiological, anatomical and molecular differences in two economically important grapevine (Vitis vinifera L.) cultivars cv. Grenache (near-isohydric) and Chardonnay (anisohydric) in their response to water-stress induced cavitation. The aim of the study was to compare organ vulnerability (petiole and stem) to cavitation by measuring ultrasonic acoustic emissions (UAE) and percent loss of conductance of potted grapevines subject to the onset of water-stress. Leaf (ψL) and stem water potential (ψS), stomatal conductance (gs), transpiration (E), petiole hydraulics (KPet), and xylem diameter were also measured. Chardonnay displayed hydraulic segmentation based on UAE, with cavitation occurring at a less negative ψL in the petiole than in the stem. Vulnerability segmentation was not observed in Grenache, with both petioles and stems equally vulnerable to cavitation. Leaf water potential that induced 50% of maximum UAE was significantly different between petioles and stems in Chardonnay (ψ50Petiole = -1.14 and ψ50Stem = -2.24 MPa) but not in Grenache (ψ50Petiole = -0.73 and ψ50Stem = -0.78 MPa). Grenache stems appeared more susceptible to water-stress induced cavitation than Chardonnay stems. Grenache displayed (on average) a higher KPet likely due to the presence of larger xylem vessels. A close relationship between petiole hydraulic properties and vine water status was observed in Chardonnay but not in Grenache. Transcriptional analysis of aquaporins in the petioles and leaves (VvPIP1;1, VvPIP2;1, VvPIP2;2 VvPIP2;3, VvTIP1;1, and VvTIP2;1) showed differential regulation diurnally and in response to water-stress. VvPIP2;1 showed strong diurnal regulation in the petioles and leaves of both cultivars with expression highest predawn. Expression of VvPIP2;1 and VvPIP2;2 responded to ψL and ψS in both cultivars indicating the expression of these two genes are closely linked to vine water status. Expression of several aquaporin genes correlated with gas exchange measurements, however, these genes differed between cultivars. In summary, the data shows two contrasting responses in petiole hydraulics and aquaporin expression between the near-isohydric cultivar, Grenache and anisohydric cultivar, Chardonnay.

Aim: To evaluate the usability of various plant water potentials in table grape irrigation management under a semiarid climate.Methods and results: Two water regimes were set up. The « control » water regime was the one usually used in the vineyard. The « 50 % Irrigation » water regime delivered only half the quantity of water to the vines. Predawn leaf (ψ L PD), predawn stem (ψ S PD), midday leaf (ψ L M), and midday stem (ψ S M) water potentials were measured during the growing season. The results show that the four water potentials can accurately measure the vine water status in table grape vineyard at a daily and seasonal time scale. But, ψ L M appeared to be the most reliable indicator to differentiate between the two water regimes with a frequency of 73 %. The « 50 % Irrigation » water regime induced in the ‘Italia’ cultivar an anisohydric behavior and a decrease of 29.4 % in vine vigor and 11.5 % in berry weight. Under the Tunisian climate, ‘Italia’ cultivar may exhibit night time transpiration that decreases ψ L PD by 19.5 %.Conclusion: Preliminary minimum ψ L M threshold to produce high quality table grape would be -0.8 and -1.1 MPa for pre- and post-veraison, respectively.Significance and impact of the study: The pressure chamber is an effective device for irrigation management in commercial table grape vineyards under semiarid conditions.

Hydraulic capacitance (C) in a plant tissue buffers the xylem tension, storing and releasing water and has been highlighted in recent years as an important factor that affects water relations such as drought tolerance and embolism formation. Aquaporins (AQPs) are well known to control leaf hydraulic resistance (Rh) but their role in the control of C is unknown. Here, we assess Rh and C on detached grapevines wild-type (WT) (cv. Brachetto) leaves and over-expressing the aquaporin gene VvPIP2;4N (OE). For this purpose, we developed a new method inspired from the pressure-volume curve technique and the rehydration-kinetic-method, which allowed us to monitor the dynamics of dehydration and rehydration in the same leaf. The recovery after dehydration was measured in dark, light non-transpirative conditions, light-transpirative conditions and light-transpirative condition adding abscisic acid. Pressurizing to dehydrate leaves in the OE line, the recorded Rh and C were respectively lower and higher than those in the WT. The same results were obtained in the dark recovery by rehydration treatment. In the presence of light, either when leaves transpired or not (by depressing vapor pressure deficit), the described effects disappeared. The change in Rh and C did not affect the kinetics of desiccation of detached leaves in dark in air, in OE plants compared to WT ones. Our study highlighted that both Rh and C were influenced by the constitutive over-expression of VvPIP2;4N. The effect of AQPs on C is reported here for the first time and may involve a modulation of cell reflection coefficient.

© 2014 Elsevier B.V. Ephedra alata subsp. alenda is the most important pioneer plant of the moving and semi-stable sand dunes in the deserts and steppes of south Tunisia and occurs naturally in the Grand Erg Oriental, one of the most extreme habitats for plant growth on the planet. A new analysis of physiological performance of this medically important and internationally threatened xerophytic shrub was conducted to assess possible mechanisms of drought tolerance and how these relate to its ecological success. Five-month old plants, grown under controlled climatic conditions, were subjected to a well-watered control treatment or progressive drought by withholding water for 14d with subsequent recovery for 7d. Soil water depletion significantly reduced stem relative water content (RWC) water potential (Ψw) and osmotic potential (Ψπ). Ephedra displayed more negative Ψw and Ψπ values of ca. -3.5 and -4.1MPa, respectively, at the end of the drought treatment, and were associated with turgor loss. Low stem Ψw reduced stomatal conductance (gs), photosynthetic CO2 assimilation rates (ACO2), transpiration (E) and internal CO2 concentration (Ci). However, instantaneous (WUE; ACO2 E-1) and intrinsic (WUEi; ACO2 gs-1) water use efficiency (WUE) increased gradually as water deficit was intensified. Stomatal closure therefore only exerted limited control against dehydration and could not compensate for decreases in soil water status, typical of anisohydric behavior. Drought-stressed stems accumulated high levels of proline up to 480% of control values, highlighting a pivotal role in osmotic adjustment during intense water deficit. In contrast, the osmotic adaptation to soluble sugars was limited. Drought-stressed plants increased ACO2, E, gs and Ci and decreased WUE and WUEi during the first 48h after re-watering, such that they reached similar values to those of control plants by the end of the experiment. Stem proline levels of drought-stressed plants returned to near control values with re-watering. Overall, rapid recovery of photosynthesis following drought-breaking moisture appears to be a critical mechanism allowing E. alata to withstand and survive dry environments.

Recent studies have reported correlations between stem sapwood capacitance (C(wood)) and xylem vulnerability to embolism, but it is unclear how C(wood) relates to the eventual ability of plants to reverse embolism. We investigated possible functional links between embolism reversal efficiency, C(wood), wood density (WD), vulnerability to xylem embolism and hydraulic safety margins in nine woody species native to dry sclerophyllous forests with different degrees of iso versus anisohydry. Substantial inter-specific differences in terms of seasonal/diurnal changes of xylem and leaf water potential, maximum diurnal values of transpiration rate and xylem vulnerability to embolism formation were recorded. Significant diurnal changes in percentage loss of hydraulic conductivity (PLC) were recorded for five species. Significant correlations were recorded between diurnal PLC changes and P50 and P88 values (i.e., xylem pressure inducing 50 and 88% PLC, respectively) as well as between diurnal PLC changes and safety margins referenced to P50 and P88. WD was linearly correlated with minimum diurnal leaf water potential, diurnal PLC changes and wood capacitance across all species. In contrast, significant relationships between P50, safety margin values referenced to P50 and WD were recorded only for the isohydric species. Functional links between diurnal changes in PLC, hydraulic strategies and WD and C(wood) are discussed.

Summary This study on a grapevine mapping population shows that isohydric or anisohydric behaviour is under genetic control and is not simply controlled by transpiration response to soil drought., In the face of water stress, plants evolved with different abilities to limit the decrease in leaf water potential, notably in the daytime (ΨM). So-called isohydric species efficiently maintain high ΨM, whereas anisohydric species cannot prevent ΨM from dropping as soil water deficit develops. The genetic and physiological origins of these differences in (an)isohydric behaviours remain to be clarified. This is of particular interest within species such as Vitis vinifera L. where continuous variation in the level of isohydry has been observed among cultivars. With this objective, a 2 year experiment was conducted on the pseudo-F1 progeny from a cross between the two widespread cultivars Syrah and Grenache using a phenotyping platform coupled to a controlled-environment chamber. Potted plants of all the progeny were analysed for ΨM, transpiration rate, and soil-to-leaf hydraulic conductance, under both well-watered and water deficit conditions. A high genetic variability was found for all the above traits. Four quantitative trait loci (QTLs) were detected for ΨM under water deficit conditions, and 28 other QTLs were detected for the different traits in either condition. Genetic variation in ΨM maintenance under water deficit weakly correlated with drought-induced reduction in transpiration rate in the progeny, and QTLs for both traits did not completely co-localize. This indicates that genetic variation in the control of ΨM under water deficit was not due simply to variation in transpiration sensitivity to soil drying. Possible origins of the diversity in (an)isohydric behaviours in grapevine are discussed on the basis of concurrent variations in soil-to-leaf hydraulic conductance and stomatal control of transpiration.

Grapevine cultivars possess large variability in their response to water availability, and are therefore considered as a good model to study plant hydraulic adjustments. The current research compared the petiole anatomy of two grapevine (Vitis vinifera L.) cultivars, Shiraz and Cabernet Sauvignon, in respect to hydraulic properties. Hydraulic differences between the cultivar petioles were tested over 3 years (2011–2013). Anatomical differences, hydraulic conductivity and embolism were tested under terminal drought conditions. Additionally, xylem differentiation under well watered (WW) and water deficit (WD) conditions was compared. Shiraz was shown to possess larger xylem vessels that resulted in a significantly higher theoretical specific hydraulic conductivity (Kts), leaf hydraulic conductivity (Kleaf) and maximal petiole hydraulic conductivity (Kpetiole). Under WD, smaller vessels were developed, more noticeably in Shiraz. Results confirmed a link between petiole hydraulic architecture and hydraulic behaviour, providing a simple mechanistic explanation for the higher transpiration rates commonly measured in Shiraz. Smaller xylem vessels in Cabernet Sauvignon could imply on its adaptation to WD, and explains its better performances under such conditions.

International audience; The classical view that the drought-related hormone ABA simply acts locally at the guard cell level to induce stomatal closure is questioned by differences between isolated epidermis and intact leaves in stomatal response to several stimuli. We tested the hypothesis that ABA mediates, in addition to a local effect, a remote effect in planta by changing hydraulic regulation in the leaf upstream of the stomata. By gravimetry, porometry to water vapour and argon, and psychrometry, we investigated the effect of exogenous ABA on transpiration, stomatal conductance and leaf hydraulic conductance of mutants described as ABA-insensitive at the guard cell level. We show that foliar transpiration of several ABA-insensitive mutants decreases in response to ABA. We demonstrate that ABA decreases stomatal conductance and down-regulates leaf hydraulic conductance in both the wildtype Col-0 and the ABA-insensitive mutant ost2-2. We propose that ABA promotes stomatal closure in a dual way via its already known biochemical effect on guard cells and a novel, indirect hydraulic effect through a decrease in water permeability within leaf vascular tissues. Variability in sensitivity of leaf hydraulic conductance to ABA among species could provide a physiological basis to the isohydric or anisohydric behaviour.

Aims: Genetic variability in grapevine cultivars may influence their strategy to cope with drought through stomatal regulation of transpiration rate. The aim of the present study was to evaluate the stomatal sensitivity of five cultivars (EkigaIna, Grenache. Marselan, Mourvedre. and Syrah) to soil water status and air water vapor pressure deficit (VPD). Methods and results: Leaf gas exchange and canopy light interception efficiency (epsilon(1)) were evaluated through a wide range of predawn leaf water potential (psi(PD)) measurements in a field experiment in Southern France. Additionally. greenhouse experiments were carried out to monitor stomatal response to increasing VPD levels. Ekigaina showed a strong isohydric behavior with the highest decrease in leaf gas exchange in response to soil water stress and VPD. Mourvedre and Grenache showed a similar but relatively less extreme behavior. These three cultivars showed a constant leaf water status during the day through stomatal regulation and a strong decrease in epsilon(1). In contrast, Syrah and Marselan displayed anisohydric behavior as they presented a less sensitive stomatal control. Both cultivars showed fluctuating midday leaf water potential and Marselan was the least affected in tem of c. Conclusion: This study demonstrated that grape cultivars differed in their stomatal response to soil water deficit and VPD. For a given cultivar, a similar stomatal behavior was found in response to both tyro and VPD. Significance and impact of the results: Adaptation to drought and viticulture viability in hot and dry environments could be achieved by identifying and breeding cultivars with drought tolerance traits, Objectifs : La variabilité génétique existante chez les cépages de vigne peut affecter leur stratégie d’adaptation à la sécheresse en régulant la transpiration au travers de la fermeture des stomates. L’objectif du travail a été d’évaluer la sensibilité des stomates de cinq cultivars (Ekigaïna, Grenache, Marselan, Mourvèdre et Syrah) à la disponibilité en eau du sol et au déficit de pression de vapeur (VPD). Méthodes et résultats : Les échanges gazeux des feuilles et l’efficience d’interception du rayonnement (ei) ont été évalués dans une large gamme des valeurs de potentiel hydrique de base (yPD) en conditions de terrain dans le Sud de la France. En outre, une expérimentation complémentaire en serre a été réalisée pour analyser la réponse des stomates face à une augmentation du VPD. L’Ekigaïna a présenté un caractère ioshydrique très marqué avec la plus grande diminution des échanges gazeux en réponse au déficit hydrique du sol et au VPD. Le Mourvèdre et le Grenache ont présenté un comportement similaire bien que moins extrême que l’Ekigaïna. Ces trois cultivars ont montré un potentiel hydrique foliaire constant pendant la journée à cause d’une forte régulation stomatique et une diminution de ei. Par contre, le Syrah et le Marselan, ont montré un comportement anisohydrique avec un control stomatique moins sensible. Les deux cultivars ont présenté une diminution du potentiel hydrique foliaire à midi tandis que le Marselan a été le moins affecté en termes de ei. Conclusions : La présente étude montre que les cultivars présentent des différences dans leur réponse au déficit hydrique du sol et au VPD. Pour un cultivar donné, un comportement stomatique similaire a été observé en réponse au yPD et au VPD. Impacts et signification des résultats : L’adaptation à la sécheresse et la viabilité de la viticulture sur de climats chauds et secs pourrait être atteint par la sélection de cultivars avec de caractères de tolérance.

Background - A large proportion of vineyards are located in regions with seasonal drought (e.g. Mediterraneantype climates) where soil and atmospheric water deficits, together with high temperatures, exert large constraints on yield and quality. The increasing demand for vineyard irrigation requires an improvement in the efficiency of water use. Deficit irrigation has emerged as a potential strategy to allow crops to withstand mild water stress with little or no decreases of yield, and potentially a positive impact on fruit quality. Understanding the physiological and molecular bases of grapevine responses to mild to moderate water deficits is fundamental to optimize deficit irrigation management and identify the most suitable varieties to those conditions. Scope - How the whole plant acclimatizes to water scarcity and how short- and long-distance chemical and hydraulic signals intervene are reviewed. Chemical compounds synthesized in drying roots are shown to act as long-distance signals inducing leaf stomatal closure and/or restricting leaf growth. This explains why some plants endure soil drying without significant changes in shoot water status. The control of plant water potential by stomatal aperture via feed-forward mechanisms is associated with ‘isohydric’ behaviour in contrast to ‘anysohydric’ behaviour in which lower plant water potentials are attained. This review discusses differences in this respect between grapevines varieties and experimental conditions. Mild water deficits also exert direct and/or indirect (via the light environment around grape clusters) effects on berry development and composition; a higher content of skin-based constituents (e.g. tannins and anthocyanins) has generally being reported. Regulation under water deficit of genes and proteins of the various metabolic pathways responsible for berry composition and therefore wine quality are reviewed.