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.

Grapevine irrigation is becoming an important practice to guarantee wine quality or even plant survival in regions affected by seasonal drought. Nevertheless, irrigation has to be controlled to optimise source to sink balance and avoid excessive vigour. The results we present here in two grapevine varieties (Moscatel and Castelão) during 3 years, indicate that we can decrease the amount of water applied by 50% (as in deficit irrigation, DI, and in partial root drying, PRD) in relation to full crop's evapotranspiration (ETc) [full irrigated (FI) vines] with no negative effects on production and even get some gains of quality (in the case of PRD). We report that in non-irrigated and in several cases in PRD vines exhibit higher concentrations of berry skin anthocyanins and total phenols than those presented by DI and FI vines. We showed that these effects on quality were mediated by a reduction in vigour, leading to an increase on light interception in the cluster zone. Because plant water status during most of the dates along the season was not significantly different between PRD and DI, and when different, PRD even exhibited a higher leaf water potential than DI vines, we conclude that growth inhibition in PRD was not a result of a hydraulic control. The gain in crop water use in DI and PRD was accompanied by an increase of the d 13C values in the berries in DI and PRD as compared to FI, suggesting that we can use this methodology to assess the integrated water-use efficiency over the growing season.

The effect of gradually-developing water-stress has been studied in Lupinus albus L., Helianthus annuus L., Vitis vinifera cv. Rosaki and Eucalyptus globulus Labill. Water was withheld and diurnal rhythms were investigated 4-8 d later, when the predawn water deficit was more negative than in watered plants, and the stomata closed almost completely early during the photoperiod. The contribution of 'stomatal' and 'non-stomatal' components to the decrease of photosynthetic rate was investigated by (1) comparing the changes of the rate of photosynthesis in air with the changes of stomatal conductance and (2) measuring photosynthetic capacity in saturating irradiance and 15% CO2. Three species (lupin, eucalyptus and sunflower) showed larger changes of stomatal conductance than photosynthesis in air, and showed little or no decrease of photosynthetic capacity in saturating CO2. Photosynthesis in air also recovered fully overnight after watering the plants in the evening. In grapevines, stomatal conductance and photosynthesis in air changed in parallel, there was a marked decrease of photosynthetic capacity, and photosynthesis and stomatal conductance did not recover overnight after watering water-stressed plants. Relative water content remained above 90% in grapevine. We conclude that non-stomatai components do not play a significant role in lupins, sunflower or eucalyptus, but could in grapevine. The effect of water-stress on partitioning of photosynthate was investigated by measuring the amounts of sucrose and starch in leaves during a diurnal rhythm, and by measuring the partitioning of "C-carbon dioxide between sucrose and starch. In all four species, starch was depleted in water-stressed leaves but sucrose was maintained at amounts similar to, or higher than, those in watered plants. Partitioning into sucrose was increased in lupins and eucalyptus, and remained unchanged in grapevine and sunflower. It is concluded that water-stressed leaves in all four species maintain high levels of soluble sugars in their Correspotidetiee: R. C. Leegood, Robert Hitt histitute. Departinent ofAtiimal and Platit .Scietiees, University of Sheffield, Sheffield SW 2TN, UK.leaves, despite having lower rates of field photosynthesis, decreased rates of export, and low amounts of starch in their leaves. in the intracellular spaces in the leaf; D, water vapour pressure deficit between the air and the leaf; DW, dry weight; FW, fresh weight; gs, stomatal conductance; RWC, relative water content; Tr, transpiration rate; ip, water potential.

Abstract. A study to assess the effects of the Partial Rootzone Drying (PRD) irrigation strategy in comparison to other irrigation systems was carried out in southern Portugal in two field-grown grapevines varieties, Moscatel and Castelão. We addressed the question of whether by regulating growth and plant water use, the PRD system would enable an equilibrated vegetative development, leading to a favourable capture of solar radiation for photoassimilate production and, at the same time to provide an optimum environment for fruit maturation. Three irrigation schemes were applied in addition to the non-irrigated (NI) vines: partial root drying (PRD), 50% of crop evapotranspiration (ETc), supplied to only one side of the root system while the other one was allowed to dry, alternating sides every 15 days; deficit irrigated (DI), 50% ETc supplied, half to each side of the root system and full irrigated (FI, 100% ETc). During the whole season FI plants of both varieties exhibited a high leaf predawn water potential (ψ pd , ca -0.2 MPa) while a progressive decline was observed in NI plants, reaching ψ pd values near -0.7 MPa at the end of August. PRD and DI presented intermediate values. PRD vines exhibited a stronger control over vegetative growth as compared with DI and FI plants. This was expressed by lower values of total leaf area at harvest, leaf layer number, canopy wideness and water shoots number, allowing a higher light interception at the cluster zone that induced an improvement in some berry quality characteristics. Watering had no significant effects on sugar accumulation in the berries but led to a favourable increase in the must titratable acidity, mainly in Castelão. Whereas in DI and FI treatments berry skin anthocyanins and phenols content were always lower than in NI, in PRD there was either no reduction or the reduction was much lower than in the other irrigation treatments. Water use efficiency (WUE) was increased by about 80% in PRD and DI when compared with FI, as a result of almost similar yields in the three treatments. Yield gains of irrigated plants in relation to NI were modest, explained by the rainy spring in both years.

Abstract. The effects of a slowly-imposed drought stress on gas-exchange, chlorophyll a fluorescence, biochemical and physiological parameters of Vitis vinifera L. leaves (cv. Aragonez, syn. Tempranillo) growing in a commercial vineyard (South Portugal) were evaluated. Relative to well-watered plants (predawn water potential, Ψ PD = -0.13 ± 0.01 MPa), drought-stressed plants (Ψ PD = -0.97 ± 0.01 MPa) had lower photosynthetic rates (ca 70%), stomatal conductance, and PSII activity (associated with a higher reduction of the quinone A pool and lower efficiency of PSII open centres). Stomatal limitation to photosynthesis was increased in drought-stressed plants relative to well-watered plants by ca 44%. Modelled responses of net photosynthesis to internal CO 2 indicated that drought-stressed plants had significant reductions in maximum Rubisco carboxylation activity (ca 32%), ribulose-1,5-bisphosphate regeneration (ca 27%), and triose phosphate (triose-P) utilization rates (ca 37%) relative to well-watered plants. There was good agreement between the effects of drought on modelled biochemical parameters, and in vitro activities of key enzymes of carbon metabolism, namely Rubisco, glyceraldehyde-3-phosphate dehydrogenase, ribulose-5-phosphate kinase and fructose-1,6-bisphosphate phosphatase. Quantum yields measured under both ambient (35 Pa) and saturating CO 2 (100 Pa) for drought-stressed plants were decreased relative to well-watered plants, as well as maximum photosynthetic rates measured at light and CO 2 saturating conditions (three times ambient CO 2 levels). Although stomatal closure was a strong limitation to CO 2 assimilation under drought, comparable reductions in electron transport, CO 2 carboxylation, and utilization of triose-P capacities were also adaptations of the photosynthetic machinery to dehydration that slowly developed under field conditions. Results presented in this study confirm that modelling photosynthetic responses based on gas-exchange data can be successfully used to predict metabolic limitations to photosynthesis.

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