Functional phenotypic plasticity mediated by water stress and [CO2] explains differences in drought tolerance of two phylogenetically close conifers.

Forests are threatened globally by increased recurrence and intensity of hot droughts. Functionally close coexisting species may exhibit differences in drought vulnerability large enough to cause niche differentiation and affect forest dynamics. The effect of rising atmospheric [CO2], which could partly alleviate the negative effects of drought, may also differ between species. We analysed functional plasticity in seedlings of two taxonomically close pine species (Pinus pinaster Ait., Pinus pinea L.) under different [CO2] and water stress levels. The multidimensional functional trait variability was more influenced by water stress (preferentially xylem traits) and [CO2] (mostly leaf traits) than by differences between species. However, we observed differences between species in the strategies followed to coordinate their hydraulic and structural traits under stress. Leaf 13C discrimination decreased with water stress and increased under elevated [CO2]. Under water stress both species increased their sapwood area to leaf area ratios, tracheid density and xylem cavitation, whereas they reduced tracheid lumen area and xylem conductivity. Pinus pinea was more anisohydric than P. pinaster. Pinus pinaster produced larger conduits under well-watered conditions than P. pinea. Pinus pinea was more tolerant to water stress and more resistant to xylem cavitation under low water potentials. The higher xylem plasticity in P. pinea, particularly in tracheid lumen area, expressed a higher capacity of acclimation to water stress than P. pinaster. In contrast, P. pinaster coped with water stress comparatively more by increasing plasticity of leaf hydraulic traits. Despite the small differences observed in the functional response to water stress and drought tolerance between species, these interspecific differences agreed with ongoing substitution of P. pinaster by P. pinea in forests where both species co-occur. Increased [CO2] had little effect on the species-specific relative performance. Thus, a competitive advantage under moderate water stress of P. pinea compared with P. pinaster is expected to continue in the future.

Stomatal responses in grapevine become increasingly more tolerant to low water potentials throughout the growing season.

The leaf of a deciduous species completes its life cycle in a few months. During leaf maturation, osmolyte accumulation leads to a significant reduction of the turgor loss point (ΨTLP ), a known marker for stomatal closure. Here we exposed two grapevine cultivars to drought at three different times during the growing season to explore if the seasonal decrease in leaf ΨTLP influences the stomatal response to drought. The results showed a significant seasonal shift in the response of stomatal conductance to stem water potential (gs ~Ψstem ), demonstrating that grapevines become increasingly tolerant to low Ψstem as the season progresses in coordination with the decrease in ΨTLP . We also used the SurEau hydraulic model to demonstrate a direct link between osmotic adjustment and the plasticity of gs ~Ψstem . To understand the possible advantages of gs ~Ψstem plasticity, we incorporated a seasonally dynamic leaf osmotic potential into the model that simulated stomatal conductance under several water availabilities and climatic scenarios. The model demonstrated that a seasonally dynamic stomatal closure threshold results in trade-offs: it reduces the time to turgor loss under sustained long-term drought, but increases overall gas exchange particularly under seasonal shifts in temperature and stochastic water availability. A projected hotter future is expected to lower the increase in gas exchange that plants gain from the seasonal shift in gs ~Ψstem . These findings show that accounting for dynamic stomatal regulation is critical for understanding drought tolerance.

Reproductive Stage Drought Tolerance in Wheat: Importance of Stomatal Conductance and Plant Growth Regulators.

Drought stress requires plants to adjust their water balance to maintain tissue water levels. Isohydric plants ('water-savers') typically achieve this through stomatal closure, while anisohydric plants ('water-wasters') use osmotic adjustment and maintain stomatal conductance. Isohydry or anisohydry allows plant species to adapt to different environments. In this paper we show that both mechanisms occur in bread wheat (Triticum aestivum L.). Wheat lines with reproductive drought-tolerance delay stomatal closure and are temporarily anisohydric, before closing stomata and become isohydric at higher threshold levels of drought stress. Drought-sensitive wheat is isohydric from the start of the drought treatment. The capacity of the drought-tolerant line to maintain stomatal conductance correlates with repression of ABA synthesis in spikes and flag leaves. Gene expression profiling revealed major differences in the drought response in spikes and flag leaves of both wheat lines. While the isohydric drought-sensitive line enters a passive growth mode (arrest of photosynthesis, protein translation), the tolerant line mounts a stronger stress defence response (ROS protection, LEA proteins, cuticle synthesis). The drought response of the tolerant line is characterised by a strong response in the spike, displaying enrichment of genes involved in auxin, cytokinin and ethylene metabolism/signalling. While isohydry may offer advantages for longer term drought stress, anisohydry may be more beneficial when drought stress occurs during the critical stages of wheat spike development, ultimately improving grain yield.

Hydroscapes: A Useful Metric for Distinguishing Iso-/Anisohydric Behavior in Almond Cultivars.

As a consequence of climate change, water scarcity has increased the use of the iso-/anisohydric concept with the aim of identifying anisohydric or drought-tolerant genotypes. Recently, Meinzer and colleagues developed a metric for discriminating between iso- and anisohydric behavior called the hydroscape, which describes a range in which stomata control leaf water potential (Ψ) with decreasing water availability, and it is linked to several water-regulation and drought-tolerance traits. Thus, our objective was to test the usefulness of the hydroscape in discriminating between iso- and anisohydric Prunus dulcis cultivars, a species that is widely cultivated in Mediterranean central Chile due to its ability to withstand water stress. Through a pot desiccation experiment, we determined that the hydroscape was able to discriminate between two contrasting Prunus cultivars; the more anisohydric cultivar had a hydroscape 4.5 times greater than that of the other cultivar, and the hydroscape correlated with other metrics of plant water-use strategies, such as the maximum range of daily Ψ variation and the Ψ at stomatal closure. Moreover, the photosynthesis rates were also differently affected between cultivars. The more isohydric cultivar, which had a smaller hydroscape, displayed a steeper photosynthesis reduction at progressively lower midday Ψ. This methodology could be further used to identify drought-tolerant anisohydric Prunus cultivars.

Water stress assessment on grapevines by using classification and regression trees.

Multiple factors, such as the vineyard environment and winemaking practices, are known to affect the development of vines as well as the final composition of grapes. Water stress promotes the synthesis of phenols and is associated with grape quality as long as it does not inhibit production. To identify the key parameters for managing water stress and grape quality, multivariate statistical analysis is essential. Classification and regression trees are methods for constructing prediction models from data, especially when data are complex and when constructing a single global model is difficult and models are challenging to interpret. The models were obtained by recursively partitioning the data space and fitting a simple prediction model within each partition. The partitioning can be represented graphically as a decision tree. This approach permitted the most decisive variables for predicting the most vulnerable vineyards and wine quality parameters associated with water stress. In Priorat AOC, Carignan grapevines had the highest water potential and abscisic acid concentration in the early growth plant stages and permitted vineyards to be classified by mesoclimate. This information is useful for identifying which measurements could most easily differentiate between early and late-ripening vineyards. LWP and Ts during an early physiological stage (pea size) permitted warm and cold areas to be differentiated.

Evaluation of morphological, physiological, and biochemical traits for assessing drought resistance in eleven tree species.

The frequency and severity of drought are expected to increase due to climate change; therefore, selection of tree species for afforestation should consider drought resistance of the species for maximum survival and conservation of natural habitats. In this study, three soil moisture regimes: control (100% precipitation), mild drought (40% reduction in precipitation), and severe drought (80% reduction in precipitation) were applied to six gymnosperm and five angiosperm species for two consecutive years. We quantified the drought resistance index based on the root collar diameter and assessed the correlation between species drought resistance and other morphological, physiological, and biochemical traits by regression analysis. The prolonged drought stress altered the morphological, physiological, and biochemical traits, but the responses were species-specific. The species with high drought resistance had high leaf mass per area (LMA), photosynthetic rate (Pn), and midday leaf water potential (ΨMD), and low carbon isotopic discrimination (δ13C), flavonoid and polyphenol content, superoxide dismutase and DPPH radical scavenging activity. The highly drought-resistant species had a relatively less decrease in leaf size, Pn, and predawn leaf water potential (ΨPD), and less increase in δ13C, abscisic acid and sucrose content, and LMA compared to the control. The interannual variation in drought resistance (∆Rd) was positively correlated with the species hydroscopic slope (isohydric and anisohydric). Korean pine was highly resistant, sawtooth oak, hinoki cypress, East Asian white birch, East Asian ash, and mono maple were highly susceptible, and Korean red pine, Japanese larch, Sargent cherry, needle fir, and black pine were moderate in drought resistance under long-term drought. These findings will help species selection for afforestation programs and establishment of sustainable forests, especially of drought-tolerant species, under increased frequency and intensity of spring and summer droughts.

Coordinated variation in stem and leaf functional traits of temperate broadleaf tree species in the isohydric-anisohydric spectrum.

Stomatal regulation serves as an important strategy for plants to adapt to drought. However, the understanding of how complexes of plant-functional traits vary along the continuum from isohydry to anisohydry remains insufficient. In this study, we investigated a proxy of the degree of iso/anisohydry-the water potential at stomatal closure-and a series of functional traits of leaves and branches in 20 temperate broadleaf species planted in an arid limestone habitat in northern China. The results showed that the water potential at stomatal closure was significantly correlated with many functional traits. At the anisohydric end of the spectrum, species had a higher leaf carbon content and vein density, a greater stomatal length, a thicker lower leaf epidermis, higher embolism resistance, higher wood density, a greater Huber value, a greater ratio of fiber wall thickness to xylem lumen diameter, a larger proportion of total fiber wall area to xylem cross-sectional area, a lower water potential at the turgor loss point (TLP), a smaller relative water content at the TLP, a lower osmotic potential at full turgor and a smaller specific leaf area. It is concluded that a continuum of coordination and trade-offs among co-evolved anatomical and physiological traits gives rise to the spectrum from isohydry to anisohydry spanned by the 20 tree species, and the anisohydric species showed stronger stress resistance, with greater investment in stems and leaves than the isohydric species to maintain stomatal opening under drought conditions.

Starch storage capacity of sapwood is related to dehydration avoidance during drought.

PREMISE: The xylem tissue of plants performs three principal functions: transport of water, support of the plant body, and nutrient storage. Tradeoffs may arise because different structural requirements are associated with different functions or because suites of traits are under selection that relate to resource acquisition, use, and turnover. The structural and functional basis of xylem storage is not well established. We hypothesized that greater starch storage would be associated with greater sapwood parenchyma and reduced fibers, which would compromise resistance to xylem tensions during dehydration. METHODS: We measured cavitation resistance, minimum water potential, starch content, and sapwood parenchyma and fiber area in 30 species of southern California chaparral shrubs (evergreen and deciduous). RESULTS: We found that species storing greater starch within their xylem tended to avoid dehydration and were less cavitation resistant, and this was supported by phylogenetic independent contrasts. Greater sapwood starch was associated with greater parenchyma area and reduced fiber area. For species without living fibers, the associations with parenchyma were stronger, suggesting that living fibers may expand starch storage capacity while also contributing to the support function of the vascular tissue. Drought-deciduous species were associated with greater dehydration avoidance than evergreens. CONCLUSIONS: Evolutionary forces have led to an association between starch storage and dehydration resistance as part of an adaptive suite of traits. We found evidence for a tradeoff between tissue mechanical traits and starch storage; moreover, the evolution of novel strategies, such as starch-storing living fibers, may mitigate the strength of this tradeoff.

Drought response strategies of deciduous and evergreen woody species in a seasonally dry neotropical forest.

Deciduous and evergreen trees are usually considered the main coexisting functional groups in seasonally dry tropical forests (SDTF). We compared leaf and stem traits of 22 woody species in the Brazilian Caatinga to investigate whether deciduous (DC) and evergreen (EV) species have divergent water-use strategies. Our hypothesis was that DC trees compensate for their short leaf longevity by being less conservative in water use and showing higher variation in the seasonal water potential after leaf shedding. Evergreen species should exhibit a highly conservative water use strategy, which reduces variations in seasonal water potential and the negative effects of desiccation. Our leaf dynamics results indicate that the crown area of DC trees is more sensitive to air and soil drought, whereas EV trees are only sensitive to soil drought. Deciduous species exhibit differences in a set of leaf traits confirming their acquisitive strategy, which contrasts with evergreen species. However, when stomatal traits are considered, we found that DC and EV have similar stomatal regulation strategies (partially isohydric). We also found divergent physiological strategies within DC. For high wood density DC, the xylem water potential (Ψxylem) continued to drop during the dry season. We also found a negative linear relationship between leaf life span (LL) and the transpiration rate per unit of hydraulic conductivity (Λ), indicating that species with high LL are less vulnerable to hydraulic conductivity loss than early-deciduous species. Collectively, our results indicate divergence in the physiology of deciduous species, which suggests that categorizing species based solely on their leaf phenology may be an oversimplification.

Comparing Hydraulics Between Two Grapevine Cultivars Reveals Differences in Stomatal Regulation Under Water Stress and Exogenous ABA Applications.

Hydraulics of plants that have different strategies of stomatal regulation under water stress are relatively poorly understood. We explore how root and shoot hydraulics, stomatal conductance (g s), leaf and root aquaporin (AQP) expression, and abscisic acid (ABA) concentration in leaf xylem sap ([ABA]xylemsap) may be coordinated under mild water stress and exogenous ABA applications in two Vitis vinifera L. cultivars traditionally classified as near-isohydric (Grenache) and near-anisohydric (Syrah). Under water stress, Grenache exhibited stronger adjustments of plant and root hydraulic conductances and greater stomatal sensitivity to [ABA]xylemsap than Syrah resulting in greater conservation of soil moisture but not necessarily more isohydric behavior. Correlations between leaf (Ψleaf) and predawn (ΨPD) water potentials between cultivars suggested a "hydrodynamic" behavior rather than a particular iso-anisohydric classification. A significant decrease of Ψleaf in well-watered ABA-fed vines supported a role of ABA in the soil-leaf hydraulic pathway to regulate g s. Correlations between leaf and root AQPs expression levels under water deficit could explain the response of leaf (K leaf) and root (Lp r) hydraulic conductances in both cultivars. Additional studies under a wider range of soil water deficits are required to explore the possible differential regulation of g s and plant hydraulics in different cultivars and experimental conditions.

Water use strategies and drought intensity define the relative contributions of hydraulic failure and carbohydrate depletion during seedling mortality.

COMBINING HYDRAULIC: and carbon-related measurements can help elucidate drought-induced plant mortality. To study drought mortality mechanisms, seedlings of two woody species, including the anisohydric Robinia pseudoacacia and isohydric Quercus acutissima, were cultivated in a greenhouse and subjected to intense drought by withholding water and mild drought by adding half of the amount of daily water lost. Patterns of leaf and root gas exchange, leaf surface areas, growth, leaf and stem hydraulics, and carbohydrate dynamics were determined in drought-stressed and control seedlings. We detected a complete loss of hydraulic conductivity and partial depletion of total nonstructural carbohydrates contents (TNC) in the dead seedlings. We also found that intense drought triggered a more rapid decrease in plant water potential and a faster drop in net photosynthesis below zero, and a greater TNC loss in dead seedlings than mild drought. Additionally, anisohydric R. pseudoacacia suffered a rapider death than the isohydric Q. acutissima. Based on these findings, we propose that hydraulic conductivity loss and carbon limitation jointly contributed to drought-induced death, while the relative contributions could be altered by drought intensity. We thus believe that it is important to illustrate the mechanistic relationships between stress intensity and carbon-hydraulics coupling in the context of isohydric vs. anisohydric hydraulic strategies.

Water use strategy affects avoidance of ozone stress by stomatal closure in Mediterranean trees-A modelling analysis.

Both ozone (O3 ) and drought can limit carbon fixation by forest trees. To cope with drought stress, plants have isohydric or anisohydric water use strategies. Ozone enters plant tissues through stomata. Therefore, stomatal closure can be interpreted as avoidance to O3 stress. Here, we applied an optimization model of stomata involving water, CO2 , and O3 flux to test whether isohydric and anisohydric strategies may affect avoidance of O3 stress by stomatal closure in four Mediterranean tree species during drought. The data suggest that stomatal closure represents a response to avoid damage to the photosynthetic mechanisms under elevated O3 depending on plant water use strategy. Under high-O3 and well-watered conditions, isohydric species limited O3 fluxes by stomatal closure, whereas anisohydric species activated a tolerance response and did not actively close stomata. Under both O3 and drought stress, however, anisohydric species enhanced the capacity of avoidance by closing stomata to cope with the severe oxidative stress. In the late growing season, regardless of the water use strategy, the efficiency of O3 stress avoidance decreased with leaf ageing. As a result, carbon assimilation rate was decreased by O3 while stomata did not close enough to limit transpirational water losses.

Non-structural carbohydrate pools not linked to hydraulic strategies or carbon supply in tree saplings during severe drought and subsequent recovery.

Non-structural carbohydrate (NSC) pools fluctuate based on the interplay between photosynthesis, demand from various carbon (C) sinks and tree hydraulic status. Thus, it has been hypothesized that tree species with isohydric stomatal control (i.e., trees that close stomata rapidly in response to drought) rely heavily on NSC pools to sustain metabolism, which can lead to negative physiological consequences such as C depletion. Here, we seek to use a species' degree of isohydry or anisohydry as a conceptual framework for understanding the interrelations between photosynthetic C supply, hydraulic damage and fluctuations in NSC pools. We conducted a 6-week experimental drought, followed by a 6-week recovery period, in a greenhouse on seven tree species that span the spectrum from isohydric to anisohydric. Throughout the experiment, we measured photosynthesis, hydraulic damage and NSC pools. Non-structural carbohydrate pools were remarkably stable across species and tissues-even highly isohydric species that drastically reduced C assimilation were able to maintain stored C. Despite these static NSC pools, we still inferred an important role for stored C during drought, as most species converted starches into sugars during water stress (and back again post-drought). Finally, we did not observe any linkages between C supply, hydraulic damage and NSC pools, indicating that NSC was maintained independent of variation in photosynthesis and hydraulic function. Our results advance the idea that C depletion is a rare phenomenon due to either active maintenance of NSC pools or sink limitation, and thus question the hypothesis that reductions in C assimilation necessarily lead to C depletion.

Interaction of stomatal behaviour and vulnerability to xylem cavitation determines the drought response of three temperate tree species.

How the mortality and growth of tree species vary with the iso-anisohydric continuum and xylem vulnerability is still being debated. We conducted a precipitation reduction experiment to create a mild drought condition in a forest in the Baotianman Mountains, China, a sub-humid region. Three main sub-canopy tree species in this region were examined. After rainfall reduction, Lindera obtusiloba showed severe dieback, but two other co-occurring species did not show dieback. The water potential at stomatal closure of Dendrobenthamia japonica, L. obtusiloba and Sorbus alnifolia was -1.70, -2.54 and -3.41 MPa, respectively, whereas the water potential at 88 % loss in hydraulic conductivity of the three species was -2.31, -2.11 and -7.01 MPa, respectively. Taken together, near-anisohydric L. obtusiloba with vulnerable xylem was highly susceptible to drought dieback. Anisohydric S. alnifolia had the most negative minimum water potential, and its xylem was the most resistant to cavitation. Isohydric D. japonica conserved water by rapidly closing its stomata. Ultimately, the hydraulic safety margin (HSM) of L. obtusiloba was the smallest among the three species, especially in precipitation-reduced plots. In terms of the stomatal safety margin (SSM), L. obtusiloba was negative, while S. alnifolia and D. japonica were positive. Of the two species without dieback, rainfall reduction decreased growth of D. japonica, but did not influence growth of S. Alnifolia; meanwhile, rainfall reduction led to a decrease of non-structural carbohydrates (NSCs) in D. japonica, but an increase in S. alnifolia. It is concluded that HSM as well as SSM allow interpreting the sensitivity of the three sub-canopy species to drought. The drought-induced dieback of L. obtusiloba is determined by the interaction of stomatal behaviour and xylem vulnerability, and the species could be sensitive to climate change-caused drought although still in sub-humid areas. The isohydric/anisohydric degree is associated with NSCs status and growth of plants.

Linking coordinated hydraulic traits to drought and recovery responses in a tropical montane cloud forest.

PREMISE: Understanding plant hydraulic functioning and water balance during drought has become key in predicting species survival and recovery. However, there are few insightful studies that couple physiological and morphological attributes for many ecosystems, such as the vulnerable Tropical Montane Cloud Forests (TMCF). In this study, we evaluated drought resistance and recovery for saplings for five tree species spanning deciduous to evergreen habits from a Mexican TMCF. METHODS: In drought simulations, water was withheld until plants reached species-specific P50 or P88 values (pressures required to induce a 50 or 88% loss in hydraulic conductivity), then they were rewatered. Drought resistance was considered within the isohydric-anisohydric framework and compared to leaf gas exchange, water status, pressure-volume curves, specific leaf area, and stomatal density. RESULTS: The TMCF species closed stomata well before significant losses in hydraulic conductivity (isohydric). Yet, despite the coordination of these traits, the traits were not useful for predicting the time needed for the species to reach critical hydraulic thresholds. Instead, maximum photosynthetic rates explained these times, reinforcing the linkage between hydraulic and carbon dynamics. Despite their varying hydraulic conductivities, stomatal responses, and times to hydraulic thresholds, 58 of the 60 study plants recovered after the rewatering. The recovery of photosynthesis and stomatal conductance can be explained by the P50 values and isohydry. CONCLUSIONS: This study raises new questions surrounding drought management strategies, recovery processes, and how lethal thresholds are defined. Further studies need to consider the role of water and carbon balance in allowing for both survival and recovery from drought.

Beyond soil water potential: An expanded view on isohydricity including land-atmosphere interactions and phenology.

Over the past decade, the concept of isohydry or anisohydry, which describes the link between soil water potential (ΨS ), leaf water potential (ΨL ), and stomatal conductance (gs ), has soared in popularity. However, its utility has recently been questioned, and a surprising lack of coordination between the dynamics of ΨL and gs across biomes has been reported. Here, we offer a more expanded view of the isohydricity concept that considers effects of vapour pressure deficit (VPD) and leaf area index (AL ) on the apparent sensitivities of ΨL and gs to drought. After validating the model with tree- and ecosystem-scale data, we find that within a site, isohydricity is a strong predictor of limitations to stomatal function, though variation in VPD and leaf area, among other factors, can challenge its diagnosis. Across sites, the theory predicts that the degree of isohydricity is a good predictor of the sensitivity of gs to declining soil water in the absence of confounding effects from other drivers. However, if VPD effects are significant, they alone are sufficient to decouple the dynamics of ΨL and gs entirely. We conclude with a set of practical recommendations for future applications of the isohydricity framework within and across sites.

Linking variation in intrinsic water-use efficiency to isohydricity: a comparison at multiple spatiotemporal scales.

Species-specific responses of plant intrinsic water-use efficiency (iWUE) to multiple environmental drivers associated with climate change, including soil moisture (θ), vapor pressure deficit (D), and atmospheric CO2 concentration (ca ), are poorly understood. We assessed how the iWUE and growth of several species of deciduous trees that span a gradient of isohydric to anisohydric water-use strategies respond to key environmental drivers (θ, D and ca ). iWUE was calculated for individual tree species using leaf-level gas exchange and tree-ring δ13 C in wood measurements, and for the whole forest using the eddy covariance method. The iWUE of the isohydric species was generally more sensitive to environmental change than the anisohydric species was, and increased significantly with rising D during the periods of water stress. At longer timescales, the influence of ca was pronounced for isohydric tulip poplar but not for others. Trees' physiological responses to changing environmental drivers can be interpreted differently depending on the observational scale. Care should be also taken in interpreting observed or modeled trends in iWUE that do not explicitly account for the influence of D.

Aquaporin regulation in roots controls plant hydraulic conductance, stomatal conductance, and leaf water potential in Pinus radiata under water stress.

Stomatal regulation is crucial for forest species performance and survival on drought-prone sites. We investigated the regulation of root and shoot hydraulics in three Pinus radiata clones exposed to drought stress and its coordination with stomatal conductance (gs ) and leaf water potential (Ψleaf ). All clones experienced a substantial decrease in root-specific root hydraulic conductance (Kroot-r ) in response to the water stress, but leaf-specific shoot hydraulic conductance (Kshoot-l ) did not change in any of the clones. The reduction in Kroot-r caused a decrease in leaf-specific whole-plant hydraulic conductance (Kplant-l ). Among clones, the larger the decrease in Kplant-l , the more stomata closed in response to drought. Rewatering resulted in a quick recovery of Kroot-r and gs . Our results demonstrated that the reduction in Kplant-l , attributed to a down regulation of aquaporin activity in roots, was linked to the isohydric stomatal behaviour, resulting in a nearly constant Ψleaf as water stress started. We concluded that higher Kplant-l is associated with water stress resistance by sustaining a less negative Ψleaf and delaying stomatal closure.

Below versus above Ground Plant Sources of Abscisic Acid (ABA) at the Heart of Tropical Forest Response to Warming.

Warming surface temperatures and increasing frequency and duration of widespread droughts threaten the health of natural forests and agricultural crops. High temperatures (HT) and intense droughts can lead to the excessive plant water loss and the accumulation of reactive oxygen species (ROS) resulting in extensive physical and oxidative damage to sensitive plant components including photosynthetic membranes. ROS signaling is tightly integrated with signaling mechanisms of the potent phytohormone abscisic acid (ABA), which stimulates stomatal closure leading to a reduction in transpiration and net photosynthesis, alters hydraulic conductivities, and activates defense gene expression including antioxidant systems. While generally assumed to be produced in roots and transported to shoots following drought stress, recent evidence suggests that a large fraction of plant ABA is produced in leaves via the isoprenoid pathway. Thus, through stomatal regulation and stress signaling which alters water and carbon fluxes, we highlight the fact that ABA lies at the heart of the Carbon-Water-ROS Nexus of plant response to HT and drought stress. We discuss the current state of knowledge of ABA biosynthesis, transport, and degradation and the role of ABA and other isoprenoids in the oxidative stress response. We discuss potential variations in ABA production and stomatal sensitivity among different plant functional types including isohydric/anisohydric and pioneer/climax tree species. We describe experiments that would demonstrate the possibility of a direct energetic and carbon link between leaf ABA biosynthesis and photosynthesis, and discuss the potential for a positive feedback between leaf warming and enhanced ABA production together with reduced stomatal conductance and transpiration. Finally, we propose a new modeling framework to capture these interactions. We conclude by discussing the importance of ABA in diverse tropical ecosystems through increases in the thermotolerance of photosynthesis to drought and heat stress, and the global importance of these mechanisms to carbon and water cycling under climate change scenarios.

Maturation of Atriplex halimus L. leaves involves changes in the molecular regulation of stomatal conductance under high evaporative demand and high but not low soil water content.

MAIN CONCLUSION: Under high water availability, the maximum gas exchange was observed at noon in the expanding and expanded leaves. The expanded leaves showed lower gas exchange capacity due to the regulation of stomatal-movement genes. Under well-watered condition, stomatal conductance (gs) and photosynthetic rate (A) of expanding and expanded leaves of Atriplex halimus peaked at noon despite the midday decline in the leaf relative water content, suggesting deviation from typical isohydric behaviour. However, the expanding leaves had higher gs and A than the expanded ones. When light intensity was temporarily increased, A and gs were enhanced in both types of leaves though to a higher level in the expanding leaves. In well-watered expanded leaves: (1) A was mainly dependent on gs rather than photosynthetic capacity; gs was controlled by internal factors, thereby limiting water loss via transpiration (E); (2) the accumulation of total soluble sugars (TSS) along with increased Rubisco protein could be a subsidiary factor limiting A; (3) TSS and ABA seem to act in co-ordination to up-regulate ABA-dependent genes controlling gs and (4) the significant induction of DREBs suggests a role in maintaining high relative water content in these leaves compared to the expanding ones. In expanding leaves of well-watered plants, high A along with Rubisco down-regulation and elevated TSS suggests that A was regulated by signals coordinating carbon and nitrogen balance and the elevated ABA could be involved in regulating the hydraulic activity to enhance cell expansion and facilitate leaf growth. Both expanded and expanding leaves behaved in typical isohydric manner under water stress, which did not involve the accumulation of ABA suggesting that stomatal closure was primarily stimulated by hydraulic rather than chemical signals.