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International audience; Water stress can cause declines in plant function that persist after rehydration. Recent work has defined ‘resilience’ traits characterizing leaf resistance to persistent damage from drought, but whether these traits predict resilience in whole-plant function is unknown. It is also unknown whether the coordination between resilience and ‘resistance’ – the ability to maintain function during drought – observed globally occurs within ecosystems. For eight rainforest species, we dehydrated and subsequently rehydrated leaves, and measured water stress thresholds for declines in rehydration capacity and maximum quantum yield of photosystem II (Fv/Fm). We tested correlations with embolism resistance and dry season water potentials (ΨMD), and calculated safety margins for damage (ΨMD – thresholds) and tested correlations with drought resilience in sap flow and growth. Ψ thresholds for persistent declines in Fv/Fm, indicating resilience, were positively correlated with ΨMD and thresholds for leaf vein embolism. Safety margins for persistent declines in Fv/Fm, but not rehydration capacity, were positively correlated with drought resilience in sap flow.Correlations between resistance and resilience suggest that species' differences in performance during drought are perpetuated after drought, potentially accelerating shifts in forest composition. Resilience to photochemical damage emerged as a promising functional trait to characterize whole-plant drought resilience.

Selection for crop cultivars has largely focused on reproductive traits, while the impacts of global change on crop productivity are expected to depend strongly on the vegetative physiology traits that drive plant resource use and stress tolerance. We evaluated relationships between physiology traits and growing season climate across wine grape cultivars to characterize trait variation across European growing regions. We compiled values from the literature for seven water use and drought tolerance traits and growing season climate. Cultivars with a lower maximum stomatal conductance were associated with regions with a higher mean temperature and mean and maximum vapor pressure deficit (r2=0.39–0.65, P

Knowledge about physiological stress thresholds provides crucial information about plant performance and survival under drought. In this study, we report on the triphasic nature of the relationship between plant water potential (Ψ) at predawn and midday and describe a method that predicts Ψ at stomatal closure and turgor loss exclusively from this water potential curve (WP curve). The method is based on a piecewise linear regression model that was developed to predict the boundaries (termed Θ1 and Θ2) separating the three phases of the curve and corresponding slope values. The method was tested for three economically important woody species. For all species, midday Ψ was much more negative than predawn Ψ during phase I (mild drought), reductions in midday Ψ were minor while predawn Ψ continued to decline during phase II (moderate drought), and midday and predawn Ψ reached similar values during phase III (severe drought). Corresponding measurement of leaf gas exchange indicated that boundary Θ1 between phases I and II coincided with Ψ at stomatal closure. Data from pressure-volume curves demonstrated that boundary Θ2 between phases II and III predicted Ψ at leaf turgor loss. The WP curve method described here is an advanced application of the Scholander-type pressure chamber to categorize plant dehydration under drought into three distinct phases and to predict Ψ thresholds of stomatal closure and turgor loss.

Drought is a major stress for plants, creating a strong selection pressure for traits that enable plant growth and survival in dry environments. Many drought responses are conserved species-wide responses, while others vary among populations distributed across heterogeneous environments. We tested how six populations of the widely distributed California valley oak (Quercus lobata) sampled from contrasting climates would differ in their response to soil drying relative to well-watered controls in a common environment by measuring ecophysiological traits in 93 individuals and gene expression (RNA-seq) in 42 individuals. Populations did not differ in their adjustment of turgor loss point during soil drying, suggesting a generalized species-wide response. Differential expression analysis identified 689 genes with a common response to treatment across populations and 470 genes with population-specific responses. Weighted gene co-expression network analysis (WGCNA) identified groups of genes with similar expression patterns that may be regulated together (gene modules). Several gene modules responded differently to water stress among populations, suggesting regional differences in gene network regulation. Populations from sites with a high mean annual temperature responded to the imposed water stress with significantly greater changes in gene module expression, indicating that these populations may be locally adapted to respond to drought. We propose that this variation among valley oak populations provides a mechanism for differential tolerance to the increasingly frequent and severe droughts in California.

Stomata, the microvalves on leaf surfaces, exert major influences across scales, from plant growth and productivity to global carbon and water cycling. Stomatal opening enables leaf photosynthesis, and plant growth and water use, whereas plant survival of drought depends on stomatal closure. Here we report that stomatal function is constrained by a safety-efficiency trade-off, such that species with greater stomatal conductance under high water availability (gmax) show greater sensitivity to closure during leaf dehydration, i.e., a higher leaf water potential at which stomatal conductance is reduced by 50% (Ψgs50). The gmax - Ψgs50 trade-off and its mechanistic basis is supported by experiments on leaves of California woody species, and in analyses of previous studies of the responses of diverse flowering plant species around the world. Linking the two fundamental key roles of stomata—the enabling of gas exchange, and the first defense against drought—this trade-off constrains the rates of water use and the drought sensitivity of leaves, with potential impacts on ecosystems., Stomata enable gas exchange for photosynthesis but close to promote survival during drought. Here, Henry et al. provide evidence for a safety-efficiency trade-off whereby plants with greater stomatal conductance under well-watered conditions are more sensitive to stomatal closure during dehydration.

Plant carbon transport is controlled by a multitude of parameters both internal and external to the sugar transporting phloem tissue. Sucrose transporter kinetics, conduit hydraulic resistance, and xylem water stress are all hypothesized to impact the amount of carbon delivered to sink tissues. However, the most important traits determining carbon export under drought are not well understood, especially for species with active molecular regulation of sucrose transport. This in turn limits our ability to assess species’ resistances to phloem dysfunction under drought. Here, we use an integrated xylem-phloem-stomatal model to calculate leaf water potential from soil dryness, which is then used to determine gas exchange and phloem pressure gradients. We quantitatively compare the impacts of phloem loading kinetics, including feedbacks between loading and phloem pressure, phloem conduit resistances, and stomatal responses to water stress, on the total carbon export to sinks during drought. Regulating sucrose transporter kinetics which downregulates loading at high phloem pressures prevented runaway viscosity in the phloem sap and was the most important determinant of export rates under drought. In contrast to previous models, we found this feedback mechanism decoupled stomatal traits from phloem export efficiency during drought and increased the operational range of phloem hydraulic resistances.

Leaf density links the resistance to mechanical and hydraulic stress in Mediterranean shrubs as it is associated with the water potential at turgor loss and the moduli of elasticity and strength. Understanding the patterns of hydraulic and mechanical trait variation in vascular plants is critical to predicting species’ stress tolerance. Although previous work has shown that hydraulic and mechanical traits are decoupled in stems, there is little information available for leaves, which are organs more diversified in structure, function, and possibly drought tolerance strategies across habitats. We tested for coordination between leaf hydraulic traits related to drought tolerance and the mechanical resistance of leaves, for 17 shrub species from the arid and semiarid vegetation of the California Floristic Province. Bayesian and phylogenetic correlations showed that across species, hydraulic and mechanical traits both had strong associations with the water potential at turgor loss, and with leaf tissue density. However, leaf maximum hydraulic conductance and the water potential at 50% and 80% loss of hydraulic conductance were statistically independent of two key mechanical traits, the leaf modulus of elasticity and leaf structural strength. Our results suggest that leaf biomechanical traits, which reflect construction costs and contribute to leaf longevity, are decoupled from hydraulic capacity and safety. The independence of hydraulic and mechanical protection in leaves enables a wide range of trait combinations in leaves, which would increase their adaptive potential across ecosystems.

The application of functional traits to predict and explain plant species’ distributions and vital rates has been a major direction in functional ecology for decades, yet numerous physiological traits have not yet been incorporated into the approach. Using commonly measured traits such as leaf mass per area (LMA) and wood density (WD), and additional traits related to water transport, gas exchange and resource economics, including leaf vein, stomatal and wilting traits, we tested hypotheses for Hawaiian wet montane and lowland dry forests (MWF and LDF, respectively): (1) Forests would differ in a wide range of traits as expected from contrasting adaptation; (2) trait values would be more convergent among dry than wet forest species due to the stronger environmental filtering; (3) traits would be intercorrelated within “modules” supporting given functions; (4) relative growth rate (RGR) and mortality rate (m) would correlate with a number of specific traits; with (5) stronger relationships when stratifying by tree size; and (6) RGR and m can be strongly explained from trait‐based models. The MWF species’ traits were associated with adaptation to high soil moisture and nutrient supply and greater shade tolerance, whereas the LDF species’ traits were associated with drought tolerance. Thus, on average, MWF species achieved higher maximum heights than LDF species and had leaves with larger epidermal cells, higher maximum stomatal conductance and CO2 assimilation rate, lower vein lengths per area, higher saturated water content and greater shrinkage when dry, lower dry matter content, higher phosphorus concentration, lower nitrogen to phosphorus ratio, high chlorophyll to nitrogen ratio, high carbon isotope discrimination, high stomatal conductance to nitrogen ratio, less negative turgor loss point and lower WD. Functional traits were more variable in the MWF than LDF, were correlated within modules, and predicted species’ RGR and m across forests, with stronger relationships when stratifying by tree size. Models based on multiple traits predicted vital rates across forests (R2 = 0.70–0.72; p

Tropical forest responses are an important feedback on global change, but changes in forest composition with projected increases in CO2 and drought are highly uncertain. Here we determine shifts in the most competitive plant hydraulic strategy (the evolutionary stable strategy or ESS) from changes in CO2 and drought frequency and intensity. Hydraulic strategies were defined along a spectrum from drought avoidance to tolerance by physiology traits. Drought impacted competition more than CO2 , with elevated CO2 reducing but not reversing drought-induced shifts in the ESS towards more tolerant strategies. Trait plasticity and/or adaptation intensified these shifts by increasing the competitive ability of the drought tolerant relative to the avoidant strategies. These findings predict losses of drought avoidant evergreens from tropical forests under global change, and point to the importance of changes in precipitation during the dry season and constraints on plasticity and adaptation in xylem traits to forest responses.

The mechanisms governing tree drought mortality and recovery remain a subject of inquiry and active debate given their role in the terrestrial carbon cycle and their concomitant impact on climate change. Counter-intuitively, many trees do not die during the drought itself. Indeed, observations globally have documented that trees often grow for several years after drought before mortality. A combination of meta-analysis and tree physiological models demonstrate that optimal carbon allocation after drought explains observed patterns of delayed tree mortality and provides a predictive recovery framework. Specifically, post-drought, trees attempt to repair water transport tissue and achieve positive carbon balance through regrowing drought-damaged xylem. Furthermore, the number of years of xylem regrowth required to recover function increases with tree size, explaining why drought mortality increases with size. These results indicate that tree resilience to drought-kill may increase in the future, provided that CO2 fertilisation facilitates more rapid xylem regrowth.

Leaf hydraulic supply is crucial to maintaining open stomata for CO2 capture and plant growth. During drought-induced dehydration, the leaf hydraulic conductance (Kleaf) declines, which contributes to stomatal closure and, eventually, to leaf death. Previous studies have tended to attribute the decline of Kleaf to embolism in the leaf vein xylem. We visualized at high resolution and quantified experimentally the hydraulic vulnerability of xylem and outside-xylem pathways and modeled their respective influences on plant water transport. Evidence from all approaches indicated that the decline of Kleaf during dehydration arose first and foremost due to the vulnerability of outside-xylem tissues. In vivo x-ray microcomputed tomography of dehydrating leaves of four diverse angiosperm species showed that, at the turgor loss point, only small fractions of leaf vein xylem conduits were embolized, and substantial xylem embolism arose only under severe dehydration. Experiments on an expanded set of eight angiosperm species showed that outside-xylem hydraulic vulnerability explained 75% to 100% of Kleaf decline across the range of dehydration from mild water stress to beyond turgor loss point. Spatially explicit modeling of leaf water transport pointed to a role for reduced membrane conductivity consistent with published data for cells and tissues. Plant-scale modeling suggested that outside-xylem hydraulic vulnerability can protect the xylem from tensions that would induce embolism and disruption of water transport under mild to moderate soil and atmospheric droughts. These findings pinpoint outside-xylem tissues as a central locus for the control of leaf and plant water transport during progressive drought.

Climate change is expected to exacerbate drought for many plants, making drought tolerance a key driver of species and ecosystem responses. Plant drought tolerance is determined by multiple traits, but the relationships among traits, either within individual plants or across species, have not been evaluated for general patterns across plant diversity. We synthesized the published data for stomatal closure, wilting, declines in hydraulic conductivity in the leaves, stems, and roots, and plant mortality for 262 woody angiosperm and 48 gymnosperm species. We evaluated the correlations among the drought tolerance traits across species, and the general sequence of water potential thresholds for these traits within individual plants. The trait correlations across species provide a framework for predicting plant responses to a wide range of water stress from one or two sampled traits, increasing the ability to rapidly characterize drought tolerance across diverse species. Analyzing these correlations also identified correlations among the leaf and stem hydraulic traits and the wilting point, or turgor loss point, beyond those expected from shared ancestry or independent associations with water stress alone. Further, on average, the angiosperm species generally exhibited a sequence of drought tolerance traits that is expected to limit severe tissue damage during drought, such as wilting and substantial stem embolism. This synthesis of the relationships among the drought tolerance traits provides crucial, empirically supported insight into representing variation in multiple traits in models of plant and ecosystem responses to drought.

Salinity tolerance in plant species varies widely due to adaptation and acclimation processes at the cellular and whole-plant scales. In mangroves, extreme substrate salinity induces hydraulic failure and ion excess toxicity and reduces growth and survival, thus suggesting a potentially critical role for physiological acclimation to salinity. We tested the hypothesis that osmotic adjustment, a key type of plasticity that mitigates salinity shock, would take place in coordination with declines in whole-plant hydraulic conductance in a common garden experiment using saplings of three mangrove species with different salinity tolerances (Avicennia germinans L., Rhizophora mangle L. and Laguncularia racemosa (L.) C.F. Gaertn., ordered from higher to lower salinity tolerance). For each mangrove species, four salinity treatments (1, 10, 30 and 50 practical salinity units) were established and the time trajectories were determined for leaf osmotic potential (Ψs), stomatal conductance (gs), whole-plant hydraulic conductance (Kplant) and predawn disequilibrium between xylem and substrate water potentials (Ψpdd). We expected that, for all three species, salinity increments would result in coordinated declines in Ψs, gs and Kplant, and that the Ψpdd would increase with substrate salinity and time of exposure. In concordance with our predictions, reductions in substrate water potential promoted a coordinated decline in Ψs, gs and Kplant, whereas the Ψpdd increased substantially during the first 4 days but dissipated after 7 days, indicating a time lag for equilibration after a change in substratum salinity. Our results show that mangroves confront and partially ameliorate acute salinity stress via simultaneous reductions in Ψs, gs and Kplant, thus developing synergistic physiological responses at the cell and whole-plant scales.

Drought-induced tree mortality has been observed globally and is expected to increase under climate change scenarios, with large potential consequences for the terrestrial carbon sink. Predicting mortality across species is crucial for assessing the effects of climate extremes on forest community biodiversity, composition, and carbon sequestration. However, the physiological traits associated with elevated risk of mortality in diverse ecosystems remain unknown, although these traits could greatly improve understanding and prediction of tree mortality in forests. We performed a meta-analysis on species' mortality rates across 475 species from 33 studies around the globe to assess which traits determine a species' mortality risk. We found that species-specific mortality anomalies from community mortality rate in a given drought were associated with plant hydraulic traits. Across all species, mortality was best predicted by a low hydraulic safety margin-the difference between typical minimum xylem water potential and that causing xylem dysfunction-and xylem vulnerability to embolism. Angiosperms and gymnosperms experienced roughly equal mortality risks. Our results provide broad support for the hypothesis that hydraulic traits capture key mechanisms determining tree death and highlight that physiological traits can improve vegetation model prediction of tree mortality during climate extremes.

Spatial patterns in trait variation reflect underlying community assembly processes, allowing us to test hypotheses about their trait and environmental drivers by identifying the strongest correlates of characteristic spatial patterns. For 43 evergreen tree species (> 1 cm dbh) in a 20-ha seasonal tropical rainforest plot in Xishuangbanna, China, we compared the ability of drought-tolerance traits, other physiological traits, and commonly measured functional traits to predict the spatial patterns expected from the assembly processes of habitat associations, niche-overlap-based competition, and hierarchical competition. We distinguished the neighborhood-scale (0-20 m) patterns expected from competition from larger-scale habitat associations with a wavelet method. Species' drought tolerance and habitat variables related to soil water supply were strong drivers of habitat associations, and drought tolerance showed a significant spatial signal for influencing competition. Overall, the traits most strongly associated with habitat, as quantified using multivariate models, were leaf density, leaf turgor loss point (π(tlp); also known as the leaf wilting point), and stem hydraulic conductivity (r2 range for the best fit models = 0.27-0.36). At neighborhood scales, species spatial associations were positively correlated with similarity in π(tlp), consistent with predictions for hierarchical competition. Although the correlation between π(tlp) and interspecific spatial associations was weak (r2 < 0.01), this showed a persistent influence of drought tolerance on neighborhood interactions and community assembly. Quantifying the full impact of traits on competitive interactions in forests may require incorporating plasticity among individuals within species, especially among specific life stages, and moving beyond individual traits to integrate the impact of multiple traits on whole-plant performance and resource demand.

International audience; Plants are enormously diverse in their traits and ecological adaptation, even within given ecosystems, such as tropical rainforests. Accounting for this diversity in vegetation models poses serious challenges. Global plant functional trait databases have highlighted general trait correlations across species that have considerably advanced this research program. However, it remains unclear whether trait correlations found globally hold within communities, and whether they extend to drought tolerance traits. For 134 individual plants spanning a range of sizes and life forms (tree, liana, understory species) within an Amazonian forest, we measured leaf drought tolerance (leaf water potential at turgor loss point, πtlp), together with 17 leaf traits related to various functions, including leaf economics traits and nutrient composition (leaf mass per area, LMA; and concentrations of C, N, P, K, Ca, and Mg per leaf mass and area), leaf area, water use efficiency (carbon isotope ratio), and time-integrated stomatal conductance and carbon assimilation rate per leaf mass and area. We tested trait coordination and the ability to estimate πtlp from the other traits through model selection. Performance and transferability of the best predictive model were assessed through cross-validation. πtlp was positively correlated with leaf area, and with N, P and K concentrations per leaf mass, but not with LMA or any other studied trait. Five axes were needed to account for > 80% of trait variation, but only three of them explained more variance than expected at random. The best model explained only 30% of the variation in πtlp, and out-sample predictive performance was variable across life forms or canopy strata, suggesting a limited transferability of the model. Synthesis. We found a weak correlation among leaf drought tolerance and other leaf traits within a forest community. We conclude that higher trait dimensionality than assumed under the leaf economics spectrum may operate among leaves within plant communities, with important implications for species coexistence and responses to changing environmental conditions, and also for the representation of community diversity in vegetation models.

Forest leaf area has enormous leverage on the carbon cycle because it mediates both forest productivity and resilience to climate extremes. Despite widespread evidence that trees are capable of adjusting to changes in environment across both space and time through modifying carbon allocation to leaves, many vegetation models use fixed carbon allocation schemes independent of environment, which introduces large uncertainties into predictions of future forest responses to atmospheric CO2 fertilization and anthropogenic climate change. Here, we develop an optimization-based model, whereby tree carbon allocation to leaves is an emergent property of environment and plant hydraulic traits. Using a combination of meta-analysis, observational datasets, and model predictions, we find strong evidence that optimal hydraulic-carbon coupling explains observed patterns in leaf allocation across large environmental and CO2 concentration gradients. Furthermore, testing the sensitivity of leaf allocation strategy to a diversity in hydraulic and economic spectrum physiological traits, we show that plant hydraulic traits in particular have an enormous impact on the global change response of forest leaf area. Our results provide a rigorous theoretical underpinning for improving carbon cycle predictions through advancing model predictions of leaf area, and underscore that tree-level carbon allocation to leaves should be derived from first principles using mechanistic plant hydraulic processes in the next generation of vegetation models.

Environmental filtering and dispersal limitation can both maintain diversity in plant communities by aggregating conspecifics, but parsing the contribution of each process has proven difficult empirically. Here, we assess the contribution of filtering and dispersal limitation to the spatial aggregation patterns of 456 tree species in a hyperdiverse Amazonian forest and find distinct functional trait correlates of interspecific variation in these processes. Spatial point process model analysis revealed that both mechanisms are important drivers of intraspecific aggregation for the majority of species. Leaf drought tolerance was correlated with species topographic distributions in this aseasonal rainforest, showing that future increases in drought severity could significantly impact community structure. In addition, seed mass was associated with the spatial scale and density of dispersal-related aggregation. Taken together, these results suggest environmental filtering and dispersal limitation act in concert to influence the spatial and functional structure of diverse forest communities.

Tropical forest responses are an important feedback on global change, but changes in forest composition with projected increases in CO

PREMISE OF THE STUDY Studies across diverse species have established theory for the contribution of leaf traits to plant drought tolerance. For example, species in more arid climates tend to have smaller leaves of higher vein density, higher leaf mass per area, and more negative osmotic potential at turgor loss point (πTLP ). However, few studies have tested these associations for species within a given lineage that have diversified across an aridity gradient. METHODS We analyzed the anatomy and physiology of 10 Ceanothus (Rhamnaceae) species grown in a common garden for variation between and within "wet" and "dry" subgenera (Ceanothus and Cerastes, respectively) and analyzed a database for 35 species for leaf size and leaf mass per area (LMA). We used a phylogenetic generalized least squares approach to test hypothesized relationships among traits, and of traits with climatic aridity in the native range. We also tested for allometric relationships among anatomical traits. KEY RESULTS Leaf form, anatomy, and drought tolerance varied strongly among species within and between subgenera. Cerastes species had specialized anatomy including hypodermis and encrypted stomata that may confer superior water storage and retention. The osmotic potentials at turgor loss point (πTLP ) and full turgor (πo ) showed evolutionary correlations with the aridity index (AI) and precipitation of the 10 species' native distributions, and LMA with potential evapotranspiration for the 35 species in the larger database. We found an allometric correlation between upper and lower epidermal cell wall thicknesses, but other anatomical traits diversified independently. CONCLUSIONS Leaf traits and drought tolerance evolved within and across lineages of Ceanothus consistently with climatic distributions. The πTLP has signal to indicate the evolution of drought tolerance within small clades.

The need for rigorous analyses of climate impacts has never been more crucial. Current textbooks state that climate directly influences ecosystem annual net primary productivity (NPP), emphasizing the urgent need to monitor the impacts of climate change. A recent paper challenged this consensus, arguing, based on an analysis of NPP for 1247 woody plant communities across global climate gradients, that temperature and precipitation have negligible direct effects on NPP and only perhaps have indirect effects by constraining total stand biomass (Mtot ) and stand age (a). The authors of that study concluded that the length of the growing season (lgs ) might have a minor influence on NPP, an effect they considered not to be directly related to climate. In this article, we describe flaws that affected that study's conclusions and present novel analyses to disentangle the effects of stand variables and climate in determining NPP. We re-analyzed the same database to partition the direct and indirect effects of climate on NPP, using three approaches: maximum-likelihood model selection, independent-effects analysis, and structural equation modeling. These new analyses showed that about half of the global variation in NPP could be explained by Mtot combined with climate variables and supported strong and direct influences of climate independently of Mtot , both for NPP and for net biomass change averaged across the known lifetime of the stands (ABC = average biomass change). We show that lgs is an important climate variable, intrinsically correlated with, and contributing to mean annual temperature and precipitation (Tann and Pann ), all important climatic drivers of NPP. Our analyses provide guidance for statistical and mechanistic analyses of climate drivers of ecosystem processes for predictive modeling and provide novel evidence supporting the strong, direct role of climate in determining vegetation productivity at the global scale.

The influence of the dynamics of leaf hydraulic conductance (K-leaf) diurnally and during dehydration on stomatal conductance and photosynthesis remains unclear. Using the model species Arabidopsis (Arabidopsis thaliana ecotype Columbia-0), we applied a multitiered approach including physiological measurements, high-resolution x-ray microcomputed tomography, and modeling at a range of scales to characterize (1) K-leaf decline during dehydration; (2) its basis in the hydraulic conductances of leaf xylem and outside-xylem pathways (K-ox); (3) the dependence of its dynamics on irradiance; (4) its impact on diurnal patterns of stomatal conductance and photosynthetic rate; and (5) its influence on gas exchange and survival under simulated drought regimes. Arabidopsis leaves showed strong vulnerability to dehydration diurnally in both gas exchange and hydraulic conductance, despite lack of xylem embolism or conduit collapse above the turgor loss point, indicating a pronounced sensitivity of K-ox to dehydration. K(leaf )increased under higher irradiance in well-hydrated leaves across the full range of water potential, but no shift in K-leaf vulnerability was observed. Modeling indicated that responses to dehydration and irradiance are likely attributable to changes in membrane permeability and that a dynamic K-ox would contribute strongly to stomatal closure, improving performance, survival, and efficient water use during drought. These findings for Columbia-0 provide a baseline for assessing variation across genotypes in hydraulic traits and their influence on gas exchange during dehydration.

1. Water availability is a key determinant of forest ecosystem function and tree species distributions. While droughts are increasing in frequency in many ecosystems, including in the tropics, plant responses to water supply vary with species and drought intensity and are therefore difficult to model. Based on physiological first principles, we hypothesized that trees with a lower turgor loss point (pi(tlp)), that is, a more negative leaf water potential at wilting, would maintain water transport for longer into a dry season. 2. We measured sapflux density of 22 mature trees of 10 species during a dry season in an Amazonian rainforest, quantified sapflux decline as soil water content decreased and tested its relationship to tree pi(tlp), size and leaf predawn and midday water potentials measured after the onset of the dry season. 3. The measured trees varied strongly in the response of water use to the seasonal drought, with sapflux at the end of the dry season ranging from 37 to 117% (on average 83 +/- 5 %) of that at the beginning of the dry season. The decline of water transport as soil dried was correlated with tree pi(tlp) (Spearman's rho >= 0.63), but not with tree size or predawn and midday water potentials. Thus, trees with more drought-tolerant leaves better maintained water transport during the seasonal drought. 4. Our study provides an explicit correlation between a trait, measurable at the leaf level, and whole-plant performance under drying conditions. Physiological traits such as pi(tlp) can be used to assess and model higher scale processes in response to drying conditions.

Many species face increasing drought under climate change. Plasticity has been predicted to strongly influence species' drought responses, but broad patterns in plasticity have not been examined for key drought tolerance traits, including turgor loss or 'wilting' point (πtlp ). As soil dries, plants shift πtlp by accumulating solutes (i.e. 'osmotic adjustment'). We conducted the first global analysis of plasticity in Δπtlp and related traits for 283 wild and crop species in ecosystems worldwide. Δπtlp was widely prevalent but moderate (-0.44 MPa), accounting for 16% of post-drought πtlp. Thus, pre-drought πtlp was a considerably stronger predictor of post-drought πtlp across species of wild plants. For cultivars of certain crops Δπtlp accounted for major differences in post-drought πtlp. Climate was correlated with pre- and post-drought πtlp, but not Δπtlp. Thus, despite the wide prevalence of plasticity, πtlp measured in one season can reliably characterise most species' constitutive drought tolerances and distributions relative to water supply.

A favorable leaf water balance can be achieved through coordinated variation in multiple leaf traits. However, the role of evolutionary history in producing this trait coordination is not well understood, especially in epiphytic species. To address these limitations, we measured 11 leaf traits for 19 Dendrobium species grown in the same environment, and used phylogenetically independent contrasts to test how the co-evolution of these traits has contributed to leaf water balance in epiphytes. Our results show that more closely correlated species had similar habitats, geographical distribution, and some leaf traits (leaf density, upper cuticle thickness, and stomatal index), whereas distantly related species did not exhibit such similarities. Species originating from forests below the altitude of 1,500 m exhibited smaller leaf area, stomatal area, and stomatal index, but thicker leaves than those from forests above the altitude of 1,500 m. Stomatal density was evolutionarily correlated with vein density, upper and lower epidermal thicknesses, and leaf density. Leaf thickness was significantly correlated with stomatal density and vein density, while leaf area was significantly correlated with epidermal cell thicknesses, stomatal density, and stomatal area. These results found that both the environment and evolutionary history significantly affected leaf functional traits and ecological characteristics in Dendrobium, and supported the hypothesis that co-evolution in leaf traits is an important contributor to leaf water balance in Dendrobium. This study provides new insights into the evolution of ecological strategies in epiphytic orchids.

Pan-tropically, liana density increases with decreasing rainfall and increasing seasonality. This pattern has led to the hypothesis that lianas display a growth advantage over trees under dry conditions. However, the physiological mechanisms underpinning this hypothesis remain elusive. A key trait influencing leaf and plant drought tolerance is the leaf water potential at turgor loss point (πtlp).πtlpadjusts under drier conditions and this contributes to improved leaf drought tolerance. For co-occurring Amazonian tree (n= 247) and liana (n= 57) individuals measured during the dry and the wet seasons, lianas showed a stronger osmotic adjustment than trees. Liana leaves were less drought-tolerant than trees in the wet season, but reached similar drought tolerances during the dry season. Stronger osmotic adjustment in lianas would contribute to turgor maintenance, a critical prerequisite for carbon uptake and growth, and to the success of lianas relative to trees in growth under drier conditions.

1. Across plant species, drought tolerance and distributions with respect to water availability are strongly correlated with two physiological traits, the leaf water potential at wilting, that is, turgor loss point (ptlp), and the cell solute potential at full hydration, that is, osmotic potential (po). We present methods to determine these parameters 30 times more rapidly than the standard pressurevolume (pv) curve approach, making feasible community-scale studies of plant drought tolerance. 2. We optimized existing methods for measurements of pi o using vapour-pressure osmometry of freeze-thawed leaf discs from 30 species growing in two precipitation regimes, and developed the first regression relationships to accurately estimate pressurevolume curve values of both pi o and pi tlp from osmometer values. 3. The pi o determined with the osmometer (pi osm) was an excellent predictor of the pi o determined from the pv curve (pi pv,r2 = 0.80). Although the correlation of pi osm and pi pv enabled prediction, the relationship departed from the 1 : 1 line. The discrepancy between the methods could be quantitatively accounted for by known sources of error in osmometer measurements, that is, dilution by the apoplastic water, and solute dissolution from destroyed cell walls. An even stronger prediction of pi pv could be made using pi osm, leaf density (rho), and their interaction (r2 = 0.85, all P l 2 x 10-10). 4. The pi osm could also be used to predict pi tlp (r2 = 0.86). Indeed, pi osm was a better predictor of pi tlp than leaf mass per unit area (LMA; r2 = 0.54), leaf thickness (T; r2 = 0.12), rho (r2 = 0.63), and leaf dry matter content (LDMC; r2 = 0.60), which have been previously proposed as drought tolerance indicators. Models combining posm with LMA, T, rho, or LDMC or other pv curve parameters (i.e. elasticity and apoplastic fraction) did not significantly improve prediction of pi tlp. 5. This osmometer method enables accurate measurements of drought tolerance traits across a wide range of leaf types and for plants with diverse habitat preferences, with a fraction of effort of previous methods. We expect it to have wide application for predicting species responses to climate variability and for assessing ecological and evolutionary variation in drought tolerance in natural populations and agricultural cultivars.

Increasing drought is one of the most critical challenges facing species and ecosystems worldwide, and improved theory and practices are needed for quantification of species tolerances. Leaf water potential at turgor loss, or wilting (π(tlp) ), is classically recognised as a major physiological determinant of plant water stress response. However, the cellular basis of π(tlp) and its importance for predicting ecological drought tolerance have been controversial. A meta-analysis of 317 species from 72 studies showed that π(tlp) was strongly correlated with water availability within and across biomes, indicating power for anticipating drought responses. We derived new equations giving both π(tlp) and relative water content at turgor loss point (RWC(tlp) ) as explicit functions of osmotic potential at full turgor (π(o) ) and bulk modulus of elasticity (e). Sensitivity analyses and meta-analyses showed that π(o) is the major driver of π(tlp) . In contrast, e plays no direct role in driving drought tolerance within or across species, but sclerophylly and elastic adjustments act to maintain RWC(tlp,) preventing cell dehydration, and additionally protect against nutrient, mechanical and herbivory stresses independent of drought tolerance. These findings clarify biogeographic trends and the underlying basis of drought tolerance parameters with applications in comparative assessments of species and ecosystems worldwide.

Strong positive correlations between the maximum rate of canopy photosynthesis, canopy-averaged foliar nitrogen concentration, and canopy albedo have been shown in previous studies. While leaf-level relationships between photosynthetic capacity and foliar nitrogen are well documented, it is not clear whether leaf-level relationships between solar-weighted reflectance and nitrogen underlie the canopy-scale patterns. Using an integrating sphere, we measured the reflectance and transmittance (350–2500 nm) of both individual leaves and multileaf stacks. Samples were collected from 12 broadleaf species at the Harvard Forest in central Massachusetts, USA. Across all species, foliar nitrogen (both mass-based nitrogen concentration and area-based nitrogen content) and leaf mass / area ratio were negatively, rather than positively, correlated with solar-weighted reflectance and transmittance in ultraviolet–visible and near-infrared wavelength bands (350–700 nm and 700–2500 nm, respectively). Leaf-level anatomy and biochemistry, therefore, do not appear to drive the canopy-level association between increasing foliar nitrogen content and increasing canopy albedo. This suggests that interactions between leaf optical properties and structural canopy-scale traits that correlate with nitrogen content (perhaps some combination of crown shape, leaf area index, leaf angular distribution, or other structural characteristics of the canopy), may instead underlie the previously observed relationship between nitrogen and canopy-level shortwave albedo.

International audience; Amazonian tree communities have already been seriously impacted by extreme natural droughts, and intense droughts are predicted to increase in frequency. However, our current knowledge of Amazonian tree species' responses to water stress remains limited, as plant trait databases include few drought tolerance traits, impeding the application and predictive power of models. Here we explored how leaf water potential at turgor loss point (π tlp), a determinant of leaf drought tolerance, varies with species life history, season, tree size and irradiance within a forest in French Guiana. First, we provided a further direct validation of a rapid method of π tlp determination based on osmometer measurements of leaf osmotic potential at full hydration for five Amazonian tree species. Next, we analysed a dataset of 131 π tlp values for a range of species, seasons, size (including saplings), and leaf exposure. We found that early-successional species had less drought-tolerant leaves than late-successional species. Species identity was the major driver of π tlp variation, whereas season, canopy tree size and leaf exposure explained little variation. Shifts in π tlp from saplings to canopy trees varied across species, and sapling leaf drought tolerance was a moderate predictor of canopy tree leaf drought tolerance. Given its low within-species variability, we propose that π tlp is a robust trait, and is useful as one index of species' drought tolerance. We also suggest that measuring this trait would considerably advance our knowledge on leaf drought tolerance in hyperdiverse communities and would thus likely shed light on the resilience of such vulnerable species-rich ecosystem.

International audience; 1. Amazonian droughts are predicted to become increasingly frequent and intense, and the vulnerability of Amazonian trees has become increasingly documented. However, little is known about the physiological mechanisms and the diversity of drought tolerance of tropical trees due to the lack of quantitative measurements. 2. Leaf water potential at wilting or turgor loss point (pi(tlp)) is a determinant of the tolerance of leaves to drought stress and contributes to plant-level physiological drought tolerance. Recently, it has been demonstrated that leaf osmotic water potential at full hydration (pi(o)) is tightly correlated with pi(tlp). Estimating pi(tlp) from osmometer measurements of pi(o) is much faster than the standard pressure-volume curve approach of pi(tlp) determination. We used this technique to estimate pi(tlp) for 165 trees of 71 species, at three sites within forests in French Guiana. Our data set represents a significant increase in available data for this trait for tropical tree species. 3. Tropical trees showed a wider range of drought tolerance than previously found in the literature, pi(tlp) ranging from -1.4 to -3.2 MPa. This range likely corresponds in part to adaptation and acclimation to occasionally extreme droughts during the dry season. 4. Leaf-level drought tolerance varied across species, in agreement with the available published observations of species variation in drought-induced mortality. On average, species with a more negative pi(tlp) (i.e. with greater leaf-level drought tolerance) occurred less frequently across the region than drought-sensitive species. 5. Across individuals, pi(tlp) correlated positively but weakly with leaf toughness (R-2 = 0.22, P = 0.04) and leaf thickness (R-2 = 0.03, P = 0.03). No correlation was detected with other functional traits (leaf mass per area, leaf area, nitrogen or carbon concentrations, carbon isotope ratio, sapwood density or bark thickness). 6. The variability in pi(tlp) among species indicates a potential for highly diverse species responses to drought within given forest communities. Given the weak correlations between pi(tlp) and traditionally measured plant functional traits, vegetation models seeking to predict forest response to drought should integrate improved quantification of comparative drought tolerance among tree species.

Increasing drought is one of the most critical challenges facing species and ecosystems worldwide, and improved theory and practices are needed for quantification of species tolerances. Leaf water potential at turgor loss, or wilting (π(tlp) ), is classically recognised as a major physiological determinant of plant water stress response. However, the cellular basis of π(tlp) and its importance for predicting ecological drought tolerance have been controversial. A meta-analysis of 317 species from 72 studies showed that π(tlp) was strongly correlated with water availability within and across biomes, indicating power for anticipating drought responses. We derived new equations giving both π(tlp) and relative water content at turgor loss point (RWC(tlp) ) as explicit functions of osmotic potential at full turgor (π(o) ) and bulk modulus of elasticity (ε). Sensitivity analyses and meta-analyses showed that π(o) is the major driver of π(tlp) . In contrast, ε plays no direct role in driving drought tolerance within or across species, but sclerophylly and elastic adjustments act to maintain RWC(tlp,) preventing cell dehydration, and additionally protect against nutrient, mechanical and herbivory stresses independent of drought tolerance. These findings clarify biogeographic trends and the underlying basis of drought tolerance parameters with applications in comparative assessments of species and ecosystems worldwide.

Foliar nitrogen has been shown to be positively correlated with midsummer canopy albedo and canopy near infrared (NIR) reflectance over a broad range of plant functional types (e.g., forests, grasslands, and agricultural lands). To date, the mechanism(s) driving the nitrogen–albedo relationship have not been established, and it is unknown whether factors affecting nitrogen availability will also influence albedo. To address these questions, we examined variation in foliar nitrogen in relation to leaf spectral properties, leaf mass per unit area, and leaf water content for three deciduous species subjected to either nitrogen (Harvard Forest, MA, and Oak Ridge, TN) or CO(2) fertilization (Oak Ridge, TN). At Oak Ridge, we also obtained canopy reflectance data from the airborne visible/infrared imaging spectrometer (AVIRIS) to examine whether canopy-level spectral responses were consistent with leaf-level results. At the leaf level, results showed no differences in reflectance or transmittance between CO(2) or nitrogen treatments, despite significant changes in foliar nitrogen. Contrary to our expectations, there was a significant, but negative, relationship between foliar nitrogen and leaf albedo, a relationship that held for both full spectrum leaf albedo as well as leaf albedo in the NIR region alone. In contrast, remote sensing data indicated an increase in canopy NIR reflectance with nitrogen fertilization. Collectively, these results suggest that altered nitrogen availability can affect canopy albedo, albeit by mechanisms that involve canopy-level processes rather than changes in leaf-level reflectance.