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Abstract Understanding the impact of altitude on leaf hydraulic, gas exchange, and economic traits is crucial for comprehending vegetation properties and ecosystem functioning. This knowledge also helps to elucidate species' functional strategies regarding their vulnerability or resilience to global change effects in alpine environments. Here, we conducted a global study of dataset encompassing leaf hydraulic, gas exchange, and economic traits for 3391 woody species. The results showed that high‐altitude species possessed greater hydraulic safety (KleafP50), higher water use efficiency (WUEi) and conservative resource use strategy such as higher leaf mass per area, longer leaf lifespan, lower area‐based leaf nitrogen and phosphorus contents, and lower rates of photosynthesis and dark respiration. Conversely, species at lower altitudes exhibited lower hydraulic safety (KleafP50), lower water use efficiency (WUEi) and an acquisitive resource use strategy. These global patterns of leaf traits in relation to altitude reveal the strategies that alpine plants employ for hydraulic safety, water use efficiency, and resource, which have important implications for predicting forest productivity and acclimation to rapid climate change.more » « lessFree, publicly-accessible full text available March 1, 2026
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Abstract Rising global temperatures and vapor pressure deficits (VPDs) are increasing plant water demand and becoming major drivers of large-scale plant mortality. Controlling transient leaf water loss after stomatal closure (minimum stomatal conductance [gmin]) is recognized as a key trait determining how long plants survive during soil drought. Yet, substantial uncertainty remains regarding how gmin responds to elevated temperatures and VPD and the underlying mechanisms. We measured gmin in 24 Quercus species from temperate and Mediterranean climates to determine whether gmin was sensitive to a coupled temperature and VPD increase. We also explored mechanistic links to phenology, climate, evolutionary history, and leaf anatomy. We found that gmin in all species exhibited a nonlinear negative temperature and VPD dependence. At 25 °C (VPD = 2.2 kPa), gmin varied from 1.19 to 8.09 mmol m−2 s−1 across species but converged to 0.57 ± 0.06 mmol m−2 s−1 at 45 °C (VPD = 6.6 kPa). In a subset of species, the effect of temperature and VPD on gmin was reversible and linked to the degree of stomatal closure, which was greater at 45 °C than at 25 °C. Our results show that gmin is dependent on temperature and VPD, is highly conserved in Quercus species, and is linked to leaf anatomy and stomatal behavior.more » « lessFree, publicly-accessible full text available December 23, 2025
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Sensitive Hydraulic and Stomatal Decline in Extreme Drought Tolerant Species of California CeanothusABSTRACT Identifying the physiological mechanisms by which plants are adapted to drought is critical to predict species responses to climate change. We measured the responses of leaf hydraulic and stomatal conductances (Kleafandgs, respectively) to dehydration, and their association with anatomy, in seven species of CaliforniaCeanothusgrown in a common garden, including some of the most drought‐tolerant species in the semi‐arid flora. We tested for matching of maximum hydraulic supply and demand and quantified the role of decline ofKleafin driving stomatal closure. AcrossCeanothusspecies, maximumKleafandgswere negatively correlated, and bothKleafandgsshowed steep declines with decreasing leaf water potential (i.e., a high sensitivity to dehydration). The leaf water potential at 50% decline ingswas linked with a low ratio of maximum hydraulic supply to demand (i.e., maximumKleaf:gs). This sensitivity ofgs, combined with low minimum epidermal conductance and water storage, could contribute to prolonged leaf survival under drought. The specialized anatomy of subg.Cerastesincludes trichomous stomatal crypts and pronounced hypodermis, and was associated with higher water use efficiency and water storage. Combining our data with comparative literature of other California species, species of subg. Cerastesshow traits associated with greater drought tolerance and reliance on leaf water storage relative to other California species. In addition to drought resistance mechanisms such as mechanical protection and resistance to embolism, drought avoidance mechanisms such as sensitive stomatal closure could contribute importantly to drought tolerance in dry‐climate adapted species.more » « lessFree, publicly-accessible full text available February 1, 2026
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Abstract Soil nutrients and water availability are strong drivers of tropical tree species distribution across scales. However, the physiological mechanisms underlying environmental filtering along these gradients remain incompletely understood. Previous studies mostly focused on univariate variation in structural traits, but a more integrative approach combining multiple physiological traits is needed to fully portray species functional strategies.We measured nine leaf functional traits related to trees' resource capture and hydraulic strategies for 552 individuals belonging to 21 tropical tree species across an environmental gradient in Amazonian forests. Our sampling included generalist and specialist species fromterra firme(TF) and seasonally flooded (SF) forests. We tested the influence of the topographic wetness index, a proxy for soil moisture and nutrient gradients, on each trait separately and on the trait integration through multivariate indices computed from the eigenvalues of a principal component analysis on the traits of the species. Finally, we evaluated intraspecific trait variability (ITV) for generalists and specialists by calculating the coefficient of variation for each trait.Results showed that (1) the environment had a greater influence on trait syndromes than single trait variation. Moreover, (2) SF specialist species expressed a stronger leaf trait coordination than TF specialist species. Furthermore, (3) the ability of generalist species to occupy a broader range of environments was not reflected by a larger ITV than specialist species but by the capacity to change trait coordination across environments.Our work highlights the need to investigate functional strategies as multidimensional syndromes in physiological trait space to fully understand and predict species distribution along environmental gradients. Read the freePlain Language Summaryfor this article on the Journal blog.more » « lessFree, publicly-accessible full text available January 1, 2026
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Abstract Leaf surface conductance to water vapor and CO2 across the epidermis (gleaf) strongly determines the rates of gas exchange. Thus, clarifying the drivers of gleaf has important implications for resolving the mechanisms of photosynthetic productivity and leaf and plant responses and tolerance to drought. It is well recognized that gleaf is a function of the conductances of the stomata (gs) and of the epidermis + cuticle (gec). Yet, controversies have arisen around the relative roles of stomatal density (d) and size (s), fractional stomatal opening (α; aperture relative to maximum), and gec in determining gleaf. Resolving the importance of these drivers is critical across the range of leaf surface conductances, from strong stomatal closure under drought (gleaf,min), to typical opening for photosynthesis (gleaf,op), to maximum achievable opening (gleaf,max). We derived equations and analyzed a compiled database of published and measured data for approximately 200 species and genotypes. On average, within and across species, higher gleaf,min was determined 10 times more strongly by α and gec than by d and negligibly by s; higher gleaf,op was determined approximately equally by α (47%) and by stomatal anatomy (45% by d and 8% by s), and negligibly by gec; and higher gleaf,max was determined entirely by d. These findings clarify how diversity in stomatal functioning arises from multiple structural and physiological causes with importance shifting with context. The rising importance of d relative to α, from gleaf,min to gleaf,op, enables even species with low gleaf,min, which can retain leaves through drought, to possess high d and thereby achieve rapid gas exchange in periods of high water availability.more » « less
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Summary Grasses are exceptionally productive, yet their hydraulic adaptation is paradoxical. Among C3grasses, a high photosynthetic rate (Aarea) may depend on higher vein density (Dv) and hydraulic conductance (Kleaf). However, the higherDvof C4grasses suggests a hydraulic surplus, given their reduced need for highKleafresulting from lower stomatal conductance (gs).Combining hydraulic and photosynthetic physiological data for diverse common garden C3and C4species with data for 332 species from the published literature, and mechanistic modeling, we validated a framework for linkages of photosynthesis with hydraulic transport, anatomy, and adaptation to aridity.C3and C4grasses had similarKleafin our common garden, but C4grasses had higherKleafthan C3species in our meta‐analysis. Variation inKleafdepended on outside‐xylem pathways. C4grasses have highKleaf : gs, which modeling shows is essential to achieve their photosynthetic advantage.Across C3grasses, higherAareawas associated with higherKleaf, and adaptation to aridity, whereas for C4species, adaptation to aridity was associated with higherKleaf : gs. These associations are consistent with adaptation for stress avoidance.Hydraulic traits are a critical element of evolutionary and ecological success in C3and C4grasses and are crucial avenues for crop design and ecological forecasting.more » « less
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Abstract While variation in mean annual precipitation (MAP) of the native habitat of a species has been shown to determine the ability of a species to resist a hydraulic decrease during drought, it remains unknown whether these variations in MAP also influence the ability of a species to recover and survive drought. Leaf hydraulic and gas exchange recovery following drought and the underlying mechanisms of these responses in sixCaraganaspecies from habitats along a large precipitation gradient were investigated during rehydration in a common garden. The gas exchange of species from arid habitats recovered more rapidly during rehydration after mild, moderate and severe drought stress treatments than species from humid habitats. The recovery of gas exchange was not associated with foliar abscisic acid concentration, but tightly related to the recovery of leaf hydraulic conductance (Kleaf). The recovery ofKleafwas associated with the loss ofKleafduring dehydration under mild and moderate drought stress, and to leaf xylem embolism formation under severe drought stress. Results pointed to the different ability to recover in gas exchange in sixCaraganaspecies post‐drought is associated with the MAP of the species in its native habitat.more » « less
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Summary A surge of papers have reported low leaf vulnerability to xylem embolism during drought. Here, we focus on the less studied, and more sensitive, outside‐xylem leaf hydraulic responses to multiple internal and external conditions. Studies of 34 species have resolved substantial vulnerability to dehydration of the outside‐xylem pathways, and studies of leaf hydraulic responses to light also implicate dynamic outside‐xylem responses. Detailed experiments suggest these dynamic responses arise at least in part from strong control of radial water movement across the vein bundle sheath. While leaf xylem vulnerability may influence leaf and plant survival during extreme drought, outside‐xylem dynamic responses are important for the control and resilience of water transport and leaf water status for gas exchange and growth.more » « less
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Abstract Allometric relationships among the dimensions of leaves and their cells hold across diverse eudicotyledons, but have remained untested in the leaves of grasses. We hypothesised that geometric (proportional) allometries of cell sizes across tissues and of leaf dimensions would arise due to the coordination of cell development and that of cell functions such as water, nutrient and energy transport, and that cell sizes across tissues would be associated with light‐saturated photosynthetic rate. We tested predictions across 27 globally distributed C3and C4grass species grown in a common garden. We found positive relationships among average cell sizes within and across tissues, and of cell sizes with leaf dimensions. Grass leaf anatomical allometries were similar to those of eudicots, with exceptions consistent with the fewer cell layers and narrower form of grass leaves, and the specialised roles of epidermis and bundle sheath in storage and leaf movement. Across species, mean cell sizes in each tissue were associated with light‐saturated photosynthetic rate per leaf mass, supporting the functional coordination of cell sizes. These findings highlight the generality of evolutionary allometries within the grass lineage and their interlinkage with coordinated development and function.more » « less
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Abstract Intra‐specific trait variation (ITV) plays a role in processes at a wide range of scales from organs to ecosystems across climate gradients. Yet, ITV remains rarely quantified for many ecophysiological traits typically assessed for species means, such as pressure volume (PV) curve parameters including osmotic potential at full turgor and modulus of elasticity, which are important in plant water relations. We defined a baseline “reference ITV” (ITVref) as the variation among fully exposed, mature sun leaves of replicate individuals of a given species grown in similar, well‐watered conditions, representing the conservative sampling design commonly used for species‐level ecophysiological traits. We hypothesized that PV parameters would show low ITVrefrelative to other leaf morphological traits, and that their intraspecific relationships would be similar to those previously established across species and proposed to arise from biophysical constraints. In a database of novel and published PV curves and additional leaf structural traits for 50 diverse species, we found low ITVreffor PV parameters relative to other morphological traits, and strong intraspecific relationships among PV traits. Simulation modeling showed that conservative ITVrefenables the use of species‐mean PV parameters for scaling up from spectroscopic measurements of leaf water content to enable sensing of leaf water potential.more » « less
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