Stomatal closure as a driver of minimum leaf conductance declines at high temperature and vapor pressure deficit in Quercus.

Rising global temperatures and vapor pressure deficits (VPD) are increasing plant water demand and becoming major drivers of large-scale plant mortality. Controlling transient leaf water loss after stomatal closure (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 if 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 non-linear 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.

Extension of Kelvin's equation to dipolar colloids.

Vapor pressure refers to the pressure exerted by the vapor phase in thermodynamic equilibrium with either its liquid or solid phase. An important class of active matter is field-driven colloids. A suspension of dipolar colloids placed in a high-frequency rotating magnetic field undergoes a nonequilibrium phase transition into a dilute and dense phase, akin to liquid­vapor coexistence in a simple fluid. Here, we compute the vapor pressure of this colloidal fluid. The number of particles that exist as the dilute bulk phase versus condensed cluster phases can be directly visualized. An exponential relationship between vapor pressure and effective temperature is determined as a function of applied field strength, analogous to the thermodynamic expression between vapor pressure and temperature found for pure liquids. Additionally, we demonstrate the applicability of Kelvin's equation to this field-driven system. In principle, this appears to be in conflict with macroscopic thermodynamic assumptions due to the nonequilibrium and discrete nature of this colloidal system. However, the curvature of the vapor­liquid interface provides a mechanical equilibrium characterized by interfacial tension that connects the condensed clusters observed with these active fluids to classical colligative fluid properties.

Suppressing Se Vacancies in Sb2Se3 Photocathode by In Situ Annealing with Moderate Se Vapor Pressure for Efficient Photoelectrochemical Water Splitting.

Sb2Se3 emerges as a promising material for solar energy conversion devices. Unfortunately, the common deep-level defect VSe (selenium vacancy) in Sb2Se3 results in a low solar conversion efficiency. The post selenization process has been widely adopted for suppressing VSe. However, the effect of selenization on suppressing VSe is often compromised and even more VSe are induced due to defect-correlation. Herein, high-quality Sb2Se3 films are prepared using an unconventional selenization process, with precisely regulating in situ annealing Se vapor pressure. It is found that moderate Se vapor pressure annealing can efficiently suppress VSe by overcoming defect-correlation, as well as promotes grain growth and forms a better heterojunction band alignment. Consequently, the Sb2Se3 photocathode shows a high-level photocurrent of 19.5 mA cm-2 at 0 VRHE, an onset potential of 0.40 VRHE and a half-cell solar-to-hydrogen conversion efficiency of 1.9%, owing to the inhibited charge recombination, excellent charge transport and interface charge extraction. This work provides a significant insight to suppress deep-level defect VSe by adjusting Se vapor pressure for efficient Sb2Se3 photocathode.

The stomatal response to vapor pressure deficit drives the apparent temperature response of photosynthesis in tropical forests.

As temperature rises, net carbon uptake in tropical forests decreases, but the underlying mechanisms are not well understood. High temperatures can limit photosynthesis directly, for example by reducing biochemical capacity, or indirectly through rising vapor pressure deficit (VPD) causing stomatal closure. To explore the independent effects of temperature and VPD on photosynthesis we analyzed photosynthesis data from the upper canopies of two tropical forests in Panama with Generalized Additive Models. Stomatal conductance and photosynthesis consistently decreased with increasing VPD, and statistically accounting for VPD increased the optimum temperature of photosynthesis (Topt) of trees from a VPD-confounded apparent Topt of c. 30-31°C to a VPD-independent Topt of c. 33-36°C, while for lianas no VPD-independent Topt was reached within the measured temperature range. Trees and lianas exhibited similar temperature and VPD responses in both forests, despite 1500 mm difference in mean annual rainfall. Over ecologically relevant temperature ranges, photosynthesis in tropical forests is largely limited by indirect effects of warming, through changes in VPD, not by direct warming effects of photosynthetic biochemistry. Failing to account for VPD when determining Topt misattributes the underlying causal mechanism and thereby hinders the advancement of mechanistic understanding of global warming effects on tropical forest carbon dynamics.

A medida que aumenta la temperatura, disminuye la absorción neta de carbono en los bosques tropicales, sin embargo, aún no se conocen bien los mecanismos que la subyacen. Las altas temperaturas pueden limitar la fotosíntesis directamente, por ejemplo, reduciendo la eficiencia de los procesos bioquímicos, pero también de forma indirecta a través del aumento del déficit de presión de vapor (DPV) que resulta en el cierre estomático. Para explorar los efectos independientes de la temperatura y el DPV en la fotosíntesis, analizamos datos de la absorción neta de carbono del dosel de dos bosques tropicales en Panamá utilizando modelos aditivos generalizados. La conductancia estomática y la fotosíntesis disminuyó consistentemente con el aumento de DPV, y considerando el DPV en modelas estadísticas, la temperatura óptima de la fotosíntesis (Topt) aumentó, de un Topt aparente influida por la DVP de c. 30­31°C a un Topt independiente del DPV de c. 33­36°C. Los árboles y las lianas mostraron respuestas similares a la temperatura y a la DVP en ambos bosques, a pesar de la diferencia de 1500 mm en la precipitación media anual. La fotosíntesis en los bosques tropicales está limitada en gran medida por los efectos indirectos del aumento de la temperatura, a través de cambios en el DPV y no por los efectos directos en los procesos bioquímicos. Si no se tiene en cuenta el DPV al determinar el Topt, se atribuye erróneamente el mecanismo causal subyacente y, por lo tanto, se obstaculiza el avance en la comprensión de los efectos del calentamiento global en la dinámica del carbono.

The intricacies of vegetation responses to changing moisture conditions.

A long-standing debate looks at whether air or soil dryness is more limiting to vegetation water use and productivity. The answer has large implications for future ecosystem functioning, as atmospheric dryness is predicted to increase globally while changes in soil moisture are predicted to be far more variable. Here, I review the complexities that contribute to this debate, including the strong coupling between atmospheric and soil dryness, and the widespread heterogeneity in vegetation hydraulic traits, acclimations, and adaptations to water stress. I discuss solutions to improve understanding and modeling of vegetation sensitivity to dryness, including how different types of observational data can be used together to gain insight into vegetation response to water stress across spatial and temporal scales.

Forest plant indicator values for moisture reflect atmospheric vapour pressure deficit rather than soil water content.

Soil moisture shapes ecological patterns and processes, but it is difficult to continuously measure soil moisture variability across the landscape. To overcome these limitations, soil moisture is often bioindicated using community-weighted means of the Ellenberg indicator values of vascular plant species. However, the ecology and distribution of plant species reflect soil water supply as well as atmospheric water demand. Therefore, we hypothesized that Ellenberg moisture values can also reflect atmospheric water demand expressed as a vapour pressure deficit (VPD). To test this hypothesis, we disentangled the relationships among soil water content, atmospheric vapour pressure deficit, and Ellenberg moisture values in the understory plant communities of temperate broadleaved forests in central Europe. Ellenberg moisture values reflected atmospheric VPD rather than soil water content consistently across local, landscape, and regional spatial scales, regardless of vegetation plot size, depth as well as method of soil moisture measurement. Using in situ microclimate measurements, we discovered that forest plant indicator values for moisture reflect an atmospheric VPD rather than soil water content. Many ecological patterns and processes correlated with Ellenberg moisture values and previously attributed to soil water supply are thus more likely driven by atmospheric water demand.

Different model assumptions about plant hydraulics and photosynthetic temperature acclimation yield diverging implications for tropical forest gross primary production under warming.

Tropical forest photosynthesis can decline at high temperatures due to (1) biochemical responses to increasing temperature and (2) stomatal responses to increasing vapor pressure deficit (VPD), which is associated with increasing temperature. It is challenging to disentangle the influence of these two mechanisms on photosynthesis in observations, because temperature and VPD are tightly correlated in tropical forests. Nonetheless, quantifying the relative strength of these two mechanisms is essential for understanding how tropical gross primary production (GPP) will respond to climate change, because increasing atmospheric CO2 concentration may partially offset VPD-driven stomatal responses, but is not expected to mitigate the effects of temperature-driven biochemical responses. We used two terrestrial biosphere models to quantify how physiological process assumptions (photosynthetic temperature acclimation and plant hydraulic stress) and functional traits (e.g., maximum xylem conductivity) influence the relative strength of modeled temperature versus VPD effects on light-saturated GPP at an Amazonian forest site, a seasonally dry tropical forest site, and an experimental tropical forest mesocosm. By simulating idealized climate change scenarios, we quantified the divergence in GPP predictions under model configurations with stronger VPD effects compared with stronger direct temperature effects. Assumptions consistent with stronger direct temperature effects resulted in larger GPP declines under warming, while assumptions consistent with stronger VPD effects resulted in more resilient GPP under warming. Our findings underscore the importance of quantifying the role of direct temperature and indirect VPD effects for projecting the resilience of tropical forests in the future, and demonstrate that the relative strength of temperature versus VPD effects in models is highly sensitive to plant functional parameters and structural assumptions about photosynthetic temperature acclimation and plant hydraulics.

Large-scale remote sensing analysis reveals an increasing coupling of grassland vitality to atmospheric water demand.

Grasslands provide important ecosystem services to society, including biodiversity, water security, erosion control, and forage production. Grasslands are also vulnerable to droughts, rendering their future vitality under climate change uncertain. Yet, the grassland response to drought is not well understood, especially for heterogeneous Central European grasslands. We here fill this gap by quantifying the spatiotemporal sensitivity of grasslands to drought using a novel remote sensing dataset from Landsat/Sentinel-2 paired with climate re-analysis data. Specifically, we quantified annual grassland vitality at fine spatial scale and national extent (Germany) from 1985 to 2021. We analyzed grassland sensitivity to drought by testing for statistically robust links between grassland vitality and common drought indices. We furthermore explored the spatiotemporal variability of drought sensitivity for 12 grassland habitat types given their different biotic and abiotic features. Grassland vitality maps revealed a large-scale reduction of grassland vitality during past droughts. The unprecedented drought of 2018-2019 stood out as the largest multi-year vitality decline since the mid-1980s. Grassland vitality was consistently coupled to drought (R2 = .09-.22) with Vapor Pressure Deficit explaining vitality best. This suggests that high atmospheric water demand, as observed during recent compounding drought and heatwave events, has major impacts on grassland vitality in Central Europe. We found a significant increase in drought sensitivity over time with highest sensitivities detected in periods of extremely high atmospheric water demand, suggesting that drought impacts on grasslands are becoming more severe with ongoing climate change. The spatial variability of grassland drought sensitivity was linked to different habitat types, with declining sensitivity from dry and mesic to wet habitats. Our study provides the first large-scale, long-term, and spatially explicit evidence of increasing drought sensitivities of Central European grasslands. With rising compound droughts and heatwaves under climate change, large-scale grassland vitality loss, as in 2018-2019, will thus become more likely in the future.

Atmospheric drying and soil drying: Differential effects on grass community composition.

Global surface temperatures are projected to increase in the future; this will modify regional precipitation regimes and increase global atmospheric drying. Despite many drought studies examining the consequences of reduced precipitation, there are few experimental studies exploring plant responses to atmospheric drying via relative humidity and vapor pressure deficit (VPD). We examined eight native California perennial grass species grown in pots in a greenhouse in Los Angeles, California for 34 weeks. All pots were well-watered for 21 weeks, at which point we reduced watering to zero and recorded daily growth and dormancy for 3 weeks. We used this information to better understand the drought tolerance of our species in a larger soil drying × atmospheric drying experiment. In this larger experiment, we grew all eight species together in outdoor mesocosms and measured changes in community composition after 4 years of growth. Soil drying in our small pot experiment mirrored compositional shifts in the larger experiment. Namely, our most drought-tolerant species in our pot experiment was Poa secunda, due to a summer dormancy strategy. Similarly, the grass community shifted toward P. secunda in the driest soils as P. secunda was mostly unaffected by either soil drying or atmospheric drying. We found that some species responded strongly to soil drying (Elymus glaucus, Festuca idahoensis, and Hordeum b. californicum), while others responded strongly to atmospheric drying (Bromus carinatus and Stipa cernua). As result, community composition shifted in different and interacting ways in response to soil drying, atmospheric drying, and their combination. Further study of community responses to increasing atmospheric aridity is an essential next step to predicting the future consequences of climate change.

Eddy covariance measurements reveal a decreased carbon sequestration strength 2010-2022 in an African semiarid savanna.

Monitoring the changes of ecosystem functioning is pivotal for understanding the global carbon cycle. Despite its size and contribution to the global carbon cycle, Africa is largely understudied in regard to ongoing changes of its ecosystem functioning and their responses to climate change. One of the reasons is the lack of long-term in situ data. Here, we use eddy covariance to quantify the net ecosystem exchange (NEE) and its components-gross primary production (GPP) and ecosystem respiration (Reco) for years 2010-2022 for a Sahelian semiarid savanna to study trends in the fluxes. Significant negative trends were found for NEE (12.7 ± 2.8 g C m2 year-1), GPP (39.6 ± 7.9 g C m2 year-1), and Reco (32.2 ± 8.9 g C m2 year-1). We found that NEE decreased by 60% over the study period, and this decrease was mainly caused by stronger negative trends in rainy season GPP than in Reco. Additionally, we observed strong increasing trends in vapor pressure deficit, but no trends in rainfall or soil water content. Thus, a proposed explanation for the decrease in carbon sink strength is increasing atmospheric dryness. The warming climate in the Sahel, coupled with increasing evaporative demand, may thus lead to decreased GPP levels across this biome, and lowering its CO2 sequestration.

Occurrence and Fate of Per- and Polyfluoroalkyl Substances (PFAS) in Atmosphere: Size-Dependent Gas-Particle Partitioning, Precipitation Scavenging, and Amplification.

The concerns about the fate of per- and polyfluoroalkyl substances (PFAS) in the atmosphere are continuously growing. In this study, size-fractionated particles, gas, and rainwater samples were simultaneously collected in Shijiazhuang, China, to investigate the multiphase distribution of PFAS in the atmosphere. Perfluoroalkyl carboxylic acids (PFCAs) dominated the total concentration of PFAS in atmospheric media. A strong positive relationship (0.79 < R2 < 0.99) was observed between the concentration of PFCAs and organic matter fraction (fOM) in different particle size fractions, while no such relationship for perfluoroalkyl sulfonic acids (PFSAs) and fOM, suggesting fOM may be an important factor influencing the size-dependent distribution of PFCAs. Temperature played a key role in the gas-particle partitioning of PFAS, while it did not significantly affect their particle-size-dependent distribution. The associative concentration fluctuation of particle and particle-bound PFAS during precipitation suggested that precipitation scavenging was an important mechanism for the removal of PFAS from the atmosphere. Furthermore, temporary increases in atmospheric PFAS concentrations were observed during the precipitation. Fugacity ratios of PFAS in rainwater and gas phase (log fR/fG ranged between 2.0 and 6.6) indicated a strong trend for PFAS to diffuse from the rainwater to the gas phase during the precipitation, which may explain that the concentration of PFAS in the gas phase continued to increase even at the end of the precipitation.

Raf-like kinases and receptor-like (pseudo)kinase GHR1 are required for stomatal vapor pressure difference response.

Stomatal pores close rapidly in response to low-air-humidity-induced leaf-to-air vapor pressure difference (VPD) increases, thereby reducing excessive water loss. The hydroactive signal-transduction mechanisms mediating high VPD-induced stomatal closure remain largely unknown. The kinetics of stomatal high-VPD responses were investigated by using time-resolved gas-exchange analyses of higher-order mutants in guard-cell signal-transduction branches. We show that the slow-type anion channel SLAC1 plays a relatively more substantial role than the rapid-type anion channel ALMT12/QUAC1 in stomatal VPD signaling. VPD-induced stomatal closure is not affected in mpk12/mpk4GC double mutants that completely disrupt stomatal CO2 signaling, indicating that VPD signaling is independent of the early CO2 signal-transduction pathway. Calcium imaging shows that osmotic stress causes cytoplasmic Ca2+ transients in guard cells. Nevertheless, osca1-2/1.3/2.2/2.3/3.1 Ca2+-permeable channel quintuple, osca1.3/1.7-channel double, cngc5/6-channel double, cngc20-channel single, cngc19/20crispr-channel double, glr3.2/3.3-channel double, cpk-kinase quintuple, cbl1/4/5/8/9 quintuple, and cbl2/3rf double mutants showed wild-type-like stomatal VPD responses. A B3-family Raf-like mitogen-activated protein (MAP)-kinase kinase kinase, M3Kδ5/RAF6, activates the OST1/SnRK2.6 kinase in plant cells. Interestingly, B3 Raf-kinase m3kδ5 and m3kδ1/δ5/δ6/δ7 (raf3/6/5/4) quadruple mutants, but not a 14-gene raf-kinase mutant including osmotic stress-linked B4-family Raf-kinases, exhibited slowed high-VPD responses, suggesting that B3-family Raf-kinases play an important role in stomatal VPD signaling. Moreover, high VPD-induced stomatal closure was impaired in receptor-like pseudokinase GUARD CELL HYDROGEN PEROXIDE-RESISTANT1 (GHR1) mutant alleles. Notably, the classical transient "wrong-way" VPD response was absent in ghr1 mutant alleles. These findings reveal genes and signaling mechanisms in the elusive high VPD-induced stomatal closing response pathway.

Estimation of the Vaporization Enthalpies and Vapor Pressures of α-Tocopherol and Δ9-Tetrahydrocannabinol via the Use of a Surrogate, Correlation Gas Chromatography, and Synthetic and Retrosynthetic Analysis.

A protocol is proposed that combines the use of the known properties of a surrogate containing various functional groups together with n-alkanes as standards to evaluate the properties of much larger related substances using correlation gas chromatography. An objective of this work is to develop options that circumvent the lack of appropriate vaporization enthalpy standards that can be used for evaluation of various thermodynamic properties of larger complex molecules using gas chromatography. The surrogate in this case is 2,2,5,7,8-pentamethylchroman-6-ol (PMC) and is used to evaluate the vaporization enthalpies and vapor pressures of α-tocopherol (α-TOC) and Δ9-tetrahydrocannabinol (Δ9-THC). The results are compared to the available literature data and to estimated properties. Vaporization enthalpies are also evaluated by a proposed method that involves the use of synthetic and retrosynthetic analysis.

Thermodynamic Study of N-Methylformamide and N,N-Dimethyl-Formamide.

An extensive thermodynamic study of N-methylformamide (CAS RN: 123-39-7) and N,N-dimethylformamide (CAS RN: 68-12-2), is presented in this work. The liquid heat capacities of N-methylformamide were measured by Tian-Calvet calorimetry in the temperature interval (250-300) K. The vapor pressures for N-methylformamide and N,N-dimethylformamide were measured using static method in the temperature range 238 K to 308 K. The ideal-gas thermodynamic properties were calculated using a combination of the density functional theory (DFT) and statistical thermodynamics. A consistent thermodynamic description was developed using the method of simultaneous correlation, where the experimental and selected literature data for vapor pressures, vaporization enthalpies, and liquid phase heat capacities and the calculated ideal-gas heat capacities were treated together to ensure overall thermodynamic consistency of the results. The resulting vapor pressure equation is valid from the triple point to the normal boiling point temperature.

Phase Transition Thermodynamics of 1,3,5-Tris-(α-naphthyl)benzene: Theory and Experiment.

1,3,5-Tris-(α-naphthyl)benzene is an organic non-electrolyte with notable stability of an amorphous phase. Its glassy and supercooled liquid states were previously studied by spectroscopic and calorimetric methods. Despite the continuing interest in its amorphous state and, particularly, vapor-deposited glasses, the thermodynamic parameters of the vaporization of 1,3,5-tris-(α-naphthyl)benzene have not been obtained yet. Likewise, the reliable evaluation of the thermodynamic parameters of fusion below the melting point, required to establish the thermodynamic state of its glass, is still an unsolved problem. In this work, the heat capacities of crystalline and liquid phases, the temperature dependence of the saturated vapor pressures, fusion and vaporization enthalpies were determined using differential and fast scanning calorimetry and were verified using the estimates based on solution calorimetry. The structural features of 1,3,5-tris-(α-naphthyl)benzene are discussed based on the computations performed and the data on the molecular refractivity. The consistency between the values obtained by independent techniques was demonstrated.

Xylem conduit deformation across vascular plants: an evolutionary spandrel or protective valve?

The hydraulic system of vascular plants and its integrity is essential for plant survival. To transport water under tension, the walls of xylem conduits must approximate rigid pipes. Against this expectation, conduit deformation has been reported in the leaves of a few species and hypothesized to function as a 'circuit breaker' against embolism. Experimental evidence is lacking, and its generality is unknown. We demonstrated the role of conduit deformation in protecting the upstream xylem from embolism through experiments on three species and surveyed a diverse selection of vascular plants for conduit deformation in leaves. Conduit deformation in minor veins occurred before embolism during slow dehydration. When leaves were exposed to transient increases in transpiration, conduit deformation was accompanied by large water potential differences from leaf to stem and minimal embolism in the upstream xylem. In the three species tested, collapsible vein endings provided clear protection of upstream xylem from embolism during transient increases in transpiration. We found conduit deformation in diverse vascular plants, including 11 eudicots, ginkgo, a cycad, a fern, a bamboo, and a grass species, but not in two bamboo and a palm species, demonstrating that the potential for 'circuit breaker' functionality may be widespread across vascular plants.

Diagnosing evapotranspiration responses to water deficit across biomes using deep learning.

Accounting for water limitation is key to determining vegetation sensitivity to drought. Quantifying water limitation effects on evapotranspiration (ET) is challenged by the heterogeneity of vegetation types, climate zones and vertically along the rooting zone. Here, we train deep neural networks using flux measurements to study ET responses to progressing drought conditions. We determine a water stress factor (fET) that isolates ET reductions from effects of atmospheric aridity and other covarying drivers. We regress fET against the cumulative water deficit, which reveals the control of whole-column moisture availability. We find a variety of ET responses to water stress. Responses range from rapid declines of fET to 10% of its water-unlimited rate at several savannah and grassland sites, to mild fET reductions in most forests, despite substantial water deficits. Most sensitive responses are found at the most arid and warm sites. A combination of regulation of stomatal and hydraulic conductance and access to belowground water reservoirs, whether in groundwater or deep soil moisture, could explain the different behaviors observed across sites. This variety of responses is not captured by a standard land surface model, likely reflecting simplifications in its representation of belowground water storage.

Limited-transpiration trait in response to high vapor pressure deficit from wild to cultivated species: study of the Lens genus.

Lentil (Lens culinaris Medik.) is commonly grown in drought-prone areas where terminal heat and drought are frequent. The limited-transpiration (TRlim) trait under high vapor pressure deficit (VPD) could be a way to conserve water and increase yield under water deficit conditions. The TRlim trait was examined in cultivated and wild lentil species together with its evolution throughout the breeding pipeline. Sixty-one accessions representing the six wild lentil species (L. orientalis, L. tomentosus, L. odemensis, L. lamottei, L. ervoides, and L. nigricans) and 13 interspecific advanced lines were evaluated in their transpiration response to high VPD. A large variation in transpiration rate (TR) response to increased VPD was recorded among wild lentil accessions, with 43 accessions exhibiting a breakpoint (BP) in their TR response to increasing VPD, with values ranging from 0.92 kPa to 3.38 kPa under greenhouse conditions. Ten genotypes for the interspecific advanced lines displayed a BP with an average of 1.95 kPa, much lower than previously reported for cultivated lentil. Results from field experiments suggest that the TRlim trait (BP=0.97 kPa) positively affected yield and yield-related parameters during the years with late-season water stress. The selection of TRlim genotypes for high VPD environments could improve lentil productivity in drought-prone areas.

Transpiration response to soil drying versus increasing vapor pressure deficit in crops: physical and physiological mechanisms and key plant traits.

The water deficit experienced by crops is a function of atmospheric water demand (vapor pressure deficit) and soil water supply over the whole crop cycle. We summarize typical transpiration response patterns to soil and atmospheric drying and the sensitivity to plant hydraulic traits. We explain the transpiration response patterns using a soil-plant hydraulic framework. In both cases of drying, stomatal closure is triggered by limitations in soil-plant hydraulic conductance. However, traits impacting the transpiration response differ between the two drying processes and act at different time scales. A low plant hydraulic conductance triggers an earlier restriction in transpiration during increasing vapor pressure deficit. During soil drying, the impact of the plant hydraulic conductance is less obvious. It is rather a decrease in the belowground hydraulic conductance (related to soil hydraulic properties and root length density) that is involved in transpiration down-regulation. The transpiration response to increasing vapor pressure deficit has a daily time scale. In the case of soil drying, it acts on a seasonal scale. Varieties that are conservative in water use on a daily scale may not be conservative over longer time scales (e.g. during soil drying). This potential independence of strategies needs to be considered in environment-specific breeding for yield-based drought tolerance.

Decoupling between stomatal conductance and photosynthesis occurs under extreme heat in broadleaf tree species regardless of water access.

High air temperatures increase atmospheric vapor pressure deficit (VPD) and the severity of drought, threatening forests worldwide. Plants regulate stomata to maximize carbon gain and minimize water loss, resulting in a close coupling between net photosynthesis (Anet ) and stomatal conductance (gs ). However, evidence for decoupling of gs from Anet under extreme heat has been found. Such a response both enhances survival of leaves during heat events but also quickly depletes available water. To understand the prevalence and significance of this decoupling, we measured leaf gas exchange in 26 tree and shrub species growing in the glasshouse or at an urban site in Sydney, Australia on hot days (maximum Tair > 40°C). We hypothesized that on hot days plants with ample water access would exhibit reduced Anet and use transpirational cooling leading to stomatal decoupling, whereas plants with limited water access would rely on other mechanisms to avoid lethal temperatures. Instead, evidence for stomatal decoupling was found regardless of plant water access. Transpiration of well-watered plants was 23% higher than model predictions during heatwaves, which effectively cooled leaves below air temperature. For hotter, droughted plants, the increase in transpiration during heatwaves was even more pronounced-gs was 77% higher than model predictions. Stomatal decoupling was found for most broadleaf evergreen and broadleaf deciduous species at the urban site, including some wilted trees with limited water access. Decoupling may simply be a passive consequence of the physical effects of high temperature on plant leaves through increased cuticular conductance of water vapor, or stomatal decoupling may be an adaptive response that is actively regulated by stomatal opening under high temperatures. This temperature response is not yet included in any land surface model, suggesting that model predictions of evapotranspiration may be underpredicted at high temperature and high VPD.