Stomatal Behaviour Research Articles

Starch biosynthesis in guard cells has features of both autotrophic and heterotrophic tissues.

The pathway of starch synthesis in guard cells (GCs), despite the crucial role starch plays in stomatal movements, is not well understood. Here, we characterized starch dynamics in GCs of Arabidopsis (Arabidopsis thaliana) mutants lacking enzymes of the phosphoglucose isomerase-phosphoglucose mutase-ADP-glucose pyrophosphorylase starch synthesis pathway in leaf mesophyll chloroplasts or sugar transporters at the plastid membrane, such as glucose-6-phosphate/phosphate translocators, which are active in heterotrophic tissues. We demonstrate that GCs have metabolic features of both photoautotrophic and heterotrophic cells. GCs make starch using different carbon precursors depending on the time of day, which can originate both from GC photosynthesis and/or sugars imported from the leaf mesophyll. Furthermore, we unravel the major enzymes involved in GC starch synthesis and demonstrate that they act in a temporal manner according to the fluctuations of stomatal aperture, which is unique for GCs. Our work substantially enhances our knowledge on GC starch metabolism and uncovers targets for manipulating GC starch dynamics to improve stomatal behavior, directly affecting plant productivity.

Unravelling the responses of different apple varieties to water constraints by continuous field thermal monitoring

This research aimed at analyzing the response of apple tree varieties subjected to soil water deficit and atmospheric drought in a field phenotyping platform located in the Mediterranean area. The main assumption of the study was that seasonal and daily stomatal behavior can be monitored by continuous measurement of canopy surface temperature (Ts) as a proxy of stomatal closure. To achieve the study objectives, thermal monitoring of 6 pre-commercial apple varieties was simultaneously carried out throughout one season by nadir-oriented thermo-radiometers placed 1.50 m over the tree top canopy. Two water regimes were applied to each variety during a 4-week summer period: normal irrigation (WW) vs progressive water deficit (WS). The maximum difference in Ts between water regimes was recorded daily between 11:00 and 14:20 GMT, with an earlier closure of stomata in WS trees. During the day, a more negative stem water potential (Ψstem) and a higher diurnal Ts (+1° to +2 °C) were observed on WS trees, resulting in a significant limitation of fruit growth. Tree water stress was caused by both edaphic and atmospheric droughts, in the medium and short terms respectively, with inter-varietal and inter-regime differences highlighting distinct stomatal closure behaviors. Results suggest that some of the varieties studied are well adapted to stressful summer conditions, as long as irrigation needs are met, while other varieties show a particular sensitivity to the mid-day evaporative demand, which may limit their extension. Although these results are not comprehensive enough to predict the optimal performance of varieties under different stress scenarios, the proposed methodology allows to assess the dynamics of tree response to water constraints using non-invasive thermal sensors. It opens up new perspectives for the phenotyping of apple cultivars under abiotic stress, achievable through the quantified study of their transpiration flux in response to stress scenarios. These prospects will require further in planta measurements to dissect varietal differences.

NaCl affects photosynthetic and stomatal dynamics by osmotic effects and reduces photosynthetic capacity by ionic effects in tomato.

NaCl stress affects stomatal behavior and photosynthesis by a combination of osmotic and ionic components, but it is unknown how these components affect stomatal and photosynthetic dynamics. Tomato (Solanum lycopersicum) plants were grown in a reference nutrient solution [control; electrical conductivity (EC)=2.3 dS m-1], a solution containing additional macronutrients (osmotic effect; EC=12.6 dS m-1), or a solution with additional 100mM NaCl (osmotic and ionic effects; EC=12.8 dS m-1). Steady-state and dynamic photosynthesis, and leaf biochemistry, were characterized throughout leaf development. The osmotic effect decreased steady-state stomatal conductance while speeding up stomatal responses to light intensity shifts. After 19 d of treatment, photosynthetic induction was reduced by the osmotic effect, which was attributable to lower initial stomatal conductance due to faster stomatal closing under low light. Ionic effects of NaCl were barely observed in dynamic stomatal and photosynthetic behavior, but led to a reduction in leaf photosynthetic capacity, CO2 carboxylation rate, and stomatal conductance in old leaves after 26 d of treatment. With increasing leaf age, rates of light-triggered stomatal movement and photosynthetic induction decreased across treatments. We conclude that NaCl impacts dynamic stomatal and photosynthetic kinetics by osmotic effects and reduces photosynthetic capacity by ionic effects.

Leaf and canopy stomatal resistance, aerodynamic resistance, and evapotranspiration of irrigated continuous no‐till and disk‐till maize

AbstractThe impact of no‐till (NT) management as compared with conventional tillage practices (e.g., disk till [DT]) on plant stomatal behavior in relation to model estimation of crop evapotranspiration (ETc) and plant physiological functions has not been sufficiently evaluated. This research investigated the effects of NT and DT management on maize (Zea mays L.) physiological parameters and ETc losses. We also tested the methodology to estimate ETc using a one‐step Penman–Monteith (PM) model with measured variable resistance from scaled‐up leaf stomatal resistance (RL) to canopy resistance (RC) with Bowen Ratio Energy Balance System (BREBS)‐measured ETc. Photosynthetic photon flux density (PPFD) explained 76 and 79% of the variability in RL for NT and DT maize, respectively, which varied with growth stage. Disk‐till maize RL was 29, 5, 25, and 4% higher than NT maize RL during the vegetative, tasseling and silking, reproductive, and near‐physiological‐maturity stages, respectively. The seasonal average scaled‐up RC values for NT and DT maize were 102.4 and 128.6 s m–1, respectively. The PM‐estimated ETc correlated strongly with the BREBS‐measured ETc, with the PM model estimates being 8 and 6% within the BREBS measurements on an hourly and daily basis, respectively. To the best of our knowledge, this research is the first to develop the response curves (PPFD vs. RL) for maize for different growth stages under NT and DT. The analyses presented here can provide invaluable data and information for plant physiology research, crop modeling, determining crop response to different management, and environmental drivers.

Stomatal behaviour and endogenous phytohormones promotes intrinsic water use efficiency differently in cotton under drought

The coordination between phytohormones regulation, stomatal behaviour (stomatal index and opening/closing) and gas exchange are potent determinants of plant survival under drought stress. However, we found a knowledge gap in the mechanism regulating the fine-tuning of these features during drought. In order to address this we evaluated gas exchange, stomatal behaviour and endogenous phytohormones content in two cotton varieties (LRA-5166 and NBRI-67) differing in drought sensitivity during water deficit conditions. Variety specific differences were recorded in net photosynthesis rate (A), transpiration rate (E) and stomatal conductance (gs) with significantly less decrease in drought tolerant LRA-5166 than drought sensitive NBRI-67. The abscisic acid (ABA) accumulation was significantly increased in LRA-5166 while reduced in NBRI-67 under water deficit, which was accompanied by relatively less reduced 6-benzylaminopurine (6-BAP) level in LRA-5166 than NBRI-67.Thus, improved ABA/6-BAP ratio was observed in both the varieties of cotton. Critically, LRA- 5166 has reduced stomatal index, aperture size and significantly higher A and intrinsic water use efficiency (WUEi), thus higher drought tolerance than NBRI-67. Furthermore, we found that endogenous ABA predominantly maintains the stomatal behaviour and regulates its physiology either by antagonizing 6-BAP or alone to coordinate with water deficit signals. Overall, our findings describe a new insight as to how drought modulates endogenous ABA and 6-BAP homeostasis in cotton leaf and the mechanism of stomatal regulation by ABA and 6-BAP in cotton.

Effects of ozone–vegetation interactions on meteorology and air quality in China using a two-way coupled land–atmosphere model

Abstract. Tropospheric ozone (O3) is one of the most important air pollutants in China and is projected to continue to increase in the near future. O3 and vegetation closely interact with each other and such interactions may not only affect plant physiology (e.g., stomatal conductance and photosynthesis) but also influence the overlying meteorology and air quality through modifying leaf stomatal behaviors. Previous studies have highlighted China as a hotspot in terms of O3 pollution and O3 damage to vegetation. Yet, few studies have investigated the effects of O3–vegetation interactions on meteorology and air quality in China, especially in the light of recent severe O3 pollution. In this study, a two-way coupled land–atmosphere model was applied to simulate O3 damage to vegetation and the subsequent effects on meteorology and air quality in China. Our results reveal that O3 causes up to 16 % enhancement in stomatal resistance, whereby large increases are found in the Henan, Hebei, and Shandong provinces. O3 damage causes more than 0.6 µmol CO2 m−2 s−1 reductions in photosynthesis rate and at least 0.4 and 0.8 g C m−2 d−1 decreases in leaf area index (LAI) and gross primary production (GPP), respectively, and hotspot areas appear in the northeastern and southern China. The associated reduction in transpiration causes a 5–30 W m−2 decrease (increase) in latent heat (sensible heat) flux, which induces a 3 % reduction in surface relative humidity, 0.2–0.8 K increase in surface air temperature, and 40–120 m increase in boundary-layer height in China. We also found that the meteorological changes further induce a 2–6 ppb increase in O3 concentration in northern and south-central China mainly due to enhanced isoprene emission following increased air temperature, demonstrating that O3–vegetation interactions can lead to strong positive feedback that can amplify O3 pollution in China. Our findings emphasize the importance of considering the effects of O3 damage and O3–vegetation interactions in air quality simulations, with ramifications for both air quality and forest management.

Scaling Up from Leaf to Whole-Plant Level for Water Use Efficiency Estimates Based on Stomatal and Mesophyll Behaviour in Platycladus orientalis

Prediction of whole-plant short-term water use efficiency (WUEs,P) is essential to indicate plant performance and facilitate comparison across different temporal and spatial scales. In this study, an isotope model was scaled up from the leaf to the whole-plant level, in order to simulate the variation in WUEs,P in response to different CO2 concentrations (Ca; 400, 600, and 800 μmol·mol−1) and soil water content (SWC; 35–100% of field capacity). For WUEs,P modelling, leaf gas exchange information, plant respiration, and “unproductive” water loss were taken into account. Specifically, in shaping the expression of the WUEs,P, we emphasized the role of both stomatal (gsw) and mesophyll conductance (gm). Simulations were compared with the measured results to check the model’s applicability. The verification showed that estimates of gsw from the coupled photosynthesis (Pn,L)-gsw model accounting for the effect of soil water stress slightly outperformed the model neglecting the soil water status effect. The established coupled Pn,L-gm model also proved more effective in estimating gm than the previously proposed model. Introducing the two diffusion control functions into the whole-plant model, the developed model for WUEs,P effectively captured its response pattern to different Ca and SWC conditions. Overall, this study confirmed that the accurate estimation of WUEs,P requires an improved predictive accuracy of gsw and gm. These results have important implications for predicting how plants respond to climate change.

Characteristic of Stomatal Conductance and Optimal Stomatal Behaviour in an Arid Oasis of Northwestern China

Stomatal conductance (gs), the process that governs plant carbon uptake and water loss, is fundamental to most Land Surface Models (LSMs). With global change accelerating, more attention should be paid to investigating stomatal behavior, especially in extremely arid areas. In this study, gas exchange measurements and environmental/biological variables observations during growing seasons in 2016 and 2017 were combined to investigate diurnal and seasonal characteristics of gs and the applicability of the optimal stomatal conductance model in a desert oasis vineyard. The results showed that the responses of gs to environmental factors (photosynthesis active radiation, PAR; vapor pressure deficit, VPD; and temperature, T) formed hysteresis loops in the daytime. The stomatal conductance slope, g1, a parameter in the unified stomatal optimal model, varied in different growing seasons and correlated with the soil-to-leaf hydraulic conductance (KL). These results indicated the potential bias when using a constant g1 value to simulate gs and highlighted that the water-use strategy of oasis plants might not be consistent throughout the entire growing season. Our findings further help to achieve a better understanding of stomata behavior in responding to climate change and encourage future efforts toward a more accurate parameterization of gs to improve the modeling of LSMs.

Syrah and Grenache (Vitis vinifera) revealed different strategies to cope with high temperature

Background and Aims Global warming will threaten vineyard sustainability in warm and arid regions. Therefore, increasing water use efficiency (WUE) is essential for long-term viticulture sustainability. Nevertheless, studies concerning the response of stomata and WUE to temperature are scarce. We evaluated the effect of sustained warming on WUE and its components in cultivars with contrasting stomatal behaviour. Methods and Results We applied a 1.1°C increase in field-grown, well-watered plants, during the growing season and compared them to Control plants. We measured net assimilation (An), transpiration rate (E), stomatal conductance (gs), stomata density and length, intrinsic and yield WUE (WUEint and WUEyield). Gas exchange and WUEint in response to heating were cultivar-dependent. Heated Grenache plants increased An, gs, E and stomatal length but decreased WUEint. In contrast, gas exchange and WUEint were similar between heated Syrah and Control plants. Both cultivars decreased WUEyield in response to heating. Conclusions An increase in air temperature decreased WUEint in Grenache, whereas Syrah could tolerate heating without modifying leaf functioning. We proposed these strategies to cope with heating as ‘avoiding’ for Grenache and ‘tolerant’ for Syrah in well-watered conditions. Significance of the Study This research contributes to an understanding of the strategies to cope with high temperature of different cultivars, which should be considered to select better-adapted genotypes for the expected warmer conditions.

Estimating evapotranspiration using remotely sensed solar-induced fluorescence measurements

Terrestrial evapotranspiration (ET) is a critical component of the land surface and its accurate estimation is crucial in understanding the global water and energy budgets. Current large-scale ET estimation models depend on remotely sensed data products of meteorological conditions and vegetation phenology, but high uncertainty remains to correctly represent ecological factors, which are regulated by biochemistry. In recent years, solar-induced fluorescence (SIF), a proxy for photosynthesis, has been extensively used in different ecological research and is conceived to be of great potential for ET estimation to constrain the transpiration flux, which is the significant component of ET. So far, most SIF-based ET estimation methods are achieved via empirical methods, which lack a solid physical foundation. In this work, we estimated ET by combining SIF and meteorological variables with Fick's law and an optimal stomatal behavior model and validated the model across different eddy-covariance flux stations around the globe. The model showed overall good performance across different ecosystems and when compared with other remote sensing-based ET models, such as the Priestley Taylor-JPL (PT-JPL) model, the Moderate Resolution Imaging Spectroradiometer (MODIS)-ET products, and a simple empirical linear SIF-ET model, the RMSE of estimated ET of the model proposed in this paper demonstrated better performance. The model we proposed here can be potentially extended for a reliable ET estimation at a global scale.

Changes in Growth, Photosynthetic Pigments, Cell Viability, Lipid Peroxidation and Antioxidant Defense System in Two Varieties of Chickpea (Cicer arietinum L.) Subjected to Salinity Stress

Salinity is one of the most severe abiotic stresses for crop production. The present study investigates the salinity-induced modulation in growth indicators, morphology and movement of stomata, photosynthetic pigments, activity of carbonic anhydrase as well as nitrate reductase, and antioxidant systems in two varieties of chickpea (Pusa-BG5023, and Pusa-BGD72). On 20th day of sowing, plants were treated with varying levels of NaCl (0, 50, 100, 150 and 200 mM) followed by sampling on 45 days of sowing. Recorded observations on both the varieties reveal that salt stress leads to a significant decline in growth, dry biomass, leaf area, photosynthetic pigments, protein content, stomatal behavior, cell viability, activity of nitrate reductase and carbonic anhydrase with the rise in the concentration of salt. However, quantitatively these changes were less in Pusa-BG5023 as compared to Pusa-BGD72. Furthermore, salinity-induced oxidative stress enhanced malondialdehyde content, superoxide radicals, foliar proline content, and the enzymatic activities of superoxide dismutase, catalase, and peroxidase. The variety Pusa-BGD72 was found more sensitive than Pusa-BG5023 to salt stress. Out of different graded concentrations (50, 100, 150 and 200 mM) of sodium chloride, 50 mM was least toxic, and 200 mM was most damaging. The differential behavior of these two varieties measured in terms of stomatal behavior, cell viability, photosynthetic pigments, and antioxidant defense system can be used as prospective indicators for selection of chickpea plants for salt tolerance and sensitivity.

Stomatal size and density trade-off varies with leaf phenology and species shade tolerance in a South Asian moist tropical forest

The density and guard cell length of stomata regulate the physiological processes in plants. Yet, the variation of stomatal characteristics among different functional groups of trees is not been well understood. Particularly, a comprehensive understanding of stomatal behaviour in Bangladeshi moist forest trees is lacking. The study investigated how abaxial stomatal density (SD) and guard cell length (GCL) vary among tree functional types and leaf phenological groups in a moist tropical forest of Bangladesh. Cluster dendrogram revealed three groups of species based on SD and GCL. The independent sample t -test showed that there was a significant difference in SD between evergreen and deciduous tree species (t =4.18, P <0.001) but no significant difference in GCL between the two phenological groups. ANOVA revealed no significant difference in SD among the light demanding, intermediate shade tolerant and shade tolerant species (F =0.76, P =0.47). However, GCL significantly differed among the three functional groups (F =3.3, P <0.05). Maximum theoretical stomatal conductance (g max ) varied between evergreen and deciduous species but did not vary with species shade tolerance. In general, there was a significant trade-off between SD and GCL. However, the inverse relationship was stronger in deciduous and shade tolerant species than in evergreen and shade intolerant species. Leaf dry matter content was positively related with SD and negatively related with GCL. Specific leaf area and leaf thickness were not related to the stomatal traits. Our analyses suggest that leaf phenology and species shade tolerance need to be considered while estimating gas exchange through the stomata in tropical moist forests.

Stomatal behaviour of four potato cultivars under repeated water stress conditions

Stomatal behaviour of four potato cultivars under repeated water stress conditions

Natural variation in stomatal dynamics drives divergence in heat stress tolerance and contributes to seasonal intrinsic water-use efficiency in Vitis vinifera (subsp. sativa and sylvestris).

Stomata control CO2 uptake for photosynthesis and water loss through transpiration, thus playing a key role in leaf thermoregulation, water-use efficiency (iWUE), and plant productivity. In this work, we investigated the relationship between several leaf traits and hypothesized that stomatal behavior to fast (i.e. minutes) environmental changes co-determines, along with steady-state traits, the physiological response of grapevine to the surrounding fluctuating environment over the growing season. No relationship between iWUE, heat stress tolerance, and stomatal traits was observed in field-grown grapevine, suggesting that other physiological mechanisms are involved in determining leaf evaporative cooling capacity and the seasonal ratio of CO2 uptake (A) to stomatal conductance (gs). Indeed, cultivars that in the field had an unexpected combination of high iWUE but low sensitivity to thermal stress displayed a quick stomatal closure to light, but a sluggish closure to increased vapor pressure deficit (VPD) levels. This strategy, aiming both at conserving water under a high to low light transition and in prioritizing evaporative cooling under a low to high VPD transition, was mainly observed in the cultivars Regina and Syrah. Moreover, cultivars with different known responses to soil moisture deficit or high air VPD (isohydric versus anisohydric) had opposite behavior under fluctuating environments, with the isohydric cultivar showing slow stomatal closure to reduced light intensity but quick temporal responses to VPD manipulation. We propose that stomatal behavior to fast environmental fluctuations can play a critical role in leaf thermoregulation and water conservation under natural field conditions in grapevine.

Stomatal regulation prevents plants from critical water potentials during drought: Result of a model linking soil–plant hydraulics to abscisic acid dynamics

AbstractUnderstanding stomatal regulation during drought is essential to correctly predict vegetation‐atmosphere fluxes. Stomatal optimization models posit that stomata maximize the carbon gain relative to a penalty caused by water loss, such as xylem cavitation. However, a mechanism that allows the stomata to behave optimally is unknown. Here, we introduce a model of stomatal regulation that results in similar stomatal behaviour without presupposing an optimality principle. By contrast, the proposed model explains stomatal closure based on a well‐known component of stomatal regulation: abscisic acid (ABA). The ABA level depends on its production rate, which is assumed to increase with declining leaf water potential, and on its degradation rate, which is assumed to increase with assimilation rate. Our model predicts that stomata open until the ratio of leaf water potential to assimilation rate, proportional to ABA level, is at a minimum. As a prerequisite, the model simulates soil–plant hydraulics and leaf photosynthesis under varying environmental conditions. The model predicts that in wet soils and at low vapour pressure deficit (VPD), when there is no water limitation, stomatal closure is controlled by the relationship between photosynthesis and stomatal conductance. In dry soils or at high VPD, when the soil hydraulic conductivity limits the water supply, stomatal closure is triggered by the sharp decline in leaf water potential as transpiration rate increases. Being adaptive to changing soil and atmospheric conditions, the proposed model can explain how plants are enabled to avoid critical water potentials during drought for varying soil properties and atmospheric conditions.

Melatonin Enhances Drought Tolerance by Regulating Leaf Stomatal Behavior, Carbon and Nitrogen Metabolism, and Related Gene Expression in Maize Plants

It is commonly known that exogenously applied melatonin can alleviate the impact of drought stress, but the mechanism used by melatonin to regulate stomatal behavior and carbon (C) and nitrogen (N) metabolism to increase drought resistance remains elusive. Herein, our aim was to investigate the influence of exogenous melatonin on the regulation of C and N metabolism in maize plants under water deficit. In this study, we analyzed stomatal behavior, the key components of C and N metabolism, and the gene expression and activity of enzymes involved in the C and N metabolism in maize plants. The results showed that the application of melatonin (100 μM) significantly increased maize growth and sustained the opening of stomata, and secondarily increased the photosynthetic capacity in maize. Under drought stress, foliar application of melatonin induced the gene transcription and activities of sucrose phosphate synthetase, ADP-glucose pyrophosphorylase, phosphoenolpyruvate carboxylase, and citrate synthase, resulting in the enhancement of sucrose and starch synthesis and the tricarboxylic acid (TCA) cycle. This enhancement in sugar biosynthesis and the TCA cycle might lead to stronger N assimilation. As anticipated, NO3– reduction and NH4+ assimilation were also strengthened after melatonin treatment under drought stress. An increase was observed in some key enzymatic activities and transcription involved in nitrogen metabolism, such as that of nitrate reductase, nitrite reductase, glutamate synthase, and glutamine synthetase, in melatonin-treated, drought-stressed maize. Moreover, melatonin attenuated the drought-induced damage by reducing protein degradation and increasing the level of proline. Conclusively, our results indicate that exogenous melatonin enhances drought tolerance in maize via promoting stomatal opening and regulating C and N metabolism and related gene expression.

Interpreting the water use strategies of plantation tree species by canopy stomatal conductance and its sensitivity to vapor pressure deficit in South China

Stomatal regulation controls tree transpiration rate and is central to plant function. However, most previous studies that investigated the stomatal regulation of forest trees mainly focused on arid and semi-arid zones, uncertainty remains about how trees adjust their stomatal response to the changing environment in subtropical moist areas. In this study, we investigated the tree transpiration (El), canopy stomatal conductance (Gs), and Gs sensitivity to vapor pressure deficit (VPD) of four typical plantation tree species (Schima wallichii, Acacia mangium, Eucalyptus urophylla, and Cunninghamia lanceolata) in hilly lands of South China. Datasets including the continuously measured sap flow and meteorological parameters, as well as the physiological traits of the four species, were collected during the year of 2018. Our results showed that the coniferous C. lanceolata had significantly lower El, Gs, and weak sensitivity to VPD than the other three broadleaved species. We attributed the different water use and stomatal behavior to the various species-specific traits. The three broadleaved species generally possessed higher leaf area, specific leaf area (SLA), the ratio of sapwood area (As) to leaf area (Al), and deeper root depth, facilitating their transpiration and gas exchange. During the experimental period, a super typhoon “Mangkhut” hit the research site and caused substantial decline in leaf area, while the whole tree transpiration experienced slighter decrease, indicating the rapid compensatory effect of stomatal regulation. Moreover, the increased Gs sensitivity to VPD (m values) and reference Gs for the introduced A. mangium and E. urophylla in the dry season highlighted their ability to function well even under the much drier condition with no strict stomatal control. Our analyses on the response of Gs to the changing environment for typical species in the subtropical moist forests will provide important insights for forest management and ecosystem stability under future climate changes.

Overexpression of MdASMT9, an N-acetylserotonin methyltransferase gene, increases melatonin biosynthesis and improves water-use efficiency in transgenic apple.

Improving apple water-use efficiency (WUE) is increasingly desirable in the face of global climate change. Melatonin is a pleiotropic molecule that functions in plant development and stress tolerance. In apple, exogenous application of melatonin has been largely investigated, but melatonin biosynthesis and its physiological roles remain elusive. In the plant biosynthetic pathway of melatonin, the last and key step is that N-acetylserotonin methyltransferase (ASMT) converts N-acetylserotonin into melatonin. Here, we identified an apple ASMT gene, MdASMT9, using homology-based cloning and in vitro enzyme assays. Overexpression of MdASMT9 significantly increased melatonin accumulation in transgenic apple lines. Moreover, an enhanced WUE was observed in the MdASMT9-overexpressing apple lines. Under well-watered conditions, this increase in WUE was attributed to an enhancement of photosynthetic rate and stomatal aperture via a reduction in abscisic acid biosynthesis. By contrast, under long-term moderate water deficit conditions, regulations in photoprotective mechanisms, stomatal behavior, osmotic adjustment and antioxidant activity enhanced the WUE in transgenic apple lines. Taken together, our findings shed light on the positive effect of MdASMT9 on improving WUE of apple by modulating melatonin biosynthesis.

Extreme heat increases stomatal conductance and drought-induced mortality risk in vulnerable plant species.

Tree mortality during global‐change‐type drought is usually attributed to xylem dysfunction, but as climate change increases the frequency of extreme heat events, it is necessary to better understand the interactive role of heat stress. We hypothesized that some drought‐stressed plants paradoxically open stomata in heatwaves to prevent leaves from critically overheating. We experimentally imposed heat (>40°C) and drought stress onto 20 broadleaf evergreen tree/shrub species in a glasshouse study. Most well‐watered plants avoided lethal overheating, but drought exacerbated thermal damage during heatwaves. Thermal safety margins (TSM) quantifying the difference between leaf surface temperature and leaf critical temperature, where photosynthesis is disrupted, identified species vulnerability to heatwaves. Several mechanisms contributed to high heat tolerance and avoidance of damaging leaf temperatures—small leaf size, low leaf osmotic potential, high leaf mass per area (i.e., thick, dense leaves), high transpirational capacity, and access to water. Water‐stressed plants had smaller TSM, greater crown dieback, and a fundamentally different stomatal heatwave response relative to well‐watered plants. On average, well‐watered plants closed stomata and decreased stomatal conductance (g s) during the heatwave, but droughted plants did not. Plant species with low g s, either due to isohydric stomatal behavior under water deficit or inherently low transpirational capacity, opened stomata and increased g s under high temperatures. The current paradigm maintains that stomata close before hydraulic thresholds are surpassed, but our results suggest that isohydric species may dramatically increase g s (over sixfold increases) even past their leaf turgor loss point. By actively increasing water loss at high temperatures, plants can be driven toward mortality thresholds more rapidly than has been previously recognized. The inclusion of TSM and responses to heat stress could improve our ability to predict the vulnerability of different tree species to future droughts.

Biophysical constraints on evapotranspiration partitioning for a conductance-based two source energy balance model

Evapotranspiration (ET) plays an indispensable role in water-carbon-energy exchange between the land surface and atmosphere. Biophysical constraints like the canopy conductance lie the key to accurate prediction of plant transpiration (T) and soil evaporation (E) in ET models. Given the lack of comprehensive evaluation for resistances network in conductance-based two source energy balance model (TSEB), three scenarios (Gc-TSEBO, Gc-TSEBOSM, and Gc-TSEBSWC) were assessed on the frame of Gc-TSEB: one relying on Jarvis canopy conductance, other on optimal stomatal behavior, and other on soil moisture content, with forcing data from MODIS datasets and in-situ FLUXNET sites over six biome types. The ET simulated by the model combined with the optimal stomatal behavior and the complementary concept had higher values of R2 and Taylor skill scores, lower values of RMSE and BIAS, than the other two scenarios, particularly over cropland sites and arid/semi-arid sites where the other two failed. The sensitivity analysis for Gc-TSEB resistances elucidated that the effect of canopy conductance is more noticeable on ET simulation than the other resistances. When effective biophysical constraints on the canopy are lacking, a large deviation exists in ET partitioning mainly due to the systematic error of soil evaporation through the partitioning algorithm for soil and plant fluxes. We assessed the performance of calibration from MODIS land surface temperature (LST) instead of flux data. The LST-calibrated model performed better in croplands and grasslands compared to forest sites, which opens a good perspective to advance the model for operational applications in agriculture and herbaceous vegetation.