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Cortical lacunae caused by drought, especially observed in hybrids originating from Vitis rupestris, disrupt the connection between roots and soil. Yet, the physiological processes behind lacuna formation during drought and its consistency across Vitis species remain unclear. Here, we used a root pressure probe to investigate fine root hydraulic and mechanical properties, in the arid‐adapted R‐65 and drought‐susceptible 101‐14Mgt cultivars. We then performed P–V curves, root sap osmolality, and electrolyte leakage (EL) and used fluorescent light microscopy techniques. Only 101‐14Mgt showed lacunae formation during drought due to its stiffer cortical tissue, unlike R‐65. Lacunae resulted in a notable decline in root hydraulic conductivity during severe drought, with increased EL and root sap osmolality, indicating potential cellular damage. R‐65 displayed different and xerophyte‐like characteristics featuring a higher turgor loss point and decreased root capacitance, essential for maintaining root structural integrity in arid conditions. Our findings highlight lacuna formation is impacted by root tissue elasticity possibly linked to specific Vitis species favoring deeper rooting. In arid‐adapted grapevines, hydraulic regulators such as reduced turgor loss point, and root capacitance could contribute to enhanced drought tolerance. [ABSTRACT FROM AUTHOR]

Drought is one of the main factors contributing to tree mortality worldwide and drought events are set to become more frequent and intense in the face of a changing climate. Quantifying water stress of forests is crucial in predicting and understanding their vulnerability to drought‐induced mortality. Here, we explore the use of high‐resolution spectroscopy in predicting water stress indicators of two native Australian tree species, Callitris rhomboidea and Eucalyptus viminalis. Specific spectral features and indices derived from leaf‐level spectroscopy were assessed as potential proxies to predict leaf water potential (Ψleaf), equivalent water thickness (EWT) and fuel moisture content (FMC) in a dedicated laboratory experiment. New spectral indices were identified that enabled very high confidence linear prediction of Ψleaf for both species (R2 > 0.85) with predictive capacity increasing when accounting for a breakpoint in the relationships using segmented regression (E. viminalis, R2 > 0.89; C. rhomboidea, R2 > 0.87). EWT and FMC were also linearly predicted to a high accuracy (E. viminalis, R2 > 0.90; C. rhomboidea, R2 > 0.80). This study highlights the potential of spectroscopy as a tool for predicting measures of plant water noninvasively, enabling broader applications for monitoring and managing plant water stress. Summary Statement: This study explored the use of proximal spectroscopy in predicting plant water stress indicators of two native Australian tree species. We derived new spectral indices that enabled very high prediction of leaf water potential, equivalent water thickness, and fuel moisture content. [ABSTRACT FROM AUTHOR]

Hydraulic failure due to xylem embolism has been identified as one of the main mechanisms involved in drought‐induced forest decline. Trees vulnerability to hydraulic failure depends on their hydraulic safety margin (HSM). While it has been shown that HSM globally converges between tree species and biomes, there is still limited knowledge regarding how HSM can adjust locally to varying drought conditions within species. In this study, we relied on three long‐term partial rainfall exclusion experiments to investigate the plasticity of hydraulic traits and HSM for three Mediterranean tree species (Quercus ilex L., Quercus pubescens Willd., and Pinus halepensis Mill.). For all species, a homeostasis of HSM in response to rainfall reduction was found, achieved through different mechanisms. For Q. ilex, the convergence in HSM is attributed to the adjustment of both the turgor loss point (Ψtlp) and the water potential at which 50% of xylem conductivity is lost due to embolism (P50). In contrast, the maintenance of HSM for P. halepensis and Q. pubescens is related to its isohydric behavior for the first and leaf area adjustment for the latter. It remains to be seen whether this HSM homeostasis can be generalized and if it will be sufficient to withstand extreme droughts expected in the Mediterranean region. Summary statement: Mediterranean species submitted to aggravated water stress have maintained their hydraulic safety margin (HSM). For Pinus halepensis and Quercus pubescens, HSM maintenance resulted from their capacity to escape water stress, while it results from the plasticity of Ψtlp and P50 for Quercus ilex. [ABSTRACT FROM AUTHOR]

ABSTRACT Plant hydraulic theory states that leaf and stem vulnerability to embolism is coordinated within individual plants. The hydraulic vulnerability segmentation hypothesis (HVSH) predicts higher vulnerability in leaves to protect the stem from hydraulic failure, preserving stem xylem, which is generally more metabolically expensive and slower to regenerate than leaf tissues. However, studies designed to test HVSH have reported wide ranges in vulnerability segmentation (VS), and patterns with the environment have been elusive. In this study, we tested HVSH in phylogenetically constrained tree species from contrasting ecosystems across the Australian landscape. In 12 species, we found no support for HVSH. While leaf vulnerability was strongly governed by climate, VS was universally absent or negative. Consistently, the onset of leaf embolism occurred after the loss of leaf turgor and seasonally low leaf water potentials, illustrating the rarity of embolism in leaves. Within the leaf, embolism primarily occurring first and last in the leaf midvein, suggesting redundancy in leaf architecture to preserve function. Overall, this multi‐ecosystem study provides a more complete picture of drought resistance mechanisms: (1) leaf safety was greatest in trees from drier ecosystems and (2) hydraulic thresholds were mostly conserved across organs indicating environmentally driven drought resistance in both leaves and stems. [ABSTRACT FROM AUTHOR]

Summary Drought predisposes forest trees to bark beetle‐induced mortality, but the physiological mechanisms remain unclear. While drought‐induced water and carbon limitations have been implicated in defensive failure and tree susceptibility, evidence demonstrating how these factors interact is scarce. We withheld water from mature, potted Pinus edulis and subsequently applied a double‐stem girdle to inhibit carbohydrate transport from the crown and roots. Within this isolated segment we then elicited a defense response by inoculating trees with a bark beetle‐fungal symbiont (Ophiostoma sp.). We quantified local mono‐ and sesquiterpenes (MST), nonstructural carbohydrates (NSC), and pressure potential of the inner bark. Both drought‐stressed and watered trees had similar NSC concentrations just before inoculation and depleted NSC similarly following inoculation, yet MST induction (i.e. increased concentration and altered composition) was constrained only in drought‐stressed trees. Thus, NSC consumption was largely unrelated to de novo MST synthesis. Instead, stoichiometric calculations show that induction originated largely from stored resin. Watered trees experiencing higher pressure potentials consistently induced higher MST concentrations. We demonstrate the importance of preformed resin toward an induced MST response in a semi‐arid conifer where drought‐constraints on defense occurred through biophysical limitations (i.e. reduced turgor hindering resin transport) rather than through substrate limitation. [ABSTRACT FROM AUTHOR]

ABSTRACT In ecophysiology leaves are frequently stored for hours after sampling before measuring their leaf water potential (Ψleaf). Here, we address a previously unidentified source of error, that metabolic heat generation can cause continuous water loss from leaves stored in impermeable bags, leading to a Ψleaf drop over time. We tested Ψleaf drop rates under various conditions: two bag materials, two species, initial Ψleaf above or below the turgor loss point (Ψtlp), and storage at 25°C versus 4°C. We partitioned leaf water loss due to condensation on the inner bag surface or permeation through the bag. We found that Ψleaf dropped by up to 0.39 MPa per hour, with 41%–89% of the water leaving the leaf condensed on the inner bag surface. Plastic bags showed higher Ψleaf drop rates than aluminium bags, and leaves above Ψtlp experienced greater drops. Storing leaves at 4°C reduced the Ψleaf drop rate by 60% compared to 25°C. Leaves were 0.2–0.3°C warmer than the bags, likely due to metabolic heating. Our energy balance model suggests that water loss is lower when storing leaves at cooler temperatures, using leaves with low stomatal conductance, deflated bags, and leaves with low Ψleaf. [ABSTRACT FROM AUTHOR]

Premise: Vascular epiphytes of tropical montane cloud forests are vulnerable to climate change, particularly as cloud bases elevate and reduce atmospheric inputs to the system. However, studies have generally focused on epiphytes in contiguous forests, with little research being done on epiphytes on isolated pasture trees. We investigated water relations of pasture‐tree epiphytes at three sites located below and above the elevation of the average cloud base in Monteverde, Costa Rica. Methods: We measured sap velocity and four microclimate variables in both the dry and wet season of 2018. We also measured functional traits, including pressure volume (PV) curves, predawn/midday water potential, and various lab‐based water relations traits. We used linear mixed models to assess the correlation between microclimate and sap velocity in both seasons and ANOVA to assess the variation in PV curve and water potential variables. Results: The turgor loss point generally increased from the wettest to driest site. However, this trend was driven primarily by the increasing prevalence of leaf succulence at drier sites. Microclimatic variables correlated strongly with sap velocity in the wet season, but low soil moisture availability caused this correlation to break down during the dry season. Conclusions: Our results emphasize the vulnerability of cloud forest epiphytes to rising cloud bases. This vulnerability may be more severe in pasture trees that lack the potential buffer of surrounding forest, but additional research that directly compares the canopy microclimate conditions between forest and pasture trees is needed to confirm this possibility. [ABSTRACT FROM AUTHOR]

ABSTRACT Determining the differences in flower hydraulic traits and structural resource allocation among closely related species adapted to low mean annual precipitation (MAP) can provide insight into plant adaptation to arid environments. Here, we measured the maximum flower hydraulic conductance (Kmax‐flower), water potential at induction 50% loss of Kmax‐flower (P50‐flower), flower pressure–volume parameters, dry mass of individual flowers and structural components (vexillum, wings, keels, stamens and sepals) of six Caragana species growing in regions ranging from 110 to 1400 mm MAP. Compared with species from high‐MAP environments, those from low‐MAP environments presented lower Kmax‐flower, more negative P50‐flower, osmotic potential at full turgor (πo) and turgor loss points (πtlp), and a greater bulk modulus of elasticity (ε). Consequently, a negative correlation between Kmax‐flower (hydraulic efficiency) and P50‐flower (hydraulic safety) was observed across Caragana species. Furthermore, the dry masses of individual flowers and structural components (vexillum, wings, keels, stamens and sepals) were greater in the species from the low‐MAP environment than in those from the high‐MAP environment. These findings suggest that greater flower hydraulic safety and drought tolerance combined with greater structural resource allocation promote drought adaptation in Caragana species to low‐MAP environments. [ABSTRACT FROM AUTHOR]

As climate change thrives, and the frequency of intense droughts is affecting many forested regions, a mechanistic understanding of the factors conferring drought tolerance in trees is increasingly important. However, studies linking the observed growth reduction to mechanistic traits are still rare. We compared the median growth anomalies of 16 native tree species, gathered across a network of study plots in Bavaria, with the mean species‐specific turgor loss point (πtlp) measured at five locations in Central Europe πtlp explained 37% of the growth anomalies observed in response to the intense droughts between 2018 and 2020 compared to the pre‐drought period between 2006 and 2017 across sites. πtlp constitutes an important leaf drought tolerance trait and influences the growth response of native tree species during extraordinary dry periods. As climate change‐induced droughts intensify, tree species with drought‐tolerant leaves will be less vulnerable to growth reductions. πtlp provides a useful indicator for selecting tree species to adapt forest management systems to climate change. [ABSTRACT FROM AUTHOR]

The African Restionaceae (Poales), the dominant graminoid layer in the megadiverse Cape Floristic Region of South Africa, are distributed across a wide range of moisture availability, yet currently there is very little known about the underlying hydraulics of this group.We tested two methods for measuring culm vulnerability to embolism, the optical and pneumatic methods, in three species of Cannomois ranging in habitat from semi‐riparian (Cannomois virgata) to dryland (Cannomois parviflora and C. congesta). Estimates of culm xylem vulnerability were coupled with measures of turgor loss point (ΨTLP) and minimum field water potential (ΨMD) to assess hydraulic safety margins.The optical and pneumatic methods produced similar estimates of P50, but differed for P12 and P88. All three species were quite vulnerable to embolism, with P50 of –1.9 MPa (C. virgata), −2.3 MPa (C. congesta), and −2.4 MPa (C. parviflora). Estimates of P50, ΨTLP and ΨMD aligned with habitat moisture stress, with highest values found in the semi‐riparian C. virgata. Consistent differences in P50, ΨMD and ΨTLP between species resulted in consistent hydraulic safety margins across species of 0.96 ± 0.1 MPa between ΨMD and P50, with onset of embolism occurring 0.43 ± 0.04 MPa after ΨTLP for all three species.Our study demonstrates that restio occupancy of dry environments involves more than the evolution of highly resistant xylem, suggesting that other aspects of water relations are key to understanding trait–environment relationships in this group. [ABSTRACT FROM AUTHOR]

Summary: Turgor loss point (TLP) is an important proxy for plant drought tolerance, species habitat suitability, and drought‐induced plant mortality risk. Thus, TLP serves as a critical tool for evaluating climate change impacts on plants, making it imperative to develop high‐throughput and in situ methods to measure TLP.We developed hyperspectral pressure–volume curves (PV curves) to estimate TLP using leaf spectral reflectance. We used partial least square regression models to estimate water potential (Ψ) and relative water content (RWC) for two species, Frangula caroliniana and Magnolia grandiflora. RWC and Ψ's model for each species had R2 ≥ 0.7 and %RMSE = 7–10. We constructed PV curves with model estimates and compared the accuracy of directly measured and spectra‐predicted TLP.Our findings indicate that leaf spectral measurements are an alternative method for estimating TLP. F. caroliniana TLP's values were −1.62 ± 0.15 (means ± SD) and −1.62 ± 0.34 MPa for observed and reflectance predicted, respectively (P > 0.05), while M. grandiflora were −1.78 ± 0.34 and −1.66 ± 0.41 MPa (P > 0.05).The estimation of TLP through leaf reflectance‐based PV curves opens a broad range of possibilities for future research aimed at understanding and monitoring plant water relations on a large scale with spectral ecophysiology. [ABSTRACT FROM AUTHOR]

Direct and non-destructive measurements of plant-water relations of plants exhibiting the Crassulacean acid metabolism (CAM) photosynthetic pathway are seldom addressed, with most findings inferred from gas exchange measurements. The main focus of this paper was to study how the water exchange between the chlorenchyma and the hydrenchyma depends on and follows the CAM photosynthetic diel pattern using non-invasive and continuous methods. Gas exchange and leaf patch clamp pressure probe (LPCP) measurements were performed on Aloe vera (L.) Burm f., a CAM species, and compared to measurements on banana (Musa acuminata Colla), a C3 species. The LPCP output pressure, Pp, of Aloe vera plants follows its diel CAM photosynthetic cycle, reversed to that observed in banana and other C3 species. The four phases of CAM photosynthesis can also be identified in the diel LPCP output pressure, Pp, cycle. The Pp values in Aloe vera are determined by the hydrenchyma turgor pressure, with both parameters being reversely related. A non-invasive and continuous assessment of the water exchange between the chlorenchyma and the hydrenchyma in CAM plants, namely, by following the changes in the hydrenchyma turgor pressure, is presented. However, showing once more how the LPCP output pressure, Pp, depends on the leaf structure, such an approach can be used to study plant-water relations in other CAM species with a leaf structure similar to Aloe vera, with the hydrenchyma composing most of the leaf volume. [ABSTRACT FROM AUTHOR]

Drought is one of the most serious abiotic stresses to land plants. Plants sense and respond to drought stress to survive under water deficiency. Scientists have studied how plants sense drought stress, or osmotic stress caused by drought, ever since Charles Darwin, and gradually obtained clues about osmotic stress sensing and signaling in plants. Osmotic stress is a physical stimulus that triggers many physiological changes at the cellular level, including changes in turgor, cell wall stiffness and integrity, membrane tension, and cell fluid volume, and plants may sense some of these stimuli and trigger downstream responses. In this review, we emphasized water potential and movements in organisms, compared putative signal inputs in cell wall‐containing and cell wall‐free organisms, prospected how plants sense changes in turgor, membrane tension, and cell fluid volume under osmotic stress according to advances in plants, animals, yeasts, and bacteria, summarized multilevel biochemical and physiological signal outputs, such as plasma membrane nanodomain formation, membrane water permeability, root hydrotropism, root halotropism, Casparian strip and suberin lamellae, and finally proposed a hypothesis that osmotic stress responses are likely to be a cocktail of signaling mediated by multiple osmosensors. We also discussed the core scientific questions, provided perspective about the future directions in this field, and highlighted the importance of robust and smart root systems and efficient source‐sink allocations for generating future high‐yield stress‐resistant crops and plants. [ABSTRACT FROM AUTHOR]

As heatwave frequency increases, they are more likely to coincide with other disturbances like insect defoliation. But it is unclear if high temperatures after defoliation impact canopy recovery or leaf traits which may affect response to further stressors like drought. To examine these stressor interactions, we subjected defoliated (DEF) and undefoliated (UNDEF) oak saplings to a simulated spring heatwave of +10°C for 25 days. We measured gas exchange, leaf area recovery, carbohydrate storage, turgor loss point (ΨTLP), and minimum leaf conductance (gmin). During the heatwave, stem respiration exhibited stronger thermal acclimation in DEF than UNDEF saplings, while stomatal conductance and net photosynthesis increased. The heatwave did not affect leaf area recovery or carbohydrate storage of DEF saplings, but reflush leaves had higher gmin than UNDEF leaves, and this was amplified by the heatwave. Across all treatments, higher gmin was associated with higher daytime stomatal conductance and a lower ΨTLP. The results suggest defoliation stress may not be exacerbated by higher temperatures. However, reflush leaves are less conservative in their water use, limiting their ability to minimise water loss. While lower ΨTLP could help DEF trees maintain gas exchange under mild drought, they may be more vulnerable to dehydration under severe drought. Summary Statement: Leaf area recovery and carbohydrate storage after spring defoliation were not impacted by a coinciding heatwave. However, reflush leaves had higher minimum rates of water loss—and this effect was exacerbated by the heatwave, suggesting reduced drought tolerance. [ABSTRACT FROM AUTHOR]

Stomata are the key nodes linking photosynthesis and transpiration. By regulating the opening degree of stomata, plants successively achieve the balance between water loss and carbon dioxide acquisition. The dynamic behaviour of stomata is an important cornerstone of plant adaptability. Though there have been miscellaneous experimental results on stomata and their constituent cells, the guard cells and the subsidiary cells, current theory of stomata regulation is far from clear and unified. In this work, we develop an integrated model to describe the stomatal dynamics of seed plants based on existing experimental results. The model includes three parts: (1) a passive mechanical model of the stomatal aperture as a bivariate function of the guard‐cell turgor and the subsidiary‐cell turgor; (2) an active regulation model with a target ion‐content in guard cells as a function of their water potential; and (3) a dynamical model for the movement of potassium ions and water content. Our model has been used to reproduce abundant experimental phenomena semi‐quantitatively. With accurately measured parameters, our model can also be used to predict stomatal responses to changes of environmental conditions. Summary statement: A stomata model is developed based on existing experiments and reproduces important stomatal behaviour. The model takes into account passive mechanical interaction between guard cells and subsidiary cells, active movement of potassium ions, and water transport driven by water potential. [ABSTRACT FROM AUTHOR]

Summary: The high productive potential, heat resilience, and greater water use efficiency of C4 over C3 plants attract considerable interest in the face of global warming and increasing population, but C4 plants are often sensitive to dehydration, questioning the feasibility of their wider adoption.To resolve the primary effect of dehydration from slower from secondary leaf responses originating within leaves to combat stress, we conducted an innovative dehydration experiment. Four crops grown in hydroponics were forced to a rapid yet controlled decrease in leaf water potential by progressively raising roots of out of the solution while measuring leaf gas exchange.We show that, under rapid dehydration, assimilation decreased more steeply in C4 maize and sorghum than in C3 wheat and sunflower. This reduction was due to a rise of nonstomatal limitation at triple the rate in maize and sorghum than in wheat and sunflower.Rapid reductions in assimilation were previously measured in numerous C4 species across both laboratory and natural conditions. Hence, we deduce that high sensitivity to rapid dehydration might stem from the disturbance of an intrinsic aspect of C4 bicellular photosynthesis. We posit that an obstruction to metabolite transport between mesophyll and bundle sheath cells could be the cause. [ABSTRACT FROM AUTHOR]

In legumes, a common phenomenon known as nyctinastic movement is observed. This movement involves the horizontal expansion of leaves during the day and relative vertical closure at night. Nyctinastic movement is driven by the pulvinus, which consists of flexor and extensor motor cells. The turgor pressure difference between these two cell types generates a driving force for the bending and deformation of the pulvinus. This review focuses on the developmental mechanisms of the pulvinus, the factors affecting nyctinastic movement, and the biological significance of this phenomenon in legumes, thus providing a reference for further research on nyctinastic movement. Summary statement: Nyctinastic movement is characterised by the horizontal expansion of leaves during the day and the vertical closure at night. In this paper, we provided an overview of the research progress and summarised the biological significance of this phenomenon for further investigation. [ABSTRACT FROM AUTHOR]

Summary: Pollen tubes have dynamic tubular vacuoles. Functional loss of AP‐3, a regulator of one vacuolar trafficking route, reduces pollen tube growth. However, the role of canonical Rab5 GTPases that are responsible for two other vacuolar trafficking routes in Arabidopsis pollen tubes is obscure.By using genomic editing, confocal microscopy, pollen tube growth assays, and transmission electron microscopy, we demonstrate that functional loss of canonical Rab5s in Arabidopsis, RHA1 and ARA7, causes the failure of pollen tubes to grow through style and thus impairs male transmission.Functional loss of canonical Rab5s compromises vacuolar trafficking of tonoplast proteins, vacuolar biogenesis, and turgor regulation. However, rha1;ara7 pollen tubes are comparable to those of wild‐type in growing through narrow passages by microfluidic assays.We demonstrate that functional loss of canonical Rab5s compromises endocytic and secretory trafficking at the plasma membrane (PM), whereas the targeting of PM‐associated ATPases is largely unaffected. Despite that, rha1;ara7 pollen tubes contain a reduced cytosolic pH and disrupted actin microfilaments, correlating with the mis‐targeting of vacuolar ATPases (VHA). These results imply a key role of vacuoles in maintaining cytoplasmic proton homeostasis and in pollen tube penetrative growth through style. [ABSTRACT FROM AUTHOR]

Tropical forests are experiencing increases in vapour pressure deficit (D), with possible negative impacts on tree growth. Tree‐growth reduction due to rising D is commonly attributed to carbon limitation, thus overlooking the potentially important mechanism of D‐induced impairment of wood formation due to an increase in turgor limitation. Here we calibrate a mechanistic tree‐growth model to simulate turgor limitation of radial stem growth in mature Toona cilitata trees in an Asian tropical forest. Hourly sap flow and dendrometer measurements were collected to simulate turgor‐driven growth during the growing season. Simulated seasonal patterns of radial stem growth matched well with growth observations. Growth mainly occurred at night and its pre‐dawn build‐up appeared to be limited under higher D. Across seasons, the night‐time turgor pressure required for growth was negatively related to previous midday D, possibly due to a relatively high canopy conductance at high D, relative to stem rehydration. These findings provide the first evidence that tropical trees grow at night and that turgor pressure limits tree growth. We suggest including turgor limitation of tree stem growth in models also for tropical forest carbon dynamics, in particular, if these models simulate effects of warming and increased frequency of droughts. [ABSTRACT FROM AUTHOR]

Allopolyploidy is a common speciation mechanism in plants; however, its physiological and ecological consequences in niche partitioning have been scarcely studied. In this sense, leaf traits are good proxies to study the adaptive capacity of allopolyploids and diploid parents to their respective environmental conditions. In the present work, leaf water relations (assessed through pressure–volume curves) and structural and anatomical traits of the allotetraploid fern Oeosporangium tinaei and its diploid parents, Oeosporangium hispanicum and Oeosporangium pteridioides, were studied under controlled conditions in response to a water stress (WS) cycle. O. hispanicum showed the lowest osmotic potential at turgor loss point (πtlp) and leaf capacitance, together with higher leaf mass per area (LMA), leaf thickness (LT), leaf density (LD), and leaf dry matter content (LDMC), whereas O. pteridioides presented the opposite set of traits (high πtlp and capacitance, and low LMA, LT, LD, and LDMC). O. tinaei showed an intermediate position for most of the studied traits. The responsiveness (osmotic and elastic adjustments) to WS was low, although most of the traits explained the segregation of the three species across a range of drought tolerance according to the rank: O. hispanicum > O. tinaei > O. pteridioides. These trait differences may underlie the niche segregation among coexisting populations of the three species in the Mediterranean basin. [ABSTRACT FROM AUTHOR]

Cavitation resistance has often been viewed as a relatively static trait, especially for stems of forest trees. Meanwhile, other hydraulic traits, such as turgor loss point (Ψtlp) and xylem anatomy, change during the season. In this study, we hypothesized that cavitation resistance is also dynamic, changing in coordination with Ψtlp. We began with a comparison of optical vulnerability (OV), microcomputed tomography (µCT) and cavitron methods. All three methods significantly differed in the slope of the curve,Ψ12 and Ψ88, but not in Ψ50 (xylem pressures that cause 12%, 88%, 50% cavitation, respectively). Thus, we followed the seasonal dynamics (across 2 years) of Ψ50 in Pinus halepensis under Mediterranean climate using the OV method. We found that Ψ50 is a plastic trait with a reduction of approximately 1 MPa from the end of the wet season to the end of the dry season, in coordination with the dynamics of the midday xylem water potential (Ψmidday) and the Ψtlp. The observed plasticity enabled the trees to maintain a stable positive hydraulic safety margin and avoid cavitation during the long dry season. Seasonal plasticity is vital for understanding the actual risk of cavitation to plants and for modeling species' ability to tolerate harsh environments. Summary Statement: Shoot xylem vulnerability of Pinus halepensis is a seasonal plastic trait, which adjusts to lower water potential during the summer in coordination with the turgor loss point. [ABSTRACT FROM AUTHOR]

Summary: Every existing optimal stomatal model uses photosynthetic carbon assimilation as a proxy for plant evolutionary fitness. However, assimilation and growth are often decoupled, making assimilation less ideal for representing fitness when optimizing stomatal conductance to water vapor and carbon dioxide. Instead, growth should be considered a closer proxy for fitness.We hypothesize stomata have evolved to maximize turgor‐driven growth, instead of assimilation, over entire plants' lifetimes, improving their abilities to compete and reproduce. We develop a stomata model that dynamically maximizes whole‐stem growth following principles from turgor‐driven growth models. Stomata open to assimilate carbohydrates that supply growth and osmotically generate turgor, while stomata close to prevent losses of turgor and growth due to negative water potentials.In steady state, the growth optimization model captures realistic stomatal, growth, and carbohydrate responses to environmental cues, reconciles conflicting interpretations within existing stomatal optimization theories, and explains patterns of carbohydrate storage and xylem conductance observed during and after drought.Our growth optimization hypothesis introduces a new paradigm for stomatal optimization models, elevates the role of whole‐plant carbon use and carbon storage in stomatal functioning, and has the potential to simultaneously predict gross productivity, net productivity, and plant mortality through a single, consistent modeling framework. See also the Commentary on this article by Buckley, 238: 457–460. [ABSTRACT FROM AUTHOR]

Summary: Growth is central to plant morphogenesis. Plant cells are encased in rigid cell walls, and they must overcome physical confinement to grow to specific sizes and shapes. Cell wall tension and turgor pressure are the main mechanical components impacting plant cell growth. Cell wall mechanics has been the focus of most plant biomechanical studies, and turgor pressure was often considered as a constant and largely passive component. Nevertheless, it is increasingly accepted that turgor pressure plays a significant role in plant growth. Numerous theoretical and experimental studies suggest that turgor pressure can be both spatially inhomogeneous and actively modulated during morphogenesis. Here, we revisit the pressure–growth relationship by reviewing recent advances in investigating the interactions between cellular/tissular pressure and growth. [ABSTRACT FROM AUTHOR]

Fruit mesocarp cracking caused by improper irrigation during development manifests at the macroscale but is ultimately the result of increasing cell turgor pressure at the microscale. Hence, a cell finite element (FE) model including shape, protoplast turgor pressure, and ripening information and a mesocarp tissue block discrete element (DE) model including the features of cell shape and number, were developed to predict the biomechanical correlation between mesocarp and its cell. The validated cell FE model with an internal turgor pressure of 12.9 kPa could reproduce the experimental force–deformation behavior of a single cell in compression up to 11% deformation with an average relative error of 5.8%. The validated mesocarp tissue block DE model could reproduce the experimental force–deformation behavior of a mesocarp block in compression up to 20% deformation with an average relative error of 9.5%. Sensitivity and regression analysis showed that turgor pressure was the most important factor affecting cell biomechanics, followed by cell shape and wall elastic modulus. Similarly, the apparent elastic modulus of the cells has the most significant effect on the mesocarp tissue biomechanics, followed by the number and shape of cells. Finally, a mathematical model was obtained to quantitatively describe the relationship between the elastic modulus of the mesocarp and its cell turgor pressure. This study contributes to a better understanding of the biomechanical mechanisms of irrigation‐caused tomato fruit cracking at the cellular level and the development of strategies to prevent fruit cracking through a combination of gene breeding and irrigation management. [ABSTRACT FROM AUTHOR]

In recent years, attempts have been made in linking pressure–volume parameters and the leaf economics spectrum to expand our knowledge of the interrelationships among leaf traits. We provide theoretical and empirical evidence for the coordination of the turgor loss point and associated traits with net CO2 assimilation (An) and leaf mass per area (LMA). We measured gas exchange, pressure–volume curves and leaf structure in 45 ferns and angiosperms, and explored the anatomical and chemical basis of the key traits. We propose that the coordination observed between mass‐based An, capacitance and the turgor loss point (πtlp) emerges from their shared link with leaf density (one of the components of LMA) and, specially, leaf saturated water content (LSWC), which in turn relates to cell size and nitrogen and carbon content. Thus, considering the components of LMA and LSWC in ecophysiological studies can provide a broader perspective on leaf structure and function. [ABSTRACT FROM AUTHOR]

Leaf hydraulic traits characterize plant drought tolerance and responses to climate change. Yet, plant hydraulics are biased towards northern hemisphere woody species. We collected rhizomes of several perennial grass species along a precipitation gradient in eastern Australia and grew them in an experimental pot study to investigate potential trade‐offs between drought tolerance and plant morphology.We measured the following leaf hydraulic traits: the leaf water potential (Ψleaf) at 50% and 88% loss of leaf hydraulic conductance (P50Kleaf and P88Kleaf), the Ψleaf at 50% loss of stomatal conductance (P50gs), leaf turgor loss point (TLP), leaf dry matter content (LDMC), leaf modulus of elasticity (ε), and the slope of the relationship between predawn and midday Ψleaf. We also measured basal area, tiller density, seed head density, root collar diameter, plant height, and aboveground biomass of each individual.As expected, grass species varied widely in leaf‐level drought tolerance, with loss of 88% hydraulic conductance occurring at a Ψleaf ranging from −1.52 to −4.01 MPa. However, all but one species lost leaf turgor, and most reached P50gs before this critical threshold. Taller more productive grass species tended to have drought vulnerable leaves characterized by low LDMC and less negative P88Kleaf. Species with greater tiller production experienced stomatal closure and lost turgor at more negative Ψleaf. Although our sample size was limited, we found no relationships between these species' traits and their climate of origin.Overall, we identified important hydraulic and morphological trade‐offs in Australian grasses that were surprisingly similar to those observed for woody plants: (1) xylem of taller species was less drought tolerant and (2) turgor loss occurs and stomatal closure begins before significant loss of Kleaf. These data build upon a small yet growing field of grass hydraulics and may be informative of species responses to further drought intensification in Australia. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

The sequence of physiological events during drought strongly impacts plants' overall performance. Here, we synthesized the global data of stomatal and hydraulic traits in leaves and stems of 202 woody species to evaluate variations in the water potentials for key physiological events and their sequence along the climatic gradient. We found that the seasonal minimum water potential, turgor loss point, stomatal closure point, and leaf and stem xylem vulnerability to embolism were intercorrelated and decreased with aridity, indicating that water stress drives trait co‐selection. In xeric regions, the seasonal minimum water potential occurred at lower water potential than turgor loss point, and the subsequent stomatal closure delayed embolism formation. In mesic regions, however, the seasonal minimum water potential did not pose a threat to the physiological functions, and stomatal closure occurred even at slightly more negative water potential than embolism. Our study demonstrates that the sequence of water potentials for physiological dysfunctions of woody plants varies with aridity, that is, xeric species adopt a more conservative sequence to prevent severe tissue damage through tighter stomatal regulation (isohydric strategy) and higher embolism resistance, while mesic species adopt a riskier sequence via looser stomatal regulation (anisohydric strategy) to maximize carbon uptake at the cost of hydraulic safety. Integrating both aridity‐dependent sequence of water potentials for physiological dysfunctions and gap between these key traits into the hydraulic framework of process‐based vegetation models would improve the prediction of woody plants' responses to drought under global climate change. [ABSTRACT FROM AUTHOR]

Summary: Evolutionary relationships are likely to play a significant role in shaping plant physiological and structural traits observed in contemporary taxa. We review research on phylogenetic signal and correlated evolution in plant–water relation traits, which play important roles in allowing plants to acquire, use, and conserve water. We found more evidence for a phylogenetic signal in structural traits (e.g. stomatal length and stomatal density) than in physiological traits (e.g. stomatal conductance and water potential at turgor loss). Although water potential at turgor loss is the most‐studied plant–water relation trait in an evolutionary context, it is the only trait consistently found to not have a phylogenetic signal. Correlated evolution was common among traits related to water movement efficiency and hydraulic safety in both leaves and stems. We conclude that evidence for phylogenetic signal varies depending on: the methodology used for its determination, that is, model‐based approaches to determine phylogenetic signal such as Blomberg's K or Pagel's λ vs statistical approaches such as ANOVAs with taxonomic classification as a factor; on the number of taxa studied (size of the phylogeny); and the setting in which plants grow (field vs common garden). More explicitly and consistently considering the role of evolutionary relationships in shaping plant ecophysiology could improve our understanding of how traits compare among species, how traits are coordinated with one another, and how traits vary with the environment. [ABSTRACT FROM AUTHOR]

Increasing frequency and severity of drought events is posing risks to trees' health, including those planted in urban settlements. Drought‐induced decline of urban trees negatively affects ecosystem services of urban green spaces and implies cost for maintenance and removal of plants. We aimed at identifying physiological traits that can explain and predict the species‐specific vulnerability to climate change in urban habitats.We assessed the relationships between long‐term risk of decline of different tree species in a medium‐sized town and their key indicators of drought stress tolerance, i.e. turgor loss point (TLP) and vulnerability to xylem embolism (P50).Starting from 2012, the study area experienced several summer seasons with positive anomalies of temperature and negative anomalies of precipitation. This trend was coupled with increasing percentages of urban trees showing signs of crown die‐back and mortality. The species‐specific risk of decline was higher for species with less negative TLP and P50 values.The relationship between species‐specific risk of climate change‐induced decline of urban trees and key physiological indicators of drought tolerance confirms findings obtained in natural forests and highlights that TLP and P50 are useful indicators for species selection for tree plantation in towns, to mitigate negative impacts of climate change. [ABSTRACT FROM AUTHOR]

Predicting the consequences of climate change is of utmost importance to mitigate impacts on vulnerable ecosystems; plant hydraulic traits are particularly useful proxies for predicting functional disruptions potentially occurring in the near future. This study assessed the current and future regional patterns of leaf water potential at turgor loss point (Ψtlp) by measuring and projecting the Ψtlp of 166 vascular plant species (159 angiosperms and 7 gymnosperms) across a large climatic range spanning from alpine to Mediterranean areas in NE Italy. For angiosperms, random forest models predicted a consistent shift toward more negative values in low‐elevation areas, whereas for gymnosperms the pattern was more variable, particularly in the alpine sector (i.e., Alps and Prealps). Simulations were also developed to evaluate the number of threatened species under two Ψtlp plasticity scenarios (low vs. high plasticity), and it was found that in the worst‐case scenario approximately 72% of the angiosperm species and 68% of gymnosperms within a location were at risk to exceed their physiological plasticity. The different responses to climate change by specific clades might produce reassembly in natural communities, undermining the resilience of natural ecosystems to climate change. [ABSTRACT FROM AUTHOR]

Summary: In higher plants, photosystems II and I are found in grana stacks and unstacked stroma lamellae, respectively. To connect them, electron carriers negotiate tortuous multi‐media paths and are subject to macromolecular blocking. Why does evolution select an apparently unnecessary, inefficient bipartition? Here we systematically explain this perplexing phenomenon. We propose that grana stacks, acting like bellows in accordions, increase the degree of ultrastructural control on photosynthesis through thylakoid swelling/shrinking induced by osmotic water fluxes. This control coordinates with variations in stomatal conductance and the turgor of guard cells, which act like an accordion's air button. Thylakoid ultrastructural dynamics regulate macromolecular blocking/collision probability, direct diffusional pathlengths, division of function of Cytochrome b6f complex between linear and cyclic electron transport, luminal pH via osmotic water fluxes, and the separation of pH dynamics between granal and lamellar lumens in response to environmental variations. With the two functionally asymmetrical photosystems located distantly from each other, the ultrastructural control, nonphotochemical quenching, and carbon‐reaction feedbacks maximally cooperate to balance electron transport with gas exchange, provide homeostasis in fluctuating light environments, and protect photosystems in drought. Grana stacks represent a dry/high irradiance adaptation of photosynthetic machinery to improve fitness in challenging land environments. Our theory unifies many well‐known but seemingly unconnected phenomena of thylakoid structure and function in higher plants. [ABSTRACT FROM AUTHOR]

The phloem is a key player in whole plant functioning—transporting carbon from sites of production to sites of demand—and is likely influenced by drought due to its dependence on water for generating pressure‐driven bulk flow transport. Yet, phloem functioning during drought remains largely unknown due to a lack of experimental studies. Here, we use a phloem‐bleeding species, Ricinus communis, to investigate phloem loss‐of‐function in the context of leaf physiological processes, the mechanisms of phloem turgor maintenance during drought, and the role of turgor in phloem loss‐of‐function. We found that the solute concentration in the phloem sap doubled over the drought, which allowed phloem turgor to be maintained past the point at which leaves have reached permanent stomatal closure. We also found that phloem turgor did not decline before bleeding ceased, which suggests that phloem bleeding ceassation (interpreted as the cessation of transport) occurred when the phloem still had turgor. In sum, our findings highlight the robustness of phloem functioning, with important implications for forecasting whole‐plant carbon dynamics and drought‐induced tree mortality. [ABSTRACT FROM AUTHOR]

Turgor pressure is an essential, but difficult to measure indicator of plant water status. Turgor has been quantified by localized compression of cells or tissues, but a simple method to perform these measurements is lacking. We hypothesized that changes in leaf turgidity can be monitored by uniaxially compressing the leaf lamina and measuring the mechanical stress under a constrained thickness (stress relaxation) and that changes in leaf water content can be monitored by measuring the leaf thickness under constant mechanical stress. Using a simple, custom‐built leaf squeeze‐flow rheometer, we performed different compression tests on leaves from 13 plant species. The mechanical stress measured during stress relaxation was correlated with leaf bulk turgor pressure (R2 > 0.95) and thus with balancing pressure (R2 > 0.94); the leaf thickness measured under constant mechanical stress was correlated with relative water content (R2 > 0.74). The coefficients of these relationships were related to the leaf bulk osmotic pressure at the turgor‐loss point. An idealized average‐cell model suggests that, under isothermal conditions, the stationary bulk modulus during compression is largely determined by the bulk osmotic pressure. Our study presents an inexpensive, accessible and automatable method to monitor plant water status noninvasively. [ABSTRACT FROM AUTHOR]

The isohydric–anisohydric continuum describes the relative stringency of stomatal control of leaf water potential (ψleaf) during drought. Hydroscape area (HA)—the water potential landscape over which stomata regulate ψleaf—has emerged as a useful metric of the iso/anisohydric continuum because it is strongly linked to several hydraulic, photosynthetic and structural traits. Previous research on HA focused on broad ecological patterns involving several plant clades. Here we investigate the relationships between HA and climatic conditions and functional traits across ecologically diverse but closely related species while accounting for phylogeny. Across a macroclimatic moisture gradient, defined by the ratio of mean annual precipitation to mean annual pan evaporation (P/Ep), HA decreased with increased P/Ep across 10 Eucalyptus species. Greater anisohydry reflects lower turgor loss points and greater hydraulic safety, mirroring global patterns. Larger HA coincides with mesophyll photosynthetic capacity that is more sensitive to ψleaf. Hydroscapes exhibit little plasticity in response to variation in water supply, and the extent of plasticity does not vary with P/Ep of native habitats. These findings strengthen the case that HA is a useful metric for characterizing drought tolerance and water‐status regulation. Smaller hydroscape area (increased isohydry) in Eucalyptus species is tied to moister climates and drought‐sensitive functional traits like higher turgor loss points and mesophyll photosynthetic sensitivity to leaf water potential. Hydroscape area shows little plasticity to water availability. [ABSTRACT FROM AUTHOR]

Increasing temperature and drought can result in leaf dehydration and defoliation even in drought‐adapted tree species such as the Mediterranean evergreen Quercus ilex L. The stomatal regulation of leaf water potential plays a central role in avoiding this phenomenon and is constrained by a suite of leaf traits including hydraulic conductance and vulnerability, hydraulic capacitance, minimum conductance to water vapour, osmotic potential and cell wall elasticity. We investigated whether the plasticity in these traits may improve leaf tolerance to drought in two long‐term rainfall exclusion experiments in Mediterranean forests. Osmotic adjustment was observed to lower the water potential at turgor loss in the rainfall‐exclusion treatments, thus suggesting a stomatal closure at more negative water potentials and a more anisohydric behaviour in drier conditions. Conversely, leaf hydraulic conductance and vulnerability did not exhibit any plasticity between treatments so the hydraulic safety margins were narrower in the rainfall‐exclusion treatments. The sequence of leaf responses to seasonal drought and dehydration was conserved among treatments and sites but trees were more likely to suffer losses of turgor and hydraulic functioning in the rainfall‐exclusion treatments. We conclude that leaf plasticity might help the trees to tolerate moderate drought but not to resist severe water stress. [ABSTRACT FROM AUTHOR]

Identifying key traits that can serve as proxies for species drought resistance is crucial for predicting and mitigating the effects of climate change in diverse plant communities. Turgor loss point (πtlp) is a recently emerged trait that has been linked to species distributions across gradients of water availability. However, a direct relationship between πtlp and species ability to survive drought has yet to be established for woody species. Using a manipulative field experiment to quantify species drought resistance (i.e., their survival response to drought), combined with measurements of πtlp for 16 tree species, we show a negative relationship between πtlp and seedling drought resistance. Using long‐term forest plot data, we also show that πtlp predicts seedling survival responses to a severe El Niño‐related drought, although additional factors are clearly also important. Our study demonstrates that species with lower πtlp exhibit higher survival under both experimental and natural drought. These results provide a missing cornerstone in the assessment of the traits underlying drought resistance in woody species and strengthen πtlp as a proxy for evaluating which species will lose or win under projections of exacerbating drought regimes. [ABSTRACT FROM AUTHOR]

Summary: Experimental evidence that nonstomatal limitations to photosynthesis (NSLs) correlate with leaf sugar and/or leaf water status suggests the possibility that stomata adjust to maximise photosynthesis through a trade‐off between leaf CO2 supply and NSLs, potentially involving source–sink interactions. However, the mechanisms regulating NSLs and sink strength, as well as their implications for stomatal control, remain uncertain.We used an analytically solvable model to explore optimal stomatal control under alternative hypotheses for source and sink regulation. We assumed that either leaf sugar concentration or leaf water potential regulates NSLs, and that either phloem turgor pressure or phloem sugar concentration regulates sink phloem unloading.All hypotheses led to realistic stomatal responses to light, CO2 and air humidity, including conservative behaviour for the intercellular‐to‐atmospheric CO2 concentration ratio. Sugar‐regulated and water‐regulated NSLs are distinguished by the presence/absence of a stomatal closure response to changing sink strength. Turgor‐regulated and sugar‐regulated phloem unloading are distinguished by the presence/absence of stomatal closure under drought and avoidance/occurrence of negative phloem turgor. Results from girdling and drought experiments on Pinus sylvestris, Betula pendula, Populus tremula and Picea abies saplings are consistent with optimal stomatal control under sugar‐regulated NSLs and turgor‐regulated unloading.Our analytical results provide a simple representation of stomatal responses to above‐ground and below‐ground environmental factors and sink activity. [ABSTRACT FROM AUTHOR]

It has been suggested that a trade‐off between hydraulic efficiency and safety is related to drought adaptation across species. However, whether leaf hydraulic efficiency is sacrificed for safety during woody resprout regrowth after crown removal is not well understood. We measured leaf water potential (ψleaf) at predawn (ψpd) and midday (ψmid), leaf maximum hydraulic conductance (Kleaf‐max), ψleaf at induction 50% loss of Kleaf‐max (KleafP50), leaf area‐specific whole‐plant hydraulic conductance (LSC), leaf vein structure and turgor loss point (πtlp) in 1‐ to 13‐year‐old resprouts of the aridland shrub (Caragana korshinskii). ψpd was similar, ψmid and KleafP50 became more negative, and Kleaf–max decreased in resprouts with the increasing age; thus, leaf hydraulic efficiency clearly traded off against safety. The difference between ψmid and KleafP50, leaf hydraulic safety margin, increased gradually with increasing resprout age. More negative ψmid and KleafP50 were closely related to decreasing LSC and more negative πtlp, respectively, and the decreasing Kleaf‐max arose from the lower minor vein density and the narrower midrib xylem vessels. Our results showed that a clear trade‐off between leaf hydraulic efficiency and safety helps C. korshinskii resprouts adapt to increasing water stress as they approach final size. A trade‐off between leaf hydraulic efficiency and safety is proposed to be related to drought adaptation, but it is still under debate. Here, leaf hydraulic efficiency clearly traded off against safety in resprouts of Caragana korshinskii, thus helping resprouts adapt to increasing water stress. [ABSTRACT FROM AUTHOR]

Summary: Despite the importance of understanding plant growth, the mechanisms underlying how plant and fruit growth declines during drought remain poorly understood. Specifically, it remains unresolved whether carbon or water factors are responsible for limiting growth as drought progresses. We examine questions regarding the relative importance of water and carbon to fruit growth depending on the water deficit level and the fruit growth stage by measuring fruit diameter, leaf photosynthesis, and a proxy of cell turgor in olive (Olea europaea). Flow cytometry was also applied to determine the fruit cell division stage. We found that photosynthesis and turgor were related to fruit growth; specifically, the relative importance of photosynthesis was higher during periods of more intense cell division, while turgor had higher relative importance in periods where cell division comes close to ceasing and fruit growth is dependent mainly on cell expansion. This pattern was found regardless of the water deficit level, although turgor and growth ceased at more similar values of leaf water potential than photosynthesis. Cell division occurred even when fruit growth seemed to stop under water deficit conditions, which likely helped fruits to grow disproportionately when trees were hydrated again, compensating for periods with low turgor. As a result, the final fruit size was not severely penalized. We conclude that carbon and water processes are able to explain fruit growth, with importance placed on the combination of cell division and expansion. However, the major limitation to growth is turgor, which adds evidence to the sink limitation hypothesis. Significance Statement: The mechanism by which whole plants/organs reduce their growth during drought is still relatively little studied, the reduction of photosynthesis or turgor being the two major limitations considered. This paper aims at adding some evidence to the current debate on growth limitation thanks to a synchronous estimation of growth, turgor, and photosynthesis in olive trees (Olea europaea) exposed or not to water deficit using sensors and modeling with implications on process‐based models of forest and agriculture productivity. [ABSTRACT FROM AUTHOR]

Summary: Photosynthetic sensitivity to drought is a fundamental constraint on land‐plant evolution and ecosystem function. However, little is known about how the sensitivity of photosynthesis to nonstomatal limitations varies among species in the context of phylogenetic relationships.Using saplings of 10 Eucalyptus species, we measured maximum CO2‐saturated photosynthesis using A–ci curves at several different leaf water potentials (ψleaf) to quantify mesophyll photosynthetic sensitivity to ψleaf (MPS), a measure of how rapidly nonstomatal limitations to carbon uptake increase with declining ψleaf. MPS was compared to the macroclimatic moisture availability of the species' native habitats, while accounting for phylogenetic relationships.We found that species native to mesic habitats have greater MPS but higher maximum photosynthetic rates during non‐water‐stressed conditions, revealing a trade‐off between maximum photosynthesis and drought sensitivity. Species with lower turgor loss points have lower MPS, indicating coordination among photosynthetic and water‐relations traits.By accounting for phylogenetic relationships among closely related species, we provide the first compelling evidence that MPS in Eucalyptus evolved in an adaptive fashion with climatically determined moisture availability, opening the way for further study of this poorly explored dimension of plant adaptation to drought. [ABSTRACT FROM AUTHOR]

Opportunistic diversification has allowed ferns to radiate into epiphytic niches in angiosperm dominated landscapes. However, our understanding of how ecophysiological function allowed establishment in the canopy and the potential transitionary role of the hemi‐epiphytic life form remain unclear. Here, we surveyed 39 fern species in Costa Rican tropical forests to explore epiphytic trait divergence in a phylogenetic context. We examined leaf responses to water deficits in terrestrial, hemi‐epiphytic and epiphytic ferns and related these findings to functional traits that regulate leaf water status. Epiphytic ferns had reduced xylem area (−63%), shorter stipe lengths (−56%), thicker laminae (+41%) and reduced stomatal density (−46%) compared to terrestrial ferns. Epiphytic ferns exhibited similar turgor loss points, higher osmotic potential at saturation and lower tissue capacitance after turgor loss than terrestrial ferns. Overall, hemi‐epiphytic ferns exhibited traits that share characteristics of both terrestrial and epiphytic species. Our findings clearly demonstrate the prevalence of water conservatism in both epiphytic and hemi‐epiphytic ferns, via selection for anatomical and structural traits that avoid leaf water stress. Even with likely evolutionarily constrained physiological function, adaptations for drought avoidance have allowed epiphytic ferns to successfully endure the stresses of the canopy habitat. We detected key trait adaptions with xylem anatomy, frond structural traits and stomatal anatomy for avoidance of dehydration in both epiphytic and hemi‐epiphytic ferns species. Each of the trait adaptations we detected are clearly evident in phylogenetic trait reconstructions, suggesting that our findings provide a foundation on which to explain the evolution of epiphytism in ferns. [ABSTRACT FROM AUTHOR]

SUMMARY: Sexual reproduction in angiosperms is siphonogamous, and the interaction between pollen tube and pistil is critical for successful fertilization. Our previous study demonstrated that mutation of the Arabidopsis turgor regulation defect 1 (TOD1) gene leads to reduced male fertility, a result of retarded pollen tube growth in the pistil. TOD1 encodes a Golgi‐localized alkaline ceramidase, a key enzyme for the production of sphingosine‐1‐phosphate (S1P), which is involved in the regulation of turgor pressure in plant cells. However, whether TOD1s play a conserved role in the innovation of siphonogamy is largely unknown. In this study, we provide evidence that OsTOD1, which is similar to AtTOD1, is also preferentially expressed in rice pollen grains and pollen tubes. OsTOD1 knockout results in reduced pollen tube growth potential in rice pistil. Both the OsTOD1 genomic sequence with its own promoter and the coding sequence under the AtTOD1 promoter can partially rescue the attod1 mutant phenotype. Furthermore, TOD1s from other angiosperm species can partially rescue the attod1 mutant phenotype, while TOD1s from gymnosperm species are not able to complement the attod1 mutant phenotype. Our data suggest that TOD1 acts conservatively in angiosperms, and this opens up an opportunity to dissect the role of sphingolipids in pollen tube growth in angiosperms. [ABSTRACT FROM AUTHOR]

Why this research Matters: The inner bark plays important roles in tree stems, including radial exchange of water with the xylem and translocation of carbohydrates. Both processes affect the water content and the thickness of the inner bark on a diurnal basis. For the first time, we simultaneously measured the diurnal variations in the inner bark thickness of hinoki cypress (Chamaecyparis obtusa) by using point dendrometers and those of local xylem potential by using stem psychrometers located next to the dendrometers to determine how these variations were related to each other, to phloem turgor and carbohydrate transport. We also estimated the axial hydrostatic pressure gradient by measuring the osmolality of the sap extracted from the inner bark. The inner bark shrunk during the day and swelled during the night with an amplitude related to day‐to‐day and seasonal variations in climate. The relationship between changes in xylem water potential and inner bark thickness exhibited a hysteresis loop during the day with a median lag of 2 h. A phloem turgor‐related signal can be retrieved from the diurnal variations in the inner bark thickness, which was higher at the upper than at the lower position along the trunk. However, a downward hydrostatic pressure gradient was only observed at dawn, suggesting diurnal variations in the phloem sap flow velocity. [ABSTRACT FROM AUTHOR]

The process of leaf elongation in grasses is characterized by the creation and transformation of distinct cell zones. The prevailing turgor pressure within these cells is one of the key drivers for the rate at which these cells divide, expand and differentiate, processes that are heavily impacted by drought stress. In this article, a turgor‐driven growth model for grass leaf elongation is presented, which combines mechanistic growth from the basis of turgor pressure with the ontogeny of the leaf. Drought‐induced reductions in leaf turgor pressure result in a simultaneous inhibition of both cell expansion and differentiation, lowering elongation rate but increasing elongation duration due to the slower transitioning of cells from the dividing and elongating zone to mature cells. Leaf elongation is, therefore, governed by the magnitude of, and time spent under, growth‐enabling turgor pressure, a metric which we introduce as turgor‐time. Turgor‐time is able to normalize growth patterns in terms of varying water availability, similar to how thermal time is used to do so under varying temperatures. Moreover, additional inclusion of temperature dependencies within our model pioneers a novel concept enabling the general expression of growth regardless of water availability or temperature. A model for grass leaf elongation is presented, which combines mechanistic growth based on turgor pressure and leaf ontogeny. An inverse relation was found between the rate and duration of the elongation rate, allowing for uniform expression of growth in terms of a new concept called 'turgor‐time'. [ABSTRACT FROM AUTHOR]

Summary: The effects of climate change on tropical forests will depend on how diverse tropical tree species respond to drought. Current distributions of evergreen and deciduous tree species across local and regional moisture gradients reflect their ability to tolerate drought stress, and might be explained by functional traits.We measured leaf water potential at turgor loss (i.e. 'wilting point'; πtlp), wood density (WD) and leaf mass per area (LMA) on 50 of the most abundant tree species in central Panama. We then tested their ability to explain distributions of evergreen and deciduous species within a 50 ha plot on Barro Colorado Island and across a 70 km rainfall gradient spanning the Isthmus of Panama.Among evergreen trees, species with lower πtlp were associated with drier habitats, with πtlp explaining 28% and 32% of habitat association on local and regional scales, respectively, greatly exceeding the predictive power of WD and LMA. In contrast, πtlp did not predict habitat associations among deciduous species.Across spatial scales, πtlp is a useful indicator of habitat preference for tropical tree species that retain their leaves during periods of water stress, and holds the potential to predict vegetation responses to climate change. [ABSTRACT FROM AUTHOR]

Summary: Under prolonged drought and reduced photosynthesis, plants consume stored nonstructural carbohydrates (NSCs). Stored NSC depletion may impair the regulation of plant water balance, but the underlying mechanisms are poorly understood, and whether such mechanisms are independent of plant water deficit is not known. If so, carbon costs of fungal symbionts could indirectly influence plant drought tolerance through stored NSC depletion.We connected well‐watered Pinus ponderosa seedling pairs via ectomycorrhizal (EM) networks where one seedling was shaded (D) and the other kept illuminated (LD) and compared responses to seedling pairs in full light (L). We measured plant NSCs, osmotic and water potential, and transfer of 13CO2 through EM to explore mechanisms linking stored NSCs to plant water balance regulation and identify potential tradeoffs between plant water retention and EM fungi under carbon‐limiting conditions.NSCs decreased from L to LD to D seedlings. Even without drought, NSC depletion impaired osmoregulation and turgor maintenance, both of which are critical for drought tolerance. Importantly, EM networks propagated NSC depletion and its negative effects on water retention from carbon stressed to nonstressed hosts.We demonstrate that NSC storage depletion influences turgor maintenance independently of plant water deficit and reveal carbon allocation tradeoffs between supporting fungal symbionts and retaining water. [ABSTRACT FROM AUTHOR]

Summary: Although xylem embolism resistance is traditionally considered as static, we hypothesized that in grapevine (Vitis vinifera) leaf xylem becomes more embolism‐resistant over the growing season.We evaluated xylem architecture, turgor loss point (ΨTLP) and water potentials leading to 25% of maximal stomatal conductance (gs25) or 50% embolism in the leaf xylem (P50) in three irrigation treatments and at three time points during the growing season, while separating the effects of leaf age and time of season.Hydraulic traits acclimated over the growing season in a coordinated manner. Without irrigation, ΨTLP, gs25, and P50 decreased between late May and late August by 0.95, 0.77 and 0.71 MPa, respectively. A seasonal shift in P50 occurred even in mature leaves, while irrigation had only a mild effect (< 0.2 MPa) on P50. Vessel size and pit membrane thickness were also seasonally dynamic, providing a plausible explanation for the shift in P50.Our findings provide clear evidence that grapevines can modify their hydraulic traits along a growing season to allow lower xylem water potential, without compromising gas exchange, leaf turgor or xylem integrity. Seasonal changes should be considered when modeling ecosystem vulnerability to drought or comparing datasets acquired at different phenological stages. [ABSTRACT FROM AUTHOR]

Summary: In the stems of terrestrial vascular plants studied to date, the diameter of xylem water‐conducting conduits D widens predictably with distance from the stem tip L approximating D ∝ Lb, with b ≈ 0.2. Because conduit diameter is central for conductance, it is essential to understand the cause of this remarkably pervasive pattern. We give reason to suspect that tip‐to‐base conduit widening is an adaptation, favored by natural selection because widening helps minimize the increase in hydraulic resistance that would otherwise occur as an individual stem grows longer and conductive path length increases. Evidence consistent with adaptation includes optimality models that predict the 0.2 exponent. The fact that this prediction can be made with a simple model of a single capillary, omitting much biological detail, itself makes numerous important predictions, e.g. that pit resistance must scale isometrically with conduit resistance. The idea that tip‐to‐base conduit widening has a nonadaptive cause, with temperature, drought, or turgor limiting the conduit diameters that plants are able to produce, is less consistent with the data than an adaptive explanation. We identify empirical priorities for testing the cause of tip‐to‐base conduit widening and underscore the need to study plant hydraulic systems leaf to root as integrated wholes. [ABSTRACT FROM AUTHOR]

Background: The research on plant salt tolerance has mainly focused on Na+, but Cl− has been relatively neglected. Previous studies have found that the xerophyte Pugionium cornutum, an important forage grass in the arid and semi‐arid regions of northwestern China, could synergistically accumulate high quintiles of Na+ and Cl− in its shoots under NaCl treatments. However, the separate effects of these ions on the adaptation of P. cornutum to saline conditions have not been investigated. Aims: In this study, the response of P. cornutum to Na+ and Cl− was analyzed. Methods: Four‐week‐old seedlings were treated with additional 50 mM NaCl, Na+‐specific solution containing 50 mM Na+ with a mix of NO3-, H2PO4-, and SO42- as counter anions, and Cl−‐specific solution containing 50 mM Cl− with a mix of K+, Ca2+, and Mg2+ as counter cations. Results: Compared with the normal growth condition irrigated with Hoagland solution, the Na+‐specific solution severely impaired the growth and photosynthesis of P. cornutum due to the high accumulation of Na+ in shoots and the deterioration of tissue K+ homeostasis; while the Cl−‐specific solution significantly increased shoot fresh and dry biomass. The Cl−‐specific solution could also increase the turgor pressure in leaves for enhancing osmotic adjustment, which should be mainly attributed to the large accumulation of Cl−, since the concentrations of other ions, including K+, Mg2+, Ca2+, H2PO4-, and SO42-, in tissues under Cl−‐specific treatment were maintained at the same levels as those observed under the normal condition. Conclusions: P. cornutum displays an excellent tolerance to moderate Cl− but not to Na+, and the large accumulation of Cl− should play a positive role in stimulating the growth of P. cornutum under salt stress. [ABSTRACT FROM AUTHOR]

Premise: Whether drought‐adaptation mechanisms tend to evolve together, evolve independently, or evolve constrained by genetic architecture is incompletely resolved, particularly for water‐relations traits besides gas exchange. We addressed this issue in two subspecies of Clarkia xantiana (Onagraceae), California winter annuals that separated approximately 65,000 years ago and are adapted, partly by differences in flowering time, to native ranges differing in precipitation. Methods: In these subspecies and in recombinant inbred lines (RILs) from a cross between them, we scored traits related to drought adaptation (timing of seed germination and of flowering, succulence, pressure–volume curve variables) in common environments. Results: The subspecies native to more arid environments (parviflora) exhibited slower seed germination in saturated conditions, earlier flowering, and greater succulence, likely indicating superior drought avoidance, drought escape, and dehydration resistance via water storage. The other subspecies (xantiana) had lower osmotic potential at full turgor and lower water potential at turgor loss, implying superior dehydration tolerance. Genetic correlations among RILs suggest facilitated evolution of some trait combinations and independence of others. Where genetic correlations exist, subspecies differences fell along them, with the exception of differences in succulence and turgor loss point. In that case, subspecies difference overcame genetic correlations, possibly reflecting strong selection and/or antagonistic genetic correlations with other traits. Conclusions: Clarkia xantiana subspecies' differ in multiple mechanisms of drought adaptation. Genetic architecture generally does not seem to have constrained the evolution of these mechanisms, and it may have facilitated the evolution of some of trait combinations. [ABSTRACT FROM AUTHOR]