Dehydration-induced corrugated folding in Rhapis excelsa plant leaves.

Plant leaves, whose remarkable ability for morphogenesis results in a wide range of petal and leaf shapes in response to environmental cues, have inspired scientific studies as well as the development of engineering structures and devices. Although some typical shape changes in plants and the driving force for such shape evolution have been extensively studied, there remain many poorly understood mechanisms, characteristics, and principles associated with the vast array of shape formation of plant leaves in nature. Here, we present a comprehensive study that combines experiment, theory, and numerical simulations of one such topic-the mechanics and mechanisms of corrugated leaf folding induced by differential shrinking in Rhapis excelsa. Through systematic measurements of the dehydration process in sectioned leaves, we identify a linear correlation between change in the leaf-folding angle and water loss. Building on experimental findings, we develop a generalized model that provides a scaling relationship for water loss in sectioned leaves. Furthermore, our study reveals that corrugated folding induced by dehydration in R. excelsa leaves is achieved by the deformation of a structural architecture-the "hinge" cells. Utilizing such connections among structure, morphology, environmental stimuli, and mechanics, we fabricate several biomimetic machines, including a humidity sensor and morphing devices capable of folding in response to dehydration. The mechanisms of corrugated folding in R. excelsa identified in this work provide a general understanding of the interactions between plant leaves and water. The actuation mechanisms identified in this study also provide insights into the rational design of soft machines.

Multi-trait selection for nutritional and physiological quality of cacao genotypes in irrigated and non-irrigated environments.

Water is a scarce, strategic resource and the most important input for economic development, especially in agricultural countries such as Brazil. Cocoa production is directly related to water availability, and, as climate changes, selecting drought-tolerant genotypes is vital to keep cacao crops sustainable. Here, we evaluated cacao genotypes under irrigated and water-stressed conditions and selected drought-tolerant ones based on nutritional and physiological traits. Thirty-nine genotypes were monitored for three years for agronomic traits and higher fruit yield. After this evaluation, the 18 most promising genotypes were evaluated in a randomized block design, under a 2 (with and without irrigation)  ×  18 (genotypes) factorial arrangement, with three replicates and five plants per plot. We evaluated seven physiological and 11 nutritional traits, selecting genotypes based on the Genotype-by-Trait Biplot approach. Significant effects (p < 0.05) were observed for the nutritional traits N, P, Mg, S, Zn, Cu, Mn and for the physiological traits CO2 assimilation rate (A), stomatal conductance (gs), transpiration (E), intercellular and atmospheric CO2 concentrations (Ci/Ca), intrinsic water use efficiency (A/gs), instantaneous water use efficiency (A/E), and instantaneous carboxylation efficiency (A/Ci), as determined by analysis of variance. The genotype  ×  irrigation treatment interaction was significant (p < 0.05) for the traits A, gs, and E. Genotypes CP 41, CP 43, and CCN 51 exhibited superior performance for both nutritional and physiological traits (A, gs, and E). In the irrigated environment, CP 41 showed superiority in traits such as P, A/E, A/gs, Mn, S, and Zn. Conversely, under non-irrigated conditions, CP 43 exhibited better performance in nutritional properties, specifically Mn, Mg, and Zn. Notably, in both irrigated and non-irrigated environments, CCN 51 excelled in key physiological traits, including A/Ci, A/E, and A/gs. This robust performance across diverse conditions suggests that these three genotypes possess physiological mechanisms to endure water-stressed conditions. Our research can generate valuable insights into these genotypes informing suitable choices for cocoa cultivation, especially in the context of global climate change.

Open field hardening improves leaf physiological drought tolerance in young plants of Sindora siamensis.

The effect of drought stress on leaf physiology was studied in 10-month-old plants of Sindora siamensis . Plants were either placed in an open greenhouse (unhardening; UH) or in an open field (open field hardening; H) for 45days. Both the UH and H plants stopped receiving water (D) until the initial drought injury and then rewatered (R) until complete recovery. Results showed necrosis in the leaves of UH+D, while H+D showed wilting at Day 7 after drought. A greater degree of necrosis was found in UH+D+R but made complete recovery in H+D+R at Day 4 after rewatering. Drought stress resulted in decreased leaf area in H, and reduced leaf and stem water status, PSII efficiency, net photosynthetic rate, stomatal conductance and transpiration rate in both UH and H. It also resulted in an increase in water use efficiency in both UH and H. Electrolyte leakage and malondialdehyde contents in UH were markedly increased due to drought stress. These results suggest that unhardened young plants of Sindora exposed to drought exhibited enhanced stomata behaviour by minimising open stomata and transpiration, resulting in high efficiency of water usage. However, there was still membrane damage from lipid peroxidation, which caused necrosis. Open field hardened plants exposed to drought demonstrated reduced open stomata and transpiration, thereby preserving leaf and soil water status and enhancing water use efficiency. This may be a reduction in lipid peroxidation though an oxidative scavenging mechanism that causes a slight alteration in membrane stability and a slight necrosis.

Potassium deficiency reduces grapevine transpiration through decreased leaf area and stomatal conductance.

Plants require potassium (K) to support growth and regulate hydraulics. Yet, K's effects on transpiration are still speculated. We hypothesized that K deficiency would limit grapevine water uptake by limiting canopy size and stomatal conductance (gs). Hence, we constructed large (2 m3) lysimeters and recorded vine transpiration for three years (2020-2022) under three fertilization application rates (8, 20, or 58 mg K L-1 in irrigation). Maximal K availability supported transpiration up to 75 L day-1, whereas K-deficient vines transpired only 60 L day-1 in midsummer. Limited vine growth and canopy size mainly accounted for reduced transpiration under low K conditions. Hence, considering K demand in addition to supply, we compared K deficiency effects on vines bearing 20 or 50 fruit clusters and found that reduced gs further limited transpiration when yields were high. Although fruits were strong K sinks, high yields did not alter K uptake because lower vegetative growth countered the additional K demands. Potassium deficiency leads to lower transpiration and productivity. Yet, internal mineral allocation compensates for fruit K uptake and masks biochemical indices or physiological proxies for K deficiency. Thus, decision support tools should integrate mineral availability, seasonal growth, and yield projections to determine grapevine water demands.

Root anatomical plasticity contributes to the different adaptive responses of two Phragmites species to water-deficit and low-oxygen conditions.

The runner reed (Phragmites japonica ) is the dominant species on riverbanks, whereas the common reed (Phragmites australis ) thrives in continuously flooded areas. Here, we aimed to identify the key root anatomical traits that determine the different adaptative responses of the two Phragmites species to water-deficit and low-oxygen conditions. Growth measurements revealed that P . japonica tolerated high osmotic conditions, whereas P . australis preferred low-oxygen conditions. Root anatomical analysis revealed that the ratios of the cortex to stele area and aerenchyma (gas space) to cortex area in both species increased under low-oxygen conditions. However, a higher ratio of cortex to stele area in P . australis resulted in a higher ratio of aerenchyma to stele, which includes xylem vessels that are essential for water and nutrient uptakes. In contrast, a lower ratio of cortex to stele area in P . japonica could be advantageous for efficient water uptake under high-osmotic conditions. In addition to the ratio of root tissue areas, rigid outer apoplastic barriers composed of a suberised exodermis may contribute to the adaptation of P . japonica and P . australis to water-deficit and low-oxygen conditions, respectively. Our results suggested that root anatomical plasticity is essential for plants to adapt and respond to different soil moisture levels.

Monitoring plant responses in field-grown peanuts exposed to exogenously applied chitosan under full and limited irrigation levels.

In recent decades, numerous studies have examined the effects of climate change on the responses of plants. These studies have primarily examined the effects of solitary stress on plants, neglecting the simultaneous effects of mixed stress, which are anticipated to transpire frequently as a result of the extreme climatic fluctuations. Therefore, this study investigated the impact of applied chitosan on boosting the resistance responses of peanuts to alkali and mixed drought-alkali stresses. Peanuts were grown in mid-alkaline soil and irrigated with full irrigation water requirements (100%IR), represented alkali condition (100% IR × alkali soil) and stress conditions (70% IR × alkali soil-represented mixed drought-alkali conditions). Additionally, the plants were either untreated or treated with foliar chitosan. The study evaluated various plant physio-chemical characteristics, including element contents (leaves and roots), seed yield, and irrigation water use efficiency (IWUE). Plants that experienced solitary alkali stress were found to be more vulnerable. However, chitosan applications were effective for reducing (soil pH and sodium absorption), alongside promoting examined physio-chemical measurements, yield traits, and IWUE. Importantly, when chitosan was applied under alkali conditions, the accumulations of (phosphorus, calcium, iron, manganese, zinc, and copper) in leaves and roots were maximized. Under mixed drought-alkali stresses, the results revealed a reduction in yield, reaching about 5.1 and 5.8% lower than under (100% IR × alkali), in the first and second seasons, respectively. Interestingly, treated plants under mixed drought-alkali stresses with chitosan recorded highest values of relative water content, proline, yield, IWUE, and nutrient uptake of (nitrogen, potassium, and magnesium) as well as the lowest sodium content in leaves and roots. Enhances the accumulation of (N, K, and Mg) instead of (phosphorus, calcium, iron, manganese, zinc, and copper) was the primary plant response to chitosan applications, which averted severe damage caused by mixed drought-alkali conditions, over time. These findings provide a framework of the nutrient homeostasis changes induced by chitosan under mixed stresses. Based on the findings, it is recommended under mixed drought-alkali conditions to treat plants with chitosan. This approach offers a promising perspective for achieving optimal yield with reduced water usage.

Finding balance: Tree-ring isotopes differentiate between acclimation and stress-induced imbalance in a long-term irrigation experiment.

Scots pine (Pinus sylvestris L.) is a common European tree species, and understanding its acclimation to the rapidly changing climate through physiological, biochemical or structural adjustments is vital for predicting future growth. We investigated a long-term irrigation experiment at a naturally dry forest in Switzerland, comparing Scots pine trees that have been continuously irrigated for 17 years (irrigated) with those for which irrigation was interrupted after 10 years (stop) and non-irrigated trees (control), using tree growth, xylogenesis, wood anatomy, and carbon, oxygen and hydrogen stable isotope measurements in the water, sugars and cellulose of plant tissues. The dendrochronological analyses highlighted three distinct acclimation phases to the treatments: irrigated trees experienced (i) a significant growth increase in the first 4 years of treatment, (ii) high growth rates but with a declining trend in the following 8 years and finally (iii) a regression to pre-irrigation growth rates, suggesting the development of a new growth limitation (i.e. acclimation). The introduction of the stop treatment resulted in further growth reductions to below-control levels during the third phase. Irrigated trees showed longer growth periods and lower tree-ring δ13 C values, reflecting lower stomatal restrictions than control trees. Their strong tree-ring δ18 O and δ2 H (O-H) relationship reflected the hydrological signature similarly to the control. On the contrary, the stop trees had lower growth rates, conservative wood anatomical traits, and a weak O-H relationship, indicating a physiological imbalance. Tree vitality (identified by crown transparency) significantly modulated growth, wood anatomical traits and tree-ring δ13 C, with low-vitality trees of all treatments performing similarly regardless of water availability. We thus provide quantitative indicators for assessing physiological imbalance and tree acclimation after environmental stresses. We also show that tree vitality is crucial in shaping such responses. These findings are fundamental for the early assessment of ecosystem imbalances and decline under climate change.

Photosynthesis of rice leaves with a parallel venation is highly tolerant to vein severing.

Vein severing in plants caused by leaf damage is common in fields where crops are cultivated. It is hypothesized that leaves with complex reticulate venation can withstand hydraulic disturbances caused by vein severing, thereby preserving leaf carbon assimilation. However, limited research focuses on vein damage of leaves with parallel venation. We studied how vein-severing affected the photosynthetic traits of rice (Oryza sativa) leaves in seconds, minutes and days, under varying water-demand conditions and differing extents of water supply disruption. Rice leaves completely lost their photosynthetic capacity within 2.5 minutes after excision. Severing the midrib resulted in reduced light-saturated photosynthetic rate (A), stomatal conductance (gsw ) and transpiration rate (E) by 2.6, 6.8 and 5.9%, respectively, already after thirty minutes. We further investigated the photosynthetic trait responses to various extents of leaf width severing, while keeping the midrib functional. Surprisingly, A, gsw and E in the downstream area of the severed leaves largely remained stable, showing minimal variation across different leaf width severing ratios. These traits declined only slightly even under increased ambient light intensity and leaf-to-air vapor pressure deficit. This sustained photosynthesis post-severing is attributed to the efficient lateral water transport. Long-term leaf damage slightly but not significantly, impacted the downstream photosynthetic traits within five days post-severing. However, a more pronounced reduction in gas exchange during leaf senescence was observed nine days after severing. These findings suggested that rice leaves can tolerate hydraulic disturbances from vein severing and maintain functionality under various conditions, which is crucial for crop yield stability. However, long-term consequences require further investigation.

Spectral ecophysiology: hyperspectral pressure-volume curves to estimate leaf turgor loss.

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.

New approaches to dissect leaf hydraulics reveal large gradients in living tissues of tomato leaves.

The water status of the living tissue in leaves is critical in determining plant function and global exchange of water and CO2. Despite significant advances in the past two decades, persistent questions remain about the tissue-specific origins of leaf hydraulic properties and their dependence on water status. We use a fluorescent nanoparticle reporter that provides water potential in the mesophyll apoplast adjacent to the epidermis of intact leaves to complement existing methods based on the Scholander Pressure Chamber (SPC). Working in tomato leaves, this approach provides access to the hydraulic conductance of the whole leaf, xylem, and outside-xylem tissues. These measurements show that, as stem water potential decreases, the water potential in the mesophyll apoplast can drop below that assessed with the SPC and can fall significantly below the turgor loss point of the leaf. We find that this drop in potential, dominated by the large loss (10-fold) of hydraulic conductance of the outside-xylem tissue, is not however strong enough to significantly limit transpiration. These observations highlight the need to reassess models of water transfer through the outside-xylem tissues, the potential importance of this tissue in regulating transpiration, and the power of new approaches for probing leaf hydraulics.

Integrative regulatory mechanisms of stomatal movements under changing climate.

Global climate change-caused drought stress, high temperatures and other extreme weather profoundly impact plant growth and development, restricting sustainable crop production. To cope with various environmental stimuli, plants can optimize the opening and closing of stomata to balance CO2 uptake for photosynthesis and water loss from leaves. Guard cells perceive and integrate various signals to adjust stomatal pores through turgor pressure regulation. Molecular mechanisms and signaling networks underlying the stomatal movements in response to environmental stresses have been extensively studied and elucidated. This review focuses on the molecular mechanisms of stomatal movements mediated by abscisic acid, light, CO2 , reactive oxygen species, pathogens, temperature, and other phytohormones. We discussed the significance of elucidating the integrative mechanisms that regulate stomatal movements in helping design smart crops with enhanced water use efficiency and resilience in a climate-changing world.

Stem heating results in hydraulic dysfunction in Symplocos tinctoria: implications for post-fire tree death.

Fire-induced heating of stems can impair plant water transport by deforming xylem and increasing vulnerability to cavitation, but it is not clear whether these effects can result in tree death, or how quickly this may occur. In field experiments, we heated stems of Symplocos tinctoria (L.) L'Hér saplings to 90 °C using a thin-film resistive heater, and we monitored stomatal conductance, leaf water potential, sap flow and hydraulic conductivity until stem death. Sap flow and stomatal conductance declined quickly after heating, while whole-plant hydraulic conductance and leaf water potential remained high for the first week. In fact, leaf water potential increased during the first days after heating, indicating that stomatal closure was not initially caused by leaf water deficit induced by impaired water transport. After 1 week, leaf water potential and whole-plant conductance declined below unheated controls, while stomatal conductance and sap flow continued declining, approaching zero after 2 weeks. To better understand the cause of these declines, we directly measured hydraulic conductivity of heated stems. Stems underwent a progressive decline in conductivity after heating, and by the time that samples were severely wilted or desiccated, the heated portion of stems had little or no conductivity. Importantly, conductivity of heated stems was not recovered by flushing stems to remove embolisms, suggesting the existence of physical occlusions. Scanning electron micrographs did not reveal deformed cell walls, nor did it identify alternative causes of blockages. These results reveal that stem heating can result in xylem dysfunction and mortality, but neither response is immediate. Dysfunction was likely caused by wound responses rather than embolism, but improved understanding of the mechanisms of heat-induced hydraulic failure is needed.

Soil warming affects sap flow and stomatal gas exchange through altering functional traits in a subtropical forest.

Climate warming influences the structure and function of ecosystems. However, the mechanisms of plant water use and gas exchange responses to climate warming have been less studied, especially from the perspective of different functional traits. We conducted a field experiment to investigate how soil warming (+2 °C) affects sap flow and stomatal gas exchange through plant functional traits and nutrient characteristics in a subtropical forest. We measured stomatal gas exchange of trees (Acacia auriculiformis and Schima superba) and shrubs (Castanea henryi and Psychotria asiatica), and monitored long-term sap flow of both tree species. Besides, plant leaf nutrient contents, functional traits, and soil nutrients were also studied. It is demonstrated that soil warming significantly increased maximum sap flow density (Js_max, 35.1 %) and whole-tree transpiration (EL, 46.0 %) of A. auriculiformis, but decreased those of S. superba (15.6 % and 14.9 %, respectively). Warming increased the photosynthetic rate of P. asiatica (18.0 %) and water use efficiency of S. superba (47.2 %). Leaf nutrients and stomatal anatomical characteristics of shrubs were less affected by soil warming. Soil warming increased (+42.7 %) leaf K content of A. auriculiformis in dry season. Decomposition of soil total carbon, total nitrogen, and available nitrogen was accelerated under soil warming, and soil exchangeable Ca2+ and Mg2+ were decreased. Trees changed stomatal and anatomic traits to adapt to soil warming, while shrubs altered leaf water content and specific leaf area under soil warming. Warming had a greater effect on sap flow of trees, as well as on their leaf gas exchange (total effect: -0.27) than on that of shrubs (total effect: 0.06). In summary, our results suggest that the combination of functional and nutrient traits can help to better understand plant water use and gas exchange responses under climate warming.

Uncoupling of stomatal conductance and photosynthesis at high temperatures: mechanistic insights from online stable isotope techniques.

The strong covariation of temperature and vapour pressure deficit (VPD) in nature limits our understanding of the direct effects of temperature on leaf gas exchange. Stable isotopes in CO2 and H2 O vapour provide mechanistic insight into physiological and biochemical processes during leaf gas exchange. We conducted combined leaf gas exchange and online isotope discrimination measurements on four common European tree species across a leaf temperature range of 5-40°C, while maintaining a constant leaf-to-air VPD (0.8 kPa) without soil water limitation. Above the optimum temperature for photosynthesis (30°C) under the controlled environmental conditions, stomatal conductance (gs ) and net photosynthesis rate (An ) decoupled across all tested species, with gs increasing but An decreasing. During this decoupling, mesophyll conductance (cell wall, plasma membrane and chloroplast membrane conductance) consistently and significantly decreased among species; however, this reduction did not lead to reductions in CO2 concentration at the chloroplast surface and stroma. We question the conventional understanding that diffusional limitations of CO2 contribute to the reduction in photosynthesis at high temperatures. We suggest that stomata and mesophyll membranes could work strategically to facilitate transpiration cooling and CO2 supply, thus alleviating heat stress on leaf photosynthetic function, albeit at the cost of reduced water-use efficiency.

Mechanistic links between physiology and spectral reflectance enable previsual detection of oak wilt and drought stress.

Tree mortality due to global change-including range expansion of invasive pests and pathogens-is a paramount threat to forest ecosystems. Oak forests are among the most prevalent and valuable ecosystems both ecologically and economically in the United States. There is increasing interest in monitoring oak decline and death due to both drought and the oak wilt pathogen (Bretziella fagacearum). We combined anatomical and ecophysiological measurements with spectroscopy at leaf, canopy, and airborne levels to enable differentiation of oak wilt and drought, and detection prior to visible symptom appearance. We performed an outdoor potted experiment with Quercus rubra saplings subjected to drought stress and/or artificially inoculated with the pathogen. Models developed from spectral reflectance accurately predicted ecophysiological indicators of oak wilt and drought decline in both potted and field experiments with naturally grown saplings. Both oak wilt and drought resulted in blocked water transport through xylem conduits. However, oak wilt impaired conduits in localized regions of the xylem due to formation of tyloses instead of emboli. The localized tylose formation resulted in more variable canopy photosynthesis and water content in diseased trees than drought-stressed ones. Reflectance signatures of plant photosynthesis, water content, and cellular damage detected oak wilt and drought 12 d before visual symptoms appeared. Our results show that leaf spectral reflectance models predict ecophysiological processes relevant to detection and differentiation of disease and drought. Coupling spectral models that detect physiological change with spatial information enhances capacity to differentiate plant stress types such as oak wilt and drought.

Unravelling the neglected role of ultraviolet radiation on stomata: A meta-analysis with implications for modelling ecosystem-climate interactions.

Stomata play a pivotal role in regulating gas exchange between plants and the atmosphere controlling water and carbon cycles. Accordingly, we investigated the impact of ultraviolet-B radiation, a neglected environmental factor varying with ongoing global change, on stomatal morphology and function by a Comprehensive Meta-Analysis. The overall UV effect at the leaf level is to decrease stomatal conductance, stomatal aperture and stomatal size, although stomatal density was increased. The significant decline in stomatal conductance is marked (6% in trees and >10% in grasses and herbs) in short-term experiments, with more modest decreases noted in long-term UV studies. Short-term experiments in growth chambers are not representative of long-term field UV effects on stomatal conductance. Important consequences of altered stomatal function are hypothesized. In the short term, UV-mediated stomatal closure may reduce carbon uptake but also water loss through transpiration, thereby alleviating deleterious effects of drought. However, in the long term, complex changes in stomatal aperture, size, and density may reduce the carbon sequestration capacity of plants and increase vegetation and land surface temperatures, potentially exacerbating negative effects of drought and/or heatwaves. Therefore, the expected future strength of carbon sink capacity in high-UV regions is likely overestimated.

Restricted internal diffusion weakens transpiration-photosynthesis coupling during heatwaves: Evidence from leaf carbonyl sulphide exchange.

Increasingly frequent and intense heatwaves threaten ecosystem health in a warming climate. However, plant responses to heatwaves are poorly understood. A key uncertainty concerns the intensification of transpiration when heatwaves suppress photosynthesis, known as transpiration-photosynthesis decoupling. Field observations of such decoupling are scarce, and the underlying physiological mechanisms remain elusive. Here, we use carbonyl sulphide (COS) as a leaf gas exchange tracer to examine potential mechanisms leading to transpiration-photosynthesis decoupling on a coast live oak in a southern California woodland in spring 2013. We found that heatwaves suppressed both photosynthesis and leaf COS uptake but increased transpiration or sustained it at non-heatwave levels throughout the day. Despite statistically significant decoupling between transpiration and photosynthesis, stomatal sensitivity to environmental factors did not change during heatwaves. Instead, midday photosynthesis during heatwaves was restricted by internal diffusion, as indicated by the lower internal conductance to COS. Thus, increased evaporative demand and nonstomatal limitation to photosynthesis act jointly to decouple transpiration from photosynthesis without altering stomatal sensitivity. Decoupling offered limited potential cooling benefits, questioning its effectiveness for leaf thermoregulation in xeric ecosystems. We suggest that adding COS to leaf and ecosystem flux measurements helps elucidate diverse physiological mechanisms underlying transpiration-photosynthesis decoupling.

Water loss after stomatal closure: quantifying leaf minimum conductance and minimal water use in nine temperate European tree species during a severe drought.

Residual canopy transpiration (Emin_canop) is a key physiological trait that determines trees' survival time under drought after stomatal closure and after trees have limited access to soil water. Emin_canop mainly depends on leaf minimum conductance (gmin) and vapor pressure deficit. Here we determined the seasonal variation of gmin and how gmin is related to interspecies variation in leaf cuticular and stomatal traits for nine European tree species in a mature forest. In addition, we determined the species-specific temperature responses of gmin. With this newly obtained insight, we calculated Emin_canop for the nine species for one day at our research site during the 2022 central European hot drought. Our results show that at ambient temperatures gmin ranged from 0.8 to 4.8 mmol m-2 s-1 across the nine species and was stable in most species throughout the growing season. The interspecies variation of gmin was associated with leaf cuticular and stomatal traits. Additionally, gmin exhibited strong temperature responses and increased, depending on species, by a factor of two to four in the range of 25-50 °C. For the studied species at the site, during a single hot drought day, Emin_canop standardized by tree size (stem basal area) ranged from 2.0 to 36.7 L m-2, and non-standardized Emin_canop for adult trees ranged from 0.3 to 5.3 L. Emin_canop also exhibited species-specific rapid increases under hotter temperatures. Our results suggest that trees, depending on species, need reasonable amounts of water during a drought, even when stomates are fully closed. Species differences in gmin and ultimately Emin_canop can, together with other traits, affect the ability of a tree to keep its tissue hydrated during a drought and is likely to contribute to species-specific differences in drought vulnerability.

Grafting seedling rootstock strengthens tolerance to drought stress in polyploid mulberry (Morus alba L.).

The economically adaptable mulberry (Morus alba L.) has a long history of grafting in China, yet the physiological mechanisms and advantages in drought tolerance remain unexplored. In our study, we investigated the responses of self-rooted 2X (diploid), 3X (triploid), and 4X (tetraploid) plants, as well as polyploid plants grafted onto diploid seedling rootstocks (2X/2X, 3X/2X, and 4X/2X) under drought stress. We found that self-rooted diploid plants exhibited the most severe phenotypic damage, lowest water retention, photosynthetic capacity, and the least effective osmotic stress adjustment compared to tetraploid and triploid plants. However, grafted diploid and triploid plants showed effective mitigation of drought-induced damage, with higher relative water content and improved soil water retention. Grafted plants also improved the photosystem response to drought stress through elevated photosynthetic potential, closed stomatal aperture, and faster recovery of chlorophyll biosynthesis in the leaves. Additionally, grafted plants altered osmotic protective compound levels, including starch, soluble sugar, and proline content, thereby enhancing drought resistance. Absolute quantification PCR indicated that the expression levels of proline synthesis-related genes in grafted plants were not influenced after drought stress, whereas they were significantly increased in self-rooted plants. Consequently, our findings support that self-rooted triploid and tetraploid mulberries exhibited superior drought resistance compared to diploid plants. Moreover, grafting onto seedling rootstocks enhanced tolerance against drought stress in diploid and triploid mulberry, but not in tetraploid. Our study provides valuable insights for a comprehensive analysis of physiological effects in response to drought stress between stem-roots and seedling rootstocks.

A trade-off between leaf hydraulic efficiency and safety across three xerophytic species in response to increased rock fragment content.

Limited information is available on the variation of plant leaf hydraulic traits in relation to soil rock fragment content (RFC), particularly for xerophytes native to rocky mountain areas. In this study, we conducted a field experiment with four gradients of RFC (0, 25, 50 and 75% ν ν-1) on three different xerophytic species (Sophora davidii, Cotinus szechuanensis and Bauhinia brachycarpa). We measured predawn and midday leaf water potential (Ψleaf), leaf hydraulic conductance (Kleaf), Ψleaf induced 50% loss of Kleaf (P50), pressure-volume curve traits and leaf structure. A consistent response of hydraulic traits to increased RFC was observed in three species. Kleaf showed a decrease, whereas P50 and turgor loss point (Ψtlp) became increasingly negative with increasing RFC. Thus, a clear trade-off between hydraulic efficiency and safety was observed in the xerophytic species. In all three species, the reduction in Kleaf was associated with an increase in leaf mass per area. In S. davidii, alterations in Kleaf and P50 were driven by leaf vein density (VLA) and Ψtlp. In C. szechuanensis, Ψtlp and VLA drove the changes in Kleaf and P50, respectively. In B. brachycarpa, changes in P50 were driven by VLA, whereas changes in both Kleaf and P50 were simultaneously influenced by Ψtlp. Our findings suggest that adaptation to increased rockiness necessarily implies a trade-off between leaf hydraulic efficiency and safety in xerophytic species. Additionally, the trade-off between leaf hydraulic efficiency and safety among xerophytic species is likely to result from processes occurring in the xylem and the outside-xylem hydraulic pathways. These findings contribute to a better understanding of the survival strategies and mechanisms of xerophytes in rocky soils, and provide a theoretical basis for the persistence of xerophytic species in areas with stony substrates.