Hydraulic Safety Research Articles

Precipitation, solar radiation, and their interaction modify leaf hydraulic efficiency-safety trade-off across angiosperms at the global scale.

In theory, there is a trade-off between hydraulic efficiency and safety. However, the strength and direction of this trade-off at the leaf level are not consistent across studies, and habitat climate may impact this trade-off. We compiled a leaf hydraulic efficiency and safety dataset for 362 species from 81 sites world-wide, with 280 paired observations of both traits, and tested whether climate was associated with departure from the proposed trade-off. The leaf hydraulic efficiency-safety trade-off was weak (R2 = 0.144) at the global scale. Mean annual precipitation and solar radiation (SR) modified the trade-off. Species from dry and high SR habitats (e.g. desert and tropical savanna) were generally located above the trade-off line, indicating that these species tended to have higher leaf hydraulic safety and efficiency than species from wet habitats with low SR (e.g. subtropical monsoon forest and montane rainforest), which were located below the trade-off line. Leaves with high vein density, dry leaf mass per area, and osmotic regulation enhanced safety without compromising hydraulic efficiency. Variation in the hydraulic efficiency-safety trade-off at the leaf level likely facilitates plant survival in specific habitats and allows for a more nuanced view of leaf hydraulic adaption strategies at the global scale.

Intensive leaf cooling promotes tree survival during a record heatwave

Increasing heatwaves are threatening forest ecosystems globally. Leaf thermal regulation and tolerance are important for plant survival during heatwaves, though the interaction between these processes and water availability is unclear. Genotypes of the widely distributed foundation tree species Populus fremontii were studied in a controlled common garden during a record summer heatwave-where air temperature exceeded 48 °C. When water was not limiting, all genotypes cooled leaves 2 to 5 °C below air temperatures. Homeothermic cooling was disrupted for weeks following a 72-h reduction in soil water, resulting in leaf temperatures rising 3 °C above air temperature and 1.3 °C above leaf thresholds for physiological damage, despite the water stress having little effect on leaf water potentials. Tradeoffs between leaf thermal safety and hydraulic safety emerged but, regardless of water use strategy, all genotypes experienced significant leaf mortality following water stress. Genotypes from warmer climates showed greater leaf cooling and less leaf mortality after water stress in comparison with genotypes from cooler climates. These results illustrate how brief soil water limitation disrupts leaf thermal regulation and potentially compromises plant survival during extreme heatwaves, thus providing insight into future scenarios in which ecosystems will be challenged with extreme heat and unreliable soil water access.

A whole-plant perspective of hydraulic strategy in temperate desert shrub species.

Desert shrubs play a crucial role in controlling desertification and promoting revegetation, but drought often hinders their growth. Investigating the hydraulic strategies of desert shrubs is important to understand their drought adaptation and predict future dynamics under climate change. In this study, we measured the hydraulic-related characteristics of roots, stems, and leaves in 19 desert shrub species from northern China. We aimed to explore the hydraulic coordination and segmentation between different plant organs. The results were as follows (1) Specific root length (SRL) was positively correlated with the water potential inducing a 50% loss in stem hydraulic conductivity (P50stem) and negatively correlated with stem hydraulic safety margin (HSMstem). This suggested that water uptake efficiency of the fine roots was traded off with stem embolism resistance and hydraulic safety. (2) The water potential inducing a 50% loss in leaf hydraulic conductance (P50leaf) was significantly less negative than P50stem, and fine root turgor loss point (TLProot) was significantly less negative than P50stem, indicating a hydraulic segmentation between the main stem and terminal organs. (3) The most negative TLPleaf indicated that leaf wilting occurred after substantial leaf and stem embolism. The high desiccation resistance of the leaves may serve as an important physiological mechanism to increase carbon gain in a relatively brief growth period. In summary, this study elucidated the hydraulic strategies employed by desert shrubs from a whole-plant perspective.

Steady-state thermal–hydraulic analysis of an NTP reactor core based on the porous medium approach

Nuclear thermal propulsion (NTP) is a promising advanced technology which has attracted wide attention in recent years. The reactor core is an essential component of an NTP system and the corresponding thermal–hydraulic analysis is necessary. In this study, the porous medium approach was applied to the simulation of a two-pass NTP reactor core which consists of the porous prismatic cermet fuel elements. The thermodynamic property models of hydrogen and the fuel element materials were implemented, as well as the empirical correlations of the heat transfer coefficient and the friction factor. The three-dimensional simulation of a single fuel element was carried out and the results were compared against another code. The code-to-code comparison verified the applicability of the porous medium approach. The three-dimensional model of the two-pass NTP reactor core was established and the steady-state simulation was carried out. The distribution patterns of the parameters are determined by the thermal–hydraulic characteristics of the reactor core, including the nonuniform heat release, contact heat conduction and folded-flow scheme. The full-core heat-flow adaptability analysis is realized, which provides a reference for the thermal–hydraulic safety analysis of the NTP reactor.

The combination of high leaf hydraulic safety and water use efficiency allows alpine shrubs to adapt to high‐altitude habitats

Abstract Leaf hydraulic traits are considered the key determinants of gas exchange and therefore affect species distributions along environmental gradients, but the patterns of leaf hydraulic traits and their associations with gas exchange across altitudinal gradients remain largely unknown. Here, we measured leaf hydraulic traits, gas exchange, leaf anatomical traits and plant size traits in two dominant alpine shrubs (Caragana jubata and Salix gilashanica) across an altitudinal gradient from 3100 to 3700 m. The findings indicated that with increasing altitude, both shrub species exhibited an increase in leaf hydraulic safety (more negative Kleaf P50), a decrease in leaf hydraulic efficiency (Kleaf‐max) and an increase in intrinsic water use efficiency (WUEi), thus allowing them to adapt to higher altitude habitats. The more negative Kleaf P50 was associated with a greater ratio of major to minor vein density (VLAmaj/VLAmin), the lower Kleaf‐max was associated with a lower minor vein density (VLAmin) and greater increase in WUEi arose from the small decrease in the photosynthetic rate relative to the stomatal conductance. However, C. jubata was consistent with the ‘hydraulic safety strategy’ with a great decrease in Kleaf P50, a small decrease in Kleaf‐max and a small increase in WUEi along with decreasing plant height and leaf area with increasing altitude. Whereas S. gilashanica was consistent with the ‘photosynthetic efficiency strategy’ with a small decrease in Kleaf P50, a greater decrease in Kleaf‐max and a greater increase in WUEi along with an increasing plant height and unchanged leaf area with increasing altitude. Overall, these findings provide new insights to improve understanding how shift in leaf hydraulic traits and their associations with gas exchange and plant size allow plants to adapt to high‐altitude habitats. Read the free Plain Language Summary for this article on the Journal blog.

Higher Flower Hydraulic Safety, Drought Tolerance and Structural Resource Allocation Provide Drought Adaptation to Low Mean Annual Precipitation in Caragana Species.

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.

Hydraulic mechanism of limiting growth and maintaining survival of desert shrubs in arid habitats.

The growth and survival of woody plant species is mainly driven by evolutionary and environmental factors. However, little is known about the hydraulic mechanisms that respond to growth limitation and enable desert shrub survival in arid habitats. To shed light on these hydraulic mechanisms, 9-, 31-, and 56-year-old Caragana korshinskii plants that had been grown under different soil water conditions at the southeast edge of the Tengger Desert, Ningxia, China were used in this study. The growth of C. korshinskii was mainly limited by soil water rather than shrub age in non-watered habitats, which indicated the importance of maintaining shrub survival prior to growth under drought. Meanwhile, higher vessel density, narrower vessels and lower xylem hydraulic conductivity indicated that shrubs enhanced hydraulic safety and reduced their hydraulic efficiency in arid conditions. Importantly, xylem hydraulic conductivity mediated by variation in xylem hydraulic architecture regulated photosynthetic carbon assimilation and growth of C. korshinskii. Our study highlights that the synergistic variation in xylem hydraulic safety and hydraulic efficiency is the hydraulic mechanism limiting growth and maintaining survival of C. korshinskii under drought, providing insights into the strategies for growth and survival of desert shrubs in arid habitats.

Seasonality in embolism resistance and hydraulic capacitance jointly mediate hydraulic safety in branches and leaves of oriental cork oak (Quercus variabilis Bl.).

Seasonality in temperate regions is prominent during the era of increased climatic variability. A hydraulic trait that can adjust to seasonally changing climatic conditions is crucial for tree safety. However, little attention has been paid to the intraspecific seasonality of drought-related traits and hydraulic safety of keystone forest trees. We examined seasonal variations in the key morphological and physiological traits as well as multiple hydraulic safety margins (SMs) at the branch and leaf levels in oriental cork oak (Quercus variabilis Bl.), which is predominant in Chinese temperate forests. Pneumatic measurements indicated that, as seasons progressed, the water potential at which 50% of branch embolisms occur (P50_branch) decreased from -3.34 to -4.23MPa, with a coefficient of variation (CV) of 9.08%. Sapwood capacitance ranged from 48.19 to 248.08kgm-3MPa-1, peaking in autumn and reaching minimum in winter (CV 60.58%). Rehydration kinetics confirmed higher leaf embolism vulnerability (P50_leaf) in spring and autumn than those in summer, with values ranging from -1.06 to -3.02MPa (CV 39.85%). All leaf pressure-volume (PV) traits shifted with growth, with CVs ranging from 6.95% to 46.69%. Sapwood density had significant negative correlations with P50_branch and hydraulic capacitance for elastic water storage, whereas leaf mass per area was linearly associated with PV traits but not with P50_leaf. Furthermore, the branch typical SMs (difference between branch midday water potential and P50_branch) were consistently >1.84MPa, and vulnerability segmentation was prevalent throughout, implying a plausible hydraulic foundation for the dominance of Q. variabilis. Diverse hydraulic response patterns existed across seasons, leading to positive SMs mediated by the aforementioned physiological traits. Although Q. variabilis exhibits a high level of hydraulic safety, its susceptibility to sudden summer droughts may increase due to global climate change.

Divergent hydraulic and gas exchange strategies in two closely-related Salix species

Abstract Understanding the physiological mechanisms that trees employ to cope with drought-induced mortality is crucial for predicting their responses to a changing climate. Salix species exhibit distinct habit distributions, with Salix babylonica growing in wet habitats and S. matsudana growing in relatively dry habitats. The objective of this study was to compare hydraulic and gas exchange traits between these two closely-related Salix species with contrasting natural habitats. S. matsudana had lower photosynthesis (Amax), lower stomatal conductance (gs), and lower stem and leaf hydraulic conductance, but it exhibited higher water use efficiency (WUEi), higher hydraulic safety and wider leaf-to-stem vulnerability segmentation as well as narrower, shorter and denser conduits and a lower ratio of leaf area to sapwood area than S. babylonica. These findings suggest that variation in hydraulic vulnerability and gas exchange traits enable closely-related Salix species to adapt to different habitats.

Integrated Effects of Soil Moisture on Wheat Hydraulic Properties and Stomatal Regulation.

The development of water-saving management relies on understanding the physiological response of crops to soil drought. The coordinated regulation of hydraulics and stomatal conductance in plant water relations has steadily received attention. However, research focusing on grain crops, such as winter wheat, remains limited. In this study, three soil water supply treatments, including high (H), moderate (M), and low (L) soil water contents, were conducted with potted winter wheat. Leaf water potential (Ψleaf), leaf hydraulic conductance (Kleaf), and stomatal conductance (gs), as well as leaf biochemical parameters and stomatal traits were measured. Results showed that, compared to H, predawn leaf water potential (ΨPD) significantly reduced by 48.10% and 47.91%, midday leaf water potential (ΨMD) reduced by 40.71% and 43.20%, Kleaf reduced by 64.80% and 65.61%, and gs reduced by 21.20% and 43.41%, respectively, under M and L conditions. Although gs showed a significant difference between M and L, Ψleaf and Kleaf did not show significant differences between these treatments. The maximum carboxylation rate (Vcmax) and maximum electron transfer rate (Jmax) under L significantly decreased by 23.11% and 28.10%, stomatal density (SD) and stomatal pore area index (SPI) under L on the abaxial side increased by 59.80% and 52.30%, respectively, compared to H. The leaf water potential at 50% hydraulic conduction loss (P50) under L was not significantly reduced. The gs was positively correlated with ΨMD and Kleaf, but it was negatively correlated with abscisic acid (ABA) and SD. A threshold relationship between gs and Kleaf was observed, with rapid and linear reduction in gs occurring only when Kleaf fell below 8.70 mmol m-2 s-1 MPa-1. Our findings demonstrate that wheat leaves adapt stomatal regulation strategies from anisohydric to isohydric in response to reduced soil water content. These results enrich the theory of trade-offs between the carbon assimilation and hydraulic safety in crops and also provide a theoretical basis for water management practices based on stomatal regulation strategies under varying soil water conditions.

The rates of starch depletion and hydraulic failure both play a role in drought-induced seedling mortality

Key messageThe elapsed times to deplete starch concentrations and to reach a null hydraulic safety margin were related to tree seedling mortality under experimental drought. Starch concentration showed an accelerated decline across all species during the early stages of dehydration, while the concentrations of soluble sugars and total nonstructural carbohydrates remained stable. Concomitant carbohydrate depletion and hydraulic failure drive seedling mortality under drought.ContextCurrent upsurges of drought events are provoking impacts on tree physiology, resulting in forest mortality. Hydraulic dysfunction and nonstructural carbohydrate (NSC) depletion have been posited as the main mechanisms leading to plant mortality under drought.AimsThis study explores the dynamics of the two mortality-inducing processes during drought stress using an experimental approach with 12 evergreen tree species.MethodsSeedlings were subjected to drought until 100% mortality was observed. Midday (ΨMD) and predawn (ΨPD) water potentials, xylem pressure leading to a 50% loss of hydraulic conductivity (Ψ50), along with NSC concentrations in different organs (leaves, stems, and roots) were measured regularly during drought.ResultsTotal NSC concentrations and soluble sugar pools did not decline during drought. However, starch pools showed strong reductions early during drought stress as ΨPD decreased, and the time leading to starch depletion emerged as a strong mortality predictor. Ψ50 alone did not provide an accurate estimate of mortality, while the elapsed time to reach a null hydraulic safety margin (ΨMD—Ψ50 = 0) was related to seedling mortality.ConclusionAdopting a dynamic approach by estimating the times to consume both starch reserves and hydraulic safety margins is highly relevant to improve predictions of tree mortality under the current context of increasing global drought.

Limited effects of xylem anatomy on embolism resistance in cycad leaves.

Drought-induced xylem embolism is a primary cause of plant mortality. Although c. 70% of cycads are threatened by extinction and extant cycads diversified during a period of increasing aridification, the vulnerability of cycads to embolism spread has been overlooked. We quantified the vulnerability to drought-induced embolism, pressure-volume curves, in situ water potentials, and a suite of xylem anatomical traits of leaf pinnae and rachises for 20 cycad species. We tested whether anatomical traits were linked to hydraulic safety in cycads. Compared with other major vascular plant clades, cycads exhibited similar embolism resistance to angiosperms and pteridophytes but were more vulnerable to embolism than noncycad gymnosperms. All 20 cycads had both tracheids and vessels, the proportions of which were unrelated to embolism resistance. Only vessel pit membrane fraction was positively correlated to embolism resistance, contrary to angiosperms. Water potential at turgor loss was significantly correlated to embolism resistance among cycads. Our results show that cycads exhibit low resistance to xylem embolism and that xylem anatomical traits - particularly vessels - may influence embolism resistance together with tracheids. This study highlights the importance of understanding the mechanisms of drought resistance in evolutionarily unique and threatened lineages like the cycads.

Hydraulic plasticity and water use regulation act to maintain the hydraulic safety margins of Mediterranean trees in rainfall exclusion experiments.

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.

Taller trees exhibit greater hydraulic vulnerability in southern Amazonian forests

Potential increases in drought frequency and vapour pressure deficit pose a risk to the future function of tropical trees. Previous studies have found that taller tropical trees show a stronger increase in mortality than shorter trees in response to dry anomalies, but the mechanisms behind this are unclear. Here we investigate whether canopy branches of taller tropical trees have different hydraulic traits compared to shorter conspecifics. We determined xylem resistance to embolism (P50), hydraulic safety margin (HSM), xylem functional traits and xylem theoretical hydraulic conductivity for canopy branches of four tree species across a range of tree heights (sapling to maximum tree height) in an ecotonal forest near the Amazonia-Cerrado transition. We found that canopy branches of taller trees within each species have lower HSM, suggesting that they are more susceptible to hydraulic failure under drought than smaller conspecifics. Height-related trends in HSM were driven by variation in P50 with height and not by variation in leaf water potential which did not vary with height. We find that canopy branches with greater xylem vessel diameters are generally more vulnerable to embolism, suggesting a potential role for a diameter-safety linkage in explaining observed patterns of decreasing HSM with height. However, we find no evidence of a branch-level trade-off between theoretical hydraulic conductivity and hydraulic vulnerability. The greater hydraulic vulnerability of larger trees provides a potential explanation for the higher drought-induced mortality observed in taller tropical trees. The consistency of the height-P50 relationship across species opens the door to a more accurate prediction of southern Amazon forest responses to future droughts. Whether the findings for forests in southern Amazonia can be generalized to other Amazonian regions remains an open question.

Linkages between stem vulnerability curves and tree demography and their implications for plant physiological modeling.

Vulnerability curves (VCs) have been measured extensively to describe the differences in plant vulnerability to cavitation. Although the roles of hydraulic conductivity (Ks,max) and hydraulic safety (P50, embolism resistance), both of which are parameters of VCs ('sigmoidal' type), in tree demography have been evaluated across different forests, the direct linkages between VCs and tree demography are rarely explored. In this study, we combined measured VCs and plot data of 16 tree species in Panamanian seasonal tropical forests to investigate the connections between VCs and tree mortality, recruitment and growth. We found that the mortality and recruitment rates of evergreen species were most significantly positively correlated with P50. However, the mortality and recruitment rates of deciduous species only exhibited significant positive correlations with parameter a, which describes the steepness of VCs and indicates the sensitivity of conductivity loss with water potential decline, but is often neglected. These differences among evergreen and deciduous species may contribute to the poor performance of existing quantitative relationships (such as the fitting relationships for all 16 species) in capturing tree mortality and recruitment dynamics. Additionally, evergreen species presented a significant positive relationship between relative growth rate (RGR) and Ks,max, while deciduous species did not display such relationship. The RGR of both evergreen and deciduous species also displayed no significant correlations with P50 and a. Further analysis demonstrated that species with steeper VCs tended to have high mortality and recruitment rates, while species with flatter VCs were usually those with low mortality and recruitment rates. Our results highlight the important role of parameter a in tree demography, especially for deciduous species. Given that VC is a key component of plant hydraulic models, integrating measured VC rather than optimizing its parameters will help improve the ability to simulate and predict forest response to water availability.

Characteristics, Relationships, and Anatomical Basis of Leaf Hydraulic Traits and Economic Traits in Temperate Desert Shrub Species.

Shrubs are a key component of desert ecosystems, playing a crucial role in controlling desertification and promoting revegetation, yet their growth is often impeded by drought. Leaf hydraulic traits and economic traits are both involved in the process of water exchange for carbon dioxide. Exploring the characteristics, relationships, and anatomical basis of these two suites of traits is crucial to understanding the mechanism of desert shrubs adapting to the desert arid environment. However, the relationship between these two sets of traits currently remains ambiguous. This study explored the leaf hydraulic, economic, and anatomical traits of 19 desert shrub species. The key findings include the following: Relatively larger LT values and smaller SLA values were observed in desert shrubs, aligning with the "slow strategy" in the leaf economics spectrum. The relatively high P50leaf, low HSMleaf, negative TLPleaf, and positive HSMtlp values indicated that severe embolism occurs in the leaves during the dry season, while most species were able to maintain normal leaf expansion. This implies a "tolerance" leaf hydraulic strategy in response to arid stress. No significant relationship was observed between P50leaf and Kmax, indicating the absence of a trade-off between hydraulic efficiency and embolism resistance. Certain coupling relationships were observed between leaf hydraulic traits and economic traits, both of which were closely tied to anatomical structures. Out of all of the leaf traits, LT was the central trait of the leaf traits network. The positive correlation between C content and WPleaf and HSMleaf, as well as the positive correlation between N content and HSMtlp, suggested that the cost of leaf construction was synergistic with hydraulic safety. The negative correlation between SLA, P content, GCL, and SAI suggested a functional synergistic relationship between water use efficiency and gas exchange rate. In summary, this research revealed that the coupling relationship between leaf hydraulic traits and economic traits was one of the important physiological and ecological mechanisms of desert shrubs for adapting to desert habitats.

Xylem cell size regulation is a key adaptive response to water deficit in Eucalyptus grandis.

Future climatic scenarios forecast increasingly frequent droughts that will pose substantial consequences on tree mortality. In light of this, drought-tolerant eucalypts have been propagated; however, the severity of these conditions will invoke adaptive responses, impacting the commercially valuable wood properties. To determine what mechanisms govern the wood anatomical adaptive response, highly controlled drought experiments were conducted in Eucalyptus grandis W. Hill ex Maiden, with the tree physiology and transcriptome closely monitored. In response to water deficit, E. grandis displays an isohydric stomatal response to conserve water and enable stem growth to continue, albeit at a reduced rate. Maintaining gaseous exchange is likely a critical short-term response that drives the formation of hydraulically safer xylem. For instance, the development of significantly smaller fibers and vessels was found to increase cellular density, thereby promoting drought tolerance through improved functional redundancy, as well as implosion and cavitation resistance. The transcriptome was explored to identify the molecular mechanisms responsible for controlling xylem cell size during prolonged water deficit. Downregulation of genes associated with cell wall remodeling and the biosynthesis of cellulose, hemicellulose and pectin appeared to coincide with a reduction in cellular enlargement during drought. Furthermore, transcript levels of NAC and MYB transcription factors, vital for cell wall component biosynthesis, were reduced, while those linked to lignification increased. The upregulation of EgCAD and various peroxidases under water deficit did not correlate with an increased lignin composition. However, with the elevated cellular density, a higher lignin content per xylem cross-sectional area was observed, potentially enhancing hydraulic safety. These results support the requirement for higher density, drought-adapted wood as a long-term adaptive response in E. grandis, which is largely influenced by the isohydric stomatal response coupled with cellular expansion-related molecular processes.

Site-specific climate sensitivity of tree-ring width and vessel anatomical features of Juglans regia L. in Bhutan Himalaya

Climate change significantly impacts tree growth in the Himalayas, however, there is little information on how broadleaved tree species in Himalayan regions respond to climate change. Juglans regia L. (Common or Persian walnut) plays a vital economic and ecological role across the Himalayas, but little is known about how environmental changes influence the growth and wood anatomical traits of this species. In this study, we developed chronologies of tree-ring width (TRW) and six quantitative vessel anatomical features of Persian walnut trees from two sites in the Bhutan Himalayas. We evaluated how TRW and vessel anatomical features respond to climatic conditions and extreme climate events for the period 1960–2020. We found the climate sensitivity of tree growth differed at the two study sites, with the southwest-facing Dodeyna site (DOD) exhibiting higher moisture sensitivity than the north-facing Chimithangka site (CHI). The climate sensitivity of vessel traits was much stronger than that of tree-ring widths, particularly at the DOD. Hydraulic efficiency-related vessel traits (Mean vessel area, MVA; theoretical hydraulic conductivity, Kh; and hydraulic diameter, Dh) were positively sensitive to moisture availability, whereas the hydraulic safety-related vessel traits (vessel density, VD; and vessel grouping index, RVGI) were negatively sensitive to moisture availability at the DOD site. We further observed that MVA and Kh were significantly higher during extreme dry events at CHI site but displayed opposite directions at DOD site. Notably higher RVGI at the DOD site during dry years indicated an enhancement of hydraulic safety against drought. These findings highlight that the climate sensitivity of tree-ring width and vessel traits of walnuts were site-specific and mediated by aspects with the drier site (south-facing slope) being more sensitive. Whilst consideration of the aspects and site-specific climate sensitivity of tree species is crucial to developing robust forest conservation strategies at a larger spatial scale in the Himalayan region which is more vulnerable to climate change.

The double-edged sword of potassium and sodium fertilization in xylem embolism resistance of two Eucalyptus species under drought stress.

Sodium (Na+) is a beneficial element for most plants and may replace potassium (K+) in osmoregulatory process to a certain extent, increasing plant water use efficiency. Thus, understanding coordinated mechanisms underlying the combined use of K+ and Na+ in tree drought tolerance is a key challenge for forestry in dealing with productivity and water limitations. A pot experiment with three ratios of K/Na (K-supplied, partial K replacement by Na, and K-deficient plants) and two water regimes, well-watered (W+) and water-stressed (W-), was conducted on saplings of two Eucalyptus species with contrasting drought sensitivities. We evaluated the point of stomatal closure (Pgs90), xylem water potential at 12, 50, and 88% embolized xylem area (P12, P50, P88), hydraulic safety margin, leaf gas exchange (A, E, gs, and dark respiration), pre-dawn and midday leaf water potential (ΨPD and ΨMD), long-term water use efficiency (WUEL) and total dry mass. Partial K replacement by Na increased leaf gas exchange, WUEL, and total dry mass, while Pgs90, P12, P50, P88, and ΨMD decreased (were more negative), compared with plants exclusively supplied with K and K-deficient plants of both species. Fertilized plants had narrower hydraulic safety margins than K-deficient plants, indicating that these Eucalyptus species adopt the functional adaptive strategy of operating close to their hydraulic limits to maximize carbon uptake while increasing the risk of hydraulic failure under drought stress.

Lianas in tropical dry seasonal forests have a high hydraulic efficiency but not always a higher embolism resistance than lianas in rainforests.

Lianas have higher relative abundance and biomass in drier seasonal forests than in rainforests, but whether this difference is associated with their hydraulic strategies is unclear. Here, we investigate whether lianas of seasonally dry forests are safer and more efficient in water transport than rainforest lianas, explaining patterns of liana abundance. We measured hydraulic traits on five pairs of congeneric lianas of the tribe Bignonieae in two contrasting forest sites: the wet 'Dense Ombrophilous Forest' in Central Amazonia (~2 dry months) and the drier 'Semideciduous Seasonal Forest' in the inland Atlantic Forest (~6 dry months). We also gathered a broader database, including 197 trees and 58 liana species from different tropical forests, to compare hydraulic safety between habits and forest types. Bignonieae lianas from both forests had high and similar hydraulic efficiency but exhibited variability in resistance to embolism across forest types when phylogenetic relationships were taken into account. Three genera had higher hydraulic safety in the seasonal forest than in the rainforest, but species across both forests had similar positive hydraulic safety margins despite lower predawn water potential values of seasonal forest lianas. We did not find the safety-efficiency trade-off. Merging our results with previously published data revealed a high variability of resistance to embolism in both trees and lianas, independent of forest types. The high hydraulic efficiency of lianas detected here probably favours their rapid growth across tropical forests, but differences in hydraulic safety highlight that some species are highly vulnerable and may rely on other mechanisms to cope with drought. Future research on the lethal dehydration threshold and the connection between hydraulic resistance strategies and liana abundance could offer further insights into tropical forest dynamics under climatic threats.