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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 (P gs90), xylem water potential at 12, 50, and 88% embolized xylem area (P 12, P 50, P 88), hydraulic safety margin, leaf gas exchange (A , E , g s, 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 P gs90, P 12, P 50, P 88, 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. [ABSTRACT FROM AUTHOR]

Increasing tree diversity is considered a key management option to adapt forests to climate change. However, the effect of species diversity on a forest's ability to cope with extreme drought remains elusive. In this study, we assessed drought tolerance (xylem vulnerability to cavitation) and water stress (water potential), and combined them into a metric of drought–mortality risk (hydraulic safety margin) during extreme 2021 or 2022 summer droughts in five European tree diversity experiments encompassing different biomes. Overall, we found that drought–mortality risk was primarily driven by species identity (56.7% of the total variability), while tree diversity had a much lower effect (8% of the total variability). This result remained valid at the local scale (i.e within experiment) and across the studied European biomes. Tree diversity effect on drought–mortality risk was mediated by changes in water stress intensity, not by changes in xylem vulnerability to cavitation. Significant diversity effects were observed in all experiments, but those effects often varied from positive to negative across mixtures for a given species. Indeed, we found that the composition of the mixtures (i.e., the identities of the species mixed), but not the species richness of the mixture per se, is a driver of tree drought–mortality risk. This calls for a better understanding of the underlying mechanisms before tree diversity can be considered an operational adaption tool to extreme drought. Forest diversification should be considered jointly with management strategies focussed on favouring drought‐tolerant species. [ABSTRACT FROM AUTHOR]

Knowledge of the physiological mechanisms underlying species vulnerability to drought is critical for better understanding patterns of tree mortality. Investigating plant adaptive strategies to drought should thus help to fill this knowledge gap, especially in tropical rainforests exhibiting high functional diversity. In a semi-controlled drought experiment using 12 rainforest tree species, we investigated the diversity in hydraulic strategies and whether they determined the ability of saplings to use stored non-structural carbohydrates during an extreme imposed drought. We further explored the importance of water- and carbon-use strategies in relation to drought survival through a modelling approach. Hydraulic strategies varied considerably across species with a continuum between dehydration tolerance and avoidance. During dehydration leading to hydraulic failure and irrespective of hydraulic strategies, species showed strong declines in whole-plant starch concentrations and maintenance, or even increases in soluble sugar concentrations, potentially favouring osmotic adjustments. Residual water losses mediated the trade-off between time to hydraulic failure and growth, indicating that dehydration avoidance is an effective drought-survival strategy linked to the 'fast–slow' continuum of plant performance at the sapling stage. Further investigations on residual water losses may be key to understanding the response of tropical rainforest tree communities to climate change. [ABSTRACT FROM AUTHOR]

Plant hydraulic traits related to leaf drought tolerance, like the water potential at turgor loss point (TLP) and the water potential inducing 50% loss of hydraulic conductance (P 50), are extremely useful to predict the potential impacts of drought on plants. While novel techniques have allowed the inclusion of TLP in studies targeting a large group of species, fast and reliable protocols to measure leaf P 50 are still lacking. Recently, the optical method coupled with the gas injection (GI) technique has been proposed as a possibility to speed up the P 50 estimation. Here, we present a comparison of leaf optical vulnerability curves (OVcs) measured in three woody species, namely Acer campestre (Ac), Ostrya carpinifolia (Oc) and Populus nigra (Pn), based on bench dehydration (BD) or GI of detached branches. For Pn, we also compared optical data with direct micro-computed tomography (micro-CT) imaging in both intact saplings and cut shoots subjected to BD. Based on the BD procedure, Ac, Oc and Pn had P 50 values of −2.87, −2.47 and −2.11 MPa, respectively, while the GI procedure overestimated the leaf vulnerability (−2.68, −2.04 and −1.54 MPa for Ac, Oc and Pn, respectively). The overestimation was higher for Oc and Pn than for Ac, likely reflecting the species-specific vessel lengths. According to micro-CT observations performed on Pn, the leaf midrib showed none or very few embolized conduits at −1.2 MPa, consistent with the OVcs obtained with the BD procedure but at odds with that derived on the basis of GI. Overall, our data suggest that coupling the optical method with GI might not be a reliable technique to quantify leaf hydraulic vulnerability since it could be affected by the 'open-vessel' artifact. Accurate detection of xylem embolism in the leaf vein network should be based on BD, preferably of intact up-rooted plants. [ABSTRACT FROM AUTHOR]

Hydraulic traits are essential functional characteristics of plants related to water absorption, transport, and loss, serving as indicators of a plant's adaptability to prevailing environmental water conditions. However, the hydraulic traits of shrub, particularly desert plants in arid and semi-arid regions, have been underexplored. In this study, we conducted a pot experiment using Caroxylon passerinum (Bunge) Akhani and Roalson as the subject. Three treatment groups were established: adequate water supply, mild drought, and severe drought. After subjecting the shrub to drought and subsequent rehydration, we measured hydraulic conductivity, net photosynthetic rate, and stomatal conductance. We found that leaf water potential decreased and stomatal conductance and net photosynthesis decreased with increasing drought intensity. We found that leaf water potential and stomatal conductance decreased with increasing drought intensity. Although there was no significant change in hydraulic conductivity in the two drought groups, the values were greater in the drought group than in the control, and greater in the mild drought group than in the severe drought group. Meanwhile, the embolism resistance decreased with increasing drought intensity. After rehydration, hydraulic conductivity did not return to control levels in the severe drought group, as did embolism resistance in the two drought groups, and leaf water potential did not recover significantly. The results showed that drought stress increased the hydraulic conductivity of C. passerinum, and this effect was more pronounced under mild drought stress. After the stress was lifted, C. passerinum continued to maintain a lower leaf water potential to promote water uptake. This result provides a reference for us to study water use of desert shrubs under different drought stresses. [ABSTRACT FROM AUTHOR]

Quantitative knowledge of xylem physical tolerance limits to dehydration is essential to understanding plant drought tolerance but is lacking in many long-vessel angiosperms. We examine the hypothesis that a fundamental association between sustained xylem water transport and downstream tissue function should select for xylem that avoids embolism in long-vessel trees by quantifying xylem capacity to withstand air entry of western North American oaks (Quercus spp.). Optical visualization showed that 50% of embolism occurs at water potentials below -2.7 MPa in all 19 species, and -6.6 MPa in the most resistant species. By mapping the evolution of xylem vulnerability to embolism onto a fossil-dated phylogeny of the western North American oaks, we found large differences between clades (sections) while closely related species within each clade vary little in their capacity to withstand air entry. Phylogenetic conservatism in xylem physical tolerance, together with a significant correlation between species distributions along rainfall gradients and their dehydration tolerance, suggests that closely related species occupy similar climatic niches and that species' geographic ranges may have shifted along aridity gradients in accordance with their physical tolerance. Such trends, coupled with evolutionary associations between capacity to withstand xylem embolism and other hydraulic-related traits, yield wide margins of safety against embolism in oaks from diverse habitats. Evolved responses of the vascular system to aridity support the embolism avoidance hypothesis and reveal the importance of quantifying plant capacity to withstand xylem embolism for understanding function and biogeography of some of the Northern Hemisphere's most ecologically and economically important plants.

Abiotic stresses could have complex and diverse effects on the growth and development of forest trees. In this review, we summarized the responses of morphological, physiological and anatomical traits in woody plants to abiotic stresses, including drought, flood, extreme temperature, salinity, heavy metal, microplastics and combined stresses, especially from the xylem perspective. Under most abiotic stress, xylem hydraulic conductivity decreases, which is associated with leaf stomatal regulation and the inhibition of aquaporin (AQP) activity. Meanwhile, woody plants regulate the size and morphology of their roots and leaves to balance water absorption and transpiration. The anatomical traits are also altered, such as denser leaf stomata, narrower conduits and thicker cell walls. In addition, different stresses have unique effects, such as flood-induced adventitious roots and aeration tissues, forest fire-induced irreversible xylem damage, low temperature-induced tissue freezing, salt stress-induced hinderance of ion absorption and heavy metal-induced biological toxicity. Under stresses of drought, flooding and heavy metals, woody plants' growth may occasionally be promoted. The effects of combined stress on the physiological, morphological and anatomical traits of woody plants are not simply additive, with the related mechanism to be further studied, especially in natural or near-natural conditions. [ABSTRACT FROM AUTHOR]

Summary: The propagation of xylem embolism throughout the root systems of drought‐affected plants remains largely unknown, despite this process being comparatively well characterized in aboveground tissues.We used optical and X‐ray imaging to capture xylem embolism propagation across the intact root systems of bread wheat (Triticum aestivum L. 'Krichauff') plants subjected to drying. Patterns in vulnerability to xylem cavitation were examined to investigate whether vulnerability may vary based on root size and placement across the entire root system.Individual plants exhibited similar mean whole root system vulnerabilities to xylem cavitation but showed enormous 6 MPa variation within their component roots (c. 50 roots per plant). Xylem cavitation typically initiated in the smallest, peripheral parts of the root system and moved inwards and upwards towards the root collar last, although this trend was highly variable.This pattern of xylem embolism spread likely results in the sacrifice of replaceable small roots while preserving function in larger, more costly central roots. A distinct pattern of embolism‐spread belowground has implications for how we understand the impact of drought in the root system as a critical interface between plant and soil. [ABSTRACT FROM AUTHOR]

In order to gain insights in the combined effects of drought and nitrogen fertilization on water storage and transportation in these plants, pot experiments were conducted. Two-year-old seedlings of 'Ningqi 1' variety of L. barbarum were subjected to three levels of drought combined with three levels of nitrogen fertilization (in the form of urea), plus controls of normal watering and no added nitrogen. Following these treatments, the water content, tissue water capacity, natural dehydration rate and hydraulic characteristics of the plant parts were determined. Under drought stress, the application of nitrogen increased the observed changes of water content in the plant's canopy was more significant than in their roots. The water capacity decreased under increasing drought stress, but when combined with nitrogen application this decrease could be slowed down. The natural dehydration rate of stems was the lowest, that of roots was higher and that of leaves was the highest. With increasing levels of nitrogen fertilization, the natural dehydration rate under drought stress was increased. The root system is the main part that affects water transport in the plants, and we show here that nitrogen application can temper the change of hydraulic conductance and reduce the xylem embolism vulnerability and water potential threshold under drought stress. The results suggest that under water-limiting conditions a proper nitrogen treatment can be beneficial to the water storage and transportation, reduce the vulnerability of xylem vessel embolism, and improve the ability of water retention in Lycium barbarum L. [ABSTRACT FROM AUTHOR]

Summary: "Carlquist's Law" is a striking pattern of association between anatomical features in the wood of vessel-bearing plants. It derives from Sherwin Carlquist's observation that xylem vessels tend to be solitary when embedded in a matrix of imperforate tracheary elements that appear to be conductive, whereas xylem vessels tend to be grouped when surrounded by seemingly non-conductive cells. Vessel-vessel contacts (vessel grouping) allow water to travel between conduits, but also provide pathways for air to propagate from embolized (air-filled) vessels into functional vessels. If the background matrix is conductive, it is conceivable that water could bypass embolized vessels, providing an alternative transport route in species with conductive backgrounds and solitary vessels. Much remains to be tested in this hypothesis, including the topology of the vessel networks in species with solitary versus grouped vessels and how conductive the different imperforate tracheary element types are. Exploring Carlquist's Law promises to provide key insight into the causes of embolism in plant conduits, the modes of embolism passage between conduits, and how vessels and the cells in which they are imbedded may interact to govern the pathways of water flow through plants. [ABSTRACT FROM AUTHOR]

The frequency and intensity of droughts combined with heatwave events have increased under climate change, increasing destruction in urban areas and leading to severe impacts on urban plants. These impacts remain poorly understood at the species level. Here, we investigate the effects of a drought–heatwave event on 12 urban woody species using in situ records of the dynamic changes in physiological traits in the field before, during and after the drought–heatwave event to assess resistance and resilience to hot drought. We found the following: (1) Hydraulic and photosynthesis traits showed an instantaneous decline during the hot drought event in the 12 species, with severe drought-induced xylem embolism in hydraulic systems and a high percentage loss of hydraulic conductivity (PLC). (2) The two conifer species were less resistant to hot droughts than broadleaves but capable of post-stress recovery, suggesting that conifers showed better resilience and that broadleaves showed better resistance under hot drought stress. (3) The evergreen species showed strong resistance, while three deciduous species showed strong resilience to hot drought stress. (4) The three shrubs may be more vulnerable to hot droughts than trees, as they showed lower resistance and were not able to recover the current year's growth. [ABSTRACT FROM AUTHOR]

Key message: Sapwood density is coordinated with hydraulic capacity and non-structural carbohydrate reserves in xylem in the seasonally dry tropical forests. Sapwood density (WD) is one characteristic underlying the divergence in life history strategies among species in seasonally dry tropical forests (SDTFs) since WD is liked to xylem hydraulic properties through its correlations with vessel anatomy, maximum photosynthetic capacity, and leaf phenology. However, it remains unclear how differences in WD contribute to the divergence in water conduction and non-structural carbohydrate (NSC) reserves during the dry season, the physiologically most severe period for woody plants. We hypothesized that heavy-wooded species maintain water conduction, photosynthesis, and translocation of carbon which result in high NSC reserves in xylem during the dry season. Using 13 deciduous tree species from a SDTF located in Northeast Thailand, we investigated the variation in WD and bark morphology in relation to hydraulic properties and NSC concentrations in xylem during the late-dry season. Percentage loss of conductivity (PLC) varied as a quadric function of WD: high PLC was observed in light- and heavy-wooded species. The maximum conductivity was not related to WD. The PLC was negatively related to the concentrations of soluble sugars and NSC in the trunk xylem, and these relationships underlined the negative association between WD and NSCs. We also found that species with thick bark showed relatively low PLC, and that dense-barked species exhibited higher NSC concentrations in branch xylem, but their linkages were generally weaker than WD. These results demonstrate that species hydraulics and NSC reserves are coordinated in SDTFs during the dry season, and that WD underlies these divergences. [ABSTRACT FROM AUTHOR]

Poplar (Populus spp.) is a high-value crop for wood and biomass production and a model organism for tree physiology and genomics. The early release, in 2006, of the complete genome sequence of P. trichocarpa was followed by a wealth of studies that significantly enriched our knowledge of complex pathways inherent to woody plants, such as lignin biosynthesis and secondary cell wall deposition. Recently, in the attempt to cope with the challenges posed by ongoing climate change, fundamental studies and breeding programs with poplar have gradually shifted their focus to address the responses to abiotic stresses, particularly drought. Taking advantage from a set of modern genomic and phenotyping tools, these studies are now shedding light on important processes, including embolism formation (the entry and expansion of air bubbles in the xylem) and repair, the impact of drought stress on biomass yield and quality, and the long-term effects of drought events. In this review, we summarize the status of the research on the molecular bases of the responses to drought in poplar. We highlight how this knowledge can be exploited to select more tolerant genotypes and how it can be translated to other tree species to improve our understanding of forest dynamics under rapidly changing environmental conditions. [ABSTRACT FROM AUTHOR]

Severe droughts have led to lower plant growth and high mortality in many ecosystems worldwide, including tropical forests. Drought vulnerability differs among species, but there is limited consensus on the nature and degree of this variation in tropical forest communities. Understanding species‐level vulnerability to drought requires examination of hydraulic traits since these reflect the different strategies species employ for surviving drought.Here, we examined hydraulic traits and growth reductions during a severe drought for 12 common woody species in a wet tropical forest community in Puerto Rico to ask: Q1. To what extent can hydraulic traits predict growth declines during drought? We expected that species with more hydraulically vulnerable xylem and narrower safety margins (SMP50) would grow less during drought. Q2. How does species successional association relate to the levels of vulnerability to drought and hydraulic strategies? We predicted that early‐ and mid‐successional species would exhibit more acquisitive strategies, making them more susceptible to drought than shade‐tolerant species. Q3. What are the different hydraulic strategies employed by species and are there trade‐offs between drought avoidance and drought tolerance? We anticipated that species with greater water storage capacity would have leaves that lose turgor at higher xylem water potential and be less resistant to embolism forming in their xylem (P50).We found a large range of variation in hydraulic traits across species; however, they did not closely capture the magnitude of growth declines during drought. Among larger trees (≥10 cm diameter at breast height—DBH), some tree species with high xylem embolism vulnerability (P50) and risk of hydraulic failure (SMP50) experienced substantial growth declines during drought, but this pattern was not consistent across species. We found a trade‐off among species between drought avoidance (capacitance) and drought tolerating (P50) in this tropical forest community. Hydraulic strategies did not align with successional associations. Instead, some of the more drought‐vulnerable species were shade‐tolerant dominants in the community, suggesting that a drying climate could lead to shifts in long‐term forest composition and function in Puerto Rico and the Caribbean. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

The pneumatic method is a novel method determining vulnerability to embolism in plants, yet it remains unclear whether this method is suitable for all species with different xylem anatomy. In this study, using six tree species with contrasting xylem anatomy, including four vessel-bearing species (diffuse-porous wood and ring-porous wood) and two tracheid-bearing species (non-porous wood), we test the reliability of the pneumatic method by comparing to hydraulic methods and also considering turgor loss point and native embolism. Vessel length distribution and cut-open vessel volume were also evaluated using the silicone injection technique. Additionally, we also synthesized published data to find out the consistency between the pneumatic method and hydraulic methods. Results showed that there was a maximum 10-folds difference in mean vessel length and mean vessel diameter varying from 30 to 56 μm among species. The estimated open vessel volume ranges from 0.064 to 0.397 mL, with a maximum of 14% of the tube vacuum reservoir. For four vessel-bearing species, the pneumatic method showed good consistency with hydraulic methods, and this consistency was evidenced by turgor loss point and native embolism. For two tracheid-bearing species, the pneumatic method significantly overestimated vulnerability because of the bad consistencies with hydraulic methods and plant water relations. Data synthesis of 56 species also suggested that the pneumatic method can accurately measure the embolism vulnerability of vessel-bearing species but not for tracheid-bearing species. Our study provided further evidence that the pneumatic method is accurate for most vessel-bearing species and thus has the potential to be widely used in the plant hydraulics field. However, we proposed that the precise calculation of air discharge volume should take into account the volume of open vessels for species with wide and long vessels. [ABSTRACT FROM AUTHOR]

How plants respond physiologically to leaf warming and low water availability may determine how they will perform under future climate change. In 2015-2016, an unprecedented drought occurred across Amazonia with record-breaking high temperatures and low soil moisture, offering a unique opportunity to evaluate the performances of Amazonian trees to a severe climatic event. We quantified the responses of leaf water potential, sap velocity, whole-tree hydraulic conductance (Kwt), turgor loss and xylem embolism, during and after the 2015-2016 El Niño for five canopy-tree species. Leaf/xylem safety margins (SMs), sap velocity and Kwt showed a sharp drop during warm periods. SMs were negatively correlated with vapour pressure deficit, but had no significant relationship with soil water storage. Based on our calculations of canopy stomatal and xylem resistances, the decrease in sap velocity and Kwt was due to a combination of xylem cavitation and stomatal closure. Our results suggest that warm droughts greatly amplify the degree of trees' physiological stress and can lead to mortality. Given the extreme nature of the 2015-2016 El Niño and that temperatures are predicted to increase, this work can serve as a case study of the possible impact climate warming can have on tropical trees.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Forest mortality during drought has been attributed to hydraulic failure, which can be challenging to measure. A limited number of alternative proxies for incipient leaf death exist. Here we investigate whether a terminal increase in abscisic acid (ABA) levels in leaves occurs across vascular land plants and is an indicator of imminent leaf death.For different species across vascular plants, we monitored ABA levels during lethal drought as well as leaf embolism resistance, across the canopy as leaves die following senescence, or when leaves are exposed to a heavy, lethal frost late in the growing season.We observed a considerable increase in foliage ABA levels once leaves showed signs of incipient leaf death. This increase in ABA levels upon incipient leaf death, could be induced by embolism during drought, by freezing or as leaves age naturally, and was observed in species spanning the phylogeny of vascular land plants as well as in an ABA biosynthetic mutant plant.A considerable increase in foliage ABA levels may act as an indicator of impending leaf death. [ABSTRACT FROM AUTHOR]

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., (© 2024 John Wiley & Sons Ltd.)

Sugar maples (Acer saccharum Marshall) develop elevated stem pressures in springtime through the compression and expansion of gas bubbles present within xylem fibres. The stability of this gas within the fibres is hypothesised to be due to the elevated sugar concentration of maple sap and the presence of an osmotic barrier between fibres and vessels. Without this osmotic barrier gas bubbles are predicted to dissolve rapidly. In this work we investigated the existence of this osmotic barrier. We quantified the fraction of the xylem occupied by gas-filled fibres using synchrotron based microCT. After imaging fresh stem segments we perfused them with either a 2% sucrose solution or water, imaging again following perfusion. In this way we directly observed how total gas present in the fibres changed when an osmotic pressure difference should be present, with the 2% sucrose solution, and when it is absent, with the water. Following a first round of perfusion we perfused stem segments with the other perfusate, repeating this multiple times to observe how switching perfusates affected gas-filled fibres. We found that perfusing stem segments with water resulted in a significant reduction in the xylem fibre gas, but perfusing stem segments with a sucrose solution did not significantly reduce the gas in the fibres. These results support the hypothesis that an osmotic barrier exists between fibres and vessels., (© The Author(s) 2024. Published by Oxford University Press.)

Subtropical tree species may experience severe drought stress due to variable rainfall under future climates. However, the capacity to restore hydraulic function post-drought might differ among co-occurring species with contrasting leaf habits (e.g., evergreen and deciduous) and have implications for future forest composition. Moreover, the links between hydraulic recovery and physiological and morphological traits related to water-carbon availability are still not well understood. Here, potted seedlings of six tree species (four evergreen and two deciduous) were grown outdoors under a rainout shelter. They grew under favorable water conditions until they were experimentally subjected to a soil water deficit leading to losses of ca. 50% of hydraulic conductivity, and then soils were re-watered to field capacity. Traits related to carbon and water relations were measured. There were differences in drought responses and recovery between species, but not as a function of evergreen or deciduous groups. Sapindus mukorossi exhibited the most rapid drought response, which was associated with a suite of physiological and morphological traits (larger plant size, the lowest hydraulic capacitance (Cbranch), higher minimum conductance (gmin) and lower HV (Huber value)). Upon re-watering, xylem water potential exhibited fast recovery in 1-3 days among species, while photosynthesis at saturating light (Asat) and stomatal conductance (gs) recovery lagged behind water potential recovery depending on species, with gs recovery being more delayed than Asat in most species. Furthermore, none of the six species exhibited significant hydraulic recovery during the 7 days re-watering period, indicating that xylem refilling was apparently limited; in addition, NSC availability had a minimal role in facilitating hydraulic recovery during this short-term period. Collectively, if water supply is limited by insignificant hydraulic recovery post-drought, the observed carbon assimilation recovery of seedlings may not be sustained over the longer term, potentially altering seedling regeneration and shifting forest species composition in subtropical China under climate change. [ABSTRACT FROM AUTHOR]

Variation in resistance of xylem to embolism among flowers, leaves, and stems strongly influences the survival and reproduction of plants. However, little is known about the vulnerability to xylem embolism under drought stress and their relationships to the anatomical traits of pits among reproductive and vegetative organs. In this study, we investigated the variation in xylem vulnerability to embolism in peduncles, petioles, and stems in a woody plant, Magnolia grandiflora. We analyzed the relationships between water potentials that induced 50% embolism (P50) in peduncles, petioles, and stems and the conduit pit traits hypothesized to influence cavitation resistance. We found that peduncles were more vulnerable to cavitation than petioles and stems, supporting the hypothesis of hydraulic vulnerability segmentation that leaves and stems are prioritized over flowers during drought stress. Moreover, P50 was significantly correlated with variation in the dimensions of inter-vessel pit apertures among peduncles, petioles and stems. These findings highlight that measuring xylem vulnerability to embolism in reproductive organs is essential for understanding the effect of drought on plant reproductive success and mortality under drought stress.

Determining the relationship between reductions in stomatal conductance (gs) and leaf water transport during dehydration is key to understanding plant drought responses. While numerous studies have analysed the hydraulic function of woody species, minimal research has been conducted on grasses. Here, we sought to characterize hydraulic vulnerability in five widely‐occurring pasture grasses (including both C3 and C4 grasses) and determine whether reductions in gs and leaf hydraulic conductance (Kleaf) during dehydration could be attributed to xylem embolism. Using the optical vulnerability (OV) technique, we found that all species were highly resistant to xylem embolism when compared to other herbaceous angiosperms, with 50% xylem embolism (PX50) occurring at xylem pressures ranging from −4.4 to −6.1 MPa. We observed similar reductions in gs and Kleaf under mild water stress for all species, occurring well before PX50. The onset of xylem embolism (PX12) occurred consistently after stomatal closure and 90% reduction of Kleaf. Our results suggest that factors other than xylem embolism are responsible for the majority of reductions in gs and Kleaf during drought and reductions in the productivity of pasture species under moderate drought may not be driven by embolism. Summary Statement: Pasture grasses are highly resistant to xylem cavitation, with cavitation occurring consistently after significant reductions in stomatal conductance (gs) and leaf hydraulic conductance (Kleaf). Our results suggest that factors other than xylem cavitation are responsible for reductions in gs and Kleaf during moderate drought with cavitation only occurring under extreme drought conditions. [ABSTRACT FROM AUTHOR]

Root pressure is a positive hydraulic pressure that generated by plant roots. As a common physiological phenomenon in many plants, root pressure can drive the sap flow from roots to canopy leaves, which alleviates plant water deficits caused by strong transpiration during the day. It also plays an important role in xylem embolism refilling. At present, there are many explanations for the generation of root pressure. The generally accepted views include the osmotic theory, the metabolic theory, and the uphill water co-transport hypothesis, but the mechanism of root pressure is still the focus of controversy among the plant scientists. There are direct and indirect methods for measuring root pressure, but the more advanced techniques of measuring root pressure still need to be improved. Under the influence of many factors such as aquaporin, genetic factors and habitats, there are differences in the magnitude of root pressure, and moderate root pressure has important significance for crops growth. Therefore, it is of great biological significance to deepen our understanding of root pressure. Here we discussed root pressure from the following perspectives: the definition and mechanism of root pressure generation; plant groups with root pressure; the methods for measuring root pressure; the main factors about influencing root pressure; the significance and implications of root pressure in various fields of plant science studies. Lastly, based on current research topics and some new results on root pressure, we discussed future perspectives on root pressure of plants. [ABSTRACT FROM AUTHOR]

In humid temperate forests, the occurrence of frequent freeze–thaw cycles (FTC) is a main factor limiting tree growth, as xylem embolism induced by FTC poses a serious threat to the hydraulic integrity of trees. A high resilience to hydraulic dysfunction involves the enhancement of embolism resistance and/or extra non‐structural carbohydrate (NSC) inputs for restoration of an impaired hydraulic system. However, potentially negative implications of such NSC allocation on tree growth have not yet been explored.At a temperate forest site of northeast China, we studied xylem hydraulics and NSC contents in relation to winter embolism resilience in 15 sympatric broadleaf tree species belonging to three genera with relatively high species richness, but having distinct strategies in maintaining hydraulic integrity over the winter, that is, Acer species that generate positive stem and root pressures contributing to winter embolism refilling, Betula species that only generate root pressure and Populus species that do not generate positive pressure.Acer and Betula species had higher soluble sugar contents in the dormant season and indeed had higher hydraulic resilience to FTC‐induced embolism but slower stem growth. Populus species had higher NSC contents during the growing season and their faster stem growth was also consistent with higher hydraulic efficiency and leaf photosynthetic rate. The positive correlation between tree trunk radial growth rate and hydraulic conductivity suggests that xylem water transport efficiency can be a fundamental basis for tree productivity due to a significant hydraulic‐photosynthetic coordination. The negative correlation between soluble sugar concentration in the dormant season and stem growth rate indicates that metabolic carbon costs for enhancing hydraulic resilience may compromise tree growth during the growing season.Comparisons among Acer, Betula and Populus and the correlation analyses based on phylogenetic independent contrasts strongly support the existence of a trade‐off between hydraulic resilience against FTC‐induced embolism and growth rate among sympatric tree species under humid temperate climate conditions. This trade‐off has likely contributed to the sorting of temperate tree species and genera to different niches along environmental gradients with respect to freezing stress and interspecific competition. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Poplar (Populus spp.) is a high-value crop for wood and biomass production and a model organism for tree physiology and genomics. The early release, in 2006, of the complete genome sequence of P. trichocarpa was followed by a wealth of studies that significantly enriched our knowledge of complex pathways inherent to woody plants, such as lignin biosynthesis and secondary cell wall deposition. Recently, in the attempt to cope with the challenges posed by ongoing climate change, fundamental studies and breeding programs with poplar have gradually shifted their focus to address the responses to abiotic stresses, particularly drought. Taking advantage from a set of modern genomic and phenotyping tools, these studies are now shedding light on important processes, including embolism formation (the entry and expansion of air bubbles in the xylem) and repair, the impact of drought stress on biomass yield and quality, and the long-term effects of drought events. In this review, we summarize the status of the research on the molecular bases of the responses to drought in poplar. We highlight how this knowledge can be exploited to select more tolerant genotypes and how it can be translated to other tree species to improve our understanding of forest dynamics under rapidly changing environmental conditions.

Drought poses a significant threat to forest survival worldwide by potentially generating air bubbles that obstruct sap transport within plants' hydraulic systems. However, the detailed mechanism of air entry and propagation at the scale of the veins remains elusive. Building upon a biomimetic model of leaf which we developed, we propose a direct comparison of the air embolism propagation in Adiantum (maidenhair fern) leaves, presented in Brodribb et al . (Brodribb TJ, Bienaimé D, Marmottant P. 2016 Revealing catastrophic failure of leaf networks under stress. Proc. Natl Acad. Sci. USA 113 , 4865-4869 (doi:10.1073/pnas.1522569113)) and in our biomimetic leaves. In particular, we evidence that the jerky dynamics of the embolism propagation observed in Adiantum leaves can be recovered through the introduction of micrometric constrictions in the section of our biomimetic veins, mimicking the nanopores present in the bordered pit membranes in real leaves. We show that the intermittency in the propagation can be retrieved by a simple model coupling the variations of pressure induced by the constrictions and the variations of the volume of the compliant microchannels. Our study marks a step with the design of a biomimetic leaf that reproduces particular aspects of embolism propagation in real leaves, using a minimal set of controllable and readily tunable components. This biomimetic leaf constitutes a promising physical analogue and sets the stage for future enhancements to fully embody the unique physical features of embolizing real leaves.

Premise: Environmental gradients of mountains are reflected in traits that are common to high‐elevation plants worldwide. Closely related species of Senecio from the equatorial Andes grow as broad‐leaved climbers in montane forests, basal broad‐leaved rosette herbs in azonal marshy habitats, and ascending, narrow‐leaved subshrubs in high‐elevation páramo. Habitat variation along the elevational gradient enables testing whether modifications in leaf and stem functional traits among species were driven by contrasting environmental conditions. Methods: We used quantitative analyses to describe changes in morphological and anatomical traits of leaves and stems in 10 species from various habitats. We applied univariate (linear regression, hierarchical ANOVA) and multivariate (NMDS ordination, permutational MANOVA) techniques to examine the correlation of traits with the species' habitats and elevation. Results: Species from the humid and frost‐free montane forest develop xylem optimized for transport efficiency by increasing the internal diameter and length of the conduits. In contrast, páramo species are optimized toward hydraulic safety by producing narrower conduits and are more likely to prevent the risk of frost‐induced cavitation. Moreover, species from the high‐elevation páramo habitats present a set of water‐transport‐related anatomical traits of leaf lamina, allowing for efficient regulation of transpiration losses. Conclusions: Morphological and anatomical traits of leaves and stems in species of Senecio inhabiting montane forests and high‐elevation páramo in the equatorial Andes demonstrate a trade‐off between hydraulic safety and efficiency of water transport. [ABSTRACT FROM AUTHOR]

Selection of high-yielding traits in cereal plants led to a continuous increase in productivity. However, less effort was made to select on adaptive traits, favorable in adverse and harsh environments. Under current climate change conditions and the knowledge that cereals are staple foods for people worldwide, it is highly important to shift focus to the selection of traits related to drought tolerance, and to evaluate new tools for efficient selection. Here, we explore the possibility to use vulnerability to drought-induced xylem embolism of wheat cultivars Excalibur and Hartog (Triticum aestivum L.), rye cultivar Duiker Max (Secale cereale L.), and triticale cultivars Dublet and US2014 (x Triticosecale Wittmack) as a proxy for their drought tolerance. Multiple techniques were combined to underpin this hypothesis. During bench-top dehydration experiments, acoustic emissions (AEs) produced by formation of air emboli were detected, and hydraulic capacitances quantified. By only looking at the AE50 values, one would classify wheat cultivar Excalibur as most tolerant and triticale cultivar Dublet as most vulnerable to drought-induced xylem embolism, though Dublet had significantly higher hydraulic capacitances, which are essential in terms of internal water storage to temporarily buffer or delay water shortage. In addition, xylem anatomical traits revealed that both cultivars have a contrasting trade-off between hydraulic safety and efficiency. This paper emphasizes the importance of including a cultivar's hydraulic capacitance when evaluating its drought response and vulnerability to drought-induced xylem embolism, instead of relying on the AE50 as the one parameter. [ABSTRACT FROM AUTHOR]

Summary: Eastern Australia was subject to its hottest and driest year on record in 2019. This extreme drought resulted in massive canopy die‐back in eucalypt forests. The role of hydraulic failure and tree size on canopy die‐back in three eucalypt tree species during this drought was examined.We measured pre‐dawn and midday leaf water potential (Ψleaf), per cent loss of stem hydraulic conductivity and quantified hydraulic vulnerability to drought‐induced xylem embolism. Tree size and tree health was also surveyed.Trees with most, or all, of their foliage dead exhibited high rates of native embolism (78–100%). This is in contrast to trees with partial canopy die‐back (30–70% canopy die‐back: 72–78% native embolism), or relatively healthy trees (little evidence of canopy die‐back: 25–31% native embolism). Midday Ψleaf was significantly more negative in trees exhibiting partial canopy die‐back (−2.7 to −6.3 MPa), compared with relatively healthy trees (−2.1 to −4.5 MPa). In two of the species the majority of individuals showing complete canopy die‐back were in the small size classes.Our results indicate that hydraulic failure is strongly associated with canopy die‐back during drought in eucalypt forests. Our study provides valuable field data to help constrain models predicting mortality risk. [ABSTRACT FROM AUTHOR]

Key message: Distributions of water- and air-filled conduits can be distinguished by observing a xylem cross-section of frozen conifer stem. This method is applicable to field observation in cold winter. Xylem embolism resulting from summer drought and frost drought is one of the critical stresses responsible for the dieback of stems and individuals of subalpine evergreen conifers. The occurrence of xylem embolism in conifers depends not only on the species but also on the plant microhabitats, causing difficulties in understanding possible adaptive strategies against xylem embolism. This study examines a simple method of using a digital camera to photograph the xylem water distribution (CXW method) in cross-sections of frozen stems. Light is transmitted through the water-filled tracheid lumen but reflects and scatters at the surface of embolized tracheids, resulting in contrast in wood color between darker (water-filled tracheids) and lighter (air-filled tracheids) colored regions. The CXW method was effective in detecting water distribution in conifers, although the colored xylem in latewood and reaction wood decreased the color contrast between air- and water-filled regions. By cutting the frozen stem with a cryostat, sequential changes in the water distribution of stem xylem were easily monitored. In the cold winter of the subalpine region, the spatial distribution of embolized conduits can be detected when a branch is collected. If a cryostat is available, this method is applicable to other tracheid-bearing wood collected in any season and does not require additional instruments or time-consuming intensive labor in the field. Information about the hydraulics of conifers growing in extreme environments contributes to the understanding of their adaptive strategy and facilitates accurate prediction of forest dynamics under future climatic conditions. [ABSTRACT FROM AUTHOR]

Soil waterlogging and drought correspond to contrasting water extremes resulting in plant dehydration. Dehydration in response to waterlogging occurs due to impairments to root water transport, but no previous study has addressed whether limitations to water transport occur beyond this organ or whether dehydration alone can explain shoot impairments. Using common bean (Phaseolus vulgaris) as a model species, we report that waterlogging also impairs water transport in leaves and stems. During the very first hours of waterlogging, leaves transiently dehydrated to water potentials close to the turgor loss point, possibly driving rapid stomatal closure and partially explaining the decline in leaf hydraulic conductance. The initial decline in leaf hydraulic conductance (occurring within 24 h), however, surpassed the levels predicted to occur based solely on dehydration. Constraints to leaf water transport resulted in a hydraulic disconnection between leaves and stems, furthering leaf dehydration during waterlogging and after soil drainage. As leaves dehydrated later during waterlogging, leaf embolism initiated and extensive embolism levels amplified leaf damage. The hydraulic disconnection between leaves and stems prevented stem water potentials from declining below the threshold for critical embolism levels in response to waterlogging. This allowed plants to survive waterlogging and soil drainage. In summary, leaf and stem dehydration are central in defining plant impairments in response to waterlogging, thus creating similarities between waterlogging and drought. Yet, our findings point to the existence of additional players (likely chemicals) partially controlling the early declines in leaf hydraulic conductance and contributing to leaf damage during waterlogging., (© The Author(s) 2024. Published by Oxford University Press on behalf of American Society of Plant Biologists.)

While plant height is the main driver of variation in mean vessel diameter at the stem base (VD) across angiosperms, climate, specifically temperature, does play an explanatory role, with vessels being wider with warmer temperature for plants of the same height. Using a comparative approach sampling 537 species of angiosperms across 19 communities, we rejected selection favouring freezing‐induced embolism resistance as being able to account for wider vessels for a given height in warmer climates. Instead, we give reason to suspect that higher vapour pressure deficit (VPD) accounts for the positive scaling of height‐standardized VD (and potential xylem conductance) with temperature. Selection likely favours conductive systems that are able to meet the higher transpirational demand of warmer climates, which have higher VPD, resulting in wider vessels for a given height. At the same time, wider vessels are likely more vulnerable to dysfunction. With future climates likely to experience ever greater extremes of VPD, future forests could be increasingly vulnerable. In angiosperms, individuals from warmer climates have wider vessels than similar‐sized plants in colder climates. This trend is consistent with scaling of vessel diameter with vapour pressure deficit (VPD). Because VPD will be higher in future climates, and because wider vessels are potentially more vulnerable to embolism, future forests could be increasingly vulnerable. [ABSTRACT FROM AUTHOR]

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The observation of a much-improved fitness of wild-type plants over abscisic acid (ABA)-deficient mutants during drought has led researchers from all over to world to perform experiments aiming at a better understanding of how this hormone modulates the physiology of plants under water-limited conditions. More recently, several promising approaches manipulating ABA biosynthesis and signaling have been explored to improve water use efficiency and confer drought tolerance to major crop species. Here, we review recent progress made in the last decade on (i) ABA biosynthesis, (ii) the roles of ABA on plant-water relations and on primary and secondary metabolisms during drought, and (iii) the regulation of ABA levels and perception to improve water use efficiency and drought tolerance in crop species. [ABSTRACT FROM AUTHOR]

Premise: The young seedling life stage is critical for reforestation after disturbance and for species migration under climate change, yet little is known regarding their basic hydraulic function or vulnerability to drought. Here, we sought to characterize responses to desiccation including hydraulic vulnerability, xylem anatomical traits, and impacts on other stem tissues that contribute to hydraulic functioning. Methods: Larix occidentalis, Pseudotsuga menziesii, and Pinus ponderosa (all ≤6 weeks old) were imaged using x‐ray computed microtomography during desiccation to assess seedling biomechanical responses with concurrently measured hydraulic conductivity (ks) and water potential (Ψ) to assess vulnerability to xylem embolism formation and other tissue damage. Results: In non‐stressed samples for all species, pith and cortical cells appeared circular and well hydrated, but they started to empty and deform with decreasing Ψ which resulted in cell tearing and eventual collapse. Despite the severity of this structural damage, the vascular cambium remained well hydrated even under the most severe drought. There were significant differences among species in vulnerability to xylem embolism formation, with 78% xylem embolism in L. occidentalis by Ψ of −2.1 MPa, but only 47.7% and 62.1% in P. ponderosa and P. menziesii at −4.27 and −6.73 MPa, respectively. Conclusions: Larix occidentalis seedlings appeared to be more susceptible to secondary xylem embolism compared to the other two species, but all three maintained hydration of the vascular cambium under severe stress, which could facilitate hydraulic recovery by regrowth of xylem when stress is relieved. [ABSTRACT FROM AUTHOR]

Summary: Flowers underpin plant evolution, genetic legacy and global food supply. They are exposed to similar evaporative conditions as leaves, yet floral physiology is a product of different selective forces. We used Tanacetum cinerariifolium, a perennial daisy, to examine the response of flowers to whole‐plant water stress, determining if flowers constitute a liability during drought, and how this species has adapted to minimize risk associated with reproduction.We determined the relative transpiration cost of flowers and leaves and confirmed that flowers in this species are xylem‐hydrated. The relative water stress tolerance of leaves and flowers then was compared using xylem vulnerability measurements linked with observed tissue damage during an acute drought treatment.Flowers were a major source of water loss during drought but the xylem supplying them was much more vulnerable to cavitation than leaves. This xylem vulnerability segmentation was confirmed by observations that most flowers died whereas leaves were minimally affected during drought.Early cavitation and hydraulic isolation of flowers during drought benefits the plant by slowing the dehydration of perennial vegetative organs and delaying systemic xylem damage. Our results highlight the need to understand flower xylem vulnerability as a means of predicting plant reproductive failure under future drought. [ABSTRACT FROM AUTHOR]

Recent findings suggest that tree mortality and post‐drought recovery of gas exchange can be predicted from loss of function within the water transport system. Understanding the susceptibility of plants to hydraulic damage requires knowledge about the vulnerability of different plant organs to stress‐induced hydraulic dysfunction. This is particularly important in the context of vulnerability segmentation between plant tissues which is believed to protect more energetically 'costly' tissues, such as woody stems, by sacrificing 'cheaper' leaves early under drought conditions.Differences in vulnerability segmentation between co‐occurring plant species could explain divergent behaviours during drought, yet there are few studies considering how this characteristic may vary within a plant community. Here we investigated community‐wide vulnerability segmentation by comparing leaf/shoot and stem vulnerability in all coexistent dominant canopy and understory woody species in a diverse dry sclerophyll woodland community, including multiple angiosperms and one gymnosperm.Previously published terminal leaf/shoot vulnerability to loss of water transport capacity was compared with stem xylem vulnerability to embolism measured on the same species at the same site. We calculated hydraulic safety margins for stems to determine variation in the risk of hydraulic failure during drought among species.The xylem of all species was found to be highly resistant to hydraulic dysfunction, with only two of the eight species exhibiting significantly different vulnerability to the overall mean. No evidence of vulnerability segmentation between shoots/leaves and stems was found in seven of the eight species.Phylogenetically diverse canopy and understory species in this evergreen sclerophyll woodland appear to have evolved similar strategies of drought resistance, including low xylem vulnerability to embolism and general lack of vulnerability segmentation. This convergence in hydraulic safety indicates a lack of hydraulic niche partitioning in this woodland community. A free plain language summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Climate change threatens food security, and plant science researchers have investigated methods of sustaining crop yield under drought. One approach has been to overproduce abscisic acid (ABA) to enhance water use efficiency. However, the concomitant effects of ABA overproduction on plant vascular system functioning are critical as it influences vulnerability to xylem hydraulic failure. We investigated these effects by comparing physiological and hydraulic responses to water deficit between a tomato (Solanum lycopersicum) wild type control (WT) and a transgenic line overproducing ABA (sp12). Under well‐watered conditions, the sp12 line displayed similar growth rate and greater water use efficiency by operating at lower maximum stomatal conductance. X‐ray microtomography revealed that sp12 was significantly more vulnerable to xylem embolism, resulting in a reduced hydraulic safety margin. We also observed a significant ontogenic effect on vulnerability to xylem embolism for both WT and sp12. This study demonstrates that the greater water use efficiency in the tomato ABA overproducing line is associated with higher vulnerability of the vascular system to embolism and a higher risk of hydraulic failure. Integrating hydraulic traits into breeding programmes represents a critical step for effectively managing a crop's ability to maintain hydraulic conductivity and productivity under water deficit. We show that an ABA overproducing tomato line exhibits greater water use efficiency than its wild type counterpart but higher vulnerability to hydraulic failure through increased vulnerability to xylem embolism and a reduced hydraulic safety margin. This suggests that breeding programmes should integrate hydraulic traits for effectively managing a crop's ability to maintain hydraulic conductivity and productivity under drought. [ABSTRACT FROM AUTHOR]

Drought and nitrogen (N) addition have been shown to affect tree hydraulic traits, but few studies have been made on their interactions across species with different wood types or leaf forms. We examined the responses of hydraulic conductance and xylem anatomical traits of Quercus mongolica (ring porous with simple leaves), Fraxinus mandshurica (ring porous with compound leaves) and Tilia amurensis (diffuse porous with simple leaves) to drought, N addition and their interactions. Drought stress decreased current-year xylem-specific conductivity in stems (K sx) and leaf hydraulic conductance (K leaf), but N addition affected K sx and K leaf differently among species and watering regimes. These divergent effects were associated with different responses of anatomical traits and leaf forms. Higher mean vessel diameter in stems and lower vessel density in leaves were observed with N addition. The three-way interactive effects of drought, N addition and tree species were significant for most values of anatomical traits. These results were also reflected in large differences in vessel diameter and density among species with different wood types or leaf forms. The two-way interactive effects of drought and N addition were significant on K leaf and predawn water potential, but not K sx, indicating that leaves were more sensitive than stems to a combination of drought stress and N addition. Our results provide mechanistic insight into the variable responses of xylem water transport to the interactions of drought and N availability. [ABSTRACT FROM AUTHOR]

Aristotelia chilensis (Molina) Stuntz is a promising species in the food industry as it provides ‘super fruits’ with remarkable antioxidant activity. However, under the predicted climate change scenario, the ongoing domestication of the species must consider selecting the most productive genotypes and be based on traits conferring drought tolerance. We assessed the vulnerability to cavitation and stomatal sensitivity to vapor pressure deficit (VPD) in A. chilensis clones originated from provenances with contrasting climates. A nursery experiment was carried out for one growing season on 2-year-old potted plants. Measurements of stomatal conductance (gs) responses to VPD were taken in spring, summer, and autumn, whereas vulnerability to cavitation was evaluated at the end of spring. Overall, the vulnerability to cavitation of the species was moderate (mean P50 of −2.2 MPa). Parameters of the vulnerability curves (Kmax, P50, P88, and S50) showed no differences among clones or when northern and southern clones were compared. Moreover, there were no differences in stomatal sensitivity to VPD at the provenance or the clonal level. However, compared with other studies, the stomatal sensitivity was considered moderately low, especially in the range of 1 to 3 kPa of VPD. The comparable performance of genotypes from contrasting provenance origins suggests low genetic variation for these traits. Further research must consider testing on diverse environmental conditions to assess the phenotypic plasticity of these types of traits.

The vulnerability to xylem embolism is a key trait underlying species-specific drought tolerance of plants, and hence is critical for screening climate-resilient crops and understanding vegetation responses to drought and heat waves. Yet, accurate determination of embolism in plant's xylem is challenging, because most traditional hydraulic techniques are destructive and prone to artefacts. Hence, direct and in vivo synchrotron-based X-ray micro-CT observation of xylem conduits has emerged as a key reference technique for accurate quantification of vulnerability to xylem embolism. Micro-CT is nowadays a fundamental tool for studies of plant hydraulic architecture, and this chapter describes the fundamentals of acquisition and processing of micro-CT images of plant xylem., (© 2024. The Author(s), under exclusive license to Springer Science+Business Media, LLC, part of Springer Nature.)

Plant responses to drought and heat stress have been extensively studied, whereas post-stress recovery, which is fundamental to understanding stress resilience, has received much less attention. Here, we present a conceptual stress-recovery framework with respect to hydraulic and metabolic functioning in woody plants. We further synthesize results from controlled experimental studies following heat or drought events and highlight underlying mechanisms that drive post-stress recovery. We find that the pace of recovery differs among physiological processes. Leaf water potential and abscisic acid concentration typically recover within few days upon rewetting, while leaf gas exchange-related variables lag behind. Under increased drought severity as indicated by a loss in xylem hydraulic conductance, the time for stomatal conductance recovery increases markedly. Following heat stress release, a similar delay in leaf gas exchange recovery has been observed, but the reasons are most likely a slow reversal of photosynthetic impairment and other temperature-related leaf damages, which typically manifest at temperatures above 40 °C. Based thereon, we suggest that recovery of gas exchange is fast following mild stress, while recovery is slow and reliant on the efficiency of repair and regrowth when stress results in functional impairment and damage to critical plant processes. We further propose that increasing stress severity, particular after critical stress levels have been reached, increases the carbon cost involved in reestablishing functionality. This concept can guide future experimental research and provides a base for modeling post-stress recovery of carbon and water relations in trees. [ABSTRACT FROM AUTHOR]

• Hydraulic failure, but not carbon starvation, occurred towards mortality in all species under both drought treatments. • Rapid xylem repair after drought was not observed. • The recovery of photosynthesis was decoupled from water potential recovery. The impact of drought stress on dominant tree species of subtropical China, and the capacity of these species to recover from drought, is relatively unknown. Three ecologically important tree species (Syzygium rehderianum , Castanopsis chinensis and Schima superba), with contrasting drought sensitivities, were grown in pots under well-watered conditions in the glasshouse before being subjected to two drought treatments (i.e. higher intensity and shorter-duration fast drought(FD);lower intensity and longer-duration slow drought(SD)) towards mortality, and then re-watered to assess the rate of physiological recovery. Dry mass production and physiological traits related to water and carbon relations were measured over the experimental period. FD led to faster declines in hydraulic and photosynthetic function than SD in all species, while hydraulic failure occurred towards mortality in all species under both drought treatments. Although NSC responses varied among drought treatments and species, carbon starvation was not observed. S. rehderianum exhibited more rapid declines in leaf water potential and gas exchange traits, and shorter time-to-mortality, followed by C. chinensis and S. superba. Following re-watering, all three species showed rapid recovery of leaf water potential, while rapid recovery in stem hydraulic conductivity was not evident in any species. Furthermore, C. chinensis exhibited full rapid recovery of photosynthesis, but photosynthetic recovery in S. rehderianum and S. superba was slower than recovery of water potential, reflecting differential stomatal and biochemical limitations. The response of the three species to drought was strongly correlated with xylem embolism resistance, but recovery following drought was complex and partially associated with stomatal and biochemical traits. Under future longer and severe droughts, these species may diverge in drought responses and recovery, perhaps driving shifts in forest community structure in subtropical China. [ABSTRACT FROM AUTHOR]

Drought‐induced tree mortality is expected to increase in future climates with the potential for significant consequences to global carbon, water, and energy cycles. Xylem embolism can accumulate to lethal levels during drought, but species that can refill embolized xylem and recover hydraulic function may be able to avoid mortality. Yet the potential controls of embolism recovery, including cross‐biome patterns and plant traits such as nonstructural carbohydrates (NSCs), hydraulic traits, and nocturnal stomatal conductance, are unknown. We exposed eight plant species, originating from mesic (tropical and temperate) and semi‐arid environments, to drought under ambient and elevated CO2 levels, and assessed recovery from embolism following rewatering. We found a positive association between xylem recovery and NSCs, and, surprisingly, a positive relationship between xylem recovery and nocturnal stomatal conductance. Arid‐zone species exhibited greater embolism recovery than mesic zone species. Our results indicate that nighttime stomatal conductance often assumed to be a wasteful use of water, may in fact be a key part of plant drought responses, and contribute to drought survival. Findings suggested distinct biome‐specific responses that partially depended on species climate‐of‐origin precipitation or aridity index, which allowed some species to recover from xylem embolism. These findings provide improved understanding required to predict the response of diverse plant communities to drought. Our results provide a framework for predicting future vegetation shifts in response to climate change. [ABSTRACT FROM AUTHOR]

Centrifuge-based techniques to assess xylem vulnerability to embolism are increasingly being used, although we are yet to reach a consensus on the nature and extent of artefactual embolism observed in some angiosperm species. In particular, there is disagreement over whether these artefacts influence both the spin (Cavitron) and static versions of the centrifuge technique equally. We tested two methods for inducing embolism: bench dehydration and centrifugation. We used three methods to measure the resulting loss of conductivity: gravimetric flow measured in bench-dehydrated and centrifuged samples (static centrifuge), in situ flow measured under tension during spinning in the centrifuge (Cavitron) and direct imaging using X-ray computed microtomography (microCT) observations in stems of two species of Hakea that differ in vessel length. Both centrifuge techniques were prone to artefactual embolism in samples with maximum vessel length longer than, or similar to, the centrifuge rotor diameter. Observations with microCT indicated that this artefactual embolism occurred in the outermost portions of samples. The artefact was largely eliminated if flow was measured in an excised central part of the segment in the static centrifuge or starting measurements with the Cavitron at pressures lower than the threshold of embolism formation in open vessels. The simulations of loss of conductivity in centrifuged samples with a new model, CAVITOPEN, confirmed that the impact of open vessels on the vulnerability to embolism curve was higher when vessels were long, samples short and when embolism is formed in open vessels at less negative pressures. This model also offers a robust and quantitative tool to test and correct for artefactual embolism at low xylem tensions. [ABSTRACT FROM AUTHOR]

• We tested ecophysiological traits in tropical dipterocarp trees and their hybrids. • Parent species traits were passed to their hybrids through introgression. • Drought tolerance was enhanced via crossing with more tolerant species. • Hybridization may enable adaptation to drought induced by climate change. Gene transfer through interspecific hybridization can enhance functions such as growth performance, although its role in the ecophysiological functions of tropical tree species remains unclear. Hybrid seedlings of genus Shorea (Dipterocarpaceae), which dominates the canopy layer, are found in fragmented tropical rainforests in Southeast Asia. To elucidate changes in dipterocarp environmental adaptability due to hybridization, we studied the ecophysiological traits of dipterocarp seedlings including leaf morphology, photosynthesis, and drought tolerance in two parent species (Shorea curtisii and Shorea leprosula) and their hybrids, including F 1 and backcross hybrids. Then, we induced drought stress to investigate drought tolerance in these seedlings. Photosynthetic ability was higher in S. leprosula than in S. curtisii , with the F 1 and backcross hybrids showing intermediate characteristics. By contrast, drought tolerance, as indicated by stem hydraulic safety, cuticle thickness, and leaf mass per area, was higher in S. curtisii , lower in S. leprosula , and intermediate in their hybrids. The leaf turgor loss point decreased in S. curtisii due to osmotic adjustment under drought conditions, which led to lower predawn leaf water potential and higher water absorption ability from drier soil. By contrast, these changes were smaller and drought tolerance was weaker in S. leprosula. The hybrids including backcrosses had higher drought tolerance than S. leprosula , suggesting that gene transfer through introgression may facilitate the acquisition of drought tolerance. Although increased drought stress due to climate change and forest degradation are likely to limit seedling regeneration in S. leprosula , drought-tolerant genes obtained through hybridization with S. curtisii may contribute to its survival. [ABSTRACT FROM AUTHOR]

The review summarizes some of our current knowledge on the phenomenon of exudation from the cut surface of detached roots with emphasis on results that were mostly established over the last fifty years. The phenomenon is quantitatively documented in the 18th century (by Hales in 1727). By the 19th century, theories mainly ascribed exudation to the secretion of living root cells. The 20th century favored the osmometer model of root exudation. Nevertheless, growing insights into the mechanisms of water transport and new or rediscovered observations stimulated the quest for a more adequate exudation model. The historical overview shows how understanding of exudation changed with time following experimental opportunities and novel ideas from different areas of knowledge. Later theories included cytoskeleton-dependent micro-pulsations of turgor in root cells to explain the observed water exudation. Recent progress in experimental biomedicine led to detailed study of channels and transporters for ion transport via cellular membranes and to the discovery of aquaporins. These universal molecular entities have been incorporated to the more complex models of water transport via plant roots. A new set of ideas and explanations was based on cellular osmoregulation by mechanosensitive ion channels. Thermodynamic calculations predicted the possibility of water transport against osmotic forces based on co-transport of water with ions via cation-chloride cotransporters. Recent observations of rhizodermis exudation, exudation of roots without an external aqueous medium, segments cut from roots, pulses of exudation, a phase shifting of water uptake and exudation, and of effects of physiologically active compounds (like ion channel blockers, metabolic agents, and cytoskeletal agents) will likely refine our understanding of the phenomenon. So far, it seems that more than one mechanism is responsible for root pressure and root exudation, processes which are important for refilling of embolized xylem vessels. However, recent advances in ion and water transport research at the molecular level suggest potential future directions to understanding of root exudation and new models awaiting experimental testing.