Search

Future climate presents conflicting implications for forest biomass. We evaluate how plant hydraulic traits, elevated CO2 levels, warming, and changes in precipitation affect forest primary productivity, evapotranspiration, and the risk of hydraulic failure. We used a dynamic vegetation model with plant hydrodynamics (FATES-HYDRO) to simulate the stand-level responses to future climate changes in a wet tropical forest in Barro Colorado Island, Panama. We calibrated the model by selecting plant trait assemblages that performed well against observations. These assemblages were run with temperature and precipitation changes for two greenhouse gas emission scenarios (2086-2100: SSP2-45, SSP5-85) and two CO2 levels (contemporary, anticipated). The risk of hydraulic failure is projected to increase from a contemporary rate of 5.7% to 10.1-11.3% under future climate scenarios, and, crucially, elevated CO2 provided only slight amelioration. By contrast, elevated CO2 mitigated GPP reductions. We attribute a greater variation in hydraulic failure risk to trait assemblages than to either CO2 or climate. Our results project forests with both faster growth (through productivity increases) and higher mortality rates (through increasing rates of hydraulic failure) in the neo-tropics accompanied by certain trait plant assemblages becoming nonviable.

This article is a Commentary on Robbins et al. (2024), 244: 2239–2250. [ABSTRACT FROM AUTHOR]

Abstract Key message Winter reddening of young Douglas-fir (Pseudotsuga menziesii Mirb. Franco), triggered by large thermal fluctuations in late winter, is a critical problem for European forestry. A literature review identified certain climatic conditions that are characteristic of ‘reddening’ years, including warm daily temperatures, high daily temperature amplitude, low relative humidity, moderate wind speeds, as well as the occurrence of freeze-thaw cycles with cold night temperatures. By describing the triggering environmental and stand factors, we propose three hypotheses for the physiological processes leading to winter reddening, namely (i) hydraulic failure due to winter drought stress, (ii) photo-oxidative stress in shade-acclimated trees, and (iii) early cold deacclimation during warm periods. i) Low soil temperature, by reducing root water uptake, combined with anticyclonic conditions, by increasing water losses, can induce hydraulic failure in the xylem. Hydraulic failure may be further accelerated by night frosts. ii) Winter reddening can occur when low temperature and high irradiance coincide, disrupting photostasis. Overwhelming of winter photo-protection may lead to photodamage and subsequent reddening. iii) Warm periods, by inducing cold deacclimation, make trees susceptible to frost damage. Finally, the three processes may interact under atypical anticyclonic conditions in late winter (e.g. cold or dry soils, warm days, high irradiance and/or freezing nights). Indeed, trees under water stress would develop a higher sensitivity to freezing night and photooxidative stress. We therefore proposed mitigation actions to avoid exposing trees to stressful conditions based on e.g. stand characteristics, understorey vegetation and planting.

Key message: Winter reddening of young Douglas-fir (Pseudotsuga menziesii Mirb. Franco), triggered by large thermal fluctuations in late winter, is a critical problem for European forestry. A literature review identified certain climatic conditions that are characteristic of 'reddening' years, including warm daily temperatures, high daily temperature amplitude, low relative humidity, moderate wind speeds, as well as the occurrence of freeze-thaw cycles with cold night temperatures. By describing the triggering environmental and stand factors, we propose three hypotheses for the physiological processes leading to winter reddening, namely (i) hydraulic failure due to winter drought stress, (ii) photo-oxidative stress in shade-acclimated trees, and (iii) early cold deacclimation during warm periods. i) Low soil temperature, by reducing root water uptake, combined with anticyclonic conditions, by increasing water losses, can induce hydraulic failure in the xylem. Hydraulic failure may be further accelerated by night frosts. ii) Winter reddening can occur when low temperature and high irradiance coincide, disrupting photostasis. Overwhelming of winter photo-protection may lead to photodamage and subsequent reddening. iii) Warm periods, by inducing cold deacclimation, make trees susceptible to frost damage. Finally, the three processes may interact under atypical anticyclonic conditions in late winter (e.g. cold or dry soils, warm days, high irradiance and/or freezing nights). Indeed, trees under water stress would develop a higher sensitivity to freezing night and photooxidative stress. We therefore proposed mitigation actions to avoid exposing trees to stressful conditions based on e.g. stand characteristics, understorey vegetation and planting. [ABSTRACT FROM AUTHOR]

Summary: Mixing species with contrasting resource use strategies could reduce forest vulnerability to extreme events. Yet, how species diversity affects seedling hydraulic responses to heat and drought, including mortality risk, is largely unknown.Using open‐top chambers, we assessed how, over several years, species interactions (monocultures vs mixtures) modulate heat and drought impacts on the hydraulic traits of juvenile European beech and pubescent oak. Using modeling, we estimated species interaction effects on timing to drought‐induced mortality and the underlying mechanisms driving these impacts.We show that mixtures mitigate adverse heat and drought impacts for oak (less negative leaf water potential, higher stomatal conductance, and delayed stomatal closure) but enhance them for beech (lower water potential and stomatal conductance, narrower leaf safety margins, faster tree mortality). Potential underlying mechanisms include oak's larger canopy and higher transpiration, allowing for quicker exhaustion of soil water in mixtures.Our findings highlight that diversity has the potential to alter the effects of extreme events, which would ensure that some species persist even if others remain sensitive. Among the many processes driving diversity effects, differences in canopy size and transpiration associated with the stomatal regulation strategy seem the primary mechanisms driving mortality vulnerability in mixed seedling plantations. See also the Commentary on this article by Wright, 241: 955–957. [ABSTRACT FROM AUTHOR]

Drought-induced plant mortality, resulting from either hydraulic failure or carbon starvation, is hypothesized to be modulated by the drought intensity. However, there is a paucity of research investigating the response strategies in desert shrubs under drought stress with different intensities. We transplanted potted Reaumuria soongorica (Pall.) Maxim. and Salsola passerina Bunge seedlings in the rain-out shelter, and implemented three water treatments: a control (well-watered, CK), a chronic drought (gradually less watered, CD), and a flash drought (not watered, FD). We then quantified plant physiological traits associated with water use and carbon assimilation. Both R. soongorica and S. passerina showed similar changes in water use and carbon characteristics under different drought treatments. Water use efficiency was not significantly changed, but embolism resistance was significantly lower in CD, and leaf specific conductivity and embolism resistance were significantly lower in FD compared to CK. Under the drought treatment, both shrubs had significantly lower hydraulic safety margins than CK, with FD being significantly lower than CD. Notably, FD had lower carbon assimilation and a lower leaf non-structural carbon concentration, but higher stem non-structural carbon concentration. The results of a principal component analysis showed that net photosynthetic rate, sapwood specific conductivity, embolism resistance, midday water potential, and leaf and stem soluble sugar concentration were the main axes of variation for R. soongorica traits. CK had the highest water use efficiency, net photosynthetic rate, and gas exchange rate, while FD had the lowest embolism resistance and highest osmoregulation. Midday water potential, leaf and stem soluble sugar concentration were the main axes of variation for S. passerina traits, and individual distribution under three water treatments was associated with drought tolerance traits. The findings suggest that species exhibit different response strategies for resistance to drought stress, with R. soongorica being drought-avoidant and S. passerina being drought-tolerant. These findings highlight the adaptive capacity of desert shrubs to water deficit and provide insights for assessing hydraulic failure and carbon starvation in desert shrubs. [ABSTRACT FROM AUTHOR]

Aim: Understanding tree mortality under climate change‐induced droughts requires knowledge of hydraulic trait distribution across environmental gradients. The spatial distribution of embolism resistance (i.e., P50), a key trait linked to hydraulic failure, is currently unknown across the Amazon. Here, we tested how precipitation, water table depth (WTD) and soil fertility interact as filters modulating the geographic hydraulic trait distribution. Location: Four sites in three key regions across the Amazon basin: Central, Western peripheral area and Brazilian Shield. Time Period: Present. Major Taxa Studied: Tree species. Method: We measured hydraulic vulnerability curves of 64 tree species (165 individuals) using an orthogonal design contrasting seasonal precipitation gradients, soil fertility and WTD. To estimate the geographical distribution of hydraulic traits for the basin scale, we use the coefficients of the best generalized linear model, projected on a 1 km grid in each environmental layer. Results: We show that WTD and soil fertility are the main drivers of embolism resistance, while precipitation has a secondary effect. Trees in shallow WTD and fertile soils had riskier hydraulic strategies. Our spatial projection identified the Amazonian southern band and valleys across the basin as the most vulnerable areas. High soil fertility in these regions increases the variability of hydraulic traits and thus uncertainty in predictions of drought responses. Main Conclusion: At the scale of the Amazon basin, we show that dominant tree species traits converge toward hydraulic resistance when resource (water and nutrients) levels are low and towards hydraulic vulnerability (but with higher trait variability) when resources increase. These results highlight the importance of WTD and soil fertility in structuring hydraulic traits and their effects on tree mortality under climate change‐induced droughts. Further empirical tests covering a wider spatial range and more tree species will help validate our biogeographical distribution model. [ABSTRACT FROM AUTHOR]

The hydraulic death hypothesis suggests that fires kill trees by damaging the plant's hydraulic continuum in addition to stem cambium. A corollary to this hypothesis is that plants that survive fires possess 'pyrohydraulic' traits that prevent heat-induced embolism formation in the xylem and aid post-fire survival. We examine whether hydraulic segmentation within stem xylem may act as such a trait. To do so, we measured the percentage loss of conductance (PLC) and vulnerability to embolism axially along segments of branches exposed to heat plumes in two differing species, fire-tolerant Eucalyptus cladocalyx F. Muell and fire-sensitive Kiggelaria africana L. testing model predictions that fire-tolerant species would exhibit higher degrees of hydraulic segmentation (greater PLC in the distal parts of the branch than the basal) than fire-intolerant species (similar PLC between segments). Following exposure to a heat plume, K. africana suffered between 73 and 84% loss of conductance in all branch segments, whereas E. cladocalyx had 73% loss of conductance in whole branches, including the distal tips, falling to 29% in the most basal part of the branch. There was no evidence for differences in resistance segmentation between the species, and there was limited evidence for differences in distal vulnerability to embolism across the branches. Hydraulic segmentation in E. cladocalyx may enable it to resprout effectively post-fire with a functional hydraulic system. The lack of hydraulic segmentation in K. africana reveals the need to understand possible trade-offs associated with hydraulic segmentation in long-lived woody species with respect to drought and fire. [ABSTRACT FROM AUTHOR]

Masson pine (Pinus massoniana Lamb.) is a dominant coniferous species in southern China, known for its rapid growth, abundant yield, and extensive utilization. Despite the robust adaptability of Masson pine and the rich annual precipitation in its distribution areas, this species still faces the mortality risk caused by the recurrent high temperatures in summer and low precipitation in subtropical regions. The mortality risk of Masson pine may increase in the future when facing a more frequent or intensive drought threat due to climate change. In this study, we conducted a manipulated drought experiment accompanying high temperature (~32.3 ± 0.7 °C in daytime and 28 °C in nighttime) to simulate a flash drought, aiming to explore the composite physiological response (hydraulic, gas exchange, and nonstructural carbon (NSC) characteristics) of Masson pine seedlings to extreme drought characterized by a high intensity and long duration. We found that, as the drought developed, the leaf water potential and gas exchange traits (net photosynthesis rate, stomatal conductance, and transpiration) significantly decreased while the percentage loss of hydraulic conductivity (PLC) significantly increased. In contrast, NSC remained a more constant trend before it was significantly reduced on day 30 after the beginning of the drought. Except for NSC, all the other traits had significant correlations between them. Additionally, hydraulic dysfunction indicated by the increasing PLC preceded the NSC depletion, which may indicate a more significant role for hydraulic failure than carbon starvation in drought-induced mortality. Conclusively, hydraulic and gas exchange traits showed a coupling response to drought, but NSC displayed an independent dynamic. The findings may improve our understanding of drought-coping strategies of Masson pine and provide some theoretical basis for Masson pine forest management. [ABSTRACT FROM AUTHOR]

The subtropical regions in China are prone to recurrent summer droughts induced by the Western Pacific Subtropical High-Pressure, which has induced the death of tens of millions of culms of Moso bamboo (Phyllostachys edulis (Carriere) J. Houzeau), a widely distributed giant bamboo with high economic and ecological values. In the future, the intensity and frequency of the summer drought are projected to increase in these areas due to global climate change, which may lead to significant age-specific mortality of Moso bamboo. So far, it is still unclear about the age-specific response mechanisms of hydraulic traits and carbon balance of Moso bamboo when it is suffering to an ongoing summer drought. This study aimed to investigate the hydraulic and photosynthetic responses of newly sprouted (1 year old) and established (2-5 years old) culms of Moso bamboo to summer drought, which was manipulated by throughfall reduction in Lin'an of Zhejiang. The results showed that both newly sprouted and established culms had a gradually weakening hydraulic conductivity and photosynthesis during the whole drought process. In the early stage of the manipulated drought, the established culms had more loss of hydraulic conductivity than the newly sprouted culms. However, the newly sprouted culms had significant more loss of hydraulic conductivity and lower photosynthetic rates and stomatal conductance in the middle and late stages of the manipulated drought. The results suggest that the newly sprouted culms were more susceptible to summer drought than established culms due to the combined effects of hydraulic damage and photosynthetic restriction, explaining why the newly sprouted culms have higher mortality than elder culms when subjected to extreme drought. These findings provided insights into the mechanisms of Moso bamboo's age-specific drought-induced mortality, which will help for the anti-drought management of bamboo. [ABSTRACT FROM AUTHOR]

Summary: Relative sea level rise (SLR) increasingly impacts coastal ecosystems through the formation of ghost forests. To predict the future of coastal ecosystems under SLR and changing climate, it is important to understand the physiological mechanisms underlying coastal tree mortality and to integrate this knowledge into dynamic vegetation models.We incorporate the physiological effect of salinity and hypoxia in a dynamic vegetation model in the Earth system land model, and used the model to investigate the mechanisms of mortality of conifer forests on the west and east coast sites of USA, where trees experience different form of sea water exposure.Simulations suggest similar physiological mechanisms can result in different mortality patterns. At the east coast site that experienced severe increases in seawater exposure, trees loose photosynthetic capacity and roots rapidly, and both storage carbon and hydraulic conductance decrease significantly within a year. Over time, further consumption of storage carbon that leads to carbon starvation dominates mortality. At the west coast site that gradually exposed to seawater through SLR, hydraulic failure dominates mortality because root loss impacts on conductance are greater than the degree of storage carbon depletion.Measurements and modeling focused on understanding the physiological mechanisms of mortality is critical to reducing predictive uncertainty. [ABSTRACT FROM AUTHOR]

Survival and growth of woody species in the Mediterranean are mainly restricted by water availability. We tested the hypothesis that Mediterranean species acclimate their xylem vulnerability and osmotic potential along a precipitation gradient.We studied five predominant co‐occurring Mediterranean species; Quercus calliprinos, Pistacia palaestina, Pistacia lentiscus, Rhamnus lycioides and Phillyrea latifolia, over two summers at three sites. The driest of the sites is the distribution edge for all the five species. We measured key hydraulic and osmotic traits related to drought resistance, including resistance to embolism (Ψ50) and the seasonal dynamics of water and osmotic potentials.The leaf water potentials (Ψl) of all species declined significantly along the summer, reaching significantly lower Ψl at the end of summer in the drier sites. Surprisingly, we did not find plasticity along the drought gradient in Ψ50 or osmotic potentials. This resulted in much narrower hydraulic safety margins (HSMs) in the drier sites, where some species experienced significant embolism.Our analysis indicates that reduction in HSM to null values put Mediterranean species in embolism risk as they approach their hydraulic limit near the geographical dry edge of their distribution. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

The characterisation of ecological strategies to predict drought response is still lacking for Caribbean dry forest seedlings. This study documents growth characteristics and tolerance to drought via xylem hydraulic and leaf cell properties of three dominant native species of the Caribbean dry forest. Twenty morphological and physiological traits were assessed in Citharexylum spinosum, Guaiacum officinale and Guapira fragrans in greenhouse conditions. The seedlings displayed contrasting growth rates, which were positively correlated with the capacity to quickly develop a large leaf area and root fraction. The three species had a similar xylem tolerance to embolism (P50: −4 MPa) but differed in leaf cell tolerance to dehydration, which was negatively correlated with RGR (R2 > 0.87). The slowest-growing, G. officinale, had high leaf tolerance to cell dehydration due to low ΨTLP and πo, but displayed a narrow hydraulic safety margin. The leaves of the fast-growing C. spinosum were sensitive to leaf dehydration but exhibited a surprisingly wide stem hydraulic safety margin. G. fragrans had intermediate traits. Our results showed that dry forest seedling growth in similar environments can exhibit distinct carbon growth strategies as well as contrasting water-use strategies, primarily as they relate to drought resistance, due to variation in root development and leaf cell resistance to dehydration. Our study thus provides an approach to estimate species performance under drought conditions. [ABSTRACT FROM AUTHOR]

Xylem anatomy may change in response to environmental or biotic stresses. Vascular occlusion, an anatomical modification of mature xylem, contributes to plant resistance and susceptibility to different stresses. In woody organs, xylem occlusions have been examined as part of the senescence process, but their presence and function in leaves remain obscure. In grapevine, many stresses are associated with premature leaf senescence inducing discolorations and scorched tissue in leaves. However, we still do not know whether the leaf senescence process follows the same sequence of physiological events and whether leaf xylem anatomy is affected in similar ways. In this study, we quantified vascular occlusions in midribs from leaves with symptoms of the grapevine disease esca, magnesium deficiency and autumn senescence. We found higher amounts of vascular occlusions in leaves with esca symptoms (in 27% of xylem vessels on average), whereas the leaves with other symptoms (as well as the asymptomatic controls) had far fewer occlusions (in 3% of vessels). Therefore, we assessed the relationship between xylem occlusions and esca leaf symptoms in four different countries (California in the USA, France, Italy and Spain) and eight different cultivars. We monitored the plants over the course of the growing season, confirming that vascular occlusions do not evolve with symptom age. Finally, we investigated the hydraulic integrity of leaf xylem vessels by optical visualization of embolism propagation during dehydration. We found that the occlusions lead to hydraulic dysfunction mainly in the peripheral veins compared with the midribs in esca symptomatic leaves. These results open new perspectives on the role of vascular occlusions during the leaf senescence process, highlighting the uniqueness of esca leaf symptoms and its consequence on leaf physiology. [ABSTRACT FROM AUTHOR]

Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

Considerable evidences highlight the occurrence of increasing widespread tree mortality as a result of global climate change-associated droughts. However, knowledge about the mechanisms underlying divergent strategies of various tree species to adapt to drought has remained remarkably insufficient. Leaf stomatal regulation and embolism resistance of stem xylem serves as two important strategies for tree species to prevent hydraulic failure and carbon starvation, as comprising interconnected physiological mechanisms underlying drought-induced tree mortality. Hence, the physiological and anatomical determinants of leaf stomatal regulation and stems xylem embolism resistance are evaluated and discussed. In addition, root properties related to drought tolerance are also reviewed. Species with greater investment in leaves and stems tend to maintain stomatal opening and resist stem embolism under drought conditions. The coordination between stomatal regulation and stem embolism resistance are summarized and discussed. Previous studies showed that hydraulic safety margin (HSM, the difference between minimum water potential and that causing xylem dysfunction) is a significant predictor of tree species mortality under drought conditions. Compared with HSM, stomatal safety margin (the difference between water potential at stomatal closure and that causing xylem dysfunction) more directly merge stomatal regulation strategies with xylem hydraulic strategies, illustrating a comprehensive framework to characterize plant response to drought. A combination of plant traits reflecting species' response and adaptation to drought should be established in the future, and we propose four specific urgent issues as future research priorities. [ABSTRACT FROM AUTHOR]

Summary: Increasing seawater exposure is killing coastal trees globally, with expectations of accelerating mortality with rising sea levels. However, the impact of concomitant changes in atmospheric CO2 concentration, temperature, and vapor pressure deficit (VPD) on seawater‐induced tree mortality is uncertain.We examined the mechanisms of seawater‐induced mortality under varying climate scenarios using a photosynthetic gain and hydraulic cost optimization model validated against observations in a mature stand of Sitka spruce (Picea sitchensis) trees in the Pacific Northwest, USA, that were dying from recent seawater exposure.The simulations matched well with observations of photosynthesis, transpiration, nonstructural carbohydrates concentrations, leaf water potential, the percentage loss of xylem conductivity, and stand‐level mortality rates. The simulations suggest that seawater‐induced mortality could decrease by c. 16.7% with increasing atmospheric CO2 levels due to reduced risk of carbon starvation. Conversely, rising VPD could increase mortality by c. 5.6% because of increasing risk of hydraulic failure.Across all scenarios, seawater‐induced mortality was driven by hydraulic failure in the first 2 yr after seawater exposure began, with carbon starvation becoming more important in subsequent years. Changing CO2 and climate appear unlikely to have a significant impact on coastal tree mortality under rising sea levels. [ABSTRACT FROM AUTHOR]

To study how gap-graded soil reinforced with fibers of different types restrains suffusion in embankment dams, the hydraulic response of fiber-reinforced soil (FRS) containing polypropylene fibers (PPF), coir fibers (CF) or basalt-tape fibers (BTF) was investigated in one-dimensional seepage tests. The critical hydraulic gradient and fine-particle loss of the FRS were analysed quantitatively by changing the fiber type, fiber content, and reinforcement-layer thickness. The test results show that the three types of FRS all had restraining effect on the development of suffusion. The netting reinforcement effect of PPF was the best, followed by the water absorption and expansion of CF to reduce pore volume, and finally the effect of BTF was to block migration channels under the same reinforcement conditions. With increasing fiber content and the reinforcement-layer thickness, the critical onset hydraulic gradient (icr,onset), and critical failure hydraulic gradient (icr,failure) increased, improving the internal erosion resistance of gap-graded soils against suffusion. The fine-particles loss decreased and the restraint rate of fine-particle loss (RF) increased meant that the permeability stability of soil was enhanced. Linear regression analysis showed that the reinforcement-layer thickness has greater influence on restraining effect parameters than fiber content.

Acoustic emission analysis is promising to investigate the physiological events leading to drought-induced injury and mortality. However, their nature and source are not fully understood, making this technique difficult to use as a direct measure of the loss of xylem hydraulic conductance. Acoustic emissions were recorded during severe dehydration in lavender plants (Lavandula angustifolia) and compared with the dynamics of embolism development and cell damage. The timing and characteristics of acoustic signals from two independent recording systems were compared by principal component analysis (PCA). Changes in water potential, branch diameter, loss of hydraulic conductance, and cellular damage were also measured to quantify drought-induced damages. Two distinct phases of acoustic emissions were observed during dehydration: the first one associated with a rapid loss of diameter and a significant increase in loss of xylem conductance (90%), and the second with slower changes in diameter and a significant increase in cellular damage. Based on PCA, a developed algorithm discriminated hydraulic-related acoustic signals from other sources, proposing a reconstruction of hydraulic vulnerability curves. Cellular damage preceded by hydraulic failure seems to lead to a lack of recovery. The second acoustic phase would allow detection of plant mortality. [ABSTRACT FROM AUTHOR]

Accurately predicting global drought-induced tree mortality remains a major challenge facing plant science and ecology. Stem hydraulic safety margin (HSM, the difference between water potential at the minimum value and the value that causes xylem vulnerability to embolism) performs as one of the best hydraulic traits in predicting global drought-induced tree mortality, however, HSM is time-consuming and very difficult to measure. We proposed to use leaf turgor loss point (TLP, the water potential at which leaves start to wilt) as a proxy for HSM because HSM may be highly correlated to TLP, as both of them are tightly linked with water potential changes after stomatal closure. Since TLP is more easy and rapid to measure than HSM, if we find strong HSM-TLP relationships at the global scale, TLP can be used in predicting global drought-induced tree mortality. However, no study has quantified the relationships between HSM and TLP at the global scale. Here we draw together published data on HSM and TLP for 1,773 species from 370 sites worldwide to check whether HSM and TLP are highly associated. We found that HSMs and TLPs are merely highly related in tropical forests, thus TLP can be a reliable surrogate of HSM to predict drought-induced tree mortality in tropical forest. Here we are certainly not advocating for the use of TLP instead of HSM to predict drought-induced tree mortality in tropical forests, but simply for predicting drought-induced tree mortality in tropical forests in supplementary of HSM in the future.

Between late 2015 and early 2016, more than 7,000 ha of mangrove forest died along the coastline of the Gulf of Carpentaria, in northern Australia. This massive die-off was preceded by a strong 2015/2016 El Niño event, resulting in lower precipitation, a drop in sea level and higher than average temperatures in northern Australia. In this study, we investigated the role of hydraulic failure in the mortality and recovery of the dominant species, Avicennia marina , 2 years after the mortality event. We measured predawn water potential (Ψpd) and percent loss of stem hydraulic conductivity (PLC) in surviving individuals across a gradient of impact. We also assessed the vulnerability to drought-induced embolism (Ψ50) for the species. Areas with severe canopy dieback had higher native PLC (39%) than minimally impacted areas (6%), suggesting that hydraulic recovery was ongoing. The high resistance of A. marina to water-stress-induced embolism (Ψ50 = −9.6 MPa), indicates that severe water stress (Ψpd < −10 MPa) would have been required to cause mortality in this species. Our data indicate that the natural gradient of water-stress enhanced the impact of El Niño, leading to hydraulic failure and mortality in A. marina growing on severely impacted (SI) zones. It is likely that lowered sea levels and less frequent inundation by seawater, combined with lower inputs of fresh water, high evaporative demand and high temperatures, led to the development of hyper-salinity and extreme water stress during the 2015/16 summer. [ABSTRACT FROM AUTHOR]

Drought-related tree mortality has become a major concern worldwide due to its pronounced negative impacts on the functioning and sustainability of forest ecosystems. However, our ability to identify the species that are most vulnerable to drought, and to pinpoint the spatial and temporal patterns of mortality events, is still limited. Model is useful tools to capture the dynamics of vegetation at spatiotemporal scales, yet contemporary land surface models (LSMs) are often incapable of predicting the response of vegetation to environmental perturbations with sufficient accuracy, especially under stressful conditions such as drought. Significant progress has been made regarding the physiological mechanisms underpinning plant drought response in the past decade, and plant hydraulic dysfunction has emerged as a key determinant for tree death due to water shortage. The identification of pivotal physiological events and relevant plant traits may facilitate forecasting tree mortality through a mechanistic approach, with improved precision. In this review, we (1) summarize current understanding of physiological mechanisms leading to tree death, (2) describe the functionality of key hydraulic traits that are involved in the process of hydraulic dysfunction, and (3) outline their roles in improving the representation of hydraulic function in LSMs. We urge potential future research on detailed hydraulic processes under drought, pinpointing corresponding functional traits, as well as understanding traits variation across and within species, for a better representation of drought-induced tree mortality in models. [ABSTRACT FROM AUTHOR]

Between late 2015 and early 2016, more than 7,000 ha of mangrove forest died along the coastline of the Gulf of Carpentaria, in northern Australia. This massive die-off was preceded by a strong 2015/2016 El Niño event, resulting in lower precipitation, a drop in sea level and higher than average temperatures in northern Australia. In this study, we investigated the role of hydraulic failure in the mortality and recovery of the dominant species, Avicennia marina, 2 years after the mortality event. We measured predawn water potential (Ψpd) and percent loss of stem hydraulic conductivity (PLC) in surviving individuals across a gradient of impact. We also assessed the vulnerability to drought-induced embolism (Ψ50) for the species. Areas with severe canopy dieback had higher native PLC (39%) than minimally impacted areas (6%), suggesting that hydraulic recovery was ongoing. The high resistance of A. marina to water-stress-induced embolism (Ψ50 = −9.6 MPa), indicates that severe water stress (Ψpd < −10 MPa) would have been required to cause mortality in this species. Our data indicate that the natural gradient of water-stress enhanced the impact of El Niño, leading to hydraulic failure and mortality in A. marina growing on severely impacted (SI) zones. It is likely that lowered sea levels and less frequent inundation by seawater, combined with lower inputs of fresh water, high evaporative demand and high temperatures, led to the development of hyper-salinity and extreme water stress during the 2015/16 summer.

Drought-related tree mortality has become a major concern worldwide due to its pronounced negative impacts on the functioning and sustainability of forest ecosystems. However, our ability to identify the species that are most vulnerable to drought, and to pinpoint the spatial and temporal patterns of mortality events, is still limited. Model is useful tools to capture the dynamics of vegetation at spatiotemporal scales, yet contemporary land surface models (LSMs) are often incapable of predicting the response of vegetation to environmental perturbations with sufficient accuracy, especially under stressful conditions such as drought. Significant progress has been made regarding the physiological mechanisms underpinning plant drought response in the past decade, and plant hydraulic dysfunction has emerged as a key determinant for tree death due to water shortage. The identification of pivotal physiological events and relevant plant traits may facilitate forecasting tree mortality through a mechanistic approach, with improved precision. In this review, we (1) summarize current understanding of physiological mechanisms leading to tree death, (2) describe the functionality of key hydraulic traits that are involved in the process of hydraulic dysfunction, and (3) outline their roles in improving the representation of hydraulic function in LSMs. We urge potential future research on detailed hydraulic processes under drought, pinpointing corresponding functional traits, as well as understanding traits variation across and within species, for a better representation of drought-induced tree mortality in models.

European beech is one of the most important deciduous tree species in natural forest ecosystems in Central Europe. Its dominance is now being questioned by the emerging drought damages due to the increased incidence of severe summer droughts. In Switzerland, Fagus sylvatica have been observed in the Intercantonal Forest Observation Program since 1984. The dataset presented here includes 179176 annual observations of beech trees on 102 plots during 37 years. The plots cover gradients in drought, nitrogen deposition, ozone, age, altitude, and soil chemistry. In dry regions of Switzerland, the dry and hot summer of 2018 caused a serious branch dieback, increased mortality in Fagus sylvatica and increased yellowing of leaves. Beech trees recovered less after 2018 than after the dry summer 2003 which had been similar in drought intensity except that the drought in 2018 started earlier in spring. Our data analyses suggest the importance of drought in subsequent years for crown transparency and mortality in beech. The drought in 2018 followed previous dry years of 2015 and 2017 which pre-weakened the trees. Our long-term data indicate that the drought from up to three previous years were significant predictors for both tree mortality and for the proportion of trees with serious (>60%) crown transparency. The delay in mortality after the weakening event suggests also the importance of weakness parasites. The staining of active vessels with safranine revealed that the cavitation caused by the low tree water potentials in 2018 persisted at least partially in 2019. Thus, the ability of the branches to conduct water was reduced and the branches dried out. Furthermore, photooxidation in light-exposed leaves has increased strongly since 2011. This phenomenon was related to low concentrations of foliar phosphorus (P) and hot temperatures before leaf harvest. The observed drought effects can be categorized as (i) hydraulic failure (branch dieback), (ii) energy starvation as a consequence of closed stomata and P deficiency (photooxidation) and (iii) infestation with weakness parasites (beech bark disease and root rots).

Hydraulic failure caused by severe drought contributes to aboveground dieback and whole‐plant death. The extent to which dieback or whole‐plant death can be predicted by plant hydraulic traits has rarely been tested among species with different leaf habits and/or growth forms. We investigated 19 hydraulic traits in 40 woody species in a tropical savanna and their potential correlations with drought response during an extreme drought event during the El Niño–Southern Oscillation in 2015. Plant hydraulic trait variation was partitioned substantially by leaf habit but not growth form along a trade‐off axis between traits that support drought tolerance versus avoidance. Semi‐deciduous species and shrubs had the highest branch dieback and top‐kill (complete aboveground death) among the leaf habits or growth forms. Dieback and top‐kill were well explained by combining hydraulic traits with leaf habit and growth form, suggesting integrating life history traits with hydraulic traits will yield better predictions. [ABSTRACT FROM AUTHOR]

Xylella fastidiosa is a xylem-limited bacterium causing the Olive Quick Decline Syndrome, which is currently devastating the agricultural landscape of Southern Italy. The bacterium is injected into the xylem vessels of leaf petioles after the penetration of the insect vector's stylet. From here, it is supposed to colonize the xylem vasculature moving against water flow inside conductive vessels. Widespread vessel clogging following the bacterial infection and causing the failure of water transport seemed not to fully supported by the recent empirical xylem anatomical observations in infected olive trees. We tested the hypothesis that the higher susceptibility to the X. fastidiosa 's infection in Cellina di Nardò compared with Leccino is associated to the higher vulnerability to air embolism of its larger vessels. Such hypothesis is motivated by the recognized ability of X. fastidiosa in degrading pit membranes and also because air embolism would possibly provide microenvironmental conditions more favourable to its more efficient aerobic metabolism. We revised the relevant literature on bacterium growth and xylem physiology, and carried out empirical field, mid-summer measurements of xylem anatomy and native embolism in olive cultivars with high (Cellina di Nardò) and low susceptibility (Leccino) to the infection by X. fastidiosa. Both cultivars had similar shoot mass traits and vessel length (~80 cm), but the highly susceptible one had larger vessels and a lower number of vessels supplying a given leaf mass. Native air embolism reduced mean xylem hydraulic conductance by ~58 % (Cellina di Nardò) and ~38 % (Leccino). The higher air-embolism vulnerability of the larger vessels in Cellina di Nardò possibly facilitates the X. fastidiosa 's infection compared to Leccino. Some important characteristics of the vector–pathogen–plant interactions still require deep investigations acknowledging both the pathogen metabolic pathways and the biophysical principles of xylem hydraulics. [ABSTRACT FROM AUTHOR]

SummaryModel-data comparisons of plant physiological processes provide an understanding of mechanisms underlying vegetation responses to climate. We simulated the physiology of a piñon pine-juniper woodland (Pinus edulis-Juniperus monosperma) that experienced mortality during a 5 yr precipitation-reduction experiment, allowing a framework with which to examine our knowledge of drought-induced tree mortality. We used six models designed for scales ranging from individual plants to a global level, all containing state-of-the-art representations of the internal hydraulic and carbohydrate dynamics of woody plants. Despite the large range of model structures, tuning, and parameterization employed, all simulations predicted hydraulic failure and carbon starvation processes co-occurring in dying trees of both species, with the time spent with severe hydraulic failure and carbon starvation, rather than absolute thresholds per se, being a better predictor of impending mortality. Model and empirical data suggest that limited carbon and water exchanges at stomatal, phloem, and below-ground interfaces were associated with mortality of both species. The model-data comparison suggests that the introduction of a mechanistic process into physiology-based models provides equal or improved predictive power over traditional process-model or empirical thresholds. Both biophysical and empirical modeling approaches are useful in understanding processes, particularly when the models fail, because they reveal mechanisms that are likely to underlie mortality. We suggest that for some ecosystems, integration of mechanistic pathogen models into current vegetation models, and evaluation against observations, could result in a breakthrough capability to simulate vegetation dynamics.

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]

Hydraulic failure has been extensively studied during drought-induced plant dieback, but its role in plant-pathogen interactions is under debate. During esca, a grapevine (Vitis vinifera) disease, symptomatic leaves are prone to irreversible hydraulic dysfunctions but little is known about the hydraulic integrity of perennial organs over the short- and long-term. We investigated the effects of esca on stem hydraulic integrity in naturally infected plants within a single season and across season(s). We coupled direct (k s ) and indirect (k th ) hydraulic conductivity measurements, and tylose and vascular pathogen detection with in vivo X-ray microtomography visualizations. Xylem occlusions (tyloses) and subsequent loss of stem hydraulic conductivity (k s ) occurred in all shoots with severe symptoms (apoplexy) and in more than 60% of shoots with moderate symptoms (tiger-stripe), with no tyloses in asymptomatic shoots. In vivo stem observations demonstrated that tyloses occurred only when leaf symptoms appeared, and resulted in more than 50% loss of hydraulic conductance in 40% of symptomatic stems, unrelated to symptom age. The impact of esca on xylem integrity was only seasonal, with no long-term impact of disease history. Our study demonstrated how and to what extent a vascular disease such as esca, affecting xylem integrity, could amplify plant mortality through hydraulic failure. [ABSTRACT FROM AUTHOR]

Understanding the vulnerability of trees to drought-induced mortality is key to predicting the fate of forests in a future climate with more frequent and intense droughts, although the underlying mechanisms are difficult to study in adult trees. Here, we explored the dynamic changes of water relations and limits of hydraulic function in dying adults of Norway spruce (Picea abies L.) during the progression of the record-breaking 2018 Central European drought. In trees on the trajectory to drought-induced mortality, we observed rapid, nonlinear declines of xylem pressure that commenced at the early onset of xylem cavitation and caused a complete loss of xylem hydraulic conductance within a very short time. We also observed severe depletions of nonstructural carbohydrates, though carbon starvation could be ruled out as the cause of the observed tree death, as both dying and surviving trees showed these metabolic limitations. Our observations provide striking field-based evidence for fast dehydration and hydraulic collapse as the cause of drought-induced mortality in adult Norway spruce. The nonlinear decline of tree water relations suggests that considering the temporal dynamics of dehydration is critical for predicting tree death. The collapse of the hydraulic system within a short time demonstrates that trees can rapidly be pushed out of the zone of hydraulic safety during the progression of a severe drought. In summary, our findings point toward a higher mortality risk for Norway spruce than previously assumed, which is in line with current reports of unprecedented levels of droughtinduced mortality in this major European tree species. [ABSTRACT FROM AUTHOR]

A recent data synthesis showed that trees dehydrated to SB lethal sb experienced a > 60% loss of xylem conductivity, suggesting a ubiquitous hydraulic failure in drought-induced tree mortality (Adams I et al i ., 2017). Water potential threshold across species and biomes Across all tree species assessed, the SB lethal sb ranged widely from -1.5 MPa ( I Populus balsamifera i , an angiosperm) to -14.7 MPa ( I Callitris columellaris i , a gymnosperm) (Fig. In sum, our synthesis of data showed that tree SB lethal sb ranged widely from -1.5 to -14.7 MPa across species and biomes, and the SB lethal sb was significantly lower for gymnosperms than angiosperms. Tree SB lethal sb of gymnosperms was close to P50, while for angiosperms, tree SB lethal sb was closer to P88. [Extracted from the article]

Summary: Plants continue to lose water from their leaves even after complete stomatal closure. Although this minimum conductance (gleaf‐res) has substantial impacts on strategies of water use and conservation, little is known about the potential drivers underlying the variability of this trait across species.We thus untangled the relative contribution of water leaks from the cuticle and stomata in order to investigate how the variability in leaf morphological and anatomical traits is related to the variation in gleaf‐res and carbon assimilation capacity across 30 diverse species from the Brazilian Cerrado.In addition to cuticle permeance, water leaks from stomata had a significant impact on gleaf‐res. The differential pattern of stomata distribution in the epidermis was a key factor driving this variation, suggesting the existence of a trade‐off between carbon assimilation and water loss through gleaf‐res. For instance, higher gleaf‐res, observed in fast‐growing species, was associated with the investment in small and numerous stomata, which allowed higher carbon assimilation rates but also increased water leaks, with negative impacts on leaf survival under drought. Variation in cuticle structural properties was not linked to gleaf‐res.Our results therefore suggest the existence of a trade‐off between carbon assimilation efficiency and dehydration tolerance at foliar level. [ABSTRACT FROM AUTHOR]

Drought limits the long-distance transport of water in the xylem due to the reduced leaf-to-soil water potential difference and possible embolism-related losses of conductance and of sugars in the phloem due to the higher viscosity of the dehydrated sugary solution. This condition can have cascading effects in water and carbon (C) fluxes that may ultimately cause tree death. We hypothesize that the maintenance of xylem and phloem conductances is fundamental for survival also under reduced resource availability, when trees may produce effective and low C cost anatomical adjustments in the xylem and phloem close to the treetop where most of the hydraulic resistance is concentrated. We analyzed the treetop xylem and phloem anatomical characteristics in coexisting Scots pine trees, symptomatic and non-symptomatic of drought-induced dieback. We selected the topmost 55 cm of the main stem and selected several sampling positions at different distances from the stem apex to test for differences in the axial patterns between the two groups of trees. We measured the annual ring area, the tracheid hydraulic diameter (Dh) and cell wall thickness (CWT), the conductive phloem area and the average lumen diameter of the 20 largest phloem sieve cells (Dph). Declining trees grew less than the non-declining ones, and despite the similar axial scaling of anatomical traits, had larger Dh and lower CWT. Moreover, declining trees had wider Dph. Our results demonstrate that even under drought stress, maintenance of xylem and phloem efficiencies is of primary importance for survival, even if producing fewer larger tracheids may lead to a xylem more vulnerable to embolism formation. [ABSTRACT FROM AUTHOR]

In 2018, Central Europe experienced one of the most severe and long-lasting summer drought and heat wave ever recorded. Before 2018, the 2003 millennial drought was often invoked as the example of a "hotter drought", and was classified as the most severe event in Europe for the last 500 years. First insights now confirm that the 2018 drought event was climatically more extreme and had a greater impact on forest ecosystems of Austria, Germany and Switzerland than the 2003 drought. Across this region, mean growing season air temperature from April to October was more than 3.3°C above the long-term average, and 1.2°C warmer than in 2003. Here, we present a first impact assessment of the severe 2018 summer drought and heatwave on Central European forests. In response to the 2018 event, most ecologically and economically important tree species in temperate forests of Austria, Germany and Switzerland showed severe signs of drought stress. These symptoms included exceptionally low foliar water potentials crossing the threshold for xylem hydraulic failure in many species and observations of widespread leaf discoloration and premature leaf shedding. As a result of the extreme drought stress, the 2018 event caused unprecedented drought-induced tree mortality in many species throughout the region. Moreover, unexpectedly strong drought-legacy effects were detected in 2019. This implies that the physiological recovery of trees was impaired after the 2018 drought event, leaving them highly vulnerable to secondary drought impacts such as insect or fungal pathogen attacks. As a consequence, mortality of trees triggered by the 2018 events is likely to continue for several years. Our assessment indicates that many common temperate European forest tree species are more vulnerable to extreme summer drought and heat waves than previously thought. As drought and heat events are likely to occur more frequently with the progression of climate change, temperate European forests might approach the point for a substantial ecological and economic transition. Our assessment also highlights the urgent need for a pan-European ground-based monitoring network suited to track individual tree mortality, supported by remote sensing products with high spatial and temporal resolution to track, analyse and forecast these transitions. [ABSTRACT FROM AUTHOR]

Plant hydraulics originates from the study of water transport through the xylem, but is now extended to consider soil-root, leaf, and whole-plant transport processes, and is even now being applied at global scales via satellite remote sensing. Experimental evidence suggests that plants can survive drought when xylem embolism remains below critical thresholds (Nardini et al., [53]). Weak tradeoff between xylem safety and xylem-specific hydraulic efficiency across the world's woody plant species. X-ray microtomography observations of xylem embolism in stems of Laurus nobilis are consistent with hydraulic measurements of percentage loss of conductance. [Extracted from the article]

Summary: Catastrophic failure of the water transport pathway in trees is a principal mechanism of mortality during extreme drought. To be able to predict the probability of mortality at an individual and landscape scale we need knowledge of the time for plants to reach critical levels of hydraulic failure.We grew plants of eight species of Eucalyptus originating from contrasting climates before allowing a subset to dehydrate. We tested whether a trait‐based model of time to plant desiccation tcrit, from stomatal closure gs90 to a critical level of hydraulic dysfunction Ψcrit is consistent with observed dry‐down times.Plant desiccation time varied among species, ranging from 96.2 to 332 h at a vapour‐pressure deficit of 1 kPa, and was highly predictable using the tcrit model in conjunction with a leaf shedding function. Plant desiccation time was longest in species with high cavitation resistance, strong vulnerability segmentation, wide stomatal‐hydraulic safety, and a high ratio of total plant water content to leaf area.Knowledge of tcrit in combination with water‐use traits that influence stomatal closure could significantly increase our ability to predict the timing of drought‐induced mortality at tree and forest scales. [ABSTRACT FROM AUTHOR]

Understanding which structural and functional traits are linked to species' vulnerability to embolism formation (P50) may provide fundamental knowledge on plant strategies to maintain an efficient water transport. We measured P50, wood density (WD), mean conduit area, conduit density, percentage areas occupied by vessels, parenchyma cells (PATOT) and fibers (FA) on branches of angiosperm and gymnosperm species. Moreover, we compiled a dataset of published hydraulic and anatomical data to be compared with our results. Species more vulnerable to embolism had lower WD. In angiosperms, the variability in WD was better explained by PATOT and FA, which were highly correlated. Angiosperms with a higher P50 (less negative) had a higher amount of PATOT and total amount of nonstructural carbohydrates. Instead, in gymnosperms, P50 vs PATOT was not significant. The correlation between PATOT and P50 might have a biological meaning and also suggests that the causality of the commonly observed relationship of WD vs P50 is indirect and dependent on the parenchyma fraction. Our study suggests that angiosperms have a potential active embolism reversal capacity in which parenchyma has an important role, while in gymnosperms this might not be the case. [ABSTRACT FROM AUTHOR]

Summary: Determining physiological mechanisms and thresholds for climate‐driven tree die‐off could help improve global predictions of future terrestrial carbon sinks. We directly tested for the lethal threshold in hydraulic failure – an inability to move water due to drought‐induced xylem embolism – in a pine sapling experiment.In a glasshouse experiment, we exposed loblolly pine (Pinus taeda) saplings (n = 83) to drought‐induced water stress ranging from mild to lethal. Before rewatering to relieve drought stress, we measured native hydraulic conductivity and foliar color change. We monitored all measured individuals for survival or mortality.We found a lethal threshold at 80% loss of hydraulic conductivity – a point of hydraulic failure beyond which it is more likely trees will die, than survive, and describe mortality risk across all levels of water stress. Foliar color changes lagged behind hydraulic failure – best predicting when trees had been dead for some time, rather than when they were dying.Our direct measurement of native conductivity, while monitoring the same individuals for survival or mortality, quantifies a continuous probability of mortality risk from hydraulic failure. Predicting tree die‐off events and understanding the mechanism involved requires knowledge not only of when trees are dead, but when they begin dying – having passed the point of no return. [ABSTRACT FROM AUTHOR]

Widespread drought-induced forest mortality (DIM) is expected to increase with climate change and drought, and is expected to have major impacts on carbon and water cycles. For large-scale assessment and management, it is critical to identify variables that integrate the physiological mechanisms of DIM and signal risk of DIM. We tested whether plant water content, a variable that can be remotely sensed at large scales, is a useful indicator of DIM risk at the population level. We subjected Pinus ponderosa Douglas ex C. Lawson seedlings to experimental drought using a point of no return experimental design. Periodically during the drought, independent sets of seedlings were sampled to measure physiological state (volumetric water content (VWC), percent loss of conductivity (PLC) and non-structural carbohydrates) and to estimate population-level probability of mortality through re-watering. We show that plant VWC is a good predictor of population-level DIM risk and exhibits a threshold-type response that distinguishes plants at no risk from those at increasing risk of mortality. We also show that plant VWC integrates the mechanisms involved in individual tree death: hydraulic failure (PLC), carbon depletion across organs and their interaction. Our results are promising for landscape-level monitoring of DIM risk. [ABSTRACT FROM AUTHOR]

A growing body of evidence highlights the occurrence of increased widespread tree mortality during climate change-associated severe droughts; however, in situ long-term drought experiments with multispecies communities for the prediction of tree mortality and exploration of related mechanisms are rather limited in natural environments. We conducted a 7-year afforestation trial with 20 drought-resistant broadleaf tree species in an arid limestone habitat in northern China, where the species displayed a broad range of survival rates. The stomatal and xylem hydraulic traits of all the species were measured. We found that species' stomatal closure points were strongly related to their xylem embolism resistance and xylem minimum water potential but not to their survival rates. Hydraulic failure of the vascular system appeared to be the main cause of tree mortality, and the stomatal safety margin was a better predictor of tree mortality than the traditionally considered xylem embolism resistance and hydraulic safety margin. We recommend the stomatal safety margin as the indicator for predicting drought-induced tree mortality and for selecting tree species in future forest restorations in arid regions. [ABSTRACT FROM AUTHOR]

Forests are sensitive to droughts, which increase the mortality rate of tree species. Various processes have been proposed to underlie drought-induced tree mortality, including hydraulic failure, carbon starvation and increased susceptibility to natural enemies. To give insights into these processes, we assessed the metabolic effects of a mortality-inducing drought on seedlings of Pinus sylvestris L. (Scots Pine), a widespread and important Eurasian species. We found divergence over time in the foliar metabolic composition of droughted vs well-watered seedlings, with the former showing increased abundance of aromatic amino acids and decreases in secondary metabolism associated with defence. We observed no significant differences amongst provenances in these effects: seedlings from drought-prone areas showed the same foliar metabolic changes under drought as seedlings from moist environments, although morphological effects of drought varied by provenance. Overall, our results demonstrate how severe drought prior to death may target particular primary and secondary metabolic pathways, weakening defences against natural enemies and contributing to the risk of drought-induced mortality in P. sylvestris. [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]