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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., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Drought combined with extreme heatwaves has been increasingly identified as the important trigger of worldwide tree mortality in the context of climate change; nonetheless, our understanding of the potential hydraulic and thermal impairments of hot droughts to trees and the subsequent post-recovery process remains limited. To investigate the response of tree water and carbon relations to drought, heatwave, and combined drought-heatwave stresses, three-year-old potted seedlings of Fraxinus mandshurica Rupr., a dominant tree species in temperate forests of northeast China, were grown under well-watered and drought-stressed conditions and exposed to a rapid, acute heatwave treatment. During the heatwave treatment with a maximum temperature exceeding 40 °C for two days, the leaf temperature of drought-stressed seedlings was, on average, 5 °C higher than that of well-watered counterparts due to less effective evaporative cooling, indicating that soil water availability influenced leaf thermoregulatory capacity during hot extremes. Consistently, more pronounced crown damage, as shown by 13 % irreversible leaf scorch, was found in seedlings under the drought-heatwave treatment relative to sole heatwave treatment, alongside the more severe stem xylem embolism and leaf electrolyte leakage. While the heatwave treatment accelerated the depletion of non-structural carbohydrates in drought-stressed seedlings, the increase of branch soluble sugar concentration in response to heatwave might be related to the requirement for maintaining hydraulic functioning via osmoregulation under high dehydration risk. The coordination between leaf stomatal conductance and total non-structural carbohydrate content during the post-heatwave recovery phase implied that plant-water relations and carbon physiology were closely coupled in coping with hot droughts. This study highlights that, under scenarios of aggravating drought co-occurring with heatwaves, tree seedlings could face a high risk of crown decline in relation to the synergistically increased hydraulic and thermal impairments., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

In recent year, widespread declines of Populus bolleana Lauche trees (P. bolleana, which dieback from the top down) and Haloxylon ammodendron shrubs (H. ammodendron, which dieback starting from their outer canopy) have occurred. To investigate how both intra-canopy hydraulic changes and plasticity in hydraulic properties create differences in vulnerability between these two species, we conducted a drought simulation field experiment. We analyzed branch hydraulic vulnerability, leaf water potential (Ψ), photosynthesis (A), stomatal conductance (gs), non-structural carbohydrate (NSCs) contents and morphological traits of the plants as the plants underwent a partial canopy dieback. Our results showed that: (1) the hydraulic architecture was very different between the two life forms; (2) H. ammodendron exhibited a drought tolerance response with weak stomatal control, and thus a sharp decline in Ψ while P. bolleana showed a drought avoidance response with tighter stomatal control that maintained a relatively stable Ψ; (3) the Ψ of H. ammodendron showed relative consistent symptoms of drought stress with increasing plant stature, but the Ψ of P. bolleana showed greater drought stress in higher portions of the crown; (4) prolonged drought caused P. bolleana to consume and H. ammodendron to accumulate NSCs in the branches of their upper canopy. Thus, the prolonged drought caused the shoots of the upper canopy of P. bolleana to experience greater vulnerability leading to dieback of the upper branches first, while all the twigs of the outer canopy of H. ammodendron experienced nearly identical degrees of vulnerability, and thus dieback occurred uniformly. Our results indicate that intra-canopy hydraulic change and their plasticity under drought was the main cause of the observed canopy dieback patterns in both species. However, more work is needed to further establish that hydraulic limitation as a function of plant stature was the sole mechanism for causing the divergent canopy dieback patterns., Competing Interests: Declaration of competing interest The authors declare that they have no conflict of interest., (Published by Elsevier B.V.)

Unprecedented tree dieback across Central Europe caused by recent global change-type drought events highlights the need for a better mechanistic understanding of drought-induced tree mortality. Although numerous physiological risk factors have been identified, the importance of two principal mechanisms, hydraulic failure and carbon starvation, is still debated. It further remains largely unresolved how the local neighborhood composition affects individual mortality risk. We studied 9435 young trees of 12 temperate species planted in a diversity experiment in 2013 to assess how hydraulic traits, carbon dynamics, pest infestation, tree height and neighborhood competition influence individual mortality risk. Following the most extreme global change-type drought since record in 2018, one third of these trees died. Across species, hydraulic safety margins (HSMs) were negatively and a shift towards a higher sugar fraction in the non-structural carbohydrate (NSC) pool positively associated with mortality risk. Moreover, trees infested by bark beetles had a higher mortality risk, and taller trees a lower mortality risk. Most neighborhood interactions were beneficial, although neighborhood effects were highly species-specific. Species that suffered more from drought, especially Larix spp. and Betula spp., tended to increase the survival probability of their neighbors and vice versa. While severe tissue dehydration marks the final stage of drought-induced tree mortality, we show that hydraulic failure is interrelated with a series of other, mutually inclusive processes. These include shifts in NSC pools driven by osmotic adjustment and/or starch depletion as well as pest infestation and are modulated by the size and species identity of a tree and its neighbors. A more holistic view that accounts for multiple causes of drought-induced tree mortality is required to improve predictions of trends in global forest dynamics and to identify mutually beneficial species combinations., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Extreme drought events are becoming frequent globally, resulting in widespread plant mortality and forest dieback. Although savanna vegetation cover ~20% of the earth's land area, their responses to extreme drought have been less studied than that of forests. Herein, we quantified branch dieback, individual mortality and the associated physiological responses of four evergreen shrubs (Tarenna depauperate Hutch., Maytenus esquirolii (H. Lév.) C.Y. Cheng, Murraya exotica L., Jasminum nudiflorum Lindl.) in a savanna ecosystem in Southwest China to an incidence of extreme drought during 2019 and 2020. We found that 80-100% of the individuals of these species exhibited branch dieback, whereas individual mortality was only found in T. depauperate (4.5%). All species showed high resistance to stem embolism (P50, water potential at 50% loss of hydraulic conductivity ranged from -5.62 to -8.6 MPa), whereas the stem minimum water potentials reached -7.6 to ca -10.0 MPa during the drought. The low water potential caused high native embolism levels (percentage loss of hydraulic conductivity (PLC) 23-65%) in terminal branches, and the remaining stems maintained 15-35% PLC at the end of the drought. Large within-individual variations in stem vulnerability to embolism were detected, and shedding of vulnerable branches could be a mechanism for shrubs to reduce water and carbon consumption. Overall, the content of total nonstructural carbohydrates (NSC) and their components in the stem were generally comparable to or higher than those in the rainy season in three of the four species. Because the leaves were turgor-less for most time during the drought, high NSC levels during the drought could be due to recycling of NSC from dead branches or translocation from roots. Our results suggest high tolerance of savanna shrub species to extreme drought, which could be facilitated by high embolism resistance in some stems and shedding of vulnerable branches to maintain individual water and carbon balance., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Drought-induced tree mortality may increase with ongoing climate change. Unraveling the links between stem hydraulics and mortality thresholds, and the effects of intraspecific variation, remain important unresolved issues. We conducted a water manipulation experiment in a rain-out shelter, using four provenances of Schima superba originating from a gradient of annual precipitation (1124-1796 mm) and temperature (16.4-22.4°C). Seedlings were droughted to three levels of percentage loss of hydraulic conductivity (i.e., P 50 , P 88  and P 99) and subsequently rewatered to field capacity for 30 days; traits related to water and carbon relations were measured. The lethal water potential associated with incipient mortality was between P 50 and P 88 . Seedlings exhibited similar drought responses in xylem water potential, hydraulic conductivity and gas exchange. Upon rehydration, patterns of gas exchange differed among provenances but were not related to the climate at the origin. The four provenances exhibited a similar degree of stem hydraulic recovery, which was correlated with the magnitude of antecedent drought and stem soluble sugar at the end of the drought. Results suggest that there were intraspecific differences in the capacity of S. superba seedlings for carbon assimilation during recovery, indicating a decoupling between gas exchange recovery and stem hydraulics across provenances., (© 2022 John Wiley & Sons Ltd.)

Mistletoes are important co-contributors to tree mortality globally, particularly during droughts. In Australia, mistletoe distributions are expanding in temperate woodlands, while their hosts have experienced unprecedented heat and drought stress in recent years. We investigated whether the excessive water use of mistletoes increased the probability of xylem emboli in a mature woodland during the recent record drought that was compounded by multiple heatwaves. We continuously recorded transpiration ($T_{SLA}$) of infected and uninfected branches from two eucalypt species over two summers, monitored stem and leaf water potentials ($\Psi $) and used hydraulic vulnerability curves to estimate percent loss in conductivity (PLC) for each species. Variations in weather (vapor pressure deficit, photosynthetically active radiation, soil water content), host species and % mistletoe foliage explained 78% of hourly $T_{SLA}$. While mistletoe acted as an uncontrollable sink for water in the host even during typical summer days, daily $T_{SLA}$ increased up to 4-fold in infected branches on hot days, highlighting the previously overlooked importance of temperature stress in amplifying water loss in mistletoes. The increased water use of mistletoes resulted in significantly decreased host $\Psi _{\rm{leaf}}$ and $\Psi _{\rm{trunk}}$. It further translated to an estimated increase of up to 11% PLC for infected hosts, confirming greater hydraulic dysfunction of infected trees that place them at higher risk of hydraulic failure. However, uninfected branches of Eucalyptus fibrosa F.Muell. had much tighter controls on water loss than uninfected branches of Eucalyptus moluccana Roxb., which shifted the risk of hydraulic failure towards an increased risk of carbon starvation for E. fibrosa. The contrasting mechanistic responses to heat and drought stress between both co-occurring species demonstrates the complexity of host-parasite interactions and highlights the challenge in predicting species-specific responses to biotic agents in a warmer and drier climate., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Ongoing global warming increases the frequency and severity of tropical typhoons and prolonged drought, leading to forest degradation. Simultaneous and/or successive masting events and climatic extremes may thus occur frequently in the near future. If these climatic extremes occur immediately after mass seed reproduction, their effects on individual trees are expected to be very severe because mass reproduction decreases carbohydrate reserves. While the effects of either a single climate extreme or masting alone on tree resilience/growth have received past research attention, understanding the cumulative effects of such multiple events remains challenging and is crucial for predicting future forest changes. Here, we report tree hazards compound by two successive climate extremes, a tropical typhoon and prolonged drought, after mass reproduction in an endemic tree species (Distylium lepidotum Nakai) on oceanic islands. Across individual trees, the starch stored within the sapwood of branchlets significantly decreased with reproductive efforts (fruit mass/shoot mass ratio). Typhoon damage significantly decreased not only the total leaf area of apical shoots but also the maximum photosynthetic rates. During the 5-month period after the typhoon, the mortality of large branchlets (8-10-mm diameter) increased with decreasing stored starch when the typhoon hit. During the prolonged summer drought in the next year, the recovery of total leaf area, stored starch, and hydraulic conductivity was negatively correlated with the stored starch at the typhoon. These data indicate that the level of stored starch within branchlets is the driving factor determining tree regrowth or dieback, and the restoration of carbohydrates after mass reproduction is synergistically delayed by such climate extremes. Stored carbohydrates are the major cumulative factor affecting individual tree resilience, resulting in their historical effects. Because of highly variable carbohydrate levels among individual trees, the resultant impacts of such successive events on forest dieback will be fundamentally different among trees., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Predicted increases in forest drought mortality highlight the need for predictors of incipient drought-induced mortality (DIM) risk that enable proactive large-scale management. Such predictors should be consistent across plants with varying morphology and physiology. Because of their integrative nature, indicators of water status are promising candidates for real-time monitoring of DIM, particularly if they standardize morphological differences among plants. We assessed the extent to which differences in morphology and physiology between Pinus ponderosa populations influence time to mortality and the predictive power of key indicators of DIM risk. Time to incipient mortality differed between populations but occurred at the same relative water content (RWC) and water potential (WP). RWC and WP were accurate predictors of drought mortality risk. These results highlight that variables related to water status capture critical thresholds during DIM and the associated dehydration processes. Both WP and RWC are promising candidates for large-scale assessments of DIM risk. RWC is of special interest because it allows comparisons across different morphologies and can be remotely sensed. Our results offer promise for real-time landscape-level monitoring of DIM and its global impacts in the near term., (© 2021 John Wiley & Sons Ltd.)

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 (Ψ leaf ), 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 Ψ leaf 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., (© 2021 The Authors New Phytologist © 2021 New Phytologist Foundation.)

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 drought-induced mortality in this major European tree species., Competing Interests: The authors declare no competing interest.

COMBINING HYDRAULIC: and carbon-related measurements can help elucidate drought-induced plant mortality. To study drought mortality mechanisms, seedlings of two woody species, including the anisohydric Robinia pseudoacacia and isohydric Quercus acutissima, were cultivated in a greenhouse and subjected to intense drought by withholding water and mild drought by adding half of the amount of daily water lost. Patterns of leaf and root gas exchange, leaf surface areas, growth, leaf and stem hydraulics, and carbohydrate dynamics were determined in drought-stressed and control seedlings. We detected a complete loss of hydraulic conductivity and partial depletion of total nonstructural carbohydrates contents (TNC) in the dead seedlings. We also found that intense drought triggered a more rapid decrease in plant water potential and a faster drop in net photosynthesis below zero, and a greater TNC loss in dead seedlings than mild drought. Additionally, anisohydric R. pseudoacacia suffered a rapider death than the isohydric Q. acutissima. Based on these findings, we propose that hydraulic conductivity loss and carbon limitation jointly contributed to drought-induced death, while the relative contributions could be altered by drought intensity. We thus believe that it is important to illustrate the mechanistic relationships between stress intensity and carbon-hydraulics coupling in the context of isohydric vs. anisohydric hydraulic strategies., Competing Interests: Declaration of competing interest No conflict of interest exits in the submission of this manuscript, and manuscript is approved by all authors for publication. We would like to declare on behalf of our co-authors that the work described was original research and has not been published previously, also not under consideration for publication elsewhere, in whole or in part., (Copyright © 2020. Published by Elsevier Masson SAS.)

The maintenance of hydraulic function during and after a drought event is crucial for tree survival, but the importance of non-structural carbohydrates (NSCs) in the recovery phase is still debated. We tested whether higher NSC availability facilitates post-drought hydraulic recovery, by applying a short-term drought (S dr ) and a long-term drought combined with shading (L dr+sh ) in Fraxinus ornus and Ostrya carpinifolia. Plants were then re-irrigated and recovery was checked 24 h later, by measuring water potential, stem percentage loss of hydraulic conductance (PLC) and NSC content. The relative magnitude of hydraulic and carbon constraints was also assessed in desiccated plants. During drought, PLC increased only in F. ornus, while it was maintained almost constant in O. carpinifolia due to tighter stomatal control of xylem pressure (i.e. more isohydric). In F. ornus, only S dr plants maintained high NSC contents at the end of drought and, when re-irrigated, recovered PLC to control values. Whereas hydraulic failure was ubiquitous, only F. ornus depleted NSC reserves at mortality. Our results suggest that preserving higher NSC content at the end of a drought can be important for the hydraulic resilience of trees., (Copyright © 2019 Elsevier Masson SAS. All rights reserved.)

Despite widespread interest in drought legacies-multiyear impacts of drought on tree growth-the key implication of reported drought legacies remains unaddressed: as impaired growth and slow recovery associated with drought legacies are pervasive across forest ecosystems, what is the impact of more frequent drought conditions? We investigated the assumption that either multiple drought years occurring during a short period (multiyear droughts), or droughts occurring during the recovery period from previous drought (compounded droughts), are detrimental to subsequent growth. There is evidence that drought responses may vary among populations of widespread species, leading us to examine regional differences in responses of the conifer Pinus ponderosa to historic drought frequency in the western United States. More frequent drought conditions incurred additional growth declines and shifts in growth-climate sensitivities in the years following drought relative to single-drought events, with 'triple-droughts' being worse than 'double-droughts'. Notably, prediction skill was not strongly reduced when ignoring compounded droughts, a consequence of the temporally comprehensive formulation of our stochastic antecedent model that accounts for the climatic memory of tree growth. We argue that incorporating drought-induced temporal variability in tree growth sensitivities can aid inference gained from statistical models, where more simplistic models could overestimate the severity of drought legacies. We also found regional differences in response to repeated drought, and suggest plastic post-drought sensitivities and climatic memory may represent beneficial physiological adjustments in interior regions. Within-species variability may thus mediate forest responses to increasing drought frequency under future climate change, but experimental approaches using more species are necessary to improve our understanding of the mechanisms that underlie drought legacy effects on tree growth., (© 2019 John Wiley & Sons Ltd.)

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 t crit , from stomatal closure g s90 to a critical level of hydraulic dysfunction Ψ crit 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 t crit 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 t crit 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., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

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., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

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., (© The Author(s) 2019. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Drought can cause major damage to plant communities, but species damage thresholds and postdrought recovery of forest productivity are not yet predictable. We used an El Niño drought event as a natural experiment to test whether postdrought recovery of gas exchange could be predicted by properties of the water transport system, or if metabolism, primarily high abscisic acid concentration, might delay recovery. We monitored detailed physiological responses, including shoot sapflow, leaf gas exchange, leaf water potential and foliar abscisic acid (ABA), during drought and through the subsequent rehydration period for a sample of eight canopy and understory species. Severe drought caused major declines in leaf water potential, elevated foliar ABA concentrations and reduced stomatal conductance and assimilation rates in our eight sample species. Leaf water potential surpassed levels associated with incipient loss of leaf hydraulic conductance in four species. Following heavy rainfall gas exchange in all species, except those trees predicted to have suffered hydraulic impairment, recovered to prestressed rates within 1 d. Recovery of plant gas exchange was rapid and could be predicted by the hydraulic safety margin, providing strong support for leaf vulnerability to water deficit as an index of damage under natural drought conditions., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

Contents 513 I. 513 II. 514 III. 517 518 References 518 SUMMARY: Global forests are experiencing rising temperatures and more severe droughts, with consistently dire forecasts for negative future impacts. Current research on the physiological mechanisms underlying drought impacts is focused on the water- and carbon-associated mechanisms. The role of nutrients is notably missing from this research agenda. Here, we investigate what role, if any, forest nutrition plays for survival and recovery of forests during and after drought. High nutrient availability may play a detrimental role in drought survival due to preferential biomass allocation aboveground that (1) predispose plants to hydraulic constraints limiting photosynthesis and promoting hydraulic failure, (2) increases carbon costs during periods of carbon starvation, and (3) promote biotic attack due to low tissue carbon: nitrogen (C : N). When nutrient uptake occurs during drought, high nutrient availability can increase water use efficiency thus minimizing negative feedbacks between carbon and nutrient balance. Nutrients are released after drought ceases, which might promote faster recovery but the temporal dynamics of microbial immobilization and nutrient leaching have a significant impact on nutrient availability. We provide a framework for understanding nutrient impacts on drought survival that allows a more complete analysis of forest ecosystem responses., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

Plants exhibit a variety of drought responses involving multiple interacting traits and processes, which makes predictions of drought survival challenging. Careful evaluation of responses within species, where individuals share broadly similar drought resistance strategies, can provide insight into the relative importance of different traits and processes. We subjected Pinus sylvestris L. saplings to extreme drought (no watering) leading to death in a greenhouse to (i) determine the relative effect of predisposing factors and responses to drought on survival time, (ii) identify and rank the importance of key predictors of time to death and (iii) compare individual characteristics of dead and surviving trees sampled concurrently. Time until death varied over 3 months among individual trees (from 29 to 147 days). Survival time was best predicted (higher explained variance and impact on the median survival time) by variables related to carbon uptake and carbon/water economy before and during drought. Trees with higher concentrations of monosaccharides before the beginning of the drought treatment and with higher assimilation rates prior to and during the treatment survived longer (median survival time increased 25-70 days), even at the expense of higher water loss. Dead trees exhibited less than half the amount of nonstructural carbohydrates (NSCs) in branches, stem and relative to surviving trees sampled concurrently. Overall, our results indicate that the maintenance of carbon assimilation to prevent acute depletion of NSC content above some critical level appears to be the main factor explaining survival time of P. sylvestris trees under extreme drought., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Establishing physiological thresholds to drought-induced mortality in a range of plant species is crucial in understanding how plants respond to severe drought. Here, five common European tree species were selected (Acer campestre L., Acer pseudoplatanus L., Carpinus betulus L., Corylus avellana L. and Fraxinus excelsior L.) to study their hydraulic thresholds to mortality. Photosynthetic parameters during desiccation and the recovery of leaf gas exchange after rewatering were measured. Stem vulnerability curves and leaf pressure-volume curves were investigated to understand the hydraulic coordination of stem and leaf tissue traits. Stem and root samples from well-watered and severely drought-stressed plants of two species were observed using transmission electron microscopy to visualize mortality of cambial cells. The lethal water potential (ψlethal) correlated with stem P99 (i.e., the xylem water potential at 99% loss of hydraulic conductivity, PLC). However, several plants that were stressed beyond the water potential at 100% PLC showed complete recovery during the next spring, which suggests that the ψlethal values were underestimated. Moreover, we observed a 1 : 1 relationship between the xylem water potential at the onset of embolism and stomatal closure, confirming hydraulic coordination between leaf and stem tissues. Finally, ultrastructural changes in the cytoplasm of cambium tissue and mortality of cambial cells are proposed to provide an alternative approach to investigate the point of no return associated with plant death., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Many species have the ability to resprout vegetatively after a substantial loss of biomass induced by environmental stress, including drought. Many of the regions characterised by ecosystems where resprouting is common are projected to experience more frequent and intense drought during the 21st Century. However, in assessments of ecosystem response to drought disturbance there has been scant consideration of the resilience and post-drought recovery of resprouting species. Systematic differences in hydraulic and allocation traits suggest that resprouting species are more resilient to drought-stress than nonresprouting species. Evidence suggests that ecosystems dominated by resprouters recover from disturbance more quickly than ecosystems dominated by nonresprouters. The ability of resprouters to avoid mortality and withstand drought, coupled with their ability to recover rapidly, suggests that the impact of increased drought stress in ecosystems dominated by these species may be small. The strategy of resprouting needs to be modelled explicitly to improve estimates of future climate-change impacts on the carbon cycle, but this will require several important knowledge gaps to be filled before resprouting can be properly implemented., (© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.)

Drought-related tree die-off episodes have been observed in all vegetated continents. Despite much research effort, however, the multiple interactions between carbon starvation, hydraulic failure and biotic agents in driving tree mortality under field conditions are still not well understood. We analysed the seasonal variability of non-structural carbohydrates (NSCs) in four organs (leaves, branches, trunk and roots), the vulnerability to embolism in roots and branches, native embolism (percentage loss of hydraulic conductivity (PLC)) in branches and the presence of root rot pathogens in defoliated and non-defoliated individuals in a declining Scots pine (Pinus sylvestris L.) population in the NE Iberian Peninsula in 2012, which included a particularly dry and warm summer. No differences were observed between defoliated and non-defoliated pines in hydraulic parameters, except for a higher vulnerability to embolism at pressures below -2 MPa in roots of defoliated pines. No differences were found between defoliation classes in branch PLC. Total NSC (TNSC, soluble sugars plus starch) values decreased during drought, particularly in leaves. Defoliation reduced TNSC levels across tree organs, especially just before (June) and during (August) drought. Root rot infection by the fungal pathogen Onnia P. Karst spp. was detected but it did not appear to be associated to tree defoliation. However, Onnia infection was associated with reduced leaf-specific hydraulic conductivity and sapwood depth, and thus contributed to hydraulic impairment, especially in defoliated pines. Infection was also associated with virtually depleted root starch reserves during and after drought in defoliated pines. Moreover, defoliated and infected trees tended to show lower basal area increment. Overall, our results show the intertwined nature of physiological mechanisms leading to drought-induced mortality and the inherent difficulty of isolating their contribution under field conditions., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

It has been reported that elevated temperature accelerates the time-to-mortality in plants exposed to prolonged drought, while elevated [CO(2)] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO(2)] and temperature on the inter-dependent carbon and hydraulic characteristics associated with drought-induced mortality in Eucalyptus radiata seedlings grown in two [CO(2)] (400 and 640 μL L(-1)) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO(2)] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO(2)], may be the primary contributors to drought-induced seedling mortality under future climates., (© 2013 John Wiley & Sons Ltd.)

Drought challenges plant water uptake and the vascular system. In the xylem it causes embolism that impairs water transport from the soil to the leaves and, if uncontrolled, may even lead to plant mortality via hydraulic failure. What happens in the phloem, however, is less clear because measuring phloem transport is still a significant challenge to plant science. In all vascular plants, phloem and xylem tissues are located next to each other, and there is clear evidence that these tissues exchange water. Therefore, drought should also lead to water shortage in the phloem. In this review, theories used in phloem transport models have been applied to drought conditions, with the goal of shedding light on how phloem transport failure might occur. The review revealed that phloem failure could occur either because of viscosity build-up at the source sites or by a failure to maintain phloem water status and cell turgor. Which one of these dominates depends on the hydraulic permeability of phloem conduit walls. Impermeable walls will lead to viscosity build-up affecting flow rates, while permeable walls make the plant more susceptible to phloem turgor failure. Current empirical evidence suggests that phloem failure resulting from phloem turgor collapse is the more likely mechanism at least in relatively isohydric plants.

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]

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]

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]

Canada's boreal forests, which occupy approximately 30% of boreal forests worldwide, play an important role in the global carbon budget. However, there is little quantitative information available regarding the spatiotemporal changes in the drought‐induced tree mortality of Canada's boreal forests overall and their associated impacts on biomass carbon dynamics. Here, we develop spatiotemporally explicit estimates of drought‐induced tree mortality and corresponding biomass carbon sink capacity changes in Canada's boreal forests from 1970 to 2020. We show that the average annual tree mortality rate is approximately 2.7%. Approximately 43% of Canada's boreal forests have experienced significantly increasing tree mortality trends (71% of which are located in the western region of the country), and these trends have accelerated since 2002. This increase in tree mortality has resulted in significant biomass carbon losses at an approximate rate of 1.51 ± 0.29 MgC ha−1 year−1 (95% confidence interval) with an approximate total loss of 0.46 ± 0.09 PgC year−1 (95% confidence interval). Under the drought condition increases predicted for this century, the capacity of Canada's boreal forests to act as a carbon sink will be further reduced, potentially leading to a significant positive climate feedback effect. [ABSTRACT FROM AUTHOR]

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]

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]

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.

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]

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]

Despite a wealth of eco-physiological assessments of plant response to extreme drought, few studies have addressed the interactive effects of global change factors on traits driving mortality. To understand the interaction between hydraulic and carbon metabolic traits influencing tree mortality, which may be independently influenced by atmospheric [CO2] and temperature, we grew Eucalyptus sideroxylon A. Cunn. ex Woolls from seed in a full-factorial [CO2] (280, 400 and 640 μmol mol−1, C p, C a and C e, respectively) and temperature (ambient and ambient +4 °C, T a and T e, respectively) experiment. Prior to drought, growth across treatment combinations resulted in significant variation in physiological and morphological traits, including photosynthesis (A sat), respiration (R d), stomatal conductance, carbohydrate storage, biomass and leaf area (LA). C e increased A sat, LA and leaf carbohydrate concentration compared with C a, while C p generated the opposite response; T e reduced R d. However, upon imposition of drought, T e hastened mortality (9 days sooner compared with T a), while C e significantly exacerbated drought stress when combined with T e. Across treatments, earlier time-to-mortality was mainly associated with lower (more negative) leaf water potential (Ψl) during the initial drought phase, along with higher water loss across the first 3 weeks of water limitation. Among many variables, Ψl was more important than carbon status in predicting time-to-mortality across treatments, yet leaf starch was associated with residual variation within treatments. These results highlight the need to carefully consider the integration, interaction and hierarchy of traits contributing to mortality, along with their responses to environmental drivers. Both morphological traits, which influence soil resource extraction, and physiological traits, which affect water-for-carbon exchange to the atmosphere, must be considered to adequately predict plant response to drought. Researchers have struggled with assessing the relative importance of hydraulic and carbon metabolic traits in determining mortality, yet an integrated trait, time-dependent framework provides considerable insight into the risk of death from drought for trees. [ABSTRACT FROM AUTHOR]

Manipulative experiments have suggested that embolism-induced hydraulic impairment underpins widespread tree mortality during extreme drought, yet in situ evidence is rare. One month after drought-induced leaf and branch dieback was observed in field populations of Eucalyptus piperita Sm. in the Blue Mountains (Australia), we measured the level of native stem embolism and characterized the extent of leaf death in co-occurring dieback and healthy (non-dieback) trees. We found that canopy dieback-affected trees showed significantly higher levels of native embolism (26%) in tertiary order branchlets than healthy trees (11%). Furthermore, there was a significant positive correlation (R 2 = 0.51) between the level of leaf death and the level of native embolism recorded in branchlets from dieback-affected trees. This retrospective study suggests that hydraulic failure was the primary mechanism of leaf and branch dieback in response to a natural drought event in the field. It also suggests that post-drought embolism refilling is minimal or absent in this species of eucalypt. [ABSTRACT FROM AUTHOR]

Projected climate change scenarios such as frequently occurring dry summer spells are an enormous threat to the health of boreal conifer forests. We identified visible features indicating wood with tracheids predisposed for hydraulic and mechanical dysfunction in Norway spruce, suggest why this is formed during severe summer drought and hypothesised on mechanism that would cause tracheid collapse and stem cracks. Trees from southern Sweden that showed signs of severe reaction to drought, i.e. stem cracks along the trunk, were compared to healthy, undamaged trees. Rings investigated included those formed in 2006, a year with an extremely dry summer season in the study region. In southern Norway, we investigated trees with and without drought-induced top dieback symptoms. We analysed anatomical features such as tracheid lumen diameter, thickness of cell wall and its various layers (S1, S2 and S3), applied Raman imaging in order to get information on the lignin distribution in the cell wall and the compound middle lamellae and performed hydraulic flow and shrinkage experiments. Although tracheids in annual rings with signs of collapse had higher tangential lumen diameters than those in “normal” annual rings, we conclude that collapse of tracheid walls depends mainly on wall thickness, which is genetically determined to a large extent. Spruce trees that produce earlywood with extremely thin cell walls can develop wall collapse and internal cracks under the impact of dry spells. We also present a new diagnostic tool for detecting individuals that are prone to cell wall collapse and stem cracks: Lucid bands, i.e. bands in the fresh sapwood with very thin cell walls and inhomogeneous lignin distribution in the S-layers and the compound middle lamellae that lost their hydraulic function due to periods of severe summer drought. The detection of genotypes with lucid bands could be useful for an early selection against individuals that are prone to stem cracks under the impact of severe summer drought, and also for early downgrading of logs prone to cracking during industrial kiln drying. [ABSTRACT FROM AUTHOR]

• We exposed seedlings of conifer species to three realistic simulated future climates. • Seedlings of conifer species experienced high mortality (75%) induced by drought. • Conifer species are highly vulnerable at the seedling stage to future drought. • Hydraulic safety margins are good indicators of mortality risk for conifer species. The future climate of northern temperate forests is projected to be drier and warmer by the end of this century. As a result, more drought-induced forest dieback events are anticipated in northeastern North America, and assessing the vulnerability of dominant tree species to drought is critical for understanding the future composition of these forests. In a greenhouse experiment, we exposed two-year-old seedlings of Picea glauca (Moench) Voss, Picea mariana (Mill.) B.S.P. and Pinus strobus L. to three future climate treatments for southern Quebec, Canada, and evaluated their mortality, growth, and foliage water status responses to soil water availability and atmospheric drought. Using a unique approach, climate treatments emulated droughts of different frequencies, durations, and intensities. Treatments closely simulated one growing season, with changes in air temperature and relative humidity every six hours and daily adjustment in the amount of water delivered to the seedlings. The three species experienced high mortality (75%) in all water-limited treatments compared to a control treatment that provided non-limiting soil moisture (0% mortality). The biomass of the seedlings that survived was 40% lower than that of control seedlings. Our results confirmed that the hydraulic safety margins, defined as the difference between seasonal minimum water potential and xylem water potential leading to 12, 50 and 88% of hydraulic conductivity loss, were good predictors of probability of tree mortality. Therefore, hydraulic safety margins are useful functional traits that can be used to compare the vulnerability of various species to drought and then provide crucial information to practitioners and policymakers to adjust forest management to climate change. We showed that three dominant conifer species of northern temperate forests were highly vulnerable to drought in future climates. Because drought is projected to be a significant threat to forests, understanding potentially adaptive physiological responses to drought, such as hydraulic safety margins of tree seedlings, is important for predicting the response of forest regeneration and composition in warmer and drier climates. [ABSTRACT FROM AUTHOR]

Intensifying drought around the globe poses strong challenges to the growth and survival of trees, emphasizing the urgency of understanding the physiological mechanisms of drought-induced tree decline and mortality and the necessity of using silvicultural approaches to mitigate drought effects on tree growth. We experimentally explored whether thinning enhances tree growth vigor and drought resilience in early aging Mongolian pine (Pinus sylvestris var. mongolica Litv.) plantations in a water-limited area of Northeast China using dendrochronological methods. Meanwhile, we compared plant hydraulic traits, water relations, and non-structural carbohydrate concentrations between trees growing in the thinned and un-thinned stands with the aim of unraveling the physiological mechanisms underlying the density effects on tree growth performance. The results showed that basal area increments of Mongolian pine trees in the high-density plantations started to decline continuously at tree cambium age of around 15 years over about a decade, while thinning effectively reversed their growth decline. Trees in thinned stands exhibited significantly higher predawn leaf water potentials, higher stem hydraulic conductivities, and larger carbon reserves than trees in un-thinned stands. Although trees growing in high-density stands maintained as low risks of xylem hydraulic dysfunction as trees growing in low-density stands, that was at the cost of substantially reducing tree crown size through branch mortality that unavoidably undermined their carbon balance at the whole-plant level. Consistently, trees growing in the thinned low-density stands showed significantly higher levels of non-structural carbohydrate reserve in stems and enhanced growth resilience to extreme drought than trees growing in the high-density reference stands. Our results suggest that thinning can effectively enhance xylem hydraulic transport efficiency and reduce the risk of carbon imbalance by increasing soil water availability and hence mitigates the early aging of Mongolian pine plantations in water-limited environments. • Thinning improved growth vigor and drought resilience of aging plantations. • Trees exhibited higher hydraulic efficiency and better water status after thinning. • Higher nonstructural carbohydrate content contributed to greater drought resilience. [ABSTRACT FROM AUTHOR]