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We related measurements of annual burned area in the southwest United States during 1984–2013 to records of climate variability. Within forests, annual burned area correlated at least as strongly with spring–summer vapour pressure deficit (VPD) as with 14 other drought-related metrics, including more complex metrics that explicitly represent fuel moisture. Particularly strong correlations with VPD arise partly because this term dictates the atmospheric moisture demand. Additionally, VPD responds to moisture supply, which is difficult to measure and model regionally due to complex micrometeorology, land cover and terrain. Thus, VPD appears to be a simple and holistic indicator of regional water balance. Coupled with the well-known positive influence of prior-year cold season precipitation on fuel availability and connectivity, VPD may be utilised for burned area forecasts and also to infer future trends, though these are subject to other complicating factors such as land cover change and management. Assuming an aggressive greenhouse gas emissions scenario, climate models predict mean spring–summer VPD will exceed the highest recorded values in the southwest in nearly 40% of years by the middle of this century. These results forewarn of continued increases in burned forest area in the southwest United States, and likely elsewhere, when fuels are not limiting.

Greenhouse gas emissions have significantly altered global climate, and will continue to do so in the future. Increases in the frequency, duration, and/or severity of drought and heat stress associated with climate change could fundamentally alter the composition, structure, and biogeography of forests in many regions. Of particular concern are potential increases in tree mortality associated with climate-induced physiological stress and interactions with other climate-mediated processes such as insect outbreaks and wildfire. Despite this risk, existing projections of tree mortality are based on models that lack functionally realistic mortality mechanisms, and there has been no attempt to track observations of climate-driven tree mortality globally. Here we present the first global assessment of recent tree mortality attributed to drought and heat stress. Although episodic mortality occurs in the absence of climate change, studies compiled here suggest that at least some of the world's forested ecosystems already may be responding to climate change and raise concern that forests may become increasingly vulnerable to higher background tree mortality rates and die-off in response to future warming and drought, even in environments that are not normally considered water-limited. This further suggests risks to ecosystem services, including the loss of sequestered forest carbon and associated atmospheric feedbacks. Our review also identifies key information gaps and scientific uncertainties that currently hinder our ability to predict tree mortality in response to climate change and emphasizes the need for a globally coordinated observation system. Overall, our review reveals the potential for amplified tree mortality due to drought and heat in forests worldwide.

Summary: Frequent observations of higher mortality in larger trees than in smaller ones during droughts have sparked an increasing interest in size‐dependent drought‐induced mortality. However, the underlying physiological mechanisms are not well understood, with height‐associated hydraulic constraints often being implied as the potential mechanism driving increased drought vulnerability. We performed a quantitative synthesis on how key traits that drive plant water and carbon economy change with tree height within species and assessed the implications that the different constraints and compensations may have on the interacting mechanisms (hydraulic failure, carbon starvation and/or biotic‐agent attacks) affecting tree vulnerability to drought. While xylem tension increases with tree height, taller trees present a range of structural and functional adjustments, including more efficient water use and transport and greater water uptake and storage capacity, that mitigate the path‐length‐associated drop in water potential. These adaptations allow taller trees to withstand episodic water stress. Conclusive evidence for height‐dependent increased vulnerability to hydraulic failure and carbon starvation, and their coupling to defence mechanisms and pest and pathogen dynamics, is still lacking. Further research is needed, particularly at the intraspecific level, to ascertain the specific conditions and thresholds above which height hinders tree survival under drought. [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]

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]

Subtropical evergreen broadleaf forests (SEBF) are experiencing and expected to suffer more frequent and severe drought events. However, how the hydraulic traits directly link to the mortality and recovery of SEBF trees remains unclear. In this study, we conducted a drought–rewatering experiment on tree seedlings of five dominant species to investigate how the hydraulic traits were related to tree mortality and the resistance and recovery of photosynthesis (A) and transpiration (E) under different drought severities. Species with greater embolism resistance (P50) survived longer than those with a weaker P50. However, there was no general hydraulic threshold associated with tree mortality, with the lethal hydraulic failure varying from 64% to 93% loss of conductance. The photosynthesis and transpiration of tree species with a greater P50 were more resistant to and recovered faster from drought than those with lower P50. Other plant traits could not explain the interspecific variation in tree mortality and drought resistance and recovery. These results highlight the unique importance of embolism resistance in driving carbon and water processes under persistent drought across different trees in SEBFs. The absence of multiple efficient drought strategies in SEBF seedlings implies the difficulty of natural seedling regeneration under future droughts, which often occurs after destructive disturbances (e.g., extreme drought events and typhoon), suggesting that this biome may be highly vulnerable to co‐occurring climate extremes. [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]

Increasing seawater exposure is causing mortality of coastal forests, yet the physiological response associated with seawater-induced tree mortality, particularly in non-halophytes, is poorly understood. We investigated the shifts in carbon and nitrogen (N) metabolism of mature Sitka-spruce trees that were dying after an ecosystem-scale manipulation of tidal seawater exposure. Soil porewater salinity and foliar ion concentrations increased after seawater exposure and were strongly correlated with the percentage of live foliated crown (PLFC; e.g. crown 'greenness', a measure of progression to death). Co-occurring with decreasing PLFC was decreasing photosynthetic capacity, N-investment into photosynthesis, N-resorption efficiency and non-structural carbohydrate (soluble sugars and starch) concentrations, with the starch reserves depleted to near zero when PLFC dropped below 5%. Combined with declining PLFC, these changes subsequently decreased total carbon gain and thus exacerbated the carbon starvation process. This study suggests that an impairment in carbon and N metabolism during the mortality process after seawater exposure is associated with the process of carbon starvation, and provides critical knowledge necessary to predict sea-level rise impacts on coastal forests. [ABSTRACT FROM AUTHOR]

Short‐term plant respiration (R) increases exponentially with rising temperature, but drought could reduce respiration by reducing growth and metabolism. Acclimation may alter these responses. We examined if species with different drought responses would differ in foliar R response to +4.8°C temperature and −45% precipitation in a field experiment with mature piñon and juniper trees, and if any differences between species were related to differences in photosynthesis rates, shoot growth and nonstructural carbohydrates (NSCs). Short‐term foliar R had a Q10 of 1.6 for piñon and 2.6 for juniper. Piñon foliar R did not respond to the +4.8°C temperatures, but R increased 1.4× for juniper. Across treatments, piñon foliage had higher growth, lower NSC content, 29% lower photosynthesis rates, and 44% lower R than juniper. Removing 45% precipitation had little impact on R for either species. Species differences in the response of R under elevated temperature were related to substrate availability and stomatal response to leaf water potential. Despite not acclimating to the higher temperature and having higher R than piñon, greater substrate availability in juniper suggests it could supply respiratory demand for much longer than piñon. Species responses will be critical in ecosystem response to a warmer climate. In a field experiment, piñon foliar R did not respond to the +4.8°C temperatures, but R increased 1.4× for juniper showing no thermal acclimation. Across treatments, piñon foliage had higher growth, lower NSC content, 29% lower photosynthesis rates and 44% lower R than juniper. Removing 45% precipitation had little impact on R for either species. Species differences in the response of R under elevated temperature were related to substrate availability and stomatal response to leaf water potential. Despite not acclimating to the higher temperature and having higher R than piñon, greater substrate availability in juniper suggests it could supply respiratory demand for much longer than piñon. Species responses will be critical in ecosystem response to a warmer climate. [ABSTRACT FROM AUTHOR]

Summary: Deep‐water access is arguably the most effective, but under‐studied, mechanism that plants employ to survive during drought. Vulnerability to embolism and hydraulic safety margins can predict mortality risk at given levels of dehydration, but deep‐water access may delay plant dehydration. Here, we tested the role of deep‐water access in enabling survival within a diverse tropical forest community in Panama using a novel data‐model approach.We inversely estimated the effective rooting depth (ERD, as the average depth of water extraction), for 29 canopy species by linking diameter growth dynamics (1990–2015) to vapor pressure deficit, water potentials in the whole‐soil column, and leaf hydraulic vulnerability curves. We validated ERD estimates against existing isotopic data of potential water‐access depths.Across species, deeper ERD was associated with higher maximum stem hydraulic conductivity, greater vulnerability to xylem embolism, narrower safety margins, and lower mortality rates during extreme droughts over 35 years (1981–2015) among evergreen species. Species exposure to water stress declined with deeper ERD indicating that trees compensate for water stress‐related mortality risk through deep‐water access.The role of deep‐water access in mitigating mortality of hydraulically‐vulnerable trees has important implications for our predictive understanding of forest dynamics under current and future climates. [ABSTRACT FROM AUTHOR]

Summary: Conifer mortality rates are increasing in western North America, but the physiological mechanisms underlying this trend are not well understood.We examined tree‐ring‐based radial growth along with stable carbon (C) and oxygen (O) isotope composition (δ13C and δ18O, respectively) of dying and surviving conifers at eight old‐growth forest sites across a strong moisture gradient in the western USA to retrospectively investigate mortality predispositions.Compared with surviving trees, lower growth of dying trees was detected at least one decade before mortality at seven of the eight sites. Intrinsic water‐use efficiency increased over time in both dying and surviving trees, with a weaker increase in dying trees at five of the eight sites.C starvation was a strong correlate of conifer mortality based on a conceptual model incorporating growth, δ13C, and δ18O. However, this approach does not capture processes that occur in the final months of survival. Ultimately, C starvation may lead to increased mortality vulnerability, but hydraulic failure or biotic attack may dominate the process during the end stages of mortality in these conifers. [ABSTRACT FROM AUTHOR]

Rising sea levels under climate change may have significant impacts on coastal vegetation dynamics, yet the response of coastal forest growth, gas exchange and survival to seawater intrusion remains poorly documented.We conducted a dendroecology study across six sites in western Washington, USA, to examine how tree growth, gas exchange (indexed by basal area increment (BAI) and wood δ13C respectively), and survival varies with seawater exposure through two approaches. First, tree core samples were collected at a site where seawater exposure started only 4 years prior to sampling, which allowed a cause‐and‐effect test of the impacts of seawater exposure on trees, and second, samples were collected at five additional sites where we compared downstream to upstream trees under current sea‐level conditions.At the seawater intrusion site, BAI and carbon isotope discrimination (Δ) decreased significantly (p < 0.01) in the year of intrusion (2014) and stayed unchanged thereafter. Four years later (2018), the percentage of recently standing dead trees in the forest was 73.0% of the basal area. Across the regional assessment, percentage of standing dead trees was significantly greater in downstream than upstream forests at five of the six sites (averaged 37.7 ± 11.0% and 4.3 ± 2.1% basal area for downstream and upstream, respectively). Growth was significantly lower (p < 0.01) at the downstream than upstream for five sites, and Δ was lower for all needle‐leaf trees (three sites) on the downstream compared to the upstream, but no difference was observed between downstream and upstream for broad‐leaf trees (three sites).Synthesis. Combined both the cause‐and‐effect manipulative study and the regional assessment demonstrate that seawater exposure drives reductions in growth, decreased Δ of needle‐leaf trees, increased mortality and greater climate sensitivity, regardless of whether the seawater exposure is recent or long‐term. [ABSTRACT FROM AUTHOR]

Summary: 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. [ABSTRACT FROM AUTHOR]

Abstract: From 2011 to 2013, Texas experienced its worst drought in recorded history. This event provided a unique natural experiment to assess species‐specific responses to extreme drought and mortality of four co‐occurring woody species: Quercus fusiformis, Diospyros texana, Prosopis glandulosa, and Juniperus ashei. We examined hypothesized mechanisms that could promote these species' diverse mortality patterns using postdrought measurements on surviving trees coupled to retrospective process modelling. The species exhibited a wide range of gas exchange responses, hydraulic strategies, and mortality rates. Multiple proposed indices of mortality mechanisms were inconsistent with the observed mortality patterns across species, including measures of the degree of iso/anisohydry, photosynthesis, carbohydrate depletion, and hydraulic safety margins. Large losses of spring and summer whole‐tree conductance (driven by belowground losses of conductance) and shallower rooting depths were associated with species that exhibited greater mortality. Based on this retrospective analysis, we suggest that species more vulnerable to drought were more likely to have succumbed to hydraulic failure belowground. [ABSTRACT FROM AUTHOR]

Contents513I.513II.514III.517518References518 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. [ABSTRACT FROM AUTHOR]

Understanding the interactions between drought and tree ontogeny or size remains an essential research priority because size-specific mortality patterns have large impacts on ecosystem structure and function, determine forest carbon storage capacity, and are sensitive to climatic change. Here we investigate a xerophytic tree species ( Haloxylon ammodendron (C.A. Mey.)) with which the changes in biomass allocation with tree size may play an important role in size-specific mortality patterns. Size-related changes in biomass allocation, root distribution, plant water status, gas exchange, hydraulic architecture and non-structural carbohydrate reserves of this xerophytic tree species were investigated to assess their potential role in the observed U-shaped mortality pattern. We found that excessively negative water potentials (<−4.7 MPa, beyond the P50 leaf of −4.1 MPa) during prolonged drought in young trees lead to hydraulic failure; while the imbalance of photoassimilate allocation between leaf and root system in larger trees, accompanied with declining C reserves (<2% dry matter across four tissues), might have led to carbon starvation. The drought-resistance strategy of this species is preferential biomass allocation to the roots to improve water capture. In young trees, the drought-resistance strategy is not well developed, and hydraulic failure appears to be the dominant driver of mortality during drought. With old trees, excess root growth at the expense of leaf area may lead to carbon starvation during prolonged drought. Our results suggest that the drought-resistance strategy of this xeric tree is closely linked to its life and death: well-developed drought-resistance strategy means life, while underdeveloped or overdeveloped drought-resistance strategy means death. [ABSTRACT FROM AUTHOR]

A simplified soil-plant-atmosphere-continuum model of carbon starvation and hydraulic failure is developed and tested against observations from a drought-manipulation experiment in a woodland dominated by piñon pine ( Pinus edulis) and juniper ( Juniperus monosperma) in New Mexico. The number of model parameters is reduced using allometric relationships. The model can represent more isohydric (piñon) and more anisohydric (juniper) responses. Analysis of the parameter space suggests four main controls on hydraulic failure and carbon starvation: xylem vulnerability curve, root:shoot area ratio, rooting depth and water use efficiency. For piñon, an intermediate optimal (1.5-2 m2 m−2) tree leaf area index reduces the risk of hydraulic failure. For both piñons and junipers, hydraulic failure was relatively insensitive to root:shoot ratio across a range of tree LAI. Higher root:shoot ratios however strongly decreased the time to carbon starvation. The hydraulic safety margin of piñons is strongly diminished by large diurnal variations in xylem/leaf water potential. Diurnal drops of water potential are mitigated by high maximum hydraulic conductivity, high root:shoot ratio and stomatal regulation (more isohydric). The safety margin of junipers is not very sensitive to diurnal drops in water potential so that there is little benefit in stomatal regulation (more anisohydric). Narrower tracheid diameter and a narrower distribution of tracheid diameters reduce the risk of hydraulic failure and carbon starvation by reducing diurnal xylem water potential drop. Simulated tree diameter-dependent mortality varies between these two species, with piñon mortality decreasing with increasing tree size, whereas juniper mortality increases with tree size. Juvenile piñons might thus be overimpacted by water stress. Copyright © 2015 John Wiley & Sons, Ltd. [ABSTRACT FROM AUTHOR]

Climate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects - bark beetles and defoliators - which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree-insect interactions will better inform projections of forest ecosystem responses to climate change. [ABSTRACT FROM AUTHOR]

Conifers have incurred high mortality during recent global-change-type drought(s) in the western USA. Mechanisms of drought-related tree mortality need to be resolved to support predictions of the impacts of future increases in aridity on vegetation. Hydraulic failure, carbon starvation and lethal biotic agents are three potentially interrelated mechanisms of tree mortality during drought. Our study compared a suite of measurements related to these mechanisms between 49 mature piñon pine (Pinus edulis Engelm.) trees that survived severe drought in 2002 (live trees) and 49 trees that died during the drought (dead trees) over three sites in Arizona and New Mexico. Results were consistent over all sites indicating common mortality mechanisms over a wide region rather than site-specific mechanisms. We found evidence for an interactive role of hydraulic failure, carbon starvation and biotic agents in tree death. For the decade prior to the mortality event, dead trees had twofold greater sapwood cavitation based on frequency of aspirated tracheid pits observed with scanning electron microscopy (SEM), smaller inter-tracheid pit diameter measured by SEM, greater diffusional constraints to photosynthesis based on higher wood δ13C, smaller xylem resin ducts, lower radial growth and more bark beetle (Coleoptera: Curculionidae) attacks than live trees. Results suggest that sapwood cavitation, low carbon assimilation and low resin defense predispose piñon pine trees to bark beetle attacks and mortality during severe drought. Our novel approach is an important step forward to yield new insights into how trees die via retrospective analysis. [ABSTRACT FROM AUTHOR]

Drought-induced forest mortality is an increasing global problem with wide-ranging consequences, yet mortality mechanisms remain poorly understood. Depletion of non-structural carbohydrate ( NSC) stores has been implicated as an important mechanism in drought-induced mortality, but experimental field tests are rare. We used an ecosystem-scale precipitation manipulation experiment to evaluate leaf and twig NSC dynamics of two co-occurring conifers that differ in patterns of stomatal regulation of water loss and recent mortality: the relatively desiccation-avoiding piñon pine ( P inus edulis) and the relatively desiccation-tolerant one-seed juniper ( J uniperus monosperma). Piñon pine experienced 72% mortality after 13-25 months of experimental drought and juniper experienced 20% mortality after 32-47 months. Juniper maintained three times more NSC in the foliage than twigs, and converted NSC to glucose and fructose under drought, consistent with osmoregulation requirements to maintain higher stomatal conductance during drought than piñon. Despite these species differences, experimental drought caused decreased leaf starch content in dying trees of both species ( P < 0.001). Average dry-season leaf starch content was also a good predictor of drought-survival time for both species ( R2 = 0.93). These results, along with observations of drought-induced reductions to photosynthesis and growth, support carbon limitation as an important process during mortality of these two conifer species. [ABSTRACT FROM AUTHOR]

Non-structural carbohydrates (NSCs) are critical to maintain plant metabolism under stressful environmental conditions, but we do not fully understand how NSC allocation and utilization from storage varies with stress. While it has become established that storage allocation is unlikely to be a mere overflow process, very little empirical evidence has been produced to support this view, at least not for trees. Here we present the results of an intensively monitored experimental manipulation of whole-tree carbon (C) balance (young Picea abies (L.) H Karst.) using reduced atmospheric [CO2] and drought to reduce C sources. We measured specific C storage pools (glucose, fructose, sucrose, starch) over 21 weeks and converted concentration measurement into fluxes into and out of the storage pool. Continuous labeling (13C) allowed us to track C allocation to biomass and non-structural C pools. Net C fluxes into the storage pool occurred mainly when the C balance was positive. Storage pools increased during periods of positive C gain and were reduced under negative C gain. 13C data showed that C was allocated to storage pools independent of the net flux and even under severe C limitation. Allocation to below-ground tissues was strongest in control trees followed by trees experiencing drought followed by those grown under low [CO2]. Our data suggest that NSC storage has, under the conditions of our experimental manipulation (e.g., strong progressive drought, no above-ground growth), a high allocation priority and cannot be considered an overflow process. While these results also suggest active storage allocation, definitive proof of active plant control of storage in woody plants requires studies involving molecular tools. [ABSTRACT FROM AUTHOR]

Non-structural carbohydrates (NSCs) are critical to maintain plant metabolism under stressful environmental conditions, but we do not fully understand how NSC allocation and utilization from storage varies with stress. While it has become established that storage allocation is unlikely to be a mere overflow process, very little empirical evidence has been produced to support this view, at least not for trees. Here we present the results of an intensively monitored experimental manipulation of whole-tree carbon (C) balance (young Picea abies (L.) H Karst.) using reduced atmospheric [CO2] and drought to reduce C sources. We measured specific C storage pools (glucose, fructose, sucrose, starch) over 21 weeks and converted concentration measurement into fluxes into and out of the storage pool. Continuous labeling (13C) allowed us to track C allocation to biomass and non-structural C pools. Net C fluxes into the storage pool occurred mainly when the C balance was positive. Storage pools increased during periods of positive C gain and were reduced under negative C gain. 13C data showed that C was allocated to storage pools independent of the net flux and even under severe C limitation. Allocation to below-ground tissues was strongest in control trees followed by trees experiencing drought followed by those grown under low [CO2]. Our data suggest that NSC storage has, under the conditions of our experimental manipulation (e.g., strong progressive drought, no above-ground growth), a high allocation priority and cannot be considered an overflow process. While these results also suggest active storage allocation, definitive proof of active plant control of storage in woody plants requires studies involving molecular tools. [ABSTRACT FROM AUTHOR]

Fundamental drivers of ecosystem processes such as temperature and precipitation are rapidly changing and creating novel environmental conditions. Forest landscape models ( FLM) are used by managers and policy-makers to make projections of future ecosystem dynamics under alternative management or policy options, but the links between the fundamental drivers and projected responses are weak and indirect, limiting their reliability for projecting the impacts of climate change. We developed and tested a relatively mechanistic method to simulate the effects of changing precipitation on species competition within the LANDIS- II FLM. Using data from a field precipitation manipulation experiment in a piñon pine ( Pinus edulis) and juniper ( Juniperus monosperma) ecosystem in New Mexico ( USA), we calibrated our model to measurements from ambient control plots and tested predictions under the drought and irrigation treatments against empirical measurements. The model successfully predicted behavior of physiological variables under the treatments. Discrepancies between model output and empirical data occurred when the monthly time step of the model failed to capture the short-term dynamics of the ecosystem as recorded by instantaneous field measurements. We applied the model to heuristically assess the effect of alternative climate scenarios on the piñon-juniper ecosystem and found that warmer and drier climate reduced productivity and increased the risk of drought-induced mortality, especially for piñon. We concluded that the direct links between fundamental drivers and growth rates in our model hold great promise to improve our understanding of ecosystem processes under climate change and improve management decisions because of its greater reliance on first principles. [ABSTRACT FROM AUTHOR]

The article focuses on the amplification role of increasing atmospheric moisture demand on tree mortality and regional die-off. It mentions that there are studies which evaluated the effects of increasing atmospheric demand for moisture or the vapour pressure deficit (VPD) to the rates of tree mortality. It states that the relationship of increased temperature and VPD have a global implications to the future of vegetation.

* Premise of the study: Climate-induced forest retreat has profound ecological and biogeochemical impacts, but the physiological mechanisms underlying past tree mortality are poorly understood, limiting prediction of vegetation shifts with climate variation. Climate, drought, fire, and grazing represent agents of tree mortality during the late Cenozoic, but the interaction between drought and declining atmospheric carbon dioxide ([CO2]a) from high to near-starvation levels ~34 million years (Ma) ago has been overlooked. Here, this interaction frames our investigation of sapling mortality through the interdependence of hydraulic function, carbon limitation, and defense metabolism. * Methods: We recreated a changing Cenozoic [CO2]a regime by growing Sequoia sempervirens trees within climate-controlled growth chambers at 1500, 500, or 200 ppm [CO2]a, capturing the decline toward minimum concentrations from 34 Ma. After 7 months, we imposed drought conditions and measured key physiological components linking carbon utilization, hydraulics, and defense metabolism as hypothesized interdependent mechanisms of tree mortality. * Key results: Catastrophic failure of hydraulic conductivity, carbohydrate starvation, and tree death occurred at 200 ppm, but not 500 or 1500 ppm [CO2]a. Furthermore, declining [CO2]a reduced investment in carbon-rich foliar defense compounds that would diminish resistance to biotic attack, likely exacerbating mortality. * Conclusions: Low-[CO2]a-driven tree mortality under drought is consistent with Pleistocene pollen records charting repeated Californian Sequoia forest contraction during glacial periods (180-200 ppm [CO2]a) and may even have contributed to forest retreat as grasslands expanded on multiple continents under low [CO2]a over the past 10 Ma. In this way, geologic intervals of low [CO2]a coupled with drought could impose a demographic bottleneck in tree recruitment, driving vegetation shifts through forest mortality. [ABSTRACT FROM AUTHOR]

ABSTRACT Drought-related tree mortality occurs globally and may increase in the future, but we lack sufficient mechanistic understanding to accurately predict it. Here we present the first field assessment of the physiological mechanisms leading to mortality in an ecosystem-scale rainfall manipulation of a piñon-juniper ( Pinus edulis- Juniperus monosperma) woodland. We measured transpiration ( E) and modelled the transpiration rate initiating hydraulic failure ( Ecrit). We predicted that isohydric piñon would experience mortality after prolonged periods of severely limited gas exchange as required to avoid hydraulic failure; anisohydric juniper would also avoid hydraulic failure, but sustain gas exchange due to its greater cavitation resistance. After 1 year of treatment, 67% of droughted mature piñon died with concomitant infestation by bark beetles ( Ips confusus) and bluestain fungus ( Ophiostoma spp.); no mortality occurred in juniper or in control piñon. As predicted, both species avoided hydraulic failure, but safety margins from Ecrit were much smaller in piñon, especially droughted piñon, which also experienced chronically low hydraulic conductance. The defining characteristic of trees that died was a 7 month period of near-zero gas exchange, versus 2 months for surviving piñon. Hydraulic limits to gas exchange, not hydraulic failure per se, promoted drought-related mortality in piñon pine. [ABSTRACT FROM AUTHOR]

Summary Stomatal closure during drought inhibits carbon uptake and may reduce a tree's defensive capacity. Limited carbon availability during drought may increase a tree's mortality risk, particularly if drought constrains trees' capacity to rapidly produce defenses during biotic attack. We parameterized a new model of conifer defense using physiological data on carbon reserves and chemical defenses before and after a simulated bark beetle attack in mature Pinus edulis under experimental drought. Attack was simulated using inoculations with a consistent bluestain fungus (Ophiostoma sp.) of Ips confusus, the main bark beetle colonizing this tree, to induce a defensive response. Trees with more carbon reserves produced more defenses but measured phloem carbon reserves only accounted for c. 23% of the induced defensive response. Our model predicted universal mortality if local reserves alone supported defense production, suggesting substantial remobilization and transport of stored resin or carbon reserves to the inoculation site. Our results show that de novo terpene synthesis represents only a fraction of the total measured phloem terpenes in P. edulis following fungal inoculation. Without direct attribution of phloem terpene concentrations to available carbon, many studies may be overestimating the scale and importance of de novo terpene synthesis in a tree's induced defense response. [ABSTRACT FROM AUTHOR]

Abstract: Widespread tree mortality events have recently been observed in several biomes. To effectively quantify the severity and extent of these events, tools that allow for rapid assessment at the landscape scale are required. Past studies using high spatial resolution satellite imagery have primarily focused on detecting green, red, and gray tree canopies during and shortly after tree damage or mortality has occurred. However, detecting trees in various stages of death is not always possible due to limited availability of archived satellite imagery. Here we assess the capability of high spatial resolution satellite imagery for tree mortality detection in a southwestern U.S. mixed species woodland using archived satellite images acquired prior to mortality and well after dead trees had dropped their leaves. We developed a multistep classification approach that uses: supervised masking of non-tree image elements; bi-temporal (pre- and post-mortality) differencing of normalized difference vegetation index (NDVI) and red:green ratio (RGI); and unsupervised multivariate clustering of pixels into live and dead tree classes using a Gaussian mixture model. Classification accuracies were improved in a final step by tuning the rules of pixel classification using the posterior probabilities of class membership obtained from the Gaussian mixture model. Classifications were produced for two images acquired post-mortality with overall accuracies of 97.9% and 98.5%, respectively. Classified images were combined with land cover data to characterize the spatiotemporal characteristics of tree mortality across areas with differences in tree species composition. We found that 38% of tree crown area was lost during the drought period between 2002 and 2006. The majority of tree mortality during this period was concentrated in piñon-juniper (Pinus edulis-Juniperus monosperma) woodlands. An additional 20% of the tree canopy died or was removed between 2006 and 2011, primarily in areas experiencing wildfire and management activity. -Our results demonstrate that unsupervised clustering of bi-temporal NDVI and RGI differences can be used to detect tree mortality resulting from numerous causes and in several forest cover types. [Copyright &y& Elsevier]