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Root respiration often exhibits a direct and immediate decline with increasing concentrations of ambient soil carbon dioxide concentration ([CO2]), and recent evidence suggests this decline may be attributable to a decline in maintenance respiration within the root. If true, this concept could provide a clue to the biochemical process underlying respiratory inhibition as well as improve our knowledge of the timing and degree to which this inhibition occurs in nature. To test the hypothesis that maintenance respiration exhibits a direct, negative response to increasing [CO2], we measured total respiration in intact root systems of western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings grown at different relative growth rates and exposed to soil [CO2]s ranging from 91 to 7008 μmol mol−1. Analysis of covariance was used to separate maintenance from total respiration. Total respiration declined exponentially with increasing [CO2]. Maintenance respiration, which comprised 85% of total respiration over all treatments, also declined exponentially with increasing [CO2]. Growth respiration was not inhibited at any [CO2]. These findings may explain why roots of some fast-growing species do not show [CO2] inhibition. [ABSTRACT FROM AUTHOR]

• Whole-tree transpiration, radial growth, and δ13C were measured in a coastal forest. • Shoreline trees had higher transpiration rates than the nearby upland trees. • Shoreline trees had similar growth and gas exchange compared to the upland trees. • Trees adapted to shoreline conditions by maintaining homeostasis in water relations and growth. Coastal shoreline forests are vulnerable to seawater exposure, the impacts of which will increase due to sea-level rise, but the long-term adaptation strategies and vulnerability of coastal forests are not well understood. We used whole-tree transpiration, leaf water potential, tree-ring width, and tree-ring δ13C (a proxy for intrinsic water use efficiency, iWUE) to examine the long-term adaptation strategies of red maple (Acer rubrum) trees at the coastal interface (i.e., shoreline) and nearby upland in Maryland, USA. Red maple trees that grew along the shoreline and were exposed to slightly saline water (up to two PSU) had higher transpiration rates than those growing in the nearby upland forest during a wet year, but these differences disappeared during a normal precipitation year. Shoreline trees grew more slowly than upland trees over the last four decades, but these growth differences have disappeared in the last six years. Shoreline and upland red maple trees had similar variation in iWUE, indicating that higher transpiration rates of the seawater-exposed trees did not translate into differences in water use efficiency. There were no differences in predawn and midday water potential between upland and shoreline trees, suggesting no additional water stress occurs in shoreline trees. These findings indicate that mature red maple in our coastal study site maintains gas exchange and growth at a consistent or homeostatic level under slight soil salinity. [ABSTRACT FROM AUTHOR]

Summary: Plant water uptake from the soil is a crucial element of the global hydrological cycle and essential for vegetation drought resilience. Yet, knowledge of how the distribution of water uptake depth (WUD) varies across species, climates, and seasons is scarce relative to our knowledge of aboveground plant functions. With a global literature review, we found that average WUD varied more among biomes than plant functional types (i.e. deciduous/evergreen broadleaves and conifers), illustrating the importance of the hydroclimate, especially precipitation seasonality, on WUD. By combining records of rooting depth with WUD, we observed a consistently deeper maximum rooting depth than WUD with the largest differences in arid regions – indicating that deep taproots act as lifelines while not contributing to the majority of water uptake. The most ubiquitous observation across the literature was that woody plants switch water sources to soil layers with the highest water availability within short timescales. Hence, seasonal shifts to deep soil layers occur across the globe when shallow soils are drying out, allowing continued transpiration and hydraulic safety. While there are still significant gaps in our understanding of WUD, the consistency across global ecosystems allows integration of existing knowledge into the next generation of vegetation process models. [ABSTRACT FROM AUTHOR]

Summary: Shifts in the age or turnover time of non‐structural carbohydrates (NSC) may underlie changes in tree growth under long‐term increases in drought stress associated with climate change. But NSC responses to drought are challenging to quantify, due in part to large NSC stores in trees and subsequently long response times of NSC to climate variation.We measured NSC age (Δ14C) along with a suite of ecophysiological metrics in Pinus edulis trees experiencing either extreme short‐term drought (−90% ambient precipitation plot, 2020–2021) or a decade of severe drought (−45% plot, 2010–2021). We tested the hypothesis that carbon starvation – consumption exceeding synthesis and storage – increases the age of sapwood NSC.One year of extreme drought had no impact on NSC pool size or age, despite significant reductions in predawn water potential, photosynthetic rates/capacity, and twig and needle growth. By contrast, long‐term drought halved the age of the sapwood NSC pool, coupled with reductions in sapwood starch concentrations (−75%), basal area increment (−39%), and bole respiration rates (−28%).Our results suggest carbon starvation takes time, as tree carbon reserves appear resilient to extreme disturbance in the short term. However, after a decade of drought, trees apparently consumed old stored NSC to support metabolism. [ABSTRACT FROM AUTHOR]

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]

Vegetation browning has progressively increased globally under a background of climate change. Under climate change, increases in the intensity and duration of disturbances (i.e., drought, wildfire, insect and pathogen outbreaks, and wind and ice storms) are the major driving factors behind vegetation browning. The weakening of ecosystem services and functions and alterations to vegetation structure and species composition will be accompanied by vegetation browning. Potential positive browning feedbacks may exacerbate climate change. As global climate conditions continue to change, disturbance regimes and environmental drivers will continue to shift, impacting global vegetation dynamics. Following a period of vegetation greening, there has been a progressive increase in remotely sensed vegetation browning globally. Given the many societal benefits that forests provide, it is critical that we understand vegetation dynamic alterations. Here, we review associative drivers, impacts, and feedbacks, revealing the complexity of browning. Concomitant increases in browning include the weakening of ecosystem services and functions and alterations to vegetation structure and species composition, as well as the development of potential positive climate change feedbacks. Also discussed are the current challenges in browning detection and understanding associated impacts and feedbacks. Finally, we outline recommended strategies. [ABSTRACT FROM AUTHOR]

Tropical forest dynamics play a crucial role in the global carbon, water, and energy cycles. However, realistically simulating the dynamics of competition and coexistence between different plant functional types (PFTs) in tropical forests remains a significant challenge. This study aims to improve the modeling of PFT coexistence in the Functionally Assembled Terrestrial Ecosystem Simulator (FATES), a vegetation demography model implemented in the Energy Exascale Earth System Model (E3SM) land model (ELM), ELM-FATES. Specifically, we explore (1) whether plant trait relationships established from field measurements can constrain ELM-FATES simulations and (2) whether machine learning (ML)-based surrogate models can emulate the complex ELM-FATES model and optimize parameter selections to improve PFT coexistence modeling. We conducted three ensembles of ELM-FATES experiments at a tropical forest site near Manaus, Brazil. By comparing the ensemble experiments without (Exp-CTR) and with (Exp-OBS) consideration of observed trait relationships, we found that accounting for these relationships slightly improves the simulations of water, energy, and carbon variables when compared to observations but degrades the simulation of PFT coexistence. Using ML-based surrogate models trained on Exp-CTR, we optimized the trait parameters in ELM-FATES and conducted another ensemble of experiments (Exp-ML) with these optimized parameters. The proportion of PFT coexistence experiments significantly increased from 21 % in Exp-CTR to 73 % in Exp-ML. After filtering the experiments that allow for PFT coexistence to agree with observations (within 15 % tolerance), 33 % of the Exp-ML experiments were retained, which is a significant improvement compared to the 1.4 % in Exp-CTR. Exp-ML also accurately reproduces the annual means and seasonal variations in water, energy, and carbon fluxes and the field inventory of aboveground biomass. This study represents a reproducible method that utilizes machine learning to identify parameter values that improve model fidelity against observations and PFT coexistence in vegetation demography models for diverse ecosystems. Our study also suggests the need for new mechanisms to enhance the robust simulation of coexisting plants in ELM-FATES and has significant implications for modeling the response and feedbacks of ecosystem dynamics to climate change. [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]

Summary: The representation of stomatal regulation of transpiration and CO2 assimilation is key to forecasting terrestrial ecosystem responses to global change. Given its importance in determining the relationship between forest productivity and climate, accurate and mechanistic model representation of the relationship between stomatal conductance (gs) and assimilation is crucial.We assess possible physiological and mechanistic controls on the estimation of the g1 (stomatal slope, inversely proportional to water use efficiency) and g0 (stomatal intercept) parameters, using diurnal gas exchange surveys and leaf‐level response curves of six tropical broadleaf evergreen tree species.g1 estimated from ex situ response curves averaged 50% less than g1 estimated from survey data. While g0 and g1 varied between leaves of different phenological stages, the trend was not consistent among species. We identified a diurnal trend associated with g1 and g0 that significantly improved model projections of diurnal trends in transpiration.The accuracy of modeled gs can be improved by accounting for variation in stomatal behavior across diurnal periods, and between measurement approaches, rather than focusing on phenological variation in stomatal behavior. Additional investigation into the primary mechanisms responsible for diurnal variation in g1 will be required to account for this phenomenon in land‐surface models. [ABSTRACT FROM AUTHOR]

Within vascular plants, the partitioning of hydraulic resistance along the soil‐to‐leaf continuum affects transpiration and its response to environmental conditions. In trees, the fractional contribution of leaf hydraulic resistance (Rleaf) to total soil‐to‐leaf hydraulic resistance (Rtotal), or fRleaf (=Rleaf/Rtotal), is thought to be large, but this has not been tested comprehensively. We compiled a multibiome data set of fRleaf using new and previously published measurements of pressure differences within trees in situ. Across 80 samples, fRleaf averaged 0.51 (95% confidence interval [CI] = 0.46−0.57) and it declined with tree height. We also used the allometric relationship between field‐based measurements of soil‐to‐leaf hydraulic conductance and laboratory‐based measurements of leaf hydraulic conductance to compute the average fRleaf for 19 tree samples, which was 0.40 (95% CI = 0.29−0.56). The in situ technique produces a more accurate descriptor of fRleaf because it accounts for dynamic leaf hydraulic conductance. Both approaches demonstrate the outsized role of leaves in controlling tree hydrodynamics. A larger fRleaf may help stems from loss of hydraulic conductance. Thus, the decline in fRleaf with tree height would contribute to greater drought vulnerability in taller trees and potentially to their observed disproportionate drought mortality. Summary Statement: In trees, we found that leaves contribute a large and varying fraction of the total hydraulic resistance within the soil‐to‐leaf continuum that feeds the transpiration stream. [ABSTRACT FROM AUTHOR]

Topographic heterogeneity and lateral subsurface flow at the hillslope scale of ≤1 km may have outsized impacts on tropical forest through their impacts on water available to plants under water-stressed conditions. However, vegetation dynamics and finer-scale hydrologic processes are not concurrently represented in Earth system models. In this study, we integrate the Energy Exascale Earth System Model (E3SM) land model (ELM) that includes the Functionally Assembled Terrestrial Ecosystem Simulator (FATES), with a three-dimensional hydrology model (ParFlow) to explicitly resolve hillslope topography and subsurface flow and perform numerical experiments to understand how hillslope-scale hydrologic processes modulate vegetation along water availability gradients at Barro Colorado Island (BCI), Panama. Our simulations show that groundwater table depth (WTD) can play a large role in governing aboveground biomass (AGB) when drought-induced tree mortality is triggered by hydraulic failure. Analyzing the simulations using random forest (RF) models, we find that the domain-wide simulated AGB and WTD can be well predicted by static topographic attributes, including surface elevation, slope, and convexity, and adding soil moisture or groundwater table depth as predictors further improves the RF models. Different model representations of mortality due to hydraulic failure can change the dominant topographic driver for the simulated AGB. Contrary to the simulations, the observed AGB in the well-drained 50 ha forest census plot within BCI cannot be well predicted by the RF models using topographic attributes and observed soil moisture as predictors, suggesting other factors such as nutrient status may have a larger influence on the observed AGB. The new coupled model may be useful for understanding the diverse impact of local heterogeneity by isolating the water availability and nutrient availability from the other external and internal factors in ecosystem modeling. [ABSTRACT FROM AUTHOR]

As CO 2 concentration in the atmosphere rises, there is a need for improved physical understanding of its impact on global plant transpiration. This knowledge gap poses a major hurdle in robustly projecting changes in the global hydrologic cycle. For this reason, here we review the different processes by which atmospheric CO 2 concentration affects plant transpiration, the several uncertainties related to the complex physiological and radiative processes involved, and the knowledge gaps which need to be filled in order to improve predictions of plant transpiration. Although there is a high degree of certainty that rising CO 2 will impact plant transpiration, the exact nature of this impact remains unclear due to complex interactions between CO 2 and climate, and key aspects of plant morphology and physiology. The interplay between these factors has substantial consequences not only for future climate and global vegetation, but also for water availability needed for sustaining the productivity of terrestrial ecosystems. Future changes in global plant transpiration in response to enhanced CO 2 are expected to be driven by water availability, atmospheric evaporative demand, plant physiological processes, emergent plant disturbances related to increasing temperatures, and the modification of plant physiology and coverage. Considering the universal sensitivity of natural and agricultural systems to terrestrial water availability we argue that reliable future projections of transpiration is an issue of the highest priority, which can only be achieved by integrating monitoring and modeling efforts to improve the representation of CO 2 effects on plant transpiration in the next generation of earth system models. [ABSTRACT FROM AUTHOR]

Plants are expected to respond to drought by maximizing the efficiency of the most limiting resource, the water use efficiency (WUE), at the expense of nitrogen and carbon use efficiencies (NUE and CUE). Therefore, plants resource use efficiencies are viewed as indicators of species drought tolerance., We tested these predictions by measuring leaf-level intrinsic WUE (WUEi, the ratio of net assimilation to stomatal conductance), photosynthetic NUE (PNUE, the ratio of daily maximum net assimilation to leaf nitrogen content) and leaf-scale CUE (approached by the ratio of night-time respiration to daytime net assimilation, Rd /An) in piñon pine and juniper, two tree species that differ in drought tolerance and vulnerability to drought-induced mortality. Variations in resource use efficiency in the two species were measured in response to seasonal drought and in response to an ecosystem-scale precipitation manipulation experiment comprising three precipitation treatments: ambient, irrigation (+30%) and partial rainfall exclusion (−45%)., Increasing water limitation, either seasonally or across treatments, resulted in increased WUE and decreased PNUE and CUE in both species. WUE, PNUE and CUE varied more strongly in response to water limitation than across species and converged to the same relationships against precipitation for piñon and juniper., Plasticity in WUE, PNUE and CUE in response to water limitation was associated, in both species, with low carbon acquisition during drought. Our results exhibited a convergence in resource use efficiency across piñon and juniper which contradicts the paradigm that resource use efficiencies are indicators of species drought tolerance and ecological strategy. [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]

Topographic heterogeneity and lateral subsurface flow at the hillslope scale of = 1km may have outsized impacts on tropical forest through their impacts on water available to plants under water stressed conditions. However, vegetation dynamics and finer-scale hydrologic processes are not concurrently represented in Earth system models. In this study, we integrate the Energy Exascale Earth System Model (E3SM) Land Model (ELM) that includes the Functionally-Assembled Terrestrial Ecosystem Simulator (FATES), with a three-dimensional hydrology model (ParFlow) to explicitly resolve hillslope topography and subsurface flow and perform numerical experiments to understand how hillslope scale hydrologic processes modulate vegetation along water availability gradients at Barro Colorado Island (BCI), Panama. Our simulations show that groundwater table depth (WTD) can play a large role in governing aboveground biomass (AGB) when drought-induced tree mortality is triggered by hydraulic failure. Analyzing the simulations using random forest (RF) models, we find that the domainwide simulated AGB and WTD can be well predicted by static topographic attributes including surface elevation, slope and convexity, and adding soil moisture or ground water table depth as predictors further improves the RF models. Different model representations of mortality due to hydraulic failure can change the dominant topographic driver for the simulated AGB. Contrary to the simulations, the observed AGB in the well-drained 50-ha forest census plot within BCI cannot be well predicted by the RF models using topographic attributes and observed soil moisture as predictors, suggesting other factors such as nutrient status may have larger influence on the observed AGB. The new coupled model may be useful for understanding the diverse impact of local heterogeneity by isolating the water availability and nutrient availability from the other external and internal factors in ecosystem modeling. [ABSTRACT FROM AUTHOR]

This article is a Commentary on Cosme et al., 215: 113–125. [ABSTRACT FROM AUTHOR]

• 2015 ENSO led to potential water supply-demand imbalance at canopy level. • In the Amazon forest VPD peaks in the early afternoon. • g s is higher during the morning period to minimize water loss. • H area effectively quantifies Amazon drought stress. • H area is shaped by temporal lags between the observed variables. Understanding forest water limitation during droughts within a warming climate is essential for accurate predictions of forest-climate interactions. In hyperdiverse ecosystems like the Amazon forest, the mechanisms shaping hysteresis patterns in transpiration relative to environmental factors are not well understood. From this perspective, we investigated these dynamics by conducting in situ leaf-level measurements throughout and after the 2015 El Niño-Southern Oscillation (ENSO) drought. Our findings indicate a substantial increase in the hysteresis area (H area) among transpiration (E), vapor pressure deficit (VPD), and stomatal conductance (g s) at canopy level during the ENSO peak, attributed to both temporal lag and differences in magnitude between g s and VPD peaks. Specifically, the canopy species Pouteria anomala exhibited an increased H area , due to earlier maximum g s rates leading to a greater temporal lag with VPD compared to the post-drought period. Additionally, leaf water potential (Ψ L) and canopy temperature (T canopy) showed larger H area during the ENSO peak compared to post-drought conditions across all studied species, suggesting that stomatal closure, particularly during the afternoon, acts to minimize water loss and may explain the counterclockwise hysteresis observed between Ψ L and T canopy. The pronounced H area during the drought points to a potential imbalance between water supply and demand, underlining the role of stomatal behavior of isohydric species in response to drought. Canopy-level transpiration during the 2015 ENSO-driven drought in the Central Amazon exhibited significant deviations due to exceptionally high Vapor Pressure Deficit (VPD) conditions and increased temporal differences between the peaks of stomatal conductance (g s) and VPD. This resulted in an increased hysteresis effect, as evidenced by the expanded hysteresis area (H area) across multiple ecophysiological variables compared to the post-drought period. [Display omitted] [ABSTRACT FROM AUTHOR]

• Inundation events introduce oxygen to our wetlands, which is then rapidly consumed. • Oxygen consumption is much more rapid in wetlands vs. upland forest soils. • Oxygen consumption in subsurface B horizon soils appears to be carbon-limited. • Integrating field, lab, and models improves our understanding of biogeochemical processes. The coastal terrestrial-aquatic interface (TAI) is a highly dynamic system characterized by strong physical, chemical, and biological gradients. In particular, shifting soil redox conditions and consumption of terminal electron acceptors, due in part to dynamic hydrologic conditions, is a strong driver of carbon availability and transformations across TAIs. However, while redox dynamics are well described, our ability to quantitatively forecast rates of oxic to anoxic shifts in soils with different characteristics and inundation regimes is limited. We integrated field measurements, laboratory incubations, and model simulations to improve mechanistic understanding of oxygen consumption dynamics in coastal soils. Continuous in situ monitoring unexpectedly revealed that flooding caused temporary spikes in subsurface dissolved oxygen followed by rapid consumption in the wetlands. To further investigate these mechanisms in a controlled setting, we performed laboratory incubations using surface and subsurface soils from a TAI gradient (defined here as upland forest to transitional forest to wetland) in Western Lake Erie to measure oxygen consumption rates in TAI soils during flood events. In our experiments, wetland soils reached anoxia the fastest, in ∼ 9 h on average, whereas upland soils turned anoxic in ∼ 18 h. Subsurface upland soils did not turn anoxic even after two weeks of saturation in the lab, and their oxygen consumption patterns suggested carbon and/or nutrient limitation. These results are consistent with in-situ groundwater redox and oxygen measurements in the field, where wetland soils exhibited the highest rates of oxygen consumption along the TAI. Model simulations of oxygen consumption suggested that oxygen consumption had stronger abiotic controls in wetland soils but stronger biotic controls in upland soils, providing a useful framework for future incubation experiments. Microbial activity is a strong driver of oxygen consumption in TAI soils, although it is constrained by the availability of dissolved carbon in subsurface soils. [ABSTRACT FROM AUTHOR]

Sensitivity of forest mortality to drought in carbon‐dense tropical forests remains fraught with uncertainty, while extreme droughts are predicted to be more frequent and intense. Here, the potential of temporal autocorrelation of high‐frequency variability in Landsat Enhanced Vegetation Index (EVI), an indicator of ecosystem resilience, to predict spatial and temporal variations of forest biomass mortality is evaluated against in situ census observations for 64 site‐year combinations in Costa Rican tropical dry forests during the 2015 ENSO drought. Temporal autocorrelation, within the optimal moving window of 24 months, demonstrated robust predictive power for in situ mortality (leave‐one‐out cross‐validation R2 = 0.54), which allows for estimates of annual biomass mortality patterns at 30 m resolution. Subsequent spatial analysis showed substantial fine‐scale heterogeneity of forest mortality patterns, largely driven by drought intensity and ecosystem properties related to plant water use such as forest deciduousness and topography. Highly deciduous forest patches demonstrated much lower mortality sensitivity to drought stress than less deciduous forest patches after elevation was controlled. Our results highlight the potential of high‐resolution remote sensing to "fingerprint" forest mortality and the significant role of ecosystem heterogeneity in forest biomass resistance to drought. [ABSTRACT FROM AUTHOR]

Recent observations of elevated tree mortality following climate extremes, like heat and drought, raise concerns about climate change risks to global forest health. We currently lack both sufficient data and understanding to identify whether these observations represent a global trend toward increasing tree mortality. Here, we document events of sudden and unexpected elevated tree mortality following heat and drought events in ecosystems that previously were considered tolerant or not at risk of exposure. These events underscore the fact that climate change may affect forests with unexpected force in the future. We use the events as examples to highlight current difficulties and challenges for realistically predicting such tree mortality events and the uncertainties about future forest condition. Advances in remote sensing technology and greater availably of high-resolution data, from both field assessments and satellites, are needed to improve both understanding and prediction of forest responses to future climate change. [ABSTRACT FROM AUTHOR]

Observations of forest mortality are increasing globally, but relatively little is known regarding the underlying mechanisms driving these events. Tree rings carry physiological signatures that may be used as a tool for retrospective analyses. We capitalized on a local soil water drainage event in 1982 that resulted in increased mortality within a stand of oak trees (Quercus robur), to examine the underlying physiological patterns associated with survival and death in response to soil water limitations. Pre-dawn water potentials showed more negative values for trees in the process of dying compared with those that survived. We used tree rings formed over the 123 years prior to mortality to estimate productivity from basal area increment (BAI, mm2), multiple xylem hydraulic parameters via anatomical measurements and crown-level gas exchange via carbon isotope discrimination (Δ, ‰). Oaks that died had significantly higher BAI values than trees that survived until the drainage event, after which the BAI of trees that died declined dramatically. Hydraulic diameter and conductivity of vessels in trees that died were higher than in surviving trees until the last 5 years prior to mortality, at which time both groups had similar values. Trees that died had consistently lower Δ values than trees that survived. Therefore, tree mortality in this stand was associated with physiological differences prior to the onset of soil water reduction. We propose that trees that died may have been hydraulically underbuilt for dry conditions, which predisposes them to severe hydraulic constraints and subsequent mortality. Measurements of above-ground/below-ground dry mass partitioning will be critical to future tests of this hypothesis. Based on these results, it is probable that pedunculate oak trees will experience greater future mortality if climate changes cause more severe droughts than the trees have experienced previously. [ABSTRACT FROM AUTHOR]

Mesophyll conductance to CO2 (gm) limits carbon assimilation and influences carbon isotope discrimination (Δ) under most environmental conditions. Current work is elucidating the environmental regulation of gm, but the influence of gm on model predictions of Δ remains poorly understood. In this study, field measurements of Δ and gm were obtained using a tunable diode laser spectroscope coupled to portable photosynthesis systems. These data were used to test the importance of gm in predicting Δ using the comprehensive Farquhar model of Δ (Δcomp), where gm was parameterized using three methods based on: (i) mean gm; (ii) the relationship between stomatal conductance (gs) and gm; and (iii) the relationship between time of day (TOD) and gm. Incorporating mean gm, gs-based gm, and TOD-based gm did not consistently improve Δcomp predictions of field-grown juniper compared with the simple model of Δ (Δsimple) that omits fractionation factors associated with gm and decarboxylation. Sensitivity tests suggest that b, the fractionation due to carboxylation, was lower (25‰) than the value commonly used in Δcomp (29‰) and Δsimple (27‰). These results demonstrate the limits of all tested models in predicting observed juniper Δ, largely due to unexplained offsets between predicted and observed values that were not reconciled in sensitivity tests of variability in gm, b, or e, the day respiratory fractionation. [ABSTRACT FROM PUBLISHER]

Heat and drought affect plant chemical defenses and thereby plant susceptibility to pests and pathogens. Monoterpenes are of particular importance for conifers as they play critical roles in defense against bark beetles. To date, work seeking to understand the impacts of heat and drought on monoterpenes has primarily focused on young potted seedlings, leaving it unclear how older age classes that are more vulnerable to bark beetles might respond to stress. Furthermore, we lack a clear picture of what carbon resources might be prioritized to support monoterpene synthesis under drought stress. To address this, we measured needle and woody tissue monoterpene concentrations and physiological variables simultaneously from mature piñon pines (Pinus edulis) from a unique temperature and drought manipulation field experiment. While heat had no effect on total monoterpene concentrations, trees under combined heat and drought stress exhibited ~ 85% and 35% increases in needle and woody tissue, respectively, over multiple years. Plant physiological variables like maximum photosynthesis each explained less than 10% of the variation in total monoterpenes for both tissue types while starch and glucose + fructose measured 1-month prior explained ~ 45% and 60% of the variation in woody tissue total monoterpene concentrations. Although total monoterpenes increased under combined stress, some key monoterpenes with known roles in bark beetle ecology decreased. These shifts may make trees more favorable for bark beetle attack rather than well defended, which one might conclude if only considering total monoterpene concentrations. Our results point to cumulative and synergistic effects of heat and drought that may reprioritize carbon allocation of specific non-structural carbohydrates toward defense. [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]

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]

Sea-level rise is one of the most critical challenges facing coastal ecosystems under climate change. Observations of elevated tree mortality in global coastal forests are increasing, but important knowledge gaps persist concerning the mechanism of salinity stress-induced nonhalophytic tree mortality. We monitored progressive mortality and associated gas exchange and hydraulic shifts in Sitka-spruce (Picea sitchensis) trees located within a salinity gradient under an ecosystem-scale change of seawater exposure in Washington State, USA. Percentage of live foliated crown (PLFC) decreased and tree mortality increased with increasing soil salinity during the study period. A strong reduction in gas exchange and xylem hydraulic conductivity (Ks) occurred during tree death, with an increase in the percentage loss of conductivity (PLC) and turgor loss point (πtlp). Hydraulic and osmotic shifts reflected that hydraulic function declined from seawater exposure, and dying trees were unable to support osmotic adjustment. Constrained gas exchange was strongly related to hydraulic damage at both stem and leaf levels. Significant correlations between foliar sodium (Na+) concentration and gas exchange and key hydraulic parameters (Ks, PLC, and πtlp) suggest that cellular injury related to the toxic effects of ion accumulation impacted the physiology of these dying trees. This study provides evidence of toxic effects on the cellular function that manifests in all aspects of plant functioning, leading to unfavourable osmotic and hydraulic conditions. [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]

Climate change is expected to pose a global threat to forest health by intensifying extreme events like drought and insect attacks. Carbon allocation is a fundamental process that determines the adaptive responses of long-lived late-maturing organisms like trees to such stresses. However, our mechanistic understanding of how trees coordinate and set allocation priorities among different sinks (e.g., growth and storage) under severe source limitation remains limited. Using flux measurements, isotopic tracing, targeted metabolomics, and transcriptomics, we investigated how limitation of source supply influences sink activity, particularly growth and carbon storage, and their relative regulation in Norway spruce (Picea abies) clones. During photosynthetic deprivation, absolute rates of respiration, growth, and allocation to storage all decline. When trees approach neutral carbon balance, i.e., daytime net carbon gain equals nighttime carbon loss, genes encoding major enzymes of metabolic pathways remain relatively unaffected. However, under negative carbon balance, photosynthesis and growth are down-regulated while sucrose and starch biosynthesis pathways are up-regulated, indicating that trees prioritize carbon allocation to storage over growth. Moreover, trees under negative carbon balance actively increase the turnover rate of starch, lipids, and amino acids, most likely to support respiration and mitigate stress. Our study provides molecular evidence that trees faced with severe photosynthetic limitation strategically regulate storage allocation and consumption at the expense of growth. Understanding such allocation strategies is crucial for predicting how trees may respond to extreme events involving steep declines in photosynthesis, like severe drought, or defoliation by heat waves, late frost, or insect attack. [ABSTRACT FROM AUTHOR]

Plants are expected to face increasing water stress under future climate change. Most land surface models, including Noah‐MP, employ an idealized "big‐leaf" concept to regulate water and carbon fluxes in response to soil moisture stress through empirical soil hydraulics schemes (SHSs). However, such schemes have been shown to cause significant uncertainties in carbon and water simulations. In this paper, we present a novel plant hydraulics scheme (PHS) for Noah‐MP (hereafter, Noah‐MP‐PHS), which employs a big‐tree rather than big‐leaf concept, wherein the whole‐plant hydraulic strategy is considered, including root‐level soil water acquisition, stem‐level hydraulic conductance and capacitance, and leaf‐level anisohydricity and hydraulic capacitance. Evaluated against plot‐level observations from a mature, mixed hardwood forest at the University of Michigan Biological Station and compared with the default Noah‐MP, Noah‐MP‐PHS better represents plant water stress and improves water and carbon simulations, especially during periods of dry soil conditions. Noah‐MP‐PHS also improves the asymmetrical diel simulation of gross primary production under low soil moisture conditions. Noah‐MP‐PHS is able to reproduce different patterns of transpiration, stem water storage and root water uptake during a 2‐week dry‐down period for two species with contrasting plant hydraulic behaviors, i.e., the "cavitation risk‐averse" red maple and the "cavitation risk‐prone" red oak. Sensitivity experiments with plant hydraulic capacitance show that the stem water storage enables nocturnal plant water recharge, affects plant water use efficiency, and provides an important buffer to relieve xylem hydraulic stress during dry soil conditions. Plain Language Summary: Plants regulate transpiration dynamically through the stomatal aperture, which, in many cases, is governed by plant water status and hydraulic properties. Plant hydraulics describes the mechanics of water movement through plant vascular systems, which is the culmination of emergent phenotypical hydraulic functional traits at the leaf, stem, and root levels. Such physiological mechanisms are excluded in most land surface models, which typically represent plant water stress through empirical soil hydraulics schemes (SHSs) based on either soil water content or soil water potential. In this study, we present a novel plant hydraulics scheme (PHS) to represent plant water stress and the regulation of stomatal conductance for use in the Noah‐MP land surface model. Our results show Noah‐MP‐PHS performs better in its water and carbon simulations than the default Noah‐MP with traditional SHSs, especially under dry soil conditions. Noah‐MP‐PHS also successfully captures the different plant hydraulic behaviors between the "cavitation risk‐averse" red maple and the "cavitation risk‐prone" red oak. Sensitivity experiments also highlight the vital role played by plant water storage in water and carbon simulations in terms of buffering xylem hydraulic stress during soil moisture dry‐down periods. The incorporation of a holistic, whole‐plant hydraulic strategy, along with hydraulic trait‐based vegetation representation, can serve to improve simulations of carbon and water fluxes, particularly in cases of drought and other related disturbances. Key Points: Noah‐MP‐PHS improves the water and carbon simulations over the default soil hydraulics schemes, especially under dry soil conditionsNoah‐MP‐PHS captures different plant hydraulic behaviors between the "cavitation risk‐averse" maple and the "cavitation risk‐prone" oakPlant water storage plays a vital role in water and carbon fluxes and relieves xylem hydraulic stress during soil moisture dry‐down periods [ABSTRACT FROM AUTHOR]

Summary: ●Plants are characterized by the iso/anisohydry continuum depending on how they regulate leaf water potential (ΨL). However, how iso/anisohydry changes over time in response to year‐to‐year variations in environmental dryness and how such responses vary across different regions remains poorly characterized.●We investigated how dryness, represented by aridity index, affects the interannual variability of ecosystem iso/anisohydry at the regional scale, estimated using satellite microwave vegetation optical depth (VOD) observations. This ecosystem‐level analysis was further complemented with published field observations of species‐level ΨL.●We found different behaviors in the directionality and sensitivity of isohydricity (σ) with respect to the interannual variation of dryness in different ecosystems. These behaviors can largely be differentiated by the average dryness of the ecosystem itself: in mesic ecosystems, σ decreases in drier years with a higher sensitivity to dryness; in xeric ecosystems, σ increases in drier years with a lower sensitivity to dryness. These results were supported by the species‐level synthesis.●Our study suggests that how plants adjust their water use across years – as revealed by their interannual variability in isohydricity – depends on the dryness of plants' living environment. This finding advances our understanding of plant responses to drought at regional scales. [ABSTRACT FROM AUTHOR]