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Summary: Intraspecific variation in functional traits may mediate tree species' drought resistance, yet whether trait variation is due to genotype (G), environment (E), or G×E interactions remains unknown. Understanding the drivers of intraspecific trait variation and whether variation mediates drought response can improve predictions of species' response to future drought.Using populations of quaking aspen spanning a climate gradient, we investigated intraspecific variation in functional traits in the field as well as the influence of G and E among propagules in a common garden. We also tested for trait‐mediated trade‐offs in growth and drought stress tolerance.We observed intraspecific trait variation among the populations, yet this variation did not necessarily translate to higher drought stress tolerance in hotter/drier populations. Additionally, plasticity in the common garden was low, especially in propagules derived from the hottest/driest population. We found no growth–drought stress tolerance trade‐offs and few traits exhibited significant relationships with mortality in the natural populations, suggesting that intraspecific trait variation among the traits measured did not strongly mediate responses to drought stress.Our results highlight the limits of trait‐mediated responses to drought stress and the complex G×E interactions that may underlie drought stress tolerance variation in forests in dry environments. [ABSTRACT FROM AUTHOR]

Climate change is stressing many forests around the globe, yet some tree species may be able to persist through acclimation and adaptation to new environmental conditions. The ability of a tree to acclimate during its lifetime through changes in physiology and functional traits, defined here as its acclimation potential, is not well known.We investigated the acclimation potential of trembling aspen Populus tremuloides and ponderosa pine Pinus ponderosa trees by examining within‐species variation in drought response functional traits across both space and time, and how trait variation influences drought‐induced tree mortality. We measured xylem tension, morphological traits and physiological traits on mature trees in southwestern Colorado, USA across a climate gradient that spanned the distribution limits of each species and 3 years with large differences in climate.Trembling aspen functional traits showed high within‐species variation, and osmotic adjustment and carbon isotope discrimination were key determinants for increased drought tolerance in dry sites and in dry years. However, trembling aspen trees at low elevation were pushed past their drought tolerance limit during the severe 2018 drought year, as elevated mortality occurred. Higher specific leaf area during drought was correlated with higher percentages of canopy dieback the following year. Ponderosa pine functional traits showed less within‐species variation, though osmotic adjustment was also a key mechanism for increased drought tolerance. Remarkably, almost all traits varied more year‐to‐year than across elevation in both species.Our results shed light on the scope and limits of intraspecific trait variation for mediating drought responses in key southwestern US tree species and will help improve our ability to model and predict forest responses to climate change. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Tree species display a wide variety of water-use strategies, growth rates and capacity to tolerate drought. However, if we want to forecast species capacity to cope with increasing aridity and drought, we need to identify which measurable traits confer resilience to drought across species. Here, we use a global tree ring network (65 species; 1931 site series of ring-width indices--RWI) to evaluate the relationship of long-term growth-drought sensitivity (RWI-SPEI drought index relationship) and short-term growth response to extreme drought episodes (resistance, recovery and resilience indices) with functional traits related to leaf, wood and hydraulic properties. Furthermore, we assess the influence of climate (temperature, precipitation and climatic water deficit) on these trait-growth relationships. We found a close correspondence between the long-term relationship between RWI and SPEI and resistance and recovery of tree growth to severe drought episodes. Species displaying a stronger RWI-SPEI relationship to drought and low resistance and high recovery to extreme drought episodes tended to have a higher wood density (WD) and more negative leaf minimum water potential (Ψmin). Such associations were largely maintained when accounting for direct climate effects. Our results indicate that, at a cross-species level and global scale, wood and hydraulic functional traits explain species' growth responses to drought at short-and long-term scales. These trait-growth response relationships can improve our understanding of the cross-species capacity to withstand climate change and inform models to better predict drought effects on forest ecosystem dynamics. [ABSTRACT FROM AUTHOR]

Trees are long-lived organisms that integrate climate conditions across years or decades to produce secondary growth. This integration process is sometimes referred to as 'climatic memory.' While widely perceived, the physiological processes underlying this temporal integration, such as the storage and remobilization of non-structural carbohydrates (NSC), are rarely explicitly studied. This is perhaps most apparent when considering drought legacies (perturbed post-drought growth responses to climate), and the physiological mechanisms underlying these lagged responses to climatic extremes. Yet, drought legacies are likely to become more common if warming climate brings more frequent drought. To quantify the linkages between drought legacies, climate memory and NSC, we measured tree growth (via tree ring widths) and NSC concentrations in three dominant species across the southwestern USA. We analyzed these data with a hierarchical mixed effects model to evaluate the time-scales of influence of past climate (memory) on tree growth. We then evaluated the role of climate memory and the degree to which variation in NSC concentrations were related to forward-predicted growth during the hot 2011–2012 drought and subsequent 4-year recovery period. Populus tremuloides exhibited longer climatic memory compared to either Pinus edulis or Juniperus osteosperma , but following the 2011–2012 drought, P. tremuloides trees with relatively longer memory of temperature conditions showed larger (more negative) drought legacies. Conversely, Pinus edulis trees with longer temperature memory had smaller (less negative) drought legacies. For both species, higher NSC concentrations followed more negative (larger) drought legacies, though the relevant NSC fraction differed between P. tremuloides and P. edulis. Our results suggest that differences in tree NSC are also imprinted upon tree growth responses to climate across long time scales, which also underlie tree resilience to increasingly frequent drought events under climate change. [ABSTRACT FROM AUTHOR]

Droughts in a warming climate have become more common and more extreme, making understanding forest responses to water stress increasingly pressing. Analysis of water stress in trees has long focused on water potential in xylem and leaves, which influences stomatal closure and water flow through the soil‐plant‐atmosphere continuum. At the same time, changes of vegetation water content (VWC) are linked to a range of tree responses, including fluxes of water and carbon, mortality, flammability, and more. Unlike water potential, which requires demanding in situ measurements, VWC can be retrieved from remote sensing measurements, particularly at microwave frequencies using radar and radiometry. Here, we highlight key frontiers through which VWC has the potential to significantly increase our understanding of forest responses to water stress. To validate remote sensing observations of VWC at landscape scale and to better relate them to data assimilation model parameters, we introduce an ecosystem‐scale analog of the pressure–volume curve, the non‐linear relationship between average leaf or branch water potential and water content commonly used in plant hydraulics. The sources of variability in these ecosystem‐scale pressure‐volume curves and their relationship to forest response to water stress are discussed. We further show to what extent diel, seasonal, and decadal dynamics of VWC reflect variations in different processes relating the tree response to water stress. VWC can also be used for inferring belowground conditions—which are difficult to impossible to observe directly. Lastly, we discuss how a dedicated geostationary spaceborne observational system for VWC, when combined with existing datasets, can capture diel and seasonal water dynamics to advance the science and applications of global forest vulnerability to future droughts. [ABSTRACT FROM AUTHOR]

Multi‐year lags in tree drought recovery, termed 'drought legacy effects', are important for understanding the impacts of drought on forest ecosystems, including carbon (C) cycle feedbacks to climate change. Despite the ubiquity of lags in drought recovery, large uncertainties remain regarding the mechanistic basis of legacy effects and their importance for the C cycle. In this review, we identify the approaches used to study legacy effects, from tree rings to whole forests. We then discuss key knowledge gaps pertaining to the causes of legacy effects, and how the various mechanisms that may contribute these lags in drought recovery could have contrasting implications for the C cycle. Furthermore, we conduct a novel data synthesis and find that legacy effects differ drastically in both size and length across the US depending on if they are identified in tree rings versus gross primary productivity. Finally, we highlight promising approaches for future research to improve our capacity to model legacy effects and predict their impact on forest health. We emphasise that a holistic view of legacy effects – from tissues to whole forests – will advance our understanding of legacy effects and stimulate efforts to investigate drought recovery via experimental, observational and modelling approaches. [ABSTRACT FROM AUTHOR]

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

Modeling stomatal control is critical for predicting forest responses to the changing environment and hence the global water and carbon cycles. A trait-based stomatal control model that optimizes carbon gain while avoiding hydraulic risk has been shown to perform well in response to drought. However, the model's performance against changes in atmospheric CO2, which is rising rapidly due to human emissions, has yet to be evaluated. The present study tested the gain–risk model's ability to predict the stomatal response to CO2 concentration with potted water birch (Betula occidentalis Hook.) saplings in a growth chamber. The model's performance in predicting stomatal response to changes in atmospheric relative humidity and soil moisture was also assessed. The gain–risk model predicted the photosynthetic assimilation, transpiration rate and leaf xylem pressure under different CO2 concentrations, having a mean absolute percentage error (MAPE) of 25%. The model also predicted the responses to relative humidity and soil drought with a MAPE of 21.9% and 41.9%, respectively. Overall, the gain–risk model had an MAPE of 26.8% compared with the 37.5% MAPE obtained by a standard empirical model of stomatal conductance. Importantly, unlike empirical models, the optimization model relies on measurable physiological traits as inputs and performs well in predicting responses to novel environmental conditions without empirical corrections. Incorporating the optimization model in larger scale models has the potential for improving the simulation of water and carbon cycles. [ABSTRACT FROM AUTHOR]

Plants with crassulacean acid metabolism (CAM) are increasing in distribution and abundance in drylands worldwide, but the underlying drivers remain unknown. We investigate the impacts of extreme drought and CO2 enrichment on the competitive relationships between seedlings of Cylindropuntia imbricata (CAM species) and Bouteloua eriopoda (C4 grass), which coexist in semiarid ecosystems across the Southwestern United States. Our experiments under altered water and CO2 water conditions show that C. imbricata positively responded to CO2 enrichment under extreme drought conditions, while B. eriopoda declined from drought stress and did not recover after the drought ended. Conversely, in well‐watered conditions B. eriopoda had a strong competitive advantage on C. imbricata such that the photosynthetic rate and biomass (per individual) of C. imbricata grown with B. eriopoda were lower relative to when growing alone. A meta‐analysis examining multiple plant families across global drylands shows a positive response of CAM photosynthesis and productivity to CO2 enrichment. Collectively, our results suggest that under drought and elevated CO2 concentrations, projected with climate change, the competitive advantage of plant functional groups may shift and the dominance of CAM plants may increase in semiarid ecosystems. [ABSTRACT FROM AUTHOR]