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Severe droughts have caused widespread tree mortality across many forest biomes with profound effects on the function of ecosystems and carbon balance. Climate change is expected to intensify regional-scale droughts, focusing attention on the physiological basis of drought-induced tree mortality. Recent work has shown that catastrophic failure of the plant hydraulic system is a principal mechanism involved in extensive crown death and tree mortality during drought, but the multi-dimensional response of trees to desiccation is complex. Here we focus on the current understanding of tree hydraulic performance under drought, the identification of physiological thresholds that precipitate mortality and the mechanisms of recovery after drought. Building on this, we discuss the potential application of hydraulic thresholds to process-based models that predict mortality.

Forest mortality is accelerating due to climate change and the largest trees may be at the greatest risk, threatening critical ecological, economic, and social benefits. Here, we combine high-resolution airborne LiDAR and optical data to track tree-level mortality rates for ~2 million trees in California over 8 years, showing that tree height is the strongest predictor of mortality during extreme drought. Large trees die at twice the rate of small trees and environmental gradients of temperature, water, and competition control the intensity of the height-mortality relationship. These findings suggest that future persistent drought may cause widespread mortality of the largest trees on Earth. Drought is intensifying due to climate change, impacting forests globally. Here, the authors track nearly 2 million trees through severe drought and show that tree height is the greatest predictor of mortality risk, suggesting that the tallest trees may be the most vulnerable.

The response of vegetation physiology to drought at large spatial scales is poorly understood due to a lack of direct observations. Here, we study vegetation drought responses related to photosynthesis, evaporation, and vegetation water content using remotely sensed data, and we isolate physiological responses using a machine learning technique. We find that vegetation functional decreases are largely driven by the downregulation of vegetation physiology such as stomatal conductance and light use efficiency, with the strongest downregulation in water-limited regions. Vegetation physiological decreases in wet regions also result in a discrepancy between functional and structural changes under severe drought. We find similar patterns of physiological drought response using simulations from a soil–plant–atmosphere continuum model coupled with a radiative transfer model. Observation-derived vegetation physiological responses to drought across space are mainly controlled by aridity and additionally modulated by abnormal hydro-meteorological conditions and vegetation types. Hence, isolating and quantifying vegetation physiological responses to drought enables a better understanding of ecosystem biogeochemical and biophysical feedback in modulating climate change. Vegetation resilience to drought is underlain by plant physiological responses. Here, the authors combine remote sensing data, explainable machine learning and model simulations to map global vegetation responses to drought linked to physiological processes such as stomatal regulation and light use efficiency.

Stomata, the microvalves on leaf surfaces, exert major influences across scales, from plant growth and productivity to global carbon and water cycling. Stomatal opening enables leaf photosynthesis, and plant growth and water use, whereas plant survival of drought depends on stomatal closure. Here we report that stomatal function is constrained by a safety-efficiency trade-off, such that species with greater stomatal conductance under high water availability ( g max ) show greater sensitivity to closure during leaf dehydration, i.e., a higher leaf water potential at which stomatal conductance is reduced by 50% (Ψ gs50 ). The g max - Ψ gs50 trade-off and its mechanistic basis is supported by experiments on leaves of California woody species, and in analyses of previous studies of the responses of diverse flowering plant species around the world. Linking the two fundamental key roles of stomata—the enabling of gas exchange, and the first defense against drought—this trade-off constrains the rates of water use and the drought sensitivity of leaves, with potential impacts on ecosystems. Stomata enable gas exchange for photosynthesis but close to promote survival during drought. Here, Henry et al. provide evidence for a safety-efficiency trade-off whereby plants with greater stomatal conductance under well-watered conditions are more sensitive to stomatal closure during dehydration.

Abstract Heatwaves exert disproportionately strong and sometimes irreversible impacts on forest ecosystems. These impacts remain poorly understood at the tree and species level and across large spatial scales. Here, we investigate the effects of the record-breaking 2018 European heatwave on tree growth and tree water status using a collection of high-temporal resolution dendrometer data from 21 species across 53 sites. Relative to the two preceding years, annual stem growth was not consistently reduced by the 2018 heatwave but stems experienced twice the temporary shrinkage due to depletion of water reserves. Conifer species were less capable of rehydrating overnight than broadleaves across gradients of soil and atmospheric drought, suggesting less resilience toward transient stress. In particular, Norway spruce and Scots pine experienced extensive stem dehydration. Our high-resolution dendrometer network was suitable to disentangle the effects of a severe heatwave on tree growth and desiccation at large-spatial scales in situ, and provided insights on which species may be more vulnerable to climate extremes.

Low soil moisture and high vapour pressure deficit (VPD) cause plant water stress and lead to a variety of drought responses, including a reduction in transpiration and photosynthesis 1 , 2 . When soils dry below critical soil moisture thresholds, ecosystems transition from energy to water limitation as stomata close to alleviate water stress 3 , 4 . However, the mechanisms behind these thresholds remain poorly defined at the ecosystem scale. Here, by analysing observations of critical soil moisture thresholds globally, we show the prominent role of soil texture in modulating the onset of ecosystem water limitation through the soil hydraulic conductivity curve, whose steepness increases with sand fraction. This clarifies how ecosystem sensitivity to VPD versus soil moisture is shaped by soil texture, with ecosystems in sandy soils being relatively more sensitive to soil drying, whereas ecosystems in clayey soils are relatively more sensitive to VPD. For the same reason, plants in sandy soils have limited potential to adjust to water limitations, which has an impact on how climate change affects terrestrial ecosystems. In summary, although vegetation–atmosphere exchanges are driven by atmospheric conditions and mediated by plant adjustments, their fate is ultimately dependent on the soil. Through their effects on soil hydraulic properties, soil texture and sand content are shown to have broad implications for the terrestrial water cycle and carbon sink, and specific implications for vital ecosystems that are vulnerable to drought, especially with changing climate.

Tropical forests face increasing climate risk 1 , 2 , yet our ability to predict their response to climate change is limited by poor understanding of their resistance to water stress. Although xylem embolism resistance thresholds (for example, Ψ 50 ) and hydraulic safety margins (for example, HSM 50 ) are important predictors of drought-induced mortality risk 3 – 5 , little is known about how these vary across Earth’s largest tropical forest. Here, we present a pan-Amazon, fully standardized hydraulic traits dataset and use it to assess regional variation in drought sensitivity and hydraulic trait ability to predict species distributions and long-term forest biomass accumulation. Parameters Ψ 50 and HSM 50 vary markedly across the Amazon and are related to average long-term rainfall characteristics. Both Ψ 50 and HSM 50 influence the biogeographical distribution of Amazon tree species. However, HSM 50 was the only significant predictor of observed decadal-scale changes in forest biomass. Old-growth forests with wide HSM 50 are gaining more biomass than are low HSM 50 forests. We propose that this may be associated with a growth–mortality trade-off whereby trees in forests consisting of fast-growing species take greater hydraulic risks and face greater mortality risk. Moreover, in regions of more pronounced climatic change, we find evidence that forests are losing biomass, suggesting that species in these regions may be operating beyond their hydraulic limits. Continued climate change is likely to further reduce HSM 50 in the Amazon 6 , 7 , with strong implications for the Amazon carbon sink. A pan-Amazon study of forests shows large variations in drought tolerance traits and finds that forests in regions of pronounced climate change are losing biomass and may be operating beyond their hydraulic limits.

Plant functional traits can predict community assembly and ecosystem functioning and are thus widely used in global models of vegetation dynamics and land–climate feedbacks. Still, we lack a global understanding of how land and climate affect plant traits. A previous global analysis of six traits observed two main axes of variation: (1) size variation at the organ and plant level and (2) leaf economics balancing leaf persistence against plant growth potential. The orthogonality of these two axes suggests they are differently influenced by environmental drivers. We find that these axes persist in a global dataset of 17 traits across more than 20,000 species. We find a dominant joint effect of climate and soil on trait variation. Additional independent climate effects are also observed across most traits, whereas independent soil effects are almost exclusively observed for economics traits. Variation in size traits correlates well with a latitudinal gradient related to water or energy limitation. In contrast, variation in economics traits is better explained by interactions of climate with soil fertility. These findings have the potential to improve our understanding of biodiversity patterns and our predictions of climate change impacts on biogeochemical cycles. The authors investigate the broad-scale climatological and soil properties that co-vary with major axes of plant functional traits. They find that variation in plant size is attributed to latitudinal gradients in water or energy limitation, while variation in leaf economics traits is attributed to both climate and soil fertility including their interaction.

Efficient nutrient recycling underpins the ecological success of cnidarian-algal symbioses in oligotrophic waters. In these symbioses, nitrogen limitation restricts the growth of algal endosymbionts in hospite and stimulates their release of photosynthates to the cnidarian host. However, the mechanisms controlling nitrogen availability and their role in symbiosis regulation remain poorly understood. Here, we studied the metabolic regulation of symbiotic nitrogen cycling in the sea anemone Aiptasia by experimentally altering labile carbon availability in a series of experiments. Combining 13 C and 15 N stable isotope labeling experiments with physiological analyses and NanoSIMS imaging, we show that the competition for environmental ammonium between the host and its algal symbionts is regulated by labile carbon availability. Light regimes optimal for algal photosynthesis increase carbon availability in the holobiont and stimulate nitrogen assimilation in the host metabolism. Consequently, algal symbiont densities are lowest under optimal environmental conditions and increase toward the lower and upper light tolerance limits of the symbiosis. This metabolic regulation promotes efficient carbon recycling in a stable symbiosis across a wide range of environmental conditions. Yet, the dependence on resource competition may favor parasitic interactions, explaining the instability of the cnidarian-algal symbiosis as environmental conditions in the Anthropocene shift towards its tolerance limits. Photosymbioses enable efficient nutrient recycling between heterotrophic and phototrophic organisms. This study shows that nutrient cycling in a cnidarian-algal symbiosis is regulated through resource competition between symbiotic partners. Mutualistic interactions can therefore emerge from mutual exploitation in nutrient–exchange symbioses.

Leaf unfolding in temperate forests is driven by spring temperature, but little is known about the spatial variance of that temperature dependency. Here we use in situ leaf unfolding observations for eight deciduous tree species to show that the two factors that control chilling (number of cold days) and heat requirement (growing degree days at leaf unfolding, GDD req ) only explain 30% of the spatial variance of leaf unfolding. Radiation and aridity differences among sites together explain 10% of the spatial variance of leaf unfolding date, and 40% of the variation in GDD req . Radiation intensity is positively correlated with GDD req and aridity is negatively correlated with GDD req spatial variance. These results suggest that leaf unfolding of temperate deciduous trees is adapted to local mean climate, including water and light availability, through altered sensitivity to spring temperature. Such adaptation of heat requirement to background climate would imply that models using constant temperature response are inherently inaccurate at local scale. Drivers of spatial differences in leaf phenology are not as widely studied as temporal differences. Here the authors show that the spatial variation of leaf unfolding in 8 deciduous tree species in Europe can be explained by local adaptation to long-term mean climate conditions.