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Climate models suggest that more frequent drought events of greater severity and length, associated with climate change, can be expected in the coming decades. Although drought-induced tree mortality has been recognized as an important factor modulating forest demography at the global scale, the mechanisms underlying drought-induced tree mortality remain contentious. Above- and below-ground growth, gas exchange, water relations and carbon reserve accumulation dynamics at the organ and whole-plant scale were quantified in Populus tremuloides and P. balsamifera seedlings in response to severe drought. Seedlings were maintained in drought conditions over one growing and one dormant winter season. Our experiment presents a detailed description of the effect of severe drought on growth and physiological variables, leading to seedling mortality after an extended period of drought and dormancy. After re-watering following the dormant period, drought-exposed seedlings did not re-flush, showing that the root system had died off. The results of this study suggest a complex series of physiological feedbacks between the measured variables in both Populus species. Further, they reveal that reduced reserve accumulation in the root system during drought decreases the conversion of starch to soluble sugars in roots, which may contribute to the root death of drought-exposed seedlings during the dormant season by compromising the frost tolerance of the root system.

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Transpiration, the dominant component of terrestrial evapotranspiration (ET), directly connects the water, energy and carbon cycles and is typically restricted by soil and atmospheric (for example, the vapour pressure deficit (VPD)) moisture stresses through plant hydraulic processes. These sources of stress are likely to diverge under climate change, with a globally enhanced VPD but more variable and uncertain changes in soil moisture. Here, using a model–data fusion approach, we demonstrate that the common empirical approach used in most Earth system models to evaluate the ET response to soil moisture and VPD, which neglects plant hydraulics, underestimates ET sensitivity to VPD and compensates by overestimating the sensitivity to soil moisture stress. A hydraulic model that describes water transport through the plant better captures ET under high VPD conditions for wide-ranging soil moisture states. These findings highlight the central role of plant hydraulics in regulating the increasing importance of atmospheric moisture stress on biosphere–atmosphere interactions under elevated temperatures. Evapotranspiration links productivity with water cycling between land and atmosphere. A model including plant hydraulics better describes the response of evapotranspiration to stress from vapour pressure deficit and soil moisture under rising temperatures than approaches common in Earth system models.

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.

Consistent and reasonable characterization of the material behavior under the coupled effects of strain, strain rate and temperature on the material flow stress is remarkably crucial in order to design as well as optimize the process parameters in the metal forming industrial practice. The objective of this work was to formulate an appropriate flow stress model to characterize the flow behavior of AISI-1045 medium carbon steel over a practical range of deformation temperatures (650⁻950 ∘ C) and strain rates (0.05⁻1.0 s - 1 ). Subsequently, the Johnson-Cook flow stress model was adopted for modeling and predicting the material flow behavior at elevated temperatures. Furthermore, surrogate models were developed based on the constitutive relations, and the model constants were estimated using the experimental results. As a result, the constitutive flow stress model was formed and the constructed model was examined systematically against experimental data by both numerical and graphical validations. In addition, to predict the material damage behavior, the failure model proposed by Johnson and Cook was used, and to determine the model parameters, seven different specimens, including flat, smooth round bars and pre-notched specimens, were tested at room temperature under quasi strain rate conditions. From the results, it can be seen that the developed model over predicts the material behavior at a low temperature for all strain rates. However, overall, the developed model can produce a fairly accurate and precise estimation of flow behavior with good correlation to the experimental data under high temperature conditions. Furthermore, the damage model parameters estimated in this research can be used to model the metal forming simulations, and valuable prediction results for the work material can be achieved.

Determining physiological mechanisms and thresholds for climate-driven tree die-off could help improve global predictions of future terrestrial carbon sinks. We directly tested for the lethal threshold in hydraulic failure – an inability to move water due to drought-induced xylem embolism – in a pine sapling experiment. In a glasshouse experiment, we exposed loblolly pine (Pinus taeda) saplings (n = 83) to drought-induced water stress ranging from mild to lethal. Before rewatering to relieve drought stress, we measured native hydraulic conductivity and foliar color change. We monitored all measured individuals for survival or mortality. We found a lethal threshold at 80% loss of hydraulic conductivity – a point of hydraulic failure beyond which it is more likely trees will die, than survive, and describe mortality risk across all levels of water stress. Foliar color changes lagged behind hydraulic failure – best predicting when trees had been dead for some time, rather than when they were dying. Our direct measurement of native conductivity, while monitoring the same individuals for survival or mortality, quantifies a continuous probability of mortality risk from hydraulic failure. Predicting tree die-off events and understanding the mechanism involved requires knowledge not only of when trees are dead, but when they begin dying – having passed the point of no return.

Model–data comparisons of plant physiological processes provide an understanding of mechanisms underlying vegetation responses to climate. We simulated the physiology of a piñon pine–juniper woodland (Pinus edulis–Juniperus monosperma) that experienced mortality during a 5 yr precipitation-reduction experiment, allowing a framework with which to examine our knowledge of drought-induced tree mortality. We used six models designed for scales ranging from individual plants to a global level, all containing state-of-the-art representations of the internal hydraulic and carbohydrate dynamics of woody plants. Despite the large range of model structures, tuning, and parameterization employed, all simulations predicted hydraulic failure and carbon starvation processes co-occurring in dying trees of both species, with the time spent with severe hydraulic failure and carbon starvation, rather than absolute thresholds per se, being a better predictor of impending mortality. Model and empirical data suggest that limited carbon and water exchanges at stomatal, phloem, and below-ground interfaces were associated with mortality of both species. The model–data comparison suggests that the introduction of a mechanistic process into physiology-based models provides equal or improved predictive power over traditional process-model or empirical thresholds. Both biophysical and empirical modeling approaches are useful in understanding processes, particularly when the models fail, because they reveal mechanisms that are likely to underlie mortality. We suggest that for some ecosystems, integration of mechanistic pathogen models into current vegetation models, and evaluation against observations, could result in a breakthrough capability to simulate vegetation dynamics.

Optimization models of stomatal conductance (g s) attempt to explain observed stomatal behaviour in terms of cost–benefit tradeoffs. While the benefit of stomatal opening through increased CO2 uptake is clear, currently the nature of the associated cost(s) remains unclear. We explored the hypothesis that g s maximizes leaf photosynthesis, where the cost of stomatal opening arises from nonstomatal reductions in photosynthesis induced by leaf water stress. We analytically solved two cases, CAP and MES, in which reduced leaf water potential leads to reductions in carboxylation capacity (CAP) and mesophyll conductance (g m) (MES). Both CAP and MES predict the same one-parameter relationship between the intercellular: atmospheric CO2 concentration ratio (c i/c a) and vapour pressure deficit (VPD, D), viz. c i / c a ≈ ξ / ( ξ + D ) , as that obtained from previous optimization models, with the novel feature that the parameter ξ is determined unambiguously as a function of a small number of photosynthetic and hydraulic variables. These include soil-to-leaf hydraulic conductance, implying a stomatal closure response to drought. MES also predicts that g s/g m is closely related to c i/c a and is similarly conservative. These results are consistent with observations, give rise to new testable predictions, and offer new insights into the covariation of stomatal, mesophyll and hydraulic conductances.

Forest ecosystems store approximately 45% of the carbon found in terrestrial ecosystems, but they are sensitive to climate-induced dieback. Forest die-off constitutes a large uncertainty in projections of climate impacts on terrestrial ecosystems, climate–ecosystem interactions, and carbon-cycle feedbacks. Current understanding of the physiological mechanisms mediating climate-induced forest mortality limits the ability to model or project these threshold events. We report here a direct and in situ study of the mechanisms underlying recent widespread and climate-induced trembling aspen (Populus tremuloides) forest mortality in western North America. We find substantial evidence of hydraulic failure of roots and branches linked to landscape patterns of canopy and root mortality in this species. On the contrary, we find no evidence that drought stress led to depletion of carbohydrate reserves. Our results illuminate proximate mechanisms underpinning recent aspen forest mortality and provide guidance for understanding and projecting forest die-offs under climate change.

Drought-induced tree mortality has been observed globally and is expected to increase under climate change scenarios, with large potential consequences for the terrestrial carbon sink. Predicting mortality across species is crucial for assessing the effects of climate extremes on forest community biodiversity, composition, and carbon sequestration. However, the physiological traits associated with elevated risk of mortality in diverse ecosystems remain unknown, although these traits could greatly improve understanding and prediction of tree mortality in forests. We performed a meta-analysis on species’ mortality rates across 475 species from 33 studies around the globe to assess which traits determine a species’ mortality risk. We found that species-specific mortality anomalies from community mortality rate in a given drought were associated with plant hydraulic traits. Across all species, mortality was best predicted by a low hydraulic safety margin—the difference between typical minimum xylem water potential and that causing xylem dysfunction—and xylem vulnerability to embolism. Angiosperms and gymnosperms experienced roughly equal mortality risks. Our results provide broad support for the hypothesis that hydraulic traits capture key mechanisms determining tree death and highlight that physiological traits can improve vegetation model prediction of tree mortality during climate extremes.