• Energy Research

Shifts in rainfall patterns and increasing temperatures associated with climate change are likely to cause widespread forest decline in regions where droughts are predicted to increase in duration and severity. One primary cause of productivity loss and plant mortality during drought is hydraulic failure. Drought stress creates trapped gas emboli in the water transport system, which reduces the ability of plants to supply water to leaves for photosynthetic gas exchange and can ultimately result in desiccation and mortality. At present we lack a clear picture of how thresholds to hydraulic failure vary across a broad range of species and environments, despite many individual experiments. Here we draw together published and unpublished data on the vulnerability of the transport system to drought-induced embolism for a large number of woody species, with a view to examining the likely consequences of climate change for forest biomes. We show that 70% of 226 forest species from 81 sites worldwide operate with narrow (<1 megapascal) hydraulic safety margins against injurious levels of drought stress and therefore potentially face long-term reductions in productivity and survival if temperature and aridity increase as predicted for many regions across the globe. Safety margins are largely independent of mean annual precipitation, showing that there is global convergence in the vulnerability of forests to drought, with all forest biomes equally vulnerable to hydraulic failure regardless of their current rainfall environment. These findings provide insight into why drought-induced forest decline is occurring not only in arid regions but also in wet forests not normally considered at drought risk.

Woody plants vary in their adaptations to drought and shade. For a better prediction of vegetation responses to drought and shade within dynamic global vegetation models, it is critical to group species into functional types with similar adaptations. One of the key challenges is that the adaptations are generally determined by a large number of plant traits that may not be available for a large number of species. In this study, we present two heuristic woody plant groups that were separated using cluster analysis in a three-dimensional trait-environment space based on three key metrics for each species: mean xylem embolism resistance, shade tolerance and habitat aridity. The two heuristic groups separate these species into tolerators and avoiders. The tolerators either rely on their high embolism resistance to tolerate drought in arid habitats (e.g., Juniperus and Prunus) or rely on high shade tolerance to withstand shaded conditions in wet habitats (e.g., Picea, Abies and Acer). In contrast, all avoiders have low embolism resistance and low shade tolerance. In arid habitats, avoiders tend to minimize catastrophic embolism (e.g., most Pinus species) while in wet habitats, they may survive despite low shade tolerance (e.g., Betula, Populus, Alnus and Salix). Because our approach links traits to the environmental conditions, we expect it could be a promising framework for predicting changes in species composition, and therefore ecosystem function, under changing environmental conditions.

AbstractStomatal response to environmental conditions forms the backbone of all ecosystem and carbon cycle models, but is largely based on empirical relationships. Evolutionary theories of stomatal behaviour are critical for guarding against prediction errors of empirical models under future climates. Longstanding theory holds that stomata maximise fitness by acting to maintain constant marginal water use efficiency over a given time horizon, but a recent evolutionary theory proposes that stomata instead maximise carbon gain minus carbon costs/risk of hydraulic damage. Using data from 34 species that span global forest biomes, we find that the recent carbon‐maximisation optimisation theory is widely supported, revealing that the evolution of stomatal regulation has not been primarily driven by attainment of constant marginal water use efficiency. Optimal control of stomata to manage hydraulic risk is likely to have significant consequences for ecosystem fluxes during drought, which is critical given projected intensification of the global hydrological cycle.

Significance Predicting the impacts of climate extremes on plant communities is a central challenge in ecology. Physiological traits may improve prediction of drought impacts on forests globally. We perform a meta-analysis across 33 studies that span all forested biomes and find that, among the examined traits, hydraulic traits explain cross-species patterns in mortality from drought. Gymnosperm and angiosperm mortality was associated with different hydraulic traits, giving insight into the relative weights of different traits and mechanisms in mortality prediction. Our results provide a foundation for more mechanistic predictions of drought-induced tree mortality across Earth’s diverse forests.

Fertilization of woody plants plays a central role in agriculture and forestry, but little is known about how plant water relations are thereby affected. Here we investigated the impact of fertilization on tree hydraulics, and xylem and pit anatomy in the high-yield apple cultivars Golden and Red Delicious. In fertilized trees of Golden Delicious, specific hydraulic conductivity of branch xylem, hydraulic conductance of the root system, and maximum stomatal conductance increased considerably. In Red Delicious, differences between fertilized and control trees were less pronounced. In both cultivars, xylem embolism resistance of fertilized trees was significantly lower and stomatal closure occurred at lower water potentials. Furthermore, water potential at turgor loss point and osmotic potential at full saturation were higher and cell wall elasticity was lower in fertilized plants, suggesting reduced drought tolerance of leaves. Anatomical differences were observed regarding conduit diameters, cell wall reinforcement, pit membrane thickness, pit chamber depth, and stomatal pore length, with more pronounced differences in Golden Delicious. The findings reveal altered hydraulic behaviour in both apple cultivars upon fertilization. The increased vulnerability to hydraulic failure might pose a considerable risk for apple productivity under a changing climate, which should be considered for future cultivation and management practices.