Contents Summary 1207 I. Introduction 1207 II. A brief history of modelling plant water fluxes 1208 III. Main components of plant water transport models 1208 IV. Stand-scale water fluxes and coupling to climate and soil 1213 V. Water fluxes in terrestrial biosphere models and feedbacks to community dynamics 1215 VI. Outstanding challenges in modelling water fluxes in the soil-plant-atmosphere continuum 1217 Acknowledgements 1218 References 1218 SUMMARY: Models of plant water fluxes have evolved from studies focussed on understanding the detailed structure and functioning of specific components of the soil-plant-atmosphere (SPA) continuum to architectures often incorporated inside eco-hydrological and terrestrial biosphere (TB) model schemes. We review here the historical evolution of this field, examine the basic structure of a simplified individual-based model of plant water transport, highlight selected applications for specific ecological problems and conclude by examining outstanding issues requiring further improvements in modelling vegetation water fluxes. We particularly emphasise issues related to the scaling from tissue-level traits to individual-based predictions of water transport, the representation of nonlinear and hysteretic behaviour in soil-xylem hydraulics and the need to incorporate knowledge of hydraulics within broader frameworks of plant ecological strategies and their consequences for predicting community demography and dynamics.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.15681

Modelling water fluxes in plants: from tissues to biosphere.

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. 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.

Meta-analysis Reveals that Hydraulic Traits Explain Cross-Species Patterns of Drought-Induced Tree Mortality across the Globe

Tropical ecosystems have the highest levels of biodiversity, cycle more water and absorb more carbon than any other terrestrial ecosystem on Earth. Consequently, these ecosystems are extremely important components of Earth's climatic system and biogeochemical cycles. Plant hydraulics is an essential discipline to understand and predict the dynamics of tropical vegetation in scenarios of changing water availability. Using published plant hydraulic data we show that the trade-off between drought avoidance (expressed as deep-rooting, deciduousness and capacitance) and hydraulic safety (P50 - the water potential when plants lose 50% of their maximum hydraulic conductivity) is a major axis of physiological variation across tropical ecosystems. We also propose a novel and independent axis of hydraulic trait variation linking vulnerability to hydraulic failure (expressed as the hydraulic safety margin (HSM)) and growth, where inherent fast-growing plants have lower HSM compared to slow-growing plants. We surmise that soil nutrients are fundamental drivers of tropical community assembly determining the distribution and abundance of the slow-safe/fast-risky strategies. We conclude showing that including either the growth-HSM or the resistance-avoidance trade-off in models can make simulated tropical rainforest communities substantially more vulnerable to drought than similar communities without the trade-off. These results suggest that vegetation models need to represent hydraulic trade-off axes to accurately project the functioning and distribution of tropical ecosystems.

Linking plant hydraulics and the fast–slow continuum to understand resilience to drought in tropical ecosystems

With ongoing global warming, the amount and frequency of precipitation in the tropics is projected to change substantially. While it has been shown that tropical forests and savannahs are sustained within the same intermediate mean annual precipitation range, the mechanisms that lead to the resilience of these ecosystems are still not fully understood. In particular, the long-term impact of rainfall variability on resilience is as yet unclear. Here we present observational evidence that both tropical forest and savannah exposed to a higher rainfall variability—in particular on interannual scales—during their long-term past are overall more resilient against climatic disturbances. Based on precipitation and tree cover data in the Brazilian Amazon basin, we constructed potential landscapes that enable us to systematically measure the resilience of the different ecosystems. Additionally, we infer that shifts from forest to savannah due to decreasing precipitation in the future are more likely to occur in regions with a precursory lower rainfall variability. Long-term rainfall variability thus needs to be taken into account in resilience analyses and projections of vegetation response to climate change. Tropical forests and savannah are more resilient to climate disturbances when they have been exposed to higher rainfall variability in the long-term past, finds an analysis of Brazilian rainfall and tree-cover observations.

Higher resilience to climatic disturbances in tropical vegetation exposed to more variable rainfall

The vast majority of measurements in the field of plant hydraulics have been on small-diameter branches from woody species. These measurements have provided considerable insight into plant functioning, but our understanding of plant physiology and ecology would benefit from a broader view, because branch hydraulic properties are influenced by many factors. Here, we discuss the influence that other components of the hydraulic network have on branch vulnerability to embolism propagation. We also modelled the impact of changes in the ratio of root-to-leaf areas and soil texture on vulnerability to hydraulic failure along the soil-to-leaf continuum and showed that hydraulic function is better maintained through changes in root vulnerability and root-to-leaf area ratio than in branch vulnerability. Differences among species in the stringency with which they regulate leaf water potential and in reliance on stored water to buffer changes in water potential also affect the need to construct embolism resistant branches. Many approaches, such as measurements on fine roots, small individuals, combining sap flow and psychrometry techniques, and modelling efforts, could vastly improve our understanding of whole-plant hydraulic functioning. A better understanding of how traits are coordinated across the whole plant will improve predictions for plant function under future climate conditions.

https://onlinelibrary.wiley.com/doi/epdf/10.1111/pce.13607

A dynamic yet vulnerable pipeline: Integration and coordination of hydraulic traits across whole plants.

Species distribution is strongly driven by local and global gradients in water availability but the underlying mechanisms are not clear. Vulnerability to xylem embolism (P50 ) is a key trait that indicates how species cope with drought and might explain plant distribution patterns across environmental gradients. Here we address its role on species sorting along a hydro-topographical gradient in a central Amazonian rainforest and examine its variance at the community scale. We measured P50 for 28 tree species, soil properties and estimated the hydrological niche of each species using an indicator of distance to the water table (HAND). We found a large hydraulic diversity, covering as much as 44% of the global angiosperm variation in P50 . We show that P50 : contributes to species segregation across a hydro-topographic gradient in the Amazon, and thus to species coexistence; is the result of repeated evolutionary adaptation within closely related taxa; is associated with species tolerance to P-poor soils, suggesting the evolution of a stress-tolerance syndrome to nutrients and drought; and is higher for trees in the valleys than uplands. The large observed hydraulic diversity and its association with topography has important implications for modelling and predicting forest and species resilience to climate change.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/nph.15463

Edher C. Cordoba

Papers

Embolism resistance drives the distribution of Amazonian rainforest tree species along hydro-topographic gradients