Hydraulic diversity of forests regulates ecosystem resilience during drought.

Plants influence the atmosphere through fluxes of carbon, water and energy1, and can intensify drought through land-atmosphere feedback effects2-4. The diversity of plant functional traits in forests, especially physiological traits related to water (hydraulic) transport, may have a critical role in land-atmosphere feedback, particularly during drought. Here we combine 352 site-years of eddy covariance measurements from 40 forest sites, remote-sensing observations of plant water content and plant functional-trait data to test whether the diversity in plant traits affects the response of the ecosystem to drought. We find evidence that higher hydraulic diversity buffers variation in ecosystem flux during dry periods across temperate and boreal forests. Hydraulic traits were the predominant significant predictors of cross-site patterns in drought response. By contrast, standard leaf and wood traits, such as specific leaf area and wood density, had little explanatory power. Our results demonstrate that diversity in the hydraulic traits of trees mediates ecosystem resilience to drought and is likely to have an important role in future ecosystem-atmosphere feedback effects in a changing climate.

Testing the effects of species interactions and water limitation on tree seedling biomass allocation and physiology.

Species interactions mediate tree responses to water limitation because competition and/or facilitation alter plant physiology and growth. However, because it is difficult to isolate the effects of plant-plant interactions and water limitation from other environmental factors, the mechanisms underlying tree physiology and growth in coexisting plants under drought are poorly understood. We investigated how species interactions and water limitation impact the physiology and growth of trembling aspen (Populus tremuloides), narrowleaf cottonwood (Populus angustifolia) and ponderosa pine (Pinus ponderosa) seedlings in a controlled environment growth chamber, using aspen as a focal species. Seedlings were grown in pots alone or with a con- or hetero-specific seedling, and were subjected to a water limitation treatment. Growth, water status and physiological traits were measured before, during and after the treatment. Under well-watered conditions, the presence of another seedling affected growth or biomass allocation in all species, but did not impact the physiological traits we measured. Under water limitation, the presence of a competing seedling had a marginal impact on seedling growth and physiological traits in all species. Throughout the study, the magnitude and direction of seedling responses were complex and often species-specific. Our study serves as an important step toward testing how species' interactions modify physiological responses and growth in well-watered and water-limited periods.

Competition and Drought Alter Optimal Stomatal Strategy in Tree Seedlings.

A better understanding of plant stomatal strategies holds strong promise for improving predictions of vegetation responses to drought because stomata are the primary mechanism through which plants mitigate water stress. It has been assumed that plants regulate stomata to maintain a constant marginal water use efficiency and forego carbon gain when water is scarce. However, recent hypotheses pose that plants maximize carbon assimilation while also accounting for the risk of hydraulic damage via cavitation and hydraulic failure. This "gain-risk" framework incorporates competition in stomatal regulation because it takes into account that neighboring plants can "steal" unused water. This study utilizes stomatal models representing both the water use efficiency and carbon-maximization frameworks, and empirical data from three species in a potted growth chamber experiment, to investigate the effects of drought and competition on seedling stomatal strategy. We found that drought and competition responses in the empirical data were best explained by the carbon-maximization hypothesis and that both drought and competition affected stomatal strategy. Interestingly, stomatal responses differed substantially by species, with seedlings employing a riskier strategy when planted with a high water use competitor, and seedlings employing a more conservative strategy when planted with a low water use competitor. Lower water users in general had less stomatal sensitivity to decreasing Ψ L compared to moderate to high water users. Repeated water stress also resulted in legacy effects on plant stomatal behavior, increasing stomatal sensitivity (i.e., conservative behavior) even when the seedling was returned to well-watered conditions. These results indicate that stomatal strategies are dynamic and change with climate and competition stressors. Therefore, incorporating mechanisms that allow for stomatal behavioral changes in response to water limitation may be an important step to improving carbon cycle projections in coupled climate-Earth system models.