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Synopsis Classic debates in community ecology focused on the complexities of considering an ecosystem as a super-organ or organism. New consideration of such perspectives could clarify mechanisms underlying the dynamics of forest carbon dioxide (CO2) uptake and water vapor loss, important for predicting and managing the future of Earth’s ecosystems and climate system. Here, we provide a rubric for considering ecosystem traits as aggregated, systemic, or emergent, i.e., representing the ecosystem as an aggregate of its individuals or as a metaphorical or literal super-organ or organism. We review recent approaches to scaling-up plant water relations (hydraulics) concepts developed for organs and organisms to enable and interpret measurements at ecosystem-level. We focus on three community-scale versions of water relations traits that have potential to provide mechanistic insight into climate change responses of forest CO2 and H2O gas exchange and productivity: leaf water potential (Ψcanopy), pressure volume curves (eco-PV), and hydraulic conductance (Keco). These analyses can reveal additional ecosystem-scale parameters analogous to those typically quantified for leaves or plants (e.g., wilting point and hydraulic vulnerability) that may act as thresholds in forest responses to drought, including growth cessation, mortality, and flammability. We unite these concepts in a novel framework to predict Ψcanopy and its approaching of critical thresholds during drought, using measurements of Keco and eco-PV curves. We thus delineate how the extension of water relations concepts from organ- and organism-scales can reveal the hydraulic constraints on the interaction of vegetation and climate and provide new mechanistic understanding and prediction of forest water use and productivity. 

Abstract The rooting-zone water-storage capacity—the amount of water accessible to plants—controls the sensitivity of land–atmosphere exchange of water and carbon during dry periods. How the rooting-zone water-storage capacity varies spatially is largely unknown and not directly observable. Here we estimate rooting-zone water-storage capacity globally from the relationship between remotely sensed vegetation activity, measured by combining evapotranspiration, sun-induced fluorescence and radiation estimates, and the cumulative water deficit calculated from daily time series of precipitation and evapotranspiration. Our findings indicate plant-available water stores that exceed the storage capacity of 2-m-deep soils across 37% of Earth’s vegetated surface. We find that biome-level variations of rooting-zone water-storage capacities correlate with observed rooting-zone depth distributions and reflect the influence of hydroclimate, as measured by the magnitude of annual cumulative water-deficit extremes. Smaller-scale variations are linked to topography and land use. Our findings document large spatial variations in the effective root-zone water-storage capacity and illustrate a tight link among the climatology of water deficits, rooting depth of vegetation and its sensitivity to water stress. 

Abstract Stomatal optimization theory is a commonly used framework for modeling how plants regulate transpiration in response to the environment. Most stomatal optimization models assume that plantsinstantaneouslyoptimize a reward function such as carbon gain. However, plants are expected to optimize over longer timescales given the rapid environmental variability they encounter. There are currently no observational constraints on these timescales. Here, a new stomatal model is developed and is used to analyze the timescales over which stomatal closure is optimized. The proposed model assumes plants maximize carbon gain subject to the constraint that they cannot draw down soil moisture below a critical value. The reward is integrated over time, after being weighted by a discount factor that represents the timescale (τ) that a plant considers when optimizing stomatal conductance to save water. The model is simple enough to be analytically solvable, which allows the value ofτto be inferred from observations of stomatal behavior under known environmental conditions. The model is fitted to eddy covariance data in a range of ecosystems, finding the value ofτthat best predicts the dynamics of evapotranspiration at each site. Across 82 sites, the climate metrics with the strongest correlation toτare measures of the average number of dry days between rainfall events. Values ofτare similar in magnitude to the longest such dry period encountered in an average year. The results here shed light on which climate characteristics shape spatial variations in ecosystem‐level water use strategy. 

Abstract Spatiotemporal patterns of plant water uptake, loss, and storage exert a first‐order control on photosynthesis and evapotranspiration. Many studies of plant responses to water stress have focused on differences between species because of their different stomatal closure, xylem conductance, and root traits. However, several other ecohydrological factors are also relevant, including soil hydraulics, topographically driven redistribution of water, plant adaptation to local climatic variations, and changes in vegetation density. Here, we seek to understand the relative importance of the dominant species for regional‐scale variations in woody plant responses to water stress. We map plant water sensitivity (PWS) based on the response of remotely sensed live fuel moisture content to variations in hydrometeorology using an auto‐regressive model. Live fuel moisture content dynamics are informative of PWS because they directly reflect vegetation water content and therefore patterns of plant water uptake and evapotranspiration. The PWS is studied using 21,455 wooded locations containing U.S. Forest Service Forest Inventory and Analysis plots across the western United States, where species cover is known and where a single species is locally dominant. Using a species‐specific mean PWS value explains 23% of observed PWS variability. By contrast, a random forest driven by mean vegetation density, mean climate, soil properties, and topographic descriptors explains 43% of observed PWS variability. Thus, the dominant species explains only 53% (23% compared to 43%) of explainable variations in PWS. Mean climate and mean NDVI also exert significant influence on PWS. Our results suggest that studies of differences between species should explicitly consider the environments (climate, soil, topography) in which observations for each species are made, and whether those environments are representative of the entire species range. 

Abstract The spatio‐temporal variation of stomatal conductance directly regulates photosynthesis, water partitioning, and biosphere‐atmosphere interactions. While many studies have focused on stomatal response to stresses, the spatial variation of unstressed stomatal conductance remains poorly determined, and is usually characterized in land surface models (LSMs) simply based on plant functional type (PFT). Here, we derived unstressed stomatal conductance at the ecosystem‐scale using observations from 115 global FLUXNET sites. When aggregated by PFTs, the across‐PFT pattern was highly consistent with the parameterizations of LSMs. However, PFTs alone captured only 17% of the variation in unstressed stomatal conductance across sites. Within the same PFT, unstressed stomatal conductance was negatively related to climate dryness and canopy height, which explained 45% of the total spatial variation. Our results highlight the importance of plant‐environment interactions in shaping stomatal traits. The trait‐environment relationship established here provides an empirical approach for improved parameterizations of stomatal conductance in LSMs.