Numerous current efforts seek to improve the representation of ecosystem ecology and vegetation demographic processes within Earth System Models (ESMs). These developments are widely viewed as an important step in developing greater realism in predictions of future ecosystem states and fluxes. Increased realism, however, leads to increased model complexity, with new features raising a suite of ecological questions that require empirical constraints. Here, we review the developments that

Intermediate disturbances shape forest structure and composition, which may in turn alter carbon, nitrogen, and water cycling. We used a large-scale experiment in a forest in northern lower Michigan where we prescribed an intermediate disturbance by stem girdling all canopy-dominant early successional trees to simulate an accelerated age-related senescence associated with natural succession. Using 3 years of eddy covariance and sap flux measurements in the disturbed area and an adjacent control plot, we analyzed disturbance-induced changes to plot level and species-specific transpiration and stomatal conductance. We found transpiration to be~15% lower in disturbed plots than in unmanipulated control plots. However, species-specific responses to changes in microclimate varied. While red oak and white pine showed increases in stomatal conductance during postdisturbance (62.5 and 132.2%, respectively), red maple reduced stomatal conductance by 36.8%. We used the hysteresis between sap flux and vapor pressure deficit to quantify diurnal hydraulic stress incurred by each species in both plots. Red oak, a ring porous anisohydric species, demonstrated the largest mean relative hysteresis, while red maple, bigtooth aspen, and paper birch, all diffuse porous species, had the lowest relative hysteresis. We employed the Penman-Monteith model for LE to demonstrate that these species-specific responses to disturbance are not well captured using current modeling strategies and that accounting for changes to leaf area index and plot microclimate are insufficient to fully describe the effects of disturbance on transpiration.

Biophysical controls on plant water status exist at the leaf, stem, and root levels. Therefore, we pose that hydraulic strategy is a combination of traits governing water use at each of these three levels. We studied sap flux, stem water storage, stomatal conductance, photosynthesis, and growth of red oaks (Quercus rubra) and red maples (Acer rubrum). These species differ in stomatal hydraulic strategy and xylem architecture and may root at different depths. Stable isotope analysis of xylem water was used to identify root water uptake depth. Oaks were shown to access a deeper water source than maples. During non‐limiting soil moisture conditions, transpiration was greater in maples than in oaks. However, during a soil dry down, transpiration and stem water storage decreased by more than 80% and 28% in maples but only by 31% and 1% in oaks. We suggest that the preferential use of deep water by red oaks allows the species to continue transpiration and growth during soil water limitations. In this case, deeper roots may provide a buffer against drought‐induced mortality. Using 14 years of growth data, we show that maple growth correlates with mean annual soil moisture at 30 cm but oak growth does not. The observed responses of oak and maple to drought were not able to be explained by leaf and xylem physiology alone. We employed the Finite‐difference Ecosystem‐scale Tree Crown Hydrodynamics model version 2 plant hydrodynamics model to demonstrate the influence of root, stem, and leaf controls on tree‐level transpiration. We conclude that all three levels of hydraulic traits are required to define hydraulic strategy.

Abstract. Hydraulic capacitance and water storage form a critical buffer against cavitation and loss of conductivity within the xylem system. Withdrawal from water storage in leaves, branches, stems, and roots significantly impacts sap flow, stomatal conductance, and transpiration. Storage quantities differ based on soil water availability, tree size, wood anatomy and density, drought tolerance, and hydraulic strategy (anisohydric or isohydric). However, the majority of studies focus on the measurement of storage in conifers or tropical tree species. We demonstrate a novel methodology using frequency domain reflectometry (FDR) to make continuous, direct measurements of wood water content in two hardwood species in a forest in Michigan. We present results of a two month study comparing the water storage dynamics between a mature red oak and red maple, two species with differing wood densities, hydraulic architecture, and hydraulic strategy. We also include results pertaining to the use of different probe lengths to sample water content only within the active sapwood and over the entire conductive sapwood and the outer portion of heartwood in red oak. Both species studied exhibited diurnal cycles of storage that aligned well with the dynamics of sap flux. Red maple, a diffuse porous, relatively isohydric species showed a strong dependence on stored water during both wet and dry periods. Red oak, a ring porous relatively anisohydric species, was less reliant on storage, and did not demonstrate a dependence on soil water potential. Comparison between long and short FDR probes in the oak revealed that oaks may utilize water stored in the innermost layers of the xylem when soil moisture conditions are limiting. We found the FDR probes to be a reliable, functional means for continuous automated measurement of wood water content in hardwoods at a fast time scale. Application of FDR technology for the measurement of tree water storage will benefit forest ecologists as well as the modeling community as we improve our understanding and simulations of plant hydrodynamic processes on a large scale.

The finite difference ecosystem‐scale tree crown hydrodynamics model version 2 (FETCH2) is a tree‐scale hydrodynamic model of transpiration. The FETCH2 model employs a finite difference numerical methodology and a simplified single‐beam conduit system to explicitly resolve xylem water potentials throughout the vertical extent of a tree. Empirical equations relate water potential within the stem to stomatal conductance of the leaves at each height throughout the crown. While highly simplified, this approach brings additional realism to the simulation of transpiration by linking stomatal responses to stem water potential rather than directly to soil moisture, as is currently the case in the majority of land surface models. FETCH2 accounts for plant hydraulic traits, such as the degree of anisohydric/isohydric response of stomata, maximal xylem conductivity, vertical distribution of leaf area, and maximal and minimal xylem water content. We used FETCH2 along with sap flow and eddy covariance data sets collected from a mixed plot of two genera (oak/pine) in Silas Little Experimental Forest, NJ, USA, to conduct an analysis of the intergeneric variation of hydraulic strategies and their effects on diurnal and seasonal transpiration dynamics. We define these strategies through the parameters that describe the genus level transpiration and xylem conductivity responses to changes in stem water potential. Our evaluation revealed that FETCH2 considerably improved the simulation of ecosystem transpiration and latent heat flux in comparison to more conventional models. A virtual experiment showed that the model was able to capture the effect of hydraulic strategies such as isohydric/anisohydric behavior on stomatal conductance under different soil‐water availability conditions.

Land-surface models use different formulations of stomatal conductance and plant hydraulics, and it is unclear which type of model best matches the observed surface-atmosphere water flux. We use the North American Carbon Program data set of latent heat flux (LE) measurements from 25 sites and predictions from 9 models to evaluate models' ability to resolve subdaily dynamics of transpiration. Despite overall good forecast at the seasonal scale, the models have difficulty resolving the dynamics of intradaily hysteresis. The majority of models tend to underestimate LE in the prenoon hours and overestimate in the evening. We hypothesize that this is a result of unresolved afternoon stomatal closure due to hydrodynamic stresses. Although no model or stomata parameterization was consistently best or worst in terms of ability to predict LE, errors in model-simulated LE were consistently largest and most variable when soil moisture was moderate and vapor pressure deficit was moderate to limiting. Nearly all models demonstrate a tendency to underestimate the degree of maximum hysteresis which, across all sites studied, is most pronounced during moisture-limited conditions. These diurnal error patterns are consistent with models' diminished ability to accurately simulate the natural hysteresis of transpiration. We propose that the lack of representation of plant hydrodynamics is, in part, responsible for these error patterns.

Trees play a key role in the global hydrological cycle and measurements performed with the thermal dissipation method (TDM) have been crucial in providing whole-tree water-use estimates. Yet, different data processing to calculate whole-tree water use encapsulates uncertainties that have not been systematically assessed. We quantified uncertainties in conifer sap flux density (F ) and stand water use caused by commonly applied methods for deriving zero-flow conditions, dampening and sensor calibration. Their contribution has been assessed using a stem segment calibration experiment and 4 yr of TDM measurements in Picea abies and Larix decidua growing in contrasting environments. Uncertainties were then projected on TDM data from different conifers across the northern hemisphere. Commonly applied methods mostly underestimated absolute F . Lacking a site- and species-specific calibrations reduced our stand water-use measurements by 37% and induced uncertainty in northern hemisphere F . Additionally, although the interdaily variability was maintained, disregarding dampening and/or applying zero-flow conditions that ignored night-time water use reduced the correlation between environment and F . The presented ensemble of calibration curves and proposed dampening correction, together with the systematic quantification of data-processing uncertainties, provide crucial steps in improving whole-tree water-use estimates across spatial and temporal scales.

Plants are expected to face increasing water stress under future climate change. Most land surface models, including Noah-MP, employ an idealized "big-leaf" concept to regulate water and carbon fluxes in response to soil moisture stress through empirical soil hydraulics schemes (SHSs). However, such schemes have been shown to cause significant uncertainties in carbon and water simulations. In this paper, we present a novel plant hydraulics scheme (PHS) for Noah-MP (hereafter, Noah-MP-PHS), which employs a big-tree rather than big-leaf concept, wherein the whole-plant hydraulic strategy is considered, including root-level soil water acquisition, stem-level hydraulic conductance and capacitance, and leaflevel anisohydricity and hydraulic capacitance. Evaluated against plot-level observations from a mature, mixed hardwood forest at the University of Michigan Biological Station and compared with the default Noah-MP, Noah-MP-PHS better represents plant water stress and improves water and carbon simulations, especially during periods of dry soil conditions. Noah-MP-PHS also improves the asymmetrical diel simulation of gross primary production under low soil moisture conditions. Noah-MP-PHS is able to reproduce different patterns of transpiration, stem water storage and root water uptake during a 2-week dry-down period for two species with contrasting plant hydraulic behaviors, i.e., the "cavitation riskaverse" red maple and the "cavitation risk-prone" red oak. Sensitivity experiments with plant hydraulic capacitance show that the stem water storage enables nocturnal plant water recharge, affects plant water use efficiency, and provides an important buffer to relieve xylem hydraulic stress during dry soil conditions.Plain Language Summary Plants regulate transpiration dynamically through the stomatal aperture, which, in many cases, is governed by plant water status and hydraulic properties. Plant hydraulics describes the mechanics of water movement through plant vascular systems, which is the culmination of emergent phenotypical hydraulic functional traits at the leaf, stem, and root levels. Such physiological mechanisms are excluded in most land surface models, which typically represent plant water stress through empirical soil hydraulics schemes (SHSs) based on either soil water content or soil water potential. In this study, we present a novel plant hydraulics scheme (PHS) to represent plant water stress and the regulation of stomatal conductance for use in the Noah-MP land surface model. Our results show Noah-MP-PHS performs better in its water and carbon simulations than the default Noah-MP with traditional SHSs, especially under dry soil conditions. Noah-MP-PHS also successfully captures the different plant hydraulic behaviors between the "cavitation risk-averse" red maple and the "cavitation risk-prone" red oak. Sensitivity experiments also highlight the vital role played by plant water storage in water and carbon simulations in terms of buffering xylem hydraulic stress during soil moisture dry-down periods. The ...