Leaf hydraulic networks play an important role not only in fluid transport but also in maintaining whole-plant water status through transient environmental changes in soil-based water supply or air humidity. Both water potential and hydraulic resistance vary spatially throughout the leaf transport network, consisting of xylem, stomata and water-storage cells, and portions of the leaf areas far from the leaf base can be disproportionately disadvantaged under water stress. Besides the suppression of transpiration and reduction of water loss caused by stomatal closure, the leaf capacitance of water storage, which can also vary locally, is thought to be crucial for the maintenance of leaf water status. In order to study the fluid dynamics in these networks, we develop a spatially explicit, capacitive model which is able to capture the local spatiotemporal changes of water potential and flow rate in monocotyledonous and dicotyledonous leaves. In electrical-circuit analogs described by Ohm's law, we implement linear capacitors imitating water storage, and we present both analytical calculations of a uniform one-dimensional model and numerical simulation methods for general spatially explicit network models, and their relation to conventional lumped-element models. Calculation and simulation results are shown for the uniform model, which mimics key properties of a monocotyledonous grass leaf. We illustrate water status of a well-watered leaf, and the lowering of water potential and transpiration rate caused by excised water source or reduced air humidity. We show that the time scales of these changes under water stress are hugely affected by leaf capacitance and resistances to capacitors, in addition to stomatal resistance. Through this modeling of a grass leaf, we confirm the presence of uneven water distribution over leaf area, and also discuss the importance of considering the spatial variation of leaf hydraulic traits in plant biology.
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Comparing Model Representations of Physiological Limits on Transpiration at a Semi‐Arid Ponderosa Pine Site
Mechanistic representations of biogeochemical processes in ecosystem models are rapidly advancing, requiring advancements in model evaluation approaches. Here we quantify multiple aspects of model functional performance to evaluate improved process representations in ecosystem models. We compare semi‐empirical stomatal models with hydraulic constraints against more mechanistic representations of stomatal and hydraulic functioning at a semi‐arid pine site using a suite of metrics and analytical tools. We find that models generally perform similarly under unstressed conditions, but performance diverges under atmospheric and soil drought. The more empirical models better capture synergistic information flows between soil water potential and vapor pressure deficit to transpiration, while the more mechanistic models are overly deterministic. Although models can be parameterized to yield similar functional performance, alternate parameterizations could not overcome structural model constraints that underestimate the unique information contained in soil water potential about transpiration. Additionally, both multilayer canopy and big‐leaf models were unable to capture the magnitude of canopy temperature divergence from air temperature, and we demonstrate that errors in leaf temperature can propagate to considerable error in simulated transpiration. This study demonstrates the value of merging underutilized observational data streams with emerging analytical tools to characterize ecosystem function and discriminate among model process representations.
more » « less- Award ID(s):
- 1802885
- NSF-PAR ID:
- 10383698
- Publisher / Repository:
- DOI PREFIX: 10.1029
- Date Published:
- Journal Name:
- Journal of Advances in Modeling Earth Systems
- Volume:
- 14
- Issue:
- 11
- ISSN:
- 1942-2466
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract Plant functional traits provide a link in process‐based vegetation models between plant‐level physiology and ecosystem‐level responses. Recent advances in physiological understanding and computational efficiency have allowed for the incorporation of plant hydraulic processes in large‐scale vegetation models. However, a more mechanistic representation of water limitation that determines ecosystem responses to plant water stress necessitates a re‐evaluation of trait‐based constraints for plant carbon allocation, particularly allocation to leaf area. In this review, we examine model representations of plant allocation to leaves, which is often empirically set by plant functional type‐specific allometric relationships. We analyze the evolution of the representation of leaf allocation in models of different scales and complexities. We show the impacts of leaf allocation strategy on plant carbon uptake in the context of recent advancements in modeling hydraulic processes. Finally, we posit that deriving allometry from first principles using mechanistic hydraulic processes is possible and should become standard practice, rather than using prescribed allometries. The representation of allocation as an emergent property of scarce resource constraints is likely to be critical to representing how global change processes impact future ecosystem dynamics and carbon fluxes and may reduce the number of poorly constrained parameters in vegetation models.
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null (Ed.)Abstract. Plant activity in semi-arid ecosystems is largely controlled by pulses of precipitation, making them particularly vulnerable to increased aridity expected with climate change. Simple bucket-model hydrology schemes in land surface models (LSMs) have had limited ability in accurately capturing semi-arid water stores and fluxes. Recent, more complex, LSM hydrology models have not been widely evaluated against semi-arid ecosystem in situ data. We hypothesize that the failure of older LSM versions to represent evapotranspiration, ET, in arid lands is because simple bucket models do not capture realistic fluctuations in upper layer soil moisture. We therefore predict that including a discretized soil hydrology scheme based on a mechanistic description of moisture diffusion will result in an improvement in model ET when compared to data because the temporal variability of upper layer soil moisture content better corresponds to that of precipitation inputs. To test this prediction, we compared ORCHIDEE LSM simulations from (1) a simple conceptual 2-layer bucket scheme with fixed hydrological parameters; and (2) a 11-layer discretized mechanistic scheme of moisture diffusion in unsaturated soil based on Richards equations against daily and monthly soil moisture and ET observations, together with data-derived transpiration / evaporation, T / ET, ratios, from six semi-arid grass, shrub and forest sites in the southwestern USA. The 11-layer scheme also has modified calculations of surface runoff, bare soil evaporation, and water limitation to be compatible with the more complex hydrology configuration. To diagnose remaining discrepancies in the 11-layer model, we tested two further configurations: (i) the addition of a term that captures bare soil evaporation resistance to dry soil; and (ii) reduced bare soil fraction. We found that the more mechanistic 11-layer model results better representation of the daily and monthly ET observations. We show that is likely because of improved simulation of soil moisture in the upper layers of soil (top 5 cm). Some discrepancies between observed and modelled soil moisture and ET may allow us to prioritize future model development. Adding a soil resistance term generally decreased simulated E and increased soil moisture content, thus increasing T and T / ET ratios and reducing the negative T / ET model-data bias. By reducing the bare soil fraction in the model, we illustrated that modelled leaf T is too low at sparsely vegetated sites. We conclude that a discretized soil hydrology scheme and associated developments improves estimates of ET by allowing the model to more closely match the pulse precipitation dynamics of these semi-arid ecosystems; however, the partitioning of T from bare soil evaporation is not solved by this modification alone.more » « less
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Abstract The cause of reduced leaf-level transpiration under elevated CO2 remains largely elusive. Here, we assessed stomatal, hydraulic, and morphological adjustments in a long-term experiment on Aleppo pine (Pinus halepensis) seedlings germinated and grown for 22–40 months under elevated (eCO2; c. 860 ppm) or ambient (aCO2; c. 410 ppm) CO2. We assessed if eCO2-triggered reductions in canopy conductance (gc) alter the response to soil or atmospheric drought and are reversible or lasting due to anatomical adjustments by exposing eCO2 seedlings to decreasing [CO2]. To quantify underlying mechanisms, we analyzed leaf abscisic acid (ABA) level, stomatal and leaf morphology, xylem structure, hydraulic efficiency, and hydraulic safety. Effects of eCO2 manifested in a strong reduction in leaf-level gc (−55%) not caused by ABA and not reversible under low CO2 (c. 200 ppm). Stomatal development and size were unchanged, while stomatal density increased (+18%). An increased vein-to-epidermis distance (+65%) suggested a larger leaf resistance to water flow. This was supported by anatomical adjustments of branch xylem having smaller conduits (−8%) and lower conduit lumen fraction (−11%), which resulted in a lower specific conductivity (−19%) and leaf-specific conductivity (−34%). These adaptations to CO2 did not change stomatal sensitivity to soil or atmospheric drought, consistent with similar xylem safety thresholds. In summary, we found reductions of gc under elevated CO2 to be reflected in anatomical adjustments and decreases in hydraulic conductivity. As these water savings were largely annulled by increases in leaf biomass, we do not expect alleviation of drought stress in a high CO2 atmosphere.
