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Medlyn, Belinda E.

Publications and source records attributed to Medlyn, Belinda E..

Carbon-phosphorus cycle models overestimate CO 2 enrichment response in a mature Eucalyptus forest

The importance of phosphorus (P) in regulating ecosystem responses to climate change has fostered P-cycle implementation in land surface models, but their CO 2 effects predictions have not been evaluated against measurements. Here, we perform a data-driven model evaluation where simulations of eight widely used P-enabled models were confronted with observations from a long-term free-air CO 2 enrichment experiment in a mature, P-limited Eucalyptus forest. We show that most models predicted the correct sign and magnitude of the CO 2 effect on ecosystem carbon (C) sequestration, but they generally overestimated the effects on plant C uptake and growth. We identify leaf-to-canopy scaling of photosynthesis, plant tissue stoichiometry, plant belowground C allocation, and the subsequent consequences for plant-microbial interaction as key areas in which models of ecosystem C-P interaction can be improved. Together, this data-model intercomparison reveals data-driven insights into the performance and functionality of P-enabled models and adds to the existing evidence that the global CO 2 -driven carbon sink is overestimated by models.

54 ENVIRONMENTAL SCIENCES↗

Optimal stomatal theory predicts CO 2 responses of stomatal conductance in both gymnosperm and angiosperm trees

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (A net ) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO 2 (eCO 2 ), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO 2 on iWUE and its components A net and stomatal conductance (g s ). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf–air vapour pressure difference (D). We expected smaller g s , but greater A net , responses to eCO 2 in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO 2 in all PFTs, and that increases in A net had stronger effects than reductions in g s . The USO model correctly captured stomatal behaviour with eCO 2 across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g 1 ) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO 2 conditions.

54 ENVIRONMENTAL SCIENCES↗

Convergence in phosphorus constraints to photosynthesis in forests around the world

Tropical forests take up more carbon (C) from the atmosphere per annum by photosynthesis than any other type of vegetation. Phosphorus (P) limitations to C uptake are paramount for tropical and subtropical forests around the globe. Yet the generality of photosynthesis-P relationships underlying these limitations are in question, and hence are not represented well in terrestrial biosphere models. Here we demonstrate the dependence of photosynthesis and underlying processes on both leaf N and P concentrations. The regulation of photosynthetic capacity by P was similar across four continents. Implementing P constraints in the ORCHIDEE-CNP model, gross photosynthesis was reduced by 36% across the tropics and subtropics relative to traditional N constraints and unlimiting leaf P. Our results provide a quantitative relationship for the P dependence for photosynthesis for the front-end of global terrestrial C models that is consistent with canopy leaf measurements.

59 BASIC BIOLOGICAL SCIENCES↗

One Stomatal Model to Rule Them All? Toward Improved Representation of Carbon and Water Exchange in Global Models

Abstract Stomatal conductance schemes that optimize with respect to photosynthetic and hydraulic functions have been proposed to address biases in land‐surface model (LSM) simulations during drought. However, systematic evaluations of both optimality‐based and alternative empirical formulations for coupling carbon and water fluxes are lacking. Here, we embed 12 empirical and optimization approaches within a LSM framework. We use theoretical model experiments to explore parameter identifiability and understand how model behaviors differ in response to abiotic changes. We also evaluate the models against leaf‐level observations of gas‐exchange and hydraulic variables, from xeric to wet forest/woody species spanning a mean annual precipitation range of 361–3,286 mm yr −1 . We find that models differ in how easily parameterized they are, due to: (a) poorly constrained optimality criteria (i.e., resulting in multiple solutions), (b) low influence parameters, (c) sensitivities to environmental drivers. In both the idealized experiments and compared to observations, sensitivities to variability in environmental drivers do not agree among models. Marked differences arise in sensitivities to soil moisture (soil water potential) and vapor pressure deficit. For example, stomatal closure rates at high vapor pressure deficit range between −45% and +70% of those observed. Although over half the new generation of stomatal schemes perform to a similar standard compared to observations of leaf‐gas exchange, two models do so through large biases in simulated leaf water potential (up to 11 MPa). Our results provide guidance for LSM development, by highlighting key areas in need for additional experimentation and theory, and by constraining currently viable stomatal hypotheses.

54 ENVIRONMENTAL SCIENCES↗

Mechanisms of woody-plant mortality under rising drought, CO 2 and vapour pressure deficit

Drought-associated woody plant mortality may have increased over the past several decades, and is projected to increase in the future, impacting terrestrial climate forcing, biodiversity, and resource availability. The mechanisms underlying such mortality, however, are debated owing to interactions between multiple drivers and mechanisms. In this work, we synthesize knowledge of drought-related tree mortality under a warming and drying atmosphere with rising atmospheric CO 2 . Drought-associated mortality results from the depletion of pools of water and carbon within a plant and declines in their fluxes relative to demand by living tissues. These pools and fluxes are interdependent and underlay plant defenses against biotic agents. Death via failure to maintain a positive water balance is particularly dependent on soil-to-root conductance, capacitance, vulnerability to hydraulic failure, cuticular water losses, and dehydration tolerance, all of which could be exacerbated by reduced carbon supply rates to support cellular survival e.g. the carbon starvation process. The depletion of plant water and carbon pools accelerates under rising vapor pressure deficit; however, increasing CO 2 can mitigate these impacts. Advancing knowledge and reducing predictive uncertainties requires studies that integrate carbon, water, and defensive processes, and utilize a range of experimental and modeling approaches.

54 ENVIRONMENTAL SCIENCES↗

Triose phosphate utilization limitation: an unnecessary complexity in terrestrial biosphere model representation of photosynthesis

Triose phosphates are the principal product of photosynthesis. They are used within the chloroplast for starch synthesis, or translocated to the cytosol where they are used to fuel sucrose synthesis. Use of triose phosphate releases inorganic phosphate, and is under strict metabolic control that matches the supply of triose phosphate from the Calvin-Benson cycle to the demand for carbon by sinks (Heldt & Heldt, 2011; McClain & Sharkey, 2019). However, a low rate of triose phosphate utilization (TPU) can deplete the phosphate pool, restrict ATP synthesis and reduce the availability of ATP to power the Calvin-Benson cycle, thereby limiting photosynthesis (Sharkey, 1985).

54 ENVIRONMENTAL SCIENCES↗