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Summary Photosynthetic capacity per unit irradiance is greater, and the marginal carbon revenue of water (∂A/∂E) is smaller, in shaded leaves than sunlit leaves, apparently contradicting optimization theory. I tested the hypothesis that these patterns arise from optimal carbon partitioning subject to biophysical constraints on leaf water potential.In a whole plant model with two canopy modules, I adjusted carbon partitioning, nitrogen partitioning and leaf water potential to maximize carbon profit or canopy photosynthesis, and recorded how gas exchange parameters compared between shaded and sunlit modules in the optimum.The model predicted that photosynthetic capacity per unit irradiance should be larger, and ∂A/∂Esmaller, in shaded modules compared to sunlit modules. This was attributable partly to radiation‐driven differences in evaporative demand, and partly to differences in hydraulic conductance arising from the need to balance marginal returns on stem carbon investment between modules. The model verified, however, that invariance in the marginal carbon revenue of N (∂A/∂N) is in fact optimal.The Cowan–Farquhar optimality solution (invariance of ∂A/∂E) does not apply to spatial variation within a canopy. The resulting variation in carbon–water economy explains differences in capacity per unit irradiance, reconciling optimization theory with observations.
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Coupled whole‐tree optimality and xylem hydraulics explain dynamic biomass partitioning
Summary Trees partition biomass in response to resource limitation and physiological activity. It is presumed that these strategies evolved to optimize some measure of fitness. If the optimization criterion can be specified, then allometry can be modeled from first principles without prescribed parameterization.We present the Tree Hydraulics and Optimal Resource Partitioning (THORP) model, which optimizes allometry by estimating allocation fractions to organs as proportional to their ratio of marginal gain to marginal cost, where gain is net canopy photosynthesis rate, and costs are senescence rates. Root total biomass and profile shape are predicted simultaneously by a unified optimization. Optimal partitioning is solved by a numerically efficient analytical solution.THORP’s predictions agree with reported tree biomass partitioning in response to size, water limitations, elevated CO2and pruning. Roots were sensitive to soil moisture profiles and grew down to the groundwater table when present. Groundwater buffered against water stress regardless of meteorology, stabilizing allometry and root profiles as deep as c. 30 m.Much of plant allometry can be explained by hydraulic considerations. However, nutrient limitations cannot be fully ignored. Rooting mass and profiles were synchronized with hydrological conditions and groundwater even at considerable depths, illustrating that the below ground shapes whole‐tree allometry.
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- PAR ID:
- 10385401
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- New Phytologist
- Volume:
- 230
- Issue:
- 6
- ISSN:
- 0028-646X
- Page Range / eLocation ID:
- p. 2226-2245
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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