The mathematics of linked optimisation for water and nitrogen use in a canopy

We develop, and discuss the implementation of, a mathematical framework for inferring optimal patterns of water and nitrogen use. Our analysis is limited to a time scale of one day and a spatial scale consisting of the green canopy of one plant, and we assume that this canopy has fixed quantities of nitrogen and water available for use in photosynthesis. The efficiencies of water and nitrogen use, and the interactions between the two, are strongly affected by physiological and physical properties that can be modeled in different ways. The thrust of this study is therefore to discuss these properties and how they affect the efficiencies of nitrogen and water use, and to demonstrate, qualitatively, the effects of different model assumptions on inferred optimal strategies. Preliminary simulations suggest that the linked optimisation of nitrogen and water use is particularly sensitive to the level of detail in canopy light penetration models (e.g., whether sunlit and shaded fractions are pooled or considered independently), and to assumptions regarding nitrogen and irradiance gradients within leaves (which determine how whole-leaf potential electron transport rate is calculated from leaf nitrogen content and incident irradiance).