An approach to modeling resource optimization for substitutable and interdependent resources

We develop a hierarchical approach to modeling organism acclimation to changing availability of and requirements for substitutable and interdependent resources. Substitutable resources are resources that fill the same metabolic or stoichiometric need of the organism. Interdependent resources are resources whose acquisition or expenditure are tightly linked (e.g., light, CO2, and water in photosynthesis and associated transpiration). We illustrate the approach by simulating the development of vegetation with four substitutable sources of N that differ only in the cost of their uptake and assimilation. As the vegetation develops, it uses the least expensive N source first then uses progressively more expensive N sources as the less expensive sources are depleted. Transition among N sources is based on the marginal yield of N per unit effort expended, including effort expended to acquire C to cover the progressively higher uptake costs. We illustrate the approach to interdependent resources by simulating the expenditure of effort to acquire light energy, CO2, and water to drive photosynthesis in vegetation acclimated to different conditions of soil water, atmospheric vapor pressure deficit, CO2 concentration, and light levels. The approach is an improvement on the resource optimization used in the earlier Multiple Element Limitation (MEL) model.