Coordination theory of leaf nitrogen distribution in a canopy.

It has long been observed that leaf nitrogen concentrations decline with depth in closed canopies in a number of plant communities. This phenomenon is generally believed to be related to a changing radiation environment and it has been suggested by some researchers that plants allocate nitrogen in order to optimize total whole canopy photosynthesis. Although optimization theory has been successfully utilized to describe a variety of physiological and ecological phenomena, it has some shortcomings that are subject to criticism (e.g., time constraints, oversimplifications, lack of insights, etc.). In this paper we present an alternative to the optimization theory of plant canopy nitrogen distribution, which we term coordination theory. We hypothesize that plants allocate nitrogen to maintain a balance between two processes, each of which is dependent on leaf nitrogen content and each of which potentially limits photosynthesis. These two processes are defined as Wc, the Rubiscolimited rate of carboxylation, and Wj, the electron transport-limited rate of carboxylation. We suggest that plants allocate nitrogen differentially to, leaves in different canopy layers in such a way that Wc and Wj remain roughly balanced. In this scheme, the driving force for the allocation of nitrogen within a canopy is the difference between the leaf nitrogen content that is required to bring Wc and Wj into balance and the current nitrogen content. We show that the daily carbon assimilation of a canopy with a nitrogen distribution resulting from this internal coordination of Wc and Wj is very similar to that obtained using optimization theory.