What does optimization theory actually predict about crown profiles of photosynthetic capacity when models incorporate greater realism?

Measured profiles of photosynthetic capacity in plant crowns typically do not match those of average irradiance: the ratio of capacity to irradiance decreases as irradiance increases. This differs from optimal profiles inferred from simple models. To determine whether this could be explained by omission of physiological or physical details from such models, we performed a series of thought experiments using a new model that included more realism than previous models. We used ray-tracing to simulate irradiance for 8000 leaves in a horizontally uniform canopy. For a subsample of 500 leaves, we simultaneously optimized both nitrogen allocation (among pools representing carboxylation, electron transport and light capture) and stomatal conductance using a transdermally explicit photosynthesis model. Few model features caused the capacity/irradiance ratio to vary systematically with irradiance. However, when leaf absorptance varied as needed to optimize distribution of light-capture N, the capacity/irradiance ratio increased up through the crown - that is, opposite to the observed pattern. This tendency was counteracted by constraints on stomatal or mesophyll conductance, which caused chloroplastic CO₂ concentration to decline systematically with increasing irradiance. Our results suggest that height-related constraints on stomatal conductance can help to reconcile observations with the hypothesis that photosynthetic N is allocated optimally.