Canopy nitrogen distribution is optimized to prevent photoinhibition throughout the canopy during sun flecks

As photoinhibition primarily reduces the photosynthetic light use efficiency at low light, sunfleck-induced photoinhibition might result in a fatal loss of carbon gain in the shade leaves within a canopy with barely positive carbon balance. We hypothesized that shade leaves at the lower canopy might retain a certain amount of leaf nitrogen (N_L) to maintain energy consumption via electron transport, which contributes to circumventing photoinhibition during sunflecks to keep efficient utilization of low light during the rest period of daytime. We investigated excess energy production, a potential measure of susceptibility to photoinhibition, as a function of N_L distribution within a Japanese oak canopy. Optimal N_L distribution, which maximizes canopy carbon gain, may lead to a higher risk of photoinhibition in shade leaves during sunflecks. Conversely, uniform N_L distribution would cause a higher risk of photoinhibition in sun leaves under the direct sunlight. Actual N_L distribution equalized the risk of photoinhibition throughout the canopy indicated by the constant excess energy production at the highest light intensities that the leaves received. Such a homeostatic adjustment as a whole canopy concerning photoinhibition would be a key factor to explain why actual N_L distribution does not maximize canopy carbon gain.