Optimum leaf size predicted by a novel leaf energy balance model incorporating dependencies of photosynthesis on light and temperature

To clarify relationships between leaf size and the environment variables, we constructed an energy balance model for a single leaf incorporating Leuning's stomatal conductance model and Farquhar's leaf photosynthesis model. We ran this model for various environmental conditions paying particular attention to the leaf boundary layer. The leaf size maximizing the rate of photosynthesis per unit leaf area (A) at a high irradiance differed depending on the air temperature. In warm environments, A increased with decrease in leaf size, whereas in cool environments, there was the leaf size maximizing A. With the increase in leaf size, the CO2 concentration inside the leaf (C (i)) decreased and the leaf temperature increased, both due to lower boundary layer conductance. At low air temperatures, the negative effect of low C (i) on A in large leaves was compensated by the increase in leaf temperature towards the optimum temperature for A. This balance determined the optimum leaf size for A at low air temperatures. With respect to water use efficiency, large leaves tended to be advantageous, especially in cool environments at low-to-medium irradiances. Some temperature-dependent trends in leaf size observed in nature are discussed based on the present results.