Why are sun leaves thicker than shade leaves?: Consideration based on analyses of CO2 diffusion in the leaf

Light-saturated rates of photosynthesis on leaf area basis (A) depend not only on photosynthetic biochemistry but also on mesophyll structure. Because resistance to CO2 diffusion from the substomatal cavity to the stroma is substantial, it is likely that mesophyll structure affects A through affecting diffusion of CO2 in the leaf. To evaluate effects of various aspects of mesophyll structure on photosynthesis, we constructed a one-dimensional model of CO2 diffusion in the leaf. When mesophyll thickness of the leaf is changed with the Rubisco content per unit leaf area kept constant, the maximum A occurs at an almost identical mesophyll thickness irrespective of the Rubisco contents per leaf area. On the other hand, with an increase in Rubisco content per leaf area, the mesophyll thickness that realizes a given photosynthetic gain per mesophyll thickness (or per leaf cost) increases This probably explains the strong relationship between A and mesephyll thickness. In these simulations, an increase in mesophyll thickness simultaneously means an increase in the diffusional resistance in the intercellular spaces (R-ias), an increase in the total surface area of chloroplasts facing the intercellular spaces per unit leaf area (S-c), and an increase in construction and maintenance cost of the leaf. Leaves can increase S-c and decrease R-ias also by decreasing cell size. Leaves with smaller cells are mechanically stronger. However,actual leaves do not have very small cells. This could be because actual leaves exhibiting considerable rates of leaf area expansion, adequate heat capacitance, high efficiency of N and/or P use, etc, are favoured. Relationships between leaf longevity and mesophyll structure are also discussed.