Effects of drought on mesophyll conductance and photosynthetic limitations at different tree canopy layers

In recent years, many studies have focused on the limiting role of mesophyll conductance (g(m)) to photosynthesis (A(n)) under water stress, but no studies have examined the effect of drought on g(m) through the forest canopy. We investigated limitations to A(n) on leaves at different heights in a mixed adult stand of sessile oak (Quercus petraea) and beech (Fagus sylvatica) trees during a moderately dry summer. Moderate drought decreased A(n) of top and lowest beech canopy leaves much more than in leaves located in the mid canopy; whereas in oak, A(n) of the lower canopy was decreased more than in sunlit leaves. The decrease of A(n) was probably not due to leaf-level biochemistry given that V-Cmax was generally unaffected by drought. The reduction in A(n) was instead associated with reduction in stomatal and mesophyll conductances. Drought-induced increases in stomatal limitations were largest in leaves from the top canopy, whereas drought-induced increases in mesophyll limitations were largest in leaves from the lowest canopy. Sensitivity analysis highlighted the need to decompose the canopy into different leaf layers and to incorporate the limitation imposed by g(m) when assessing the impact of drought on the gas exchange of tree canopies.