Leaf physiological responses to extreme droughts in Mediterranean Quercus ilex forest

Global climate change is expected to result in more frequent and intense droughts in the Mediterranean region. To understand forest response to severe drought, we used a mobile rainfall shelter to examine the impact of spring and autumn rainfall exclusion on stomatal (S(L) ) and non-stomatal (NS(L) ) limitations of photosynthesis in a Quercus ilex ecosystem. Spring rainfall exclusion, carried out during increasing atmospheric demand and leaf development, had a larger impact on photosynthesis than autumn exclusion, conducted at a time of mature foliage and decreasing vapour pressure deficit. The relative importance of NS(L) increased with drought intensity. S(L) and NS(L) were equal once total limitation (T(L) ) reached 60%, but NS(L) greatly exceeded S(L) during severe drought, with 76% NS(L) partitioned equally between mesophyll conductance (MC(L) ) and biochemical (B(L) ) limitations when T(L) reached 100%. Rainfall exclusion altered the relationship between leaf water potential and photosynthesis. In response to severe mid-summer drought stress, A(n) and V(cmax) were 75% and 72% lower in the spring exclusion plot than in the control plot at the same pre-dawn leaf water potential. Our results revealed changes in the relationship between photosynthetic parameters and water stress that are not currently included in drought parameterizations for modelling applications.