Patterns of spatio-temporal distribution of winter chronic photoinhibition in leaves of three evergreen Mediterranean species with contrasting acclimation responses.

High irradiance and relatively low temperature, which characterize Mediterranean winters, cause chilling stress in plants. Downregulation of photosynthetic efficiency is a mechanism that allows plants to survive these conditions. This study aims to address whether this process shows a regular spatial pattern across leaf surface or not. Three species (Buxus sempervirens, Cistus albidus and Arctostaphylos uva-ursi) with contrasting responses to winter stress were studied. During 7 days, macro and micro Fv/Fm spatial patterns were monitored by the use of chlorophyll fluorescence imaging techniques. In the field, the strongest photoinhibition was found in B. sempervirens, while there was almost no chronic photoinhibition in C. albidus. In leaves of the first species, Fv/Fm decreased from base to tip while in C. albidus it was uniform over the leaf lamina. An intermediate behavior is shown by A. uva-ursi leaves. Spatial heterogeneity distribution of Fv/Fm was found inside the leaves, resulting in greater Fv/Fm values in the inner layers than in the outer ones. Neither xanthophyll-linked downregulation of Fv/Fm nor protein remobilization were the reasons for such spatial patterns since pigment composition and nitrogen content did not reveal tip-base differences. During recovery from winter, photoinhibition changes occurred in Fv/Fm, pigments and chloroplast ultrastructure. This work shows for the first time that irrespective of physiological mechanisms responsible for development of winter photoinhibition, there is an acclimation response with strong spatio-temporal variability at leaf level in some species. This observation should be taken into account when modeling or scaling up photosynthetic responses.