Foliar photochemical processes and carbon metabolism under favourable and adverse winter conditions in a Mediterranean mixed forest, Catalonia (Spain).

Evergreen trees in the Mediterranean region must cope with a wide range of environmental stresses from summer drought to winter cold. The mildness of Mediterranean winters can periodically lead to favourable environmental conditions above the threshold for a positive carbon balance, benefitting evergreen woody species more than deciduous ones. The comparatively lower solar energy input in winter decreases the foliar light saturation point. This leads to a higher susceptibility to photoinhibitory stress especially when chilly (<12°C) or freezing temperatures (<0°C) coincide with clear skies and relatively high solar irradiances. Nonetheless, the advantage of evergreen species that are able to photosynthesize all year round where a significant fraction can be attributed to winter months, compensates for the lower carbon uptake during spring and summer in comparison to deciduous species. We investigated the ecophysiological behaviour of three co-occurring mature evergreen tree species (Quercus ilex L., Pinus halepensis Mill., and Arbutus unedo L.) during a period of mild winter conditions and their responses to a sudden cold period. The state of the photosynthetic machinery in both periods was thus tested by estimating the foliar photosynthetic potential with CO₂ response curves in parallel with chlorophyll fluorescence measurements. The studied evergreen tree species benefited strongly from mild winter conditions by exhibiting extraordinarily high photosynthetic potentials similar to those under spring conditions. A sudden period of frost, however, negatively affected the photosynthetic apparatus, leading to significant decreases in key physiological parameters such as the maximum carboxylation velocity (V_{c, max}), the maximum photosynthetic electron transport rate (J_max), and the optimal fluorometric quantum yield of photosystem II (F_v/F_m). This change persisted for several weeks after the cold period despite the recovery of the temperature to the conditions previous to the frost event. The responses of V_{c, max} and J_max were highly species-specific, where Q. ilex exhibited the highest and P. halepensis the lowest reductions. In contrast, the optimal fluorometric quantum yield of photosystem II (F_v/F_m) was significantly lower in A. unedo after the cold period. The leaf position played an important role in Q. ilex showing a comparatively stronger winter effect on sunlit leaves. Our results generally agreed with the previous classifications of photoinhibition-avoiding (P. halepensis) and photoinhibition-tolerant (Q. ilex) species on the basis of their susceptibility to dynamic photoinhibition, whereas A. unedo was the least tolerant to photoinhibition, which was chronic in this species. Q. ilex and P. halepensis seem to follow contrasting photoprotective strategies which are, however, equally successful under the prevailing conditions exhibiting an adaptive advantage over A. unedo in our study site. These results show that our understanding of the dynamics of interspecific competition in Mediterranean ecosystems requires consideration of the physiological behaviour during winter which may have important implications for long-term carbon budgets and growth trends. [ABSTRACT FROM AUTHOR]