Seasonal changes in photosynthesis and photoprotection in a Quercus ilex subsp. ballota woodland located in its upper altitudinal extreme in the Iberian Peninsula

Quercus ilex L. subsp. ballota (Desf.) Samp., a Mediterranean evergreen species growing in a continental Mediterranean climate, did not experience water stress and showed greater sensitivity to winter stress than to summer stress over a 12-month period. Net CO2 assimilation rates and photosystem II (PSII) efficiency decreased markedly during the cold months and recovered completely in spring. Lutein, neoxanthin and beta-carotene to chlorophyll (Chl) molar ratios all showed the same trend throughout the year, increasing from September to March. This increase was a result of increases in carotenoid concentrations, because Chl concentration per unit leaf area remained stable, and was higher at the end than at the beginning of the first growing season. Lutein-epoxide was a minor component of the total lutein pool. Thermal energy dissipation and non-photochemical quenching (NPQ) were associated with the de-epoxidated forms of the xanthophyll cycle pigments in the warm months. Photosynthetic rates decreased slightly at midday in summer. These changes were accompanied by decreases in maximum potential PSII efficiency (which recovered during the night), actual and intrinsic PSII efficiencies, photochemical quenching and increases in NPQ. Overall, our data indicate down-regulation of photosynthesis during the summer. The diurnal de-epoxidation of violaxanthin to antheraxanthin and zeaxanthin occurred throughout the year, except in January. Antioxidant enzymatic activity increased in the winter months, especially during the coldest months, highlighting its key role in photoprotection against photo-oxidation. Structural and functional modifications protected PSII from permanent damage and allowed 1-year-old leaves to photosynthesize at high rates when temperatures increased in spring.