Plants, algae and cyanobacteria need to regulate photosynthetic light harvesting in response to the constantly changing light environment. Rapid adjustments are required to maintain fitness because of a tradeoff between efficient solar energy conversion and photoprotection. The xanthophyll cycle, in which the carotenoid pigment violaxanthin is reversibly converted into zeaxanthin, is ubiquitous among green algae and plants and is necessary for the regulation of light harvesting, protection from oxidative stress, and adaptation to different light conditions1,2. Violaxanthin de-epoxidase (VDE) is the key enzyme responsible for zeaxanthin synthesis from violaxanthin under excess light. Here we show that the CVDE gene from the model green alga Chlamydomonas reinhardtii encodes an atypical VDE. This protein is not homologous to the VDE found in plants and is instead related to a lycopene cyclase from photosynthetic bacteria3. Unlike the plant-type VDE that is located in the thylakoid lumen, the Chlamydomonas CVDE protein is located on the stromal side of the thylakoid membrane. Phylogenetic analysis suggests that CVDE evolved from an ancient de-epoxidase that was present in the common ancestor of green algae and plants, providing evidence of unexpected diversity in photoprotection in the green lineage.

Species composition and photosynthetic characteristics of dominant species of ungrazed plot (UG), overgrazed plot (OG), and restored grazed plot (RG) were determined in the Xilin River Basin, Inner Mongolia, China. Both heavily grazing and restoration significantly affected the composition of different species and life forms. Leymus chinensis, Stipa grandis, and Cleistogenes polyphylla, three dominant perennial grasses in UG plot, contributed 58.9 % aboveground biomass to that of whole community, and showed higher net photosynthetic rate (P N ), transpiration rate (E), and intrinsic water-use efficiency (WUE). In OG plot, relative biomass of L. chinensis and S. grandis significantly decreased, while relative biomass of three shrubs/sub-shrubs, Caragana microphylla, Artemisia frigida, and Kochia prostrata, obviously increased. Heavy grazing significantly decreased P N , E, and WUE of L. chinensis and S. grandis, while shrubs/subshrubs showed significantly higher photosynthetic activity and WUE than the grasses. After 18-year restoration, photosynthetic activities of L. chinensis and S. grandis were significantly higher than those in the OG plot. The proportion of L. chinensis, S. grandis, and C. microphylla significantly increased, and relative biomass of C. polyphylla, A. frigida, and K. prostrata markedly declined in RG plot. We found close relationships between physiological properties of species and their competitive advantage in different land use types. Higher photosynthetic capability means more contribution to total biomass. The variations in physiological characteristics of plants could partly explain the changes in species composition during degrading and restoring processes of Inner Mongolia typical steppes.

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