Molecular genetics of xanthophyll–dependent photoprotection in green algae and plants

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Abstract
The involvement of excited and highly reactive intermediates in oxygenic photosynthesis inevitably results in the generation of reactive oxygen species. To protect the photosynthetic apparatus from oxidative damage, xanthophyll pigments are involved in the quenching of excited chlorophyll and reactive oxygen species, namely 1 Chl*, 3 Chl*, and 1 1O 2 *. Quenching of 1 Chl* results in harmless dissipation of excitation energy as heat and is measured as non–photochemical quenching (NPQ) of chlorophyll fluorescence. The multiple roles of xanthophylls in photoprotection are being addressed by characterizing mutants of Chlamydomonas reinhardtii and Arabidopsis thaliana . Analysis of Arabidopsis mutants that are defective in 1 Chl* quenching has shown that, in addition to specific xanthophylls, the psbS gene is necessary for NPQ. Double mutants of Chlamydomonas and Arabidopsis that are deficient in zeaxanthin, lutein and NPQ undergo photo–oxidative bleaching in high light. Extragenic suppressors of the Chlamydomonas npq1 lor1 double mutant identify new mutations that restore varying levels of zeaxanthin accumulation and allow survival in high light.