Lichens and their algal partners are desiccation-tolerant organisms and as such survive after the complete loss of water. This trait is the consequence of several physiological, biochemical and structural features, including specific mechanisms dissipating excess light to avoid photooxidative stress. The maximum quantum yield of photosystem II (PSII; F v /F m ) is widely used as a sensitive indicator of photosynthetic performance and is calculated after complete relaxation in darkness of the fluorescence quenching associated with active light energy dissipation mechanisms. Unexpectedly, we observed that lichens and isolated chlorobionts (chlorophyte symbionts in lichen) maintained in darkness for several hours showed a strong decrease in the ratio F v /F m , which was reversible after re-illumination. We analyzed this dark-induced F v /F m decay in the chlorobiont Asterochloris erici through steady-state and fast-induction kinetics of chlorophyll a fluorescence and simultaneous P700 oxidation measurements. We found that the gradual decay of F v /F m in darkness was caused by reversible dark-induced inactivation of some PSII reaction centers that was accompanied by a decrease in the flux of electrons to PSI. Darkness induced the plastoquinone-reductase activity associated with chlororespiration and the phosphorylation of light harvesting complex (LHC). We propose that upon phosphorylation the LHC detaches from PSII, resulting in a decrease of exciton-trapping by PSII reaction centers and, consequently, an increased dissipation of light energy. This mechanism probably serves an ecophysiological function in lichens to prevent the damage at dawn or under strong fluctuating light conditions when lichens are in a hydrated state.Abbreviations -Chl, chlorophyll; F v /F m , maximum quantum yield of PSII; F o , minimal fluorescence emission; F m , maximal fluorescence emission; FR, far-red light; F v , variable fluorescence; LHCII, light-harvesting complex of PSII; NPQ, non-photochemical quenching; OM, oligotrophic medium; PG, propyl-gallate; P l , P700 signal under illumination; P m , maximal P700 signal observed upon full oxidation; P m ′ , maximal P700 signal induced by combined actinic illumination plus SP; P o , signal when P700 is completely reduced; PQ, plastoquinone; PSII, photosystem II; Q A , primary electron acceptor quinone of PSII; q P , photochemical quenching; PTOX, plastoquinone-oxidoreductase and quinol oxidase; RC, reaction center; ROS, reactive oxygen species; SM, supplemented medium; SP, saturation pulse; TRo/RC, capacity of electron trapping per reaction center; V t , normalice fluorescence; Y(I), quantum yield of photochemical energy conversion in photosystem I; Y(NA), quantum yield of non-photochemical energy dissipation due to acceptor side limitation of photosystem I; Y(ND), quantum yield of non-photochemical energy dissipation due to photosystem I donor side limitation. † These authors equally contributed to this work.
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