Complex Roles of PsbS and Xanthophylls in the Regulation of Nonphotochemical Quenching in under Fluctuating Light.

Protection of photosystem II against damage from excess light by nonphotochemical quenching (NPQ) includes responses on a wide range of timescales. The onset of the various phases of NPQ overlap in time making it difficult to discern if they influence each other or involve different photophysical mechanisms. To unravel the complex relationship of the known actors in NPQ, we perform fluorescence lifetime snapshot measurements throughout multiple cycles of alternating 2 min periods of high light and darkness. By comparing the data with an empirically based mathematical model that describes both fast and slow quenching responses, we suggest that the rapidly reversible quenching response depends on the state of the slower response. By studying a series of mutants, we find that removing zeaxanthin (Zea) or enhancing PsbS concentration, for example, influences the amplitudes of the slow quenching induction and recovery, but not the timescales. The plants' immediate response to high light appears independent of the illumination history, while PsbS and Zea have distinct roles in both quenching and recovery. We further identify two parameters in our model that predominately influence the recovery amplitude and propose that our approach may prove useful for screening new mutants or overexpressors with enhanced biomass yields under field conditions.