Plants cope with fluctuating light by frequency-dependent non-photochemical quenching and cyclic electron transport

In natural environments, plants are exposed to rapidly changing light. Maintaining photosynthetic efficiency while avoiding photodamage requires equally rapid regulation of photoprotective mechanisms. We asked what the operation frequency range of regulation is in which plants can efficiently respond to varying light. Chlorophyll fluorescence, P700, plastocyanin, and ferredoxin responses of wild-type Arabidopsis thaliana were measured in oscillating light of various frequencies. We also investigated the npq1 mutant lacking violaxanthin de-epoxidase, the npq4 mutant lacking PsbS-protein, and the mutants crr2-2, and pgrl1ab impaired in different pathways of the cyclic electron transport. The fastest was the PsbS-regulation responding to oscillation periods longer than 10s. Processes involving violaxanthin de-epoxidase dampened changes of chlorophyll fluorescence in oscillation periods of 2min or longer. Knocking out the PGRL1-PGR5 pathway strongly reduced variations of all monitored parameters, probably due to congestion in the electron transport. Incapacitating the NDH-like pathway only slightly changed the photosynthetic dynamics. Our observations are consistent with the assumption that non-photochemical quenching in slow light oscillations involves violaxanthin de-epoxidase to produce, presumably, a stationary, non-oscillating level of zeaxanthin. We interpret the observed dynamics of Photosystem I components as being formed in slow light oscillations partially by thylakoid remodeling that modulates the redox rates.