The Xanthophyll Cycle, Protein Turnover, and the High Light Tolerance of Sun-Acclimated Leaves

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Abstract
Changes in photosynthesis rate and photochemical characteristics in response to high irradiance, followed by recovery at low irradiance, were determined in four groups of sun-acclimated leaves of spinach (Spinacia oleracea L.). These four groups were untreated control leaves, leaves treated with either an inhibitor of energy dissipation associated with the xanthophyll cycle (dithiothreitol, DTT) or an inhibitor of chloroplast-encoded protein synthesis (chloramphenicol, CAP), as well as leaves treated with a combination of DTT + CAP. In these sun leaves, treatment with either CAP or DTT alone did not result in an inhibition of the recovery from high-light-induced decreases in photochemical efficiency. Only the treatment with a combination of CAP + DTT caused a strong and irreversible depression of photochemical efficiency. We suggest that in the presence of DTT (and in the absence of xanthophyll cycle-associated energy dissipation), protein turnover may be involved in the recovery process. We further suggest that the reversible depression of photochemical efficiency in CAP-treated sun leaves reflects xanthophyll cycle-associated energy dissipation. In the leaves treated with CAP + DTT a slowly developing decrease in the maximal yield of chlorophyll fluorescence in high light may indicate an alternative, xanthophyll cycle-independent dissipation process in the photochemical system. Moreover, CAP treatments did not cause any changes in the deepoxidation state of the xanthophyll cycle. However, CAP-treated leaves, but not those treated with CAP + DTT, exhibited some decrease in the pool size of the xanthophyll cycle during the exposure to high light.