Exposure of the shaded side of apple fruit to full sun leads to up-regulation of both the xanthophyll cycle and the ascorbate–glutathione cycle

Abstract

Approximately 80 days after full bloom, well-exposed fruit on the south part of the canopy of mature Liberty/M9 apple ( Malus domestica Borkh.) trees were randomly assigned to one of the following two treatments. Some fruits were turned about 180° to expose the original shaded side to full sun, whereas the rest served as untreated controls. On days 0, 1, 2, 4, 7 and 10 after treatment, fruit peel samples were taken from the original shaded side of the treated fruit and both the sun-exposed side and the shaded side of the control fruit at midday, to determine photosynthetic pigments, enzymatic and non-enzymatic antioxidants. Maximum photosystem II (PSII) efficiency, measured as F v/ F m (maximum fluorescence) of chlorophyll fluorescence at pre-dawn, was higher in the shaded side than the sun-exposed side of the control fruit. F v/ F m of the original shaded side decreased sharply after 1 day exposure to full sun, and then gradually recovered to a similar value of the sun-exposed side of the control fruit by day 10. The shaded side of the control fruit had much lower xanthophyll cycle pool size, conversion and antioxidant enzymes and also soluble antioxidants of the ascorbate–glutathione cycle than the sun-exposed side. Xanthophyll cycle pool size of the original shaded side increased in response to full sun exposure, reaching a similar value of the sun-exposed side by day 10. Almost all the xanthophyll cycle pool was converted to zeaxanthin and antheraxanthin starting from the first day of exposure to full sun. In contrast, chlorophyll content decreased whereas lutein or neoxanthin content remained unchanged. In response to full sun exposure, ascorbate peroxidase, monodehydroascorbate reductase, dehydroascorbate reductase and glutathione reductase and total pool size and reduction state of both ascorbate and glutathione, all increased to the corresponding values found in the sun-exposed side of the control fruit over a 10-day period. It is concluded that both xanthophyll cycle and the ascorbate–glutathione cycle in the original shaded side are up-regulated in response to full sun exposure to minimize photo-oxidative damage and contributes to its acclimation to full sun.