To identify factors governing peel-color development in mandarins, we examined carotenoid content and composition and the expression of carotenoid-related genes during four stages of ripening (i.e., green, breaker, yellow, and orange) in two varieties: 'Ora', which has orange fruit, and 'Shani', which has orange-reddish fruit. The two varieties had different carotenoid compositions, and 'Shani' had a significantly higher level of total carotenoid pigments. 'Shani' was rich in the deep orange β-cryptoxanthin and the orange-reddish β-citraurin, whereas 'Ora' was rich in the orange violaxanthin. RNA-Seq analysis revealed significantly greater expression of the carotenoid-biosynthesis genes PSY, βLCY, βCHX, and CCD4b, as well as MEP-pathway genes and several ethylene-biosynthesis and -signaling genes in 'Shani' fruit. In contrast, the expression levels of genes involved in the synthesis of α-branch carotenoids (i.e., εLCY and εCHX) and ZEP, which is involved in the formation of violaxanthin, were significantly higher in the 'Ora' fruit.
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