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Crocin biosynthesis in C. sativus stigmas. A, C. sativus flower at anthesis with red stigmas. B, Proposed scheme of crocin biosynthesis (Frusciante et al., 2014). The CsCCD2 enzyme cleaves zeaxanthin at the 7,8 and 7′,8′ positions, producing 3-OH-β-cyclocitral and crocetin dialdehyde. Crocetin dialdehyde is oxidized to crocetin by an aldehyde dehydrogenase (ALDH), then glycosylated to crocins by UGT enzymes. C, Sugar moieties of the four most abundant C. sativus crocins (crocins 1–4).
