Carotenoid metabolism, regulation and genetic manipulation in higher plants. Names of bacterial enzymes are in italics. Candidate genes for carotenoid accumulation obtained by QTL analysis are displayed in parentheses and in red. Red upward pointing arrows, gene expression positively correlated with carotenoid biosynthesis; red downward pointing arrows, gene expression negatively correlated with carotenoid biosynthesis; green asterisk, main genetic manipulation nodes in staple cereals. Other MEP isoprenoid-derived metabolites and carotenoid cleavage products apocarotenoids are shown in the green box. ABA, abscisic acid; AO, aldehyde oxidase; CCD, carotenoid cleavage dioxygenase; CKX, cytokinin oxidase/dehydrogenase; CrtB, bacterial phytoene synthase; CrtE, bacterial GGPP synthase; CrtI, bacterial phytoene desaturase/isomerase; CRTISO, carotene isomerase; CrtY, bacterial lycopene β-cyclase; CrtZ, bacterial β-carotene hydroxylase; CYP97C, heme-containing cytochrome P450 carotene ε-ring hydroxylase; DCS, delta-cadinene synthase; DMADP, dimethylallyl diphosphate; DXP, 1-deoxy-D-xylulose 5-phosphate; DXR, 1-deoxy-D-xylulose 5-phosphate reductoisomerase; DXS, 1-deoxyxylulose-5-phosphate synthase; GA3P, D-glyceraldehyde-3-phosphate; GGPP, geranylgeranyl diphosphate; GGPPS, GGPP synthase; HDR, 1-hydroxy-2-methyl-2-(E)-butenyl 4-diphosphate reductase; HYDB, β-carotene hydroxylase [also known as non-heme di-iron β-carotene hydroxylase (BCH) and heme-containing cytochrome P450 β-ring hydroxylase (CYP97A and CYP97B)]; IPP, isopentenyl diphosphate; IPPI, IPP isomerase; KARI, ketol-acid reductoisomerase; LCYB, lycopene β-cyclase; LCYE, lycopene ε-cyclase; MEP, 2C-methyl-D-erythritol-4-phosphate; MGAT1, alpha-1, 3-mannosyl-glycoprotein 2-beta-N-acetylglucosaminyltransferase; MK, mevalonate kinase; NCED, 9-cis-epoxycarotenoid dioxygenase; NXS, neoxanthin synthase; PDS, phytoene desaturase; PSY, phytoene synthase; RAP2.2, a member of the APETALA2 (AP2)/ethylene-responsive element-binding protein transcription factor family; RIN, MADS-box transcription factor RIPENING INHIBITOR; SDG8, SET2 histone methyltransferase; SLC, secologanin synthase; VDE, violaxanthin de-epoxidase; ZDS, ζ-carotene desaturase; ZEP, zeaxanthin epoxidase; ZISO, ζ-carotene isomerase.