Functional complementation tests of individually expressed CYP97 and HYD enzymes in cells accumulating α- and β-carotene. E. coli cells accumulating both α-carotene and β-carotene were transformed with test plasmids encoding HYD4, CYP97A4, or CYP97C2. Isolated pigments were separated by reverse-phase HPLC, with spectra extracted at 450 nm. Control, Empty pCOLADuet; Z, zeaxanthin; Zei, zeinoxanthin; αcr, α-cryptoxanthin; βcr, β-cryptoxanthin; cβ, 13-cis-β-carotene; αc; α-carotene; βc, β-carotene.