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erythrose 4-phosphate erythrose sugar intermediate pentose phosphate pathway calvin cycle precursor biosynthesis aromatic amino acid tyrosine phenylalanine tryptophan shikimate vitamin b6 dxp-dependent erythrose-4-phosphate dehydrogenase rendering metabolite metabolism mechanism path catabolism biosynthetic biogenesis substrate synthesis derivative bioprecursor metabolic enzyme endogenous contractor human metabolome database hmdb