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propionyl-coa coenzyme derivative propionic acid amino acids fatty oxidation propionyl-coa carboxylase propanoyl propionyl acetyl-coa cobalamin methylcitrate cycle citric acid mycobacterium tuberculosis metabolism biogenesis escherichia coli rhodobacter sphaeroides plant isobutyryl-coa catabolism thioesterifcation aspergillus nidulans protein propionylation acidemia gen5 3d metabolite biosynthesis mechanism pathway path biosynthetic intermediate substrate precursor synthesis bioprecursor metabolic enzyme endogenous contractor