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porphobilinogen pbg organic compound intermediate biosynthesis porphyrin hemoglobin chlorophyll pyrrole porphyrin pathway aminolevulinate enzyme ala dehydratase hydroxymethylbilane synthase porphobilinogen deaminase acute intermittent porphyria urinary rendering metabolite metabolism cycle mechanism path catabolism biosynthetic biogenesis substrate precursor synthesis derivative bioprecursor metabolic endogenous contractor human metabolome database hmdb bioactive cell biochemistry biomolecule biology molecular formula