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urocanic acid metabolite metabolism biosynthesis cycle mechanism pathway path catabolism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous contractor human metabolome database hmdb l-histidine histidine ammonialyase histidase histidinase urocanate hydratase urocanase 4-imidazolone-5-propionic glutamic aciduria atopic dermatitis strongyloides stercoralis attractant nematode regulatory cells photoprotectant function sunscreen dna damage filaggrin 3d molecule molecular formula skeletal structure geometry