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orotidine 5-monophosphate omp orotidylic acid pyrimidine nucleotide intermediate biosynthesis uridine monophosphate orotate phosphoribosyl pyrophosphate enzyme orotate phosphoribosyltransferase ump synthase orotic aciduria rendering metabolite metabolism cycle mechanism pathway path catabolism biosynthetic biogenesis substrate precursor synthesis derivative bioprecursor metabolic endogenous contractor human metabolome database hmdb bioactive cell biochemistry biology biomolecule molecule molecular structural structure formula skeletal geometry ball stick