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hmg-coa 3-hydroxy-3-methylglutaryl coa coenzyme intermediate mevalonate ketogenesis pathway hmg-coa synthase metabolic metabolism branched-chain amino acid mg-coa hmb-coa reductase mevalonic biosynthesis rendering metabolite cycle mechanism path catabolism biosynthetic biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous contractor human metabolome database hmdb atom molecule molecular structural structure formula skeletal geometry ball stick