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beta-hydroxy beta-methylbutyryl-coa hmb-coa 3-hydroxyisovaleryl-coa metabolite l-leucine human precursor hmb mc-coa rendering substrate mechanism hmdb biochemistry organic biomolecule biosynthesis pathway compound intermediate endogenous path scientific catabolism substance derivative biosynthetic biology chemical human metabolome database science cell bioprecursor cycle synthesis contractor enzyme material bioactive research metabolic matter biogenesis metabolism