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succinic acid butanedioic dicarboxylic acid amber metabolic intermediate succinate fumarate metabolite metabolism biosynthesis cycle mechanism pathway path catabolism biosynthetic biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous contractor electron transport chain atp signaling molecule food additive e363 supplement biological activity bioactive nutrient prescription drug nutraceutica dietary mitochondria tricarboxylic tca hormone-like signaling 3d molecular formula skeletal