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malic acid dicarboxylic hydroxybutanedioic 2-hydroxysuccinic 2-hydroxybutanedioic food additive malates citric acid cycle e296 food adjuvants ingredient agent ins number preservative biochemistry metabolite metabolism biosynthesis mechanism pathway path catabolism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous fruits malolactic fermentation molecule molecular formula skeletal structural structure 3d biomolecule bioactive biochemical