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isocitric acid structural isomer hydroxypropane tricarboxylic isocitrates citric acid cycle citrate enzyme aconitase isocitrate dehydrogenase parkinson's disease metabolite metabolism biosynthesis mechanism pathway path catabolism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous contractor cell biology biochemistry biomolecule bioactive molecule molecular formula skeletal structural structure geometry ball stick 3d substance matter material compound component