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phosphoenolpyruvic acid pep 2-phosphoenolpyruvate phosphoenolpyruvate anion biochemistry highest-energy phosphate glycolysis gluconeogenesis phosphotransferase system carbon fixation energy source enzyme enolase plants metabolite metabolism biosynthesis cycle mechanism pathway path catabolism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous contractor molecule molecular formula skeletal structural structure ball stick cell biology biomolecule bioactive biochemical 3d