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The universality of many pathways of core metabolism suggests a strong rolefor evolutionary selection, but it remains unclear whether existing pathwayshave been selected from a large or small set of biochemical possibilities. Toaddress this question, we construct "in silico" all possible biochemicallyfeasible alternatives to the trunk pathway of glycolysis and gluconeogenesis,one of the most highly conserved pathways in metabolism. We show that, eventhough a large number of alternative pathways exist, the alternatives carrylower flux than the real pathway under typical physiological conditions.Alternative pathways that could potentially carry higher flux often lead toinfeasible intermediate metabolite concentrations. We also find that ifphysiological conditions were different, different pathways could outperformthose found in nature. Our results demonstrate how the rules of biochemistryrestrict the alternatives that are open to evolution, and suggest that theexisting trunk pathway of glycolysis and gluconeogenesis represents a maximalflux solution.

Lower glycolysis carries a higher flux than any biochemically possible alternative