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Six‐Carbon Reactions

Six-carbon reactions of glycolysis represent an energy investment of two high-energy phosphate bonds. Glucose enters glycolysis in a phosphorylated form, as glucose-6-phosphate:

When the glucose originates by breakdown of its polymeric forms, starch or glycogen, it is already phosphorylated, as glucose-1- phosphate, and the initial reaction is catalyzed by the enzyme phosphoglucomutase.

When glucose is present in its unphosphorylated form, the first reaction of glycolysis is a phosphorylation. Although the goal of glycolysis is the synthesis of ATP, a high-energy phosphate must be invested first, catalyzed by the enzyme hexokinase:



The end result of either of these reaction schemes is glucose-6- phosphate, which is now isomerized to fructose-6-phosphate by the enzyme phosphoglucose isomerase:



This is a reversible reaction, both in vitro and in vivo. Fructose-6- phosphate is then phosphorylated by the enzyme phosphofructokinase, in a second energy investment involving ATP:



Fructose-1,6-bisphosphate, the product of this reaction, is then cleaved by aldolase to two 3-carbon compounds:



These two products, glyceraldehyde-3-phosphate and dihydroxyacetone phosphate, are rapidly interconverted by triose phosphate isomerase.

The Keq of the aldolase reaction favors dihydroxyacetone phosphate; however, glyceraldehyde-3-phosphate is drained off for the further reactions of glycolysis, while dihydroxyacetone phosphate is not. This means that the concentration of glyceraldehyde-3-phosphate is very low during active metabolism and dihydroxyacetone phosphate is converted into glyceraldehyde-3-phospate.

The production of the triose phosphates represents the end of the investment. The reactions so far can be summarized as:




In shorthand:




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