Fermentation glyceraldehyde-3-phosphate dehydrogenase

The most confusing aspect of the pathway proposed by Ochoa and his group now rests with the NAD requirement. In proceeding from L-malic acid to L-lactic acid, there is no net change in oxidation state. Yet in whole cells or cell-free extracts, the malo-lactic fermentation will not proceed in the absence of NAD. Therefore, by the proposed mechanism, one is unable to demonstrate the appearance of reduced cofactor, and the NAD specificity cannot be explained as a redox requirement. However, in the time since this mechanism was proposed, an NAD dependent enzyme (glyceraldehyde-3-phosphate dehydrogenase) has been described which requires NAD in a non-redox capacity (29), and it is possible that the same is true for the enzyme causing the malic acid-lactic acid transformation. 

A requirement for all fermentations is the existence of a mechanism for coupling ATP synthesis to the fermentation reactions. In the lactic acid and ethanol fermentations this coupling mechanism consists of the formation of the intermediate 1,3-bisphosphoglycerate by the glyceraldehyde 3-phosphate dehydrogenase (Fig. 10-3, step a). This intermediate contains parts of both the products ATP and lactate or ethanol. 

Glyceraldehyde-3-Phosphate Dehydrogenase and the Generation of ATP in Fermentation Reactions... 

This metabolic scheme, which is called lactate fermentation, is shown in Fig. 11-7. The coreactant cycle between the two dehydrogenase enzymes, glyceraldehyde-3-phosphate dehydrogenase (Step 6) and lactate dehydrogenase, ensures that there is regeneration of NAD+ in this particular oxidation state so that glycolysis, lactate fermentation, and the production of ATP can continue. 

Figure 16.12. Maintaining Redox Balance. The NADH produced by the glyceraldehyde 3-phosphate dehydrogenase reaction must be reoxidized to NAD+ for the glycolytic pathway to continue. In alcoholic fermentation, alcohol dehydrogenase oxidizes NADH and generates ethanol. In lactic acid fermentation (not shovm), lactate dehydrogenase oxidizes NADH while generating lactic acid.

Fig. 1.6. An example of the reaction which yields energy by the oxidation of an organic compound by an organic compound in the alcohol fermentation. Circled numbers 1, glyceraldehyde-3-phosphate dehydrogenase (phosphorylating) 2, alcohol dehydrogenase 3, phosphoglycerate kinase.

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