The University of Arizona
   s





16.1.9. Maintaining Redox Balance: The Diverse Fates of Pyruvate

The conversion of glucose into two molecules of pyruvate has resulted in the net synthesis of ATP. However, an energy-converting pathway that stopped at pyruvate would not proceed for long, because redox balance has not been maintained. As we have seen, the activity of glyceraldehyde 3-phosphate dehydrogenase, in addition to generating a compound with high phosphoryl-transfer potential, of necessity leads to the reduction of NAD+ to NADH. There are limited amounts of NAD+ in the cell, which is derived from the vitamin niacin, a dietary requirement in human beings. Consequently, NAD+ must be regenerated for glycolysis to proceed. Thus, the final process in the pathway is the regeneration of NAD+ through the metabolism of pyruvate. The sequence of reactions from glucose to pyruvate is similar in most organisms and most types of cells. In contrast, the fate of pyruvate is variable. Three reactions of pyruvate are of prime importance: conversion into ethanol, lactic acid, or carbon dioxide (Figure 16.10).


1. Ethanol is formed from pyruvate in yeast and several other microorganisms. The first step is the decarboxylation of pyruvate. This reaction is catalyzed by pyruvate decarboxylase, which requires the coenzyme thiamine pyrophosphate. This coenzyme, derived from the vitamin thiamine (B1), also participates in reactions catalyzed by other enzymes. The second step is the reduction of acetaldehyde to ethanol by NADH, in a reaction catalyzed by alcohol dehydrogenase. This process regenerates NAD+.


The conversion of glucose into ethanol is an example of alcoholic fermentation. The net result of this anaerobic process is:

Note that NAD+ and NADH do not appear in this equation, even though they are crucial for the overall process. NADH generated by the oxidation of glyceraldehyde 3-phosphate is consumed in the reduction of acetaldehyde to ethanol. Thus, there is no net oxidation-reduction in the conversion of glucose into ethanol (Figure 16.12). The ethanol formed in alcoholic fermentation provides a key ingredient for brewing and winemaking.



2. Lactate is formed from pyruvate in a variety of microorganisms in a process called lactic acid fermentation. The reaction also takes place in the cells of higher organisms when the amount of oxygen is limiting, as in muscle during intense activity. The reduction of pyruvate by NADH to form lactate is catalyzed by lactate dehydrogenase.

The overall reaction in the conversion of glucose into lactate is:

As in alcoholic fermentation, there is no net oxidation-reduction. The NADH formed in the oxidation of glyceraldehyde 3-phosphate is consumed in the reduction of pyruvate. The regeneration of NAD+in the reduction of pyruvate to lactate or ethanol sustains the continued operation of glycolysis under anaerobic conditions.



3. Only a fraction of the energy of glucose is released in its anaerobic conversion into ethanol or lactate. Much more energy can be extracted aerobically by means of the citric acid cycle and the electron-transport chain. The entry point to this oxidative pathway is acetyl coenzyme A (acetyl CoA), which is formed inside mitochondria by the oxidative decarboxylation of pyruvate.



© 2002 by W. H. Freeman and Company.