Alcohol fermentation pathway

Reactions involve several enzymes, which have to follow in sequence for lactic acid and alcohol fermentation. This is known as the glucose catabolism pathway, with emphasis on energetic and energy carrier molecules such as ATP, ADP, NAD+ and NADH. In this pathway the six-carbon substrate yields two three-carbon intermediates, each of which passes through a sequence of reactions to the stable end product of pyruvic acid. 

Hexose diphosphate was found by Harden and Young69 in cell-free alcoholic-fermentation liquors. In 1930, it was observed that addition of fluoride to fermenting-yeast extracts leads to an accumulation of 0-phospho-D-glyceronic acid,60 which is also a metabolite of muscle extracts.61 Attention was turned, therefore, to the pathway from hexose diphosphate to 0-phos-pho-D-glyceronic acid. In 1932, Fischer and Baer62 described the synthesis of D-glycerose 3-phosphate, and, in 1933, Smythe and Gerischer63 noted... 

For discussion of three aspects of malo-lactic fermentation not presented here, see Refs. 4 and 96. These are the relationship between malo-lactic fermentation and the yeast strain used in the alcoholic fermentation inducibility of the malic decomposing enzyme(s) and the role of oxaloacetic acid in the pathway of conversion of malic acid to lactic acid (cf. Ref. 76). 

Most known thiamin diphosphate-dependent reactions (Table 14-2) can be derived from the five halfreactions, a through e, shown in Fig. 14-3. Each halfreaction is an a cleavage which leads to a thiamin- bound enamine (center, Fig. 14-3) The decarboxylation of an a-oxo acid to an aldehyde is represented by step b followed by a in reverse. The most studied enzyme catalyzing a reaction of this type is yeast pyruvate decarboxylase, an enzyme essential to alcoholic fermentation (Fig. 10-3). There are two 250-kDa isoenzyme forms, one an a4 tetramer and one with an ( P)2 quaternary structure. The isolation of ohydroxyethylthiamin diphosphate from reaction mixtures of this enzyme with pyruvate52 provided important verification of the mechanisms of Eqs. 14-14,14-15. Other decarboxylases produce aldehydes in specialized metabolic pathways indolepyruvate decarboxylase126 in the biosynthesis of the plant hormone indoIe-3-acetate and ben-zoylformate decarboxylase in the mandelate pathway of bacterial metabolism (Chapter 25).1243/127... 

Anaerobic Oxidation of ducose. Historically, the first system of carbohydrate metabolism to be studied was the conversion by yeast of glucose to alcohol (fermentation) according to the equation CnH,Of,2CH)CH,OH + 2CO . The biochemical process is complex, involving the successive catalytic actions of 12 enzymes and known as the Emhden-Meyerhof pathway This series of reactions is summarized in the entry on Glycolysis. 

The biochemistry of alcoholic fermentation involves a series of internal enzyme-mediated oxidation-reduction reactions m which glucose is degraded via the Embden-Meyerhof-Parnas pathway See also Carbohydrates and Glycolysis. 

When plants experience anoxic conditions there is a shift in carbohydrate metabolism from an oxidative to a fermentative pathway (Fig. 1). In the absence of oxygen, ATP is generated not by the Krebs cycle but by alcoholic fermentation, i.e. glycolysis and ethanol synthesis. 

Fig. 1. The pathway of carbon in alcohol fermentation. Enzymes which have been identified as being anaerobically induced are boxed.

Ribereau-Gayon, P, Glories, Y, Maujean, A., Dubourdieu, D. (2000c) Biochemistry of alcoholic fermentation and metabolic pathways of wine yeasts. In P. Ribereau-Gayon (Ed.), Handbook ofEnology, Vol 1, (pp. 51-74). Chichester John Wiley sons. Ltd. 

The first metabolic pathway that we encounter is glycolysis, an ancient pathway employed by a host of organisms. Glycolysis is the sequence of reactions that metabolizes one molecule of glucose to two molecules ofpyruvate with the concomitant net production of two molecules of ATP. This process is anaerobic (i.e., it does not require O2) inasmuch as it evolved before the accumulation of substantial amounts of oxygen in the atmosphere. Pyruvate can be further processed anaerobically (fermented) to lactate (lactic acidfermentation) or ethanol (alcoholic fermentation). Under aerobic conditions, pyruvate can be completely oxidized to CO2, generating much more ATP, as will be discussed in Chapters 17 and 18. 

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.

Figure 1 Fermentation pathways of Escherichia coli. Gene symbols ackA, acetate kinase adhE, alcohol dehydrogenase fdh, formate dehydrogenase of FHL complex fhn, nitrate-inducible formate dehydrogenase hyd, hydrogenase 3 of FHL complex IdhA, NAD+-dependent D-(-)-lactate dehydrogenase pfl, pyruvate formate-lyase pta, phosphotransacetylase. FHL, formate hydrogenlyase. Redrawn from reference 29...

In the overview of glycolysis we noted that the pyruvate produced must be used up in some way so that the pathway will continue to produce ATP. Similarly, the NADH produced by glycolysis in step 6 (see Figure 21.8) must be reoxidized at a later time, or glycolysis will grind to a halt as the available NAD+ is used up. If the cell is functioning under aerobic conditions, NADH will be reoxidized, and pyruvate will be completely oxidized by aerobic respiration. Under anaerobic conditions, however, different types of fermentation reactions accomplish these purposes. Fermentations are catabolic reactions that occur with no net oxidation. Pyruvate or an organic compound produced from pyruvate is reduced as NADH is oxidized. We will examine two types of fermentation pathways in detail lactate fermentation and alcohol fermentation. 

Pyruvate decarboxylase is also part of the pathway involved in alcoholic fermentation in yeast. In the reaction, pyruvate is converted to acetaldehyde, which is subsequently converted to ethanol by action of the enzyme alcohol dehydrogenase ... 

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