Fermentation pentose phosphate pathway

ATP-proton motive force interconversion Electron transport Entner-Doudoroff Fermentation Glycolysis/gluconeogenesis Pentose phosphate pathway Pyruvate dehydrogenase Sugars TCA cycle Methanogenesis Polysaccharides Other... 

In this chapter we describe the individual reactions of glycolysis, gluconeogenesis, and the pentose phosphate pathway and the functional significance of each pathway. We also describe the various fates of the pyruvate produced by glycolysis they include the fermentations that are used by many organisms in anaerobic niches to produce ATP and that are exploited industrially as sources of ethanol, lactic acid, and other... 

Some lactic acid bacteria of the genus Lactobacillus, as well as Leuconostoc mesenteroides and Zymomonas mobilis, carry out the heterolactic fermentation (Eq. 17-33) which is based on the reactions of the pentose phosphate pathway. These organisms lack aldolase, the key enzyme necessary for cleavage of fructose 1,6-bisphosphate to the triose phosphates. Glucose is converted to ribulose 5-P using the oxidative reactions of the pentose phosphate pathway. The ribulose-phosphate is cleaved by phosphoketolase (Eq. 14-23) to acetyl-phosphate and glyceraldehyde 3-phosphate, which are converted to ethanol and lactate, respectively. The overall yield is only one ATP per glucose fermented. 

Some bacteria that lack the usual aldolase produce ethanol and lactic acid in a 1 1 molar ratio via the "heterolactic fermentation." Glucose is converted to ribulose 5-phosphate via the pentose phosphate pathway enzymes. A thiamin diphosphate-dependent "phosphoketolase" cleaves xylulose 5-phosphate in the presence of inorganic phosphate to acetyl phosphate and glyceraldehyde 3-phosphate. 

The five E. coli genes inserted in Z. mobilis allowed the entry of arabinose into the nonoxidative phase of the pentose phosphate pathway (Fig. 14-22), where it was converted to glucose 6-phosphate and fermented to ethanol. 

Perhaps the acid-tolerant, thermophilic Bacillus coagulans is the only known biocatalyst that naturally produces lactic acid from xylose via the pentose phosphate pathway, not the phosphoketolase pathway (Patel et al., 2006). Three strains, 17C5, P4-102B, and 36D1, can ferment both hexoses and pentoses to pure L(+)-lactic acid at 50 °C and pFI 5.0, an optimal environment... 

Ethanol fermentation from xylose by yeasts can be divided into four distinctive steps. The first step is the reduction of xylose to xylitol mediated by NADPH/ NADH-linked xylose reductase (XR). This is followed by the oxidation of xylitol to xylulose, mediated by NAD-linked xylitol dehydrogenase (XDH). Xylulose-5-phosphate, the key intermediate, is generated from the phosphorylation of xylulose by xylulose kinase. Xylulose-5-phosphate is then channeled into the pentose phosphate pathway for further metabolism (Fig. 9). 

Ten molecules of ethanol can be produced from six molecules of xylose by using a combination of fermentative and pentose phosphate pathways. The net... 

Guanosine triphosphate and ribulose-5-phosphate are recruited in a 1 2 stoichiometric ratio by GTP cyclohydrolase II and DHBP synthase, respectively, for riboflavin biosynthesis. Since at substrate saturation the activity of B. subtilis DHBP is twice the activity of B. suhtilis cyclohydrolase II (DSM, unpublished observations) and since both enzymatic activities are associated with the same bifunctional protein encoded by rihA, the balanced formation of the pyrimidinedione and the dihydroxybutanone intermediates is ensured. However, the ifg.s constant of DHBP synthase ( 1 mmol is about 100-fold higher than the ifg.s constant of GTP cyclohydrolase II imposing the risk of excessive synthesis of the pyrimidinone and pyrimidinedione intermediates in case of reduced intracellular concentrations of pentose phosphate pathway intermediates. This can be expected, for instance, in glucose-limited fed-batch fermentations, which are frequentiy used in industrial applications. The pyrimidinone and pyrimidinedione intermediates are highly reactive, oxidative compounds, which can do serious damage on the bacteria. 

Johansson, B., Hahn-Gaegerdal, B. (2002). The non-oxidative pentose phosphate pathway controls the fermentation rate of xylulose, but not of xylose in Saccharomyces cerevisiae. FEMS Yeast Research, 2, 211 282. 

Li, Y.-H., Ou-Yang, F.-Y, Yang, C.-H., and Li, S.-Y. (2015) The coupling of glycolysis and the Rubisco-based pathway through the non-oxidative pentose phosphate pathway to achieve low carbon dioxide emission fermentation. Bioresour, Technol, 187, 189-197. 

Okano, K., Yoshida, S., Tanaka, T., Ogino, C. et al (2009) Homo-d-lactic acid fermentation from arabinose by redirection of the phosphoketolase pathway to the pentose phosphate pathway in 1-lactate dehydrogenase gene-deficient Lactobacillus plan-tarum. Appl Environ. Microbiol, 75, 5175-5178,... 

Figure 2. Redox cofactor requirement in L-arjabinose catabolism. L-Arabinose conversion to equimolar amounts of CO2 and ethanol is redox neutral, i- -anaerobic fermentation to ethanol should be possible. However, the conversion of L-arabinose to D-xylulose requires NADPH and NAD and produces NADH and NADP. NADPH is mainly regenerated in the oxidative part of the pentose phosphate pathway, where the reduction ofNADP is coupled to C02 production. The abbreviations are G6p, glucose 6-phosphate F6P, fructose 6-phosphate X5P, D-Xylulose 5-phosphate GAP, D-glyceraldehyde 3-phosphate. (Reproduced from Ref. 165 with permission from Elsevier Science)...

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