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Likewise, the enzymes needed for the β oxidation of fatty acid derivatives, when isolated from mitochondria, catalyze formation of fatty acyl-CoA derivatives from acetyl-CoA and a reducing agent such as NADH. However, reactant concentrations within cells are rarely appropriate for reversal of a catabolic sequence. For catabolic sequences the Gibbs energy change is usually distinctly negative and reversal requires high concentrations of end products. However, the latter are often removed promptly from the cells.
The stoichiometry is variable but the fermentation can be described in an idealized form as follows: Ch 17IND page - E - 2 CO2 2 HCOOH 2. 2 From Tempest, D. W. and Neijssel, O. 141 2/14/03, 11:17 AM F. Fermentation: “Life without Oxygen” 1 2 Glucose Glycogen R Glycerol-P 1 2 ATP 1 2 ATP O Triose-P 1,3-Bisphosphoglycerate a b Pi Pi ATP Glycerol PEP HCO3– R ATP f ATP Pyruvate HCO3– O CO2 Ethanol R Lactate (2 mol) CO2 e d Acetaldehyde c g CO2 O Acetolactate Acetyl-CoA Oxaloacetate R R HSO3– Formate CO2 (2 mol) Acetyl-P H2 O Malate Acetate ATP Succinate Acetoacetate Butyryl-CoA R R CO2 ATP E Acetone 1-Butanol Biotin R ATP R Succinyl-CoA Acetoin 2,3-Butanediol Acetoacetyl-CoA R CoA transfer Methylmalonyl-CoA CO2 CoA Adduct R 969 Butyrate R 2-Propanol CO2 Propionyl-CoA R R Propionate 1-Propanol Figure 17-9 Reaction sequences in fermentation based on the Embden–Meyerhof–Parnas pathway.
Coupling by Phosphorylation and Subsequent Cleavage by a Phosphatase A third general method for coupling the hydrolysis of ATP to drive a synthetic sequence is to transfer the terminal phospho group from ATP to a hydroxyl group somewhere on a substrate. Then, after the substrate has undergone a synthetic reaction, the phosphate is removed by action of a phosphatase. For example, in the activation of sulfate (Eq. 17-38),159 the overall standard Gibbs energy change for steps a (catalyzed by ATP sulfurylase160,161) and b is distinctly positive (+12 kJ mol –1).