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The Complete Citric Acid Cycle - Part 2 ©1998 by Alberts, Bray, Johnson, Lewis, Raff, Roberts, Walter . http://www.essentialcellbiology.com Published by Garland Publishing, a member of the Taylor & Francis Group. COO– Step 3 In the first of four oxidation steps in the cycle, the carbon carrying the hydroxyl group is converted to a carbonyl group. The immediate product is unstable, losing CO2 while still bound to the enzyme. H C H H C COO– C H HO COO– isocitrate dehydrogenase NAD+ NADH + H COO– C H H C H H C COO– H C H C O C O H COO– + a-ketoglutarate C H H C H C O COO– a-ketoglutarate dehydrogenase complex + HS CoA NAD+ COO– NADH + H C H H C H C succinyl-CoA synthetase C H H C H COO– H 2O GDP Pi GTP succinate COO– H C H C H COO– COO– succinate dehydrogenase C H FAD COO– Step 7 C H COO– fumarase HO C H COO– C H C H COO– H 2O fumarate Step 8 C fumarate H COO– malate COO– malate dehydrogenase HO C H C H C H CH2 COO– malate H COO– FADH2 succinate In the last of four oxidation steps in the cycle, the carbon carrying the hydroxyl group is converted to a carbonyl group, regenerating the oxaloacetate needed for step 1. H O H H C O H COO– S CoA Step 6 C succinyl-CoA COO– H C H S CoA CO2 a-ketoglutarate H + succinyl-CoA The addition of water to fumarate places a hydroxyl group next to a carbonyl carbon. COO– oxalosuccinate intermediate H Step 5 In the third oxidation step in the cycle, FAD removes two hydrogen atoms from succinate. CO2 COO– Step 4 A phosphate molecule from solution displaces the CoA, forming a high-energy phosphate linkage to succinate. This phosphate is then passed to GDP to form GTP. (In bacteria and plants, ATP is formed instead.) H + isocitrate The aketoglutarate dehydrogenase complex closely resembles the large enzyme complex that converts pyruvate to acetyl CoA (pyruvate dehydrogenase). It likewise catalyzes an oxidation that produces NADH, CO2, and a high-energy thioester bond to coenzyme A (CoA). COO– NAD+ NADH + H + O COO– oxaloacetate + HS CoA