Gluconeogenesis; Regulation of Glycolysis & Gluconeogenesis
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1 Gluconeogenesis; Regulation of Glycolysis & Gluconeogenesis
2 Gluconeogenesis occurs mainly in liver. Gluconeogenesis occurs to a more limited extent in kidney & small intestine under some conditions. Synthesis of glucose from pyruvate utilizes many of the same enzymes as Glycolysis. Three Glycolysis reactions have such a large negative DG that they are essentially irreversible. exokinase (or Glucokinase) Phosphofructokinase Pyruvate Kinase. These steps must be bypassed in Gluconeogenesis. Two of the bypass reactions involve simple hydrolysis reactions.
3 P glucose-6-phosphate Glucose-6-phosphatase glucose + P i exokinase or Glucokinase (Glycolysis) catalyzes: glucose + ATP glucose-6-phosphate + ADP Glucose-6-Phosphatase (Gluconeogenesis) catalyzes: glucose-6-phosphate + 2 glucose + P i
4 6 2 2 P 3 5 Phosphofructokinase P 3 ATP ADP P P i fructose-6-phosphate fructose-1,6-bisphosphate Fructose-1,6-biosphosphatase Phosphofructokinase (Glycolysis) catalyzes: fructose-6-p + ATP fructose-1,6-bisp + ADP Fructose-1,6-bisphosphatase (Gluconeogenesis) catalyzes: fructose-1,6-bisp + 2 fructose-6-p + P i
5 Pyruvate arboxylase 3 ATP ADP + P i 3 2 PEP arboxykinase GTP GDP 2 2 pyruvate oxaloacetate PEP 2 P 3 Bypass of Pyruvate Kinase (2 enzymes): Pyruvate arboxylase (Gluconeogenesis) catalyzes: pyruvate ATP oxaloacetate + ADP + P i PEP arboxykinase (Gluconeogenesis) catalyzes: oxaloacetate + GTP PEP + GDP + 2
6 Pyruvate arboxylase (pyruvate oxaloactate) is allosterically activated by acetyl oa. [xaloacetate] tends to be limiting for Krebs cycle. Gluconeogenesis oxaloacetate Glucose-6-phosphatase glucose-6-p glucose pyruvate acetyl oa Krebs ycle Glycolysis citrate fatty acids ketone bodies When gluconeogenesis is active in liver, oxaloacetate is diverted to form glucose. xaloacetate depletion hinders acetyl oa entry into Krebs ycle. The increase in [acetyl oa] activates Pyruvate arboxylase to make oxaloacetate.
7 PEP arboxykinase Reaction GTP GDP 2 P oxaloacetate PEP PEP arboxykinase catalyzes GTP-dependent oxaloacetate PEP. It is thought to proceed in 2 steps: xaloacetate is first decarboxylated to yield a pyruvate enolate anion intermediate. Phosphate transfer from GTP then yields phosphoenolpyruvate (PEP).
8 The source of pyruvate and oxaloacetate for gluconeogenesis during fasting or carbohydrate starvation is mainly amino acid catabolism. Some amino acids are catabolized to pyruvate, oxaloacetate, or precursors of these. Muscle proteins may break down to supply amino acids. These are transported to liver where they are deaminated and converted to gluconeogenesis inputs. Glycerol, derived from hydrolysis of triacylglycerols in fat cells, is also a significant input to gluconeogenesis.
9 glyceraldehyde-3-phosphate Summary of Gluconeogenesis Pathway: Gluconeogenesis enzyme names in red. Glycolysis enzyme names in blue. NAD + + P i NAD + + Glyceraldehyde-3-phosphate Dehydrogenase 1,3-bisphosphoglycerate ADP Phosphoglycerate Kinase ATP 3-phosphoglycerate 2-phosphoglycerate 2 Enolase phosphoenolpyruvate Phosphoglycerate Mutase 2 + GDP PEP arboxykinase GTP oxaloacetate P i + ADP 3 + ATP pyruvate Pyruvate arboxylase Gluconeogenesis
10 glucose Gluconeogenesis P i Glucose-6-phosphatase 2 glucose-6-phosphate fructose-6-phosphate P i Phosphoglucose Isomerase Fructose-1,6-bisphosphatase 2 fructose-1,6-bisphosphate Aldolase glyceraldehyde-3-phosphate + dihydroxyacetone-phosphate Triosephosphate Isomerase (continued)
11 Glycolysis & Gluconeogenesis are both spontaneous. If both pathways were simultaneously active in a cell, it would constitute a "futile cycle" that would waste energy. Glycolysis: glucose + 2 NAD ADP + 2 P i 2 pyruvate + 2 NAD + 2 ATP Gluconeogenesis: 2 pyruvate + 2 NAD + 4 ATP + 2 GTP glucose + 2 NAD ADP + 2 GDP + 6 P i Questions: 1. Glycolysis yields how many ~P? 2 2. Gluconeogenesis expends how many ~P? 6 3. A futile cycle of both pathways would waste how many ~P per cycle? 4
12 Reciprocal regulation by fructose-2,6-bisphosphate: Fructose-2,6-bisphosphate stimulates Glycolysis. Fructose-2,6-bisphosphate allosterically activates the Glycolysis enzyme Phosphofructokinase. Fructose-2,6-bisphosphate also activates transcription of the gene for Glucokinase, the liver variant of exokinase that phosphorylates glucose to glucose-6-phosphate, the input to Glycolysis. Fructose-2,6-bisphosphate allosterically inhibits the gluconeogenesis enzyme Fructose-1,6-bisphosphatase.
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