Carbohydrate. Metabolism

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1 Carbohydrate Metabolism

2 Dietary carbohydrates (starch, glycogen, sucrose, lactose Mouth salivary amylase Summary of Carbohydrate Utilization Utilization for energy (glycolysis) ligosaccharides and disaccharides Storage (glycogenesis) Stomach, non-enzymatic hydrolysis Small intestine, pancreatic amylase Conversion to other carbohydrates Monosaccharides Excretion Small intestinal lining Monosaccharides in bloodstream Undesired biochemical processes

3 Glycolysis Glycolysis is a nine step biochemical pathway that oxidizes glucose into two molecules of pyruvic acid (pyruvate). During this process, energy is released and some of it is captured in the form of ATP. H H H H H C6H126 glucose The electrons removed from glucose are captured in the form of NADH. + 2 NAD Pi C ATP + 2 NADH + 2 H H2 H 3 C C C3H33 - pyruvate

4 Glycolysis The pyruvate formed may be or oxidized to carbon dioxide and water in the citric acid cycle may be reduced to L-lactic acid C NADH,H+ NAD+ C C H C H CH 3 CH 3 Lactate Dehydrogenase Reaction

5 Glycolysis Proceeds in Two Stages

6 Stage 1 of Glycolysis: Glucose and other hexoses are converted into glyceraldehyde-3-phosphate.

7 Stage 1 of Glycolysis: glucose ATP hexokinase glucose-6-phosphate 6 C 6 C - P phosphohexose isomerase fructose-6-phosphate ATP phosphofructo kinase fructose-1,6-bisphosphate 6 C - P P - 6 C - P aldolase glyceraldehyde -3-phosphate triosephosphate isomerase dihydroxyacetone phosphate 3 C - P 3 C - P

8 Stage 2 of Glycolysis: 2 molecules of glyceraldehyde-3-phosphate are oxidized and converted into two molecules of pyruvate. The NADH generated during stage 2 may be used to reduce the pyruvate molecules to lactate molecules.

9 Stage 2 of Glycolysis: glyceraldehyde-3-phosphate dehydrogenase glyceraldehyde-3-phosphate phosphoglcerate kinase phosphoglycerate mutase 1,3-bisphosphoglycerate 3-phosphoglycerate NAD+, Pi NADH, H+ ATP 3 C - P P - 3 C - P* 3 C - P 2-phosphoglycerate 3 C - P enolase phosphoenolpyruvate P - 3 C - P* pyruvate kinase pyruvate ATP 3 C - P

10 Stage 1 Stage 2 GLU G3P G6P BPG F6P 3PG 2PG FBP PEP G3P DHAP PYR Energy consumed Energy produced

11 The Individual Chemical Transformations of Glycolysis Distinguish Between the Two Stages Count the Carbons Identify High Energy Compounds Understand the verall Transformation

12 Chemical Transformations of Glycolysis The overall reaction of steps 1-10 is: Glucose Pi 6C 2 lactate + 2 ATP 3C In cells where a high rate of ATP production is not needed, the pyruvate formed by glycolysis is not reduced to lactate, but instead is oxidized to carbon dioxide and water in the mitochondria. This allows the cell to capture a greater amount of energy from the original glucose molecule. Note:In some microorganisms, such as yeast, pyruvate is first decarboxylated and the resulting acetaldehyde is reduced to ethanol, regenerating NAD +. C C CH 3 pyruvate decarboxylase C 2 H C CH 3 NADH,H+ NAD+ alcohol dehydrogenase H HC CH 3 H

13 The Pentose Phosphate Pathway Glucose-6-P N + 6C ribose-5-p NH + 2 H + + C2 5C The pentose phosphate pathway utilizes glucose to produce pentoses (for DNA, RNA) and NH (for reductive biosynthesis). This pathway includes additional enzymes that allow the interconversion of pentoses and hexoses. This allows the cell four alternatives: Produce both pentoses and NH. Produce only NH when pentoses are not required. Produce only pentoses when NH is not required. Produce ATP (glycolysis) and NH when both are required.

14 Metabolic Pathways of Glucose Name Glycolysis Pentose Phosphate Pathway Gluconeogenesis Glycogenesis Glycogenolysis Derivation of Name glyco-, glucose sweet pentose-, five carbon sugar neo-, new genesis-, creation glyco (gen)-glycogen genesis-, creation glyco (gen)-glycogen -lysis, breakdown Function glucose--->pyruvate glucose---> 5-carbon sugars small molecules---> glucose glucose--->glycogen glycogen--->glucose

15 Gluconeogenesis Gluconeogenesis is the biosynthesis of glucose from lactate and certain other small molecules. Gluconeogenesis occurs in the liver and kidney. Gluconeogenesis reverses many of the reactions in glycolysis; however, three irreversible reactions must be bypassed using different reactions and enzymes.

16 Glycolysis ATP glucose glyceraldehyde-3-phosphate NAD+, Pi glucose-6-phosphate fructose-6-phosphate ATP fructose-1,6-bisphosphate 1,3-bisphosphoglycerate 3-phosphoglycerate 2-phosphoglycerate phosphoenolpyruvate NADH, H+ ATP glyceraldehyde-3-phosphate pyruvate ATP

17 Free Energy Changes for Glycolysis Reactions Steps* Glycolytic Reaction Step ΔG(kJ/mol) 1 glucose + ATP glucose-6-phosphate glucose-6-phosphate fructose-6-phosphate 0 to fructose-6-phosphate + ATP fructose-1,6-bisphosphate fructose-1,6-bisphosphate DHP + G3P 0 to -6 dihydroxyacetone phosphate glyceraldehyde-3-phosphate 0 to 4 5 glyceraldehyde-3-phosphate 1,3-bisphosphoglycerate -2 to 2 6 1,3-bisphosphoglycerate + 3-phosphoglycerate + ATP 0 to phosphoglycerate 2-phosphoglycerate 0 to phosphoglycerate phosphoenolpyruvate 0 to phosphoenolpyruvate + pyruvate + ATP -17 *calculated from the actual physiological concentrations in erythrocytes

18 Glycolysis ATP glucose glyceraldehyde-3-phosphate NAD+, Pi glucose-6-phosphate fructose-6-phosphate ATP fructose-1,6-bisphosphate 1,3-bisphosphoglycerate 3-phosphoglycerate 2-phosphoglycerate phosphoenolpyruvate NADH, H+ ATP glyceraldehyde-3-phosphate pyruvate ATP

19 Glycolysis glyceraldehyde-3-phosphate NAD+, Pi Gluconeogenesis pyruvate 1,3-bisphosphoglycerate NADH, H+ phosphoenolpyruvate 3-phosphoglycerate ATP 2-phosphoglycerate 2-phosphoglycerate NAD+, Pi 3-phosphoglycerate phosphoenolpyruvate pyruvate ATP NADH, H+ 1,3-bisphosphoglycerate ATP glyceraldehyde-3-phosphate

20 pyruvate oxaloacetate ATP, HC3 - NADH, H+ Gluconeogenesis pyruvate phosphoenolpyruvate malate NAD+ mitochondria cytosol 2-phosphoglycerate malate NAD+ 3-phosphoglycerate NAD+, Pi oxaloacetate NADH, H+ GTP NADH, H+ 1,3-bisphosphoglycerate phosphoenolpyruvate GDP, HC3 - ATP glyceraldehyde-3-phosphate

21 pyruvate carboxylase (requires biotin) pyruvate oxaloacetate ATP, HC3 - NADH, H+ phosphoenolpyruvate carboxykinase malate malate oxaloacetate phosphoenolpyruvate NAD+ NAD+ NADH, H+ GTP GDP, HC3 - mitochondria GDP cytosol ATP only occurs when there is an abundance of ATP

22 Glycolysis Gluconeogenesis ATP glucose glyceraldehyde -3-phosphate dihydroxyacetone -3-phosphate glucose-6-phosphate fructose-6-phosphate ATP fructose-1,6-bisphosphate fructose-1,6- bisphosphatase fructose-6-phosphate H2 Pi fructose-1,6-bisphosphate glucose-6-phosphate glyceraldehyde-3-phosphate glucose- 6-phosphatase glucose H2 Pi

23 Dietary carbohydrates (starch, glycogen, sucrose, lactose Mouth salivary amylase Summary of Carbohydrate Utilization Utilization for energy (glycolysis) ligosaccharides and disaccharides Storage (glycogenesis) Stomach, non-enzymatic hydrolysis Small intestine, pancreatic amylase Conversion to other carbohydrates Monosaccharides Excretion Small intestinal lining Monosaccharides in bloodstream Undesired biochemical processes

24 Glycogenesis and Glycogenolysis

25 gluconeogenesis Glucose Glucose-6-phosphate glycolysis Glucose-1-phosphate (Glucose)n-1 UTP 2 Pi glycogenolysis Glucose-UDP glycogenesis HP3 2- (Glucose)n glycogen UDP (Glucose)n-1

26 Glycogenesis When the concentration of ATP is high, glycogen is synthesized and stored in liver and muscle cells as glycogen granules. The addition of a molecule of glucose to a growing α 1 4 glycogen strand involves these reactions: 1) Glucose-6-phosphate Glucose-1-phosphate 2) Glucose-1-phosphate + UTP UDP-glucose + PP, PPi + H2 2 Pi 3) UDP-glucose + (glucose)n-1 UDP + (glucose)n

27 H H Glycogen Synthase Reaction H H P UDP-Glucose P N NH H H H H H H H H H non-reducing end of glycogen P P H H N NH UDP H H H H H H H H H H elongated glycogen

28 Glycogenesis When a growing α 1 4 glycogen strand reaches a certain length, a branching enzyme breaks the α 1 4 chain and remakes an α 1 6 linkage. this results in branches, each of which can be extended by further α 1 4 linkages

29 H H H H H H H H H H H H H H H H H H glycogen branching enzyme H H H H H H H H H H H H H glycogen synthase autocatalysis H H H H H H H H H H H H H H CH 2 H H H H H H H H H H H H H H H H H H H Further elongation and branching CH H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H glycogenin glycogenin glycogenin glycogenin

30 Glycogenesis Glycogen synthesis is stimulated by insulin through a covalent modification of the glycogen synthase kinase enzyme. Glycogen synthesis is inhibited by glucagon and epinephrine through a covalent modification of the glycogen synthase phosphatase enzyme.

31 Glycogenolysis When the blood glucose concentration drops below about 5 mm, glycogen in the liver is degraded to free glucose which is released into the bloodstream. Glycogenolysis proceeds by a different route from glycogen biosynthesis. Glucose, as glucose-1-phosphate, is released from glycogen by the active form of the enzyme: glycogen phosphorylase.

32 Glycogenolysis H P H H H H H H H Glycogen Phosphorylase H H H non-reducing end of glycogen H H H H P H H H H H H H shortened glycogen Called a phosphorolysis reaction

33 Glycogenolysis When a shrinking α 1 4 glycogen strand approaches a branching point, a debranching enzyme transfers a portion of the α 1 6 branching chain and remakes an α 1 4 linkage. A second enzyme (an α 1 6 glucosidase) removes the remaining glucose residue from the branching point, and degradation continues by the phosphorylase.

34 H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H P H H H H P H H H H H H H H H H H H H H H H H H H H H H H H H phosphorylase H H H H H H H H H H H H H H H glycogenin H H H H H H H P H P H H H transferase H H H H H H H glucosidase H H H H H H H H H H H H H H H H H H H H H H H H glycogenin H H H glycogenin H H H H H H H P H P H H H H H H H H phosphorylase H H H H H Further chain cleavage H H H H H H H H H H H H H H H H H H H H H glycogenin

35 Glycogenolysis Glycogenolysis is stimulated by glucagon and epinephrine through covalent modification of a phosphorylase kinase enzyme.

36 Hormonal Regulation of Blood Glucose Insulin INACTIVE Glycogen Synthase kinase ACTIVE Glycogen Synthase phosphorylase ACTIVE Glycogen Synthase a Storage of glucose Glucagon, epinephrine ACTIVE Phosphorylase kinase ACTIVE Glycogen Phosphorylase a Release of glucose P

37 Inactive phosphorylase kinase Active phosphorylase kinase Inactive phosphorylase Active phosphorylase (Glucose)n (Glucose)n-1 + Glucose-1-P4 2- Glucagon, epinephrine Glycolysis Liver and Muscle Glucose-6-P4 2- Liver Bloodstream Glucose

38 Inactive G-protein 1x * Active G-protein Primary messenger (glucagon, epinephrine) binds to cell surface. Inactive adenyl cyclase Active adenyl cyclase Secondary messenger ATP 40x camp Inactive protein kinase 10x Active protein kinase Glucagon, epinephrine Inactive phosphorylase kinase 100x Inactive phosphorylase Active phosporylase kinase 1000x Active phosphorylase (Glucose)n 10,000x (Glucose)n-1 + Glucose-1-P4 2-

39 Amplification Mechanisms The glycogen cascade is one example of a general biochemical scheme for the use of relatively small numbers of secondary messenger molecules to produce general cellular and physiological responses.

40 primary messenger receptor proteins active relay molecule (second messenger) Inactive protein kinase Active protein kinase General Cellular Response Inactive protein kinase Active protein kinase Inactive protein kinase Active protein kinase Inactive protein Active protein

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