How Cells Release Chemical Energy. Chapter 8

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1 How Cells Release Chemical Energy Chapter 8

2 Impacts, Issues: When Mitochondria Spin Their Wheels More than forty disorders related to defective mitochondria are known (such as Friedreich s ataxia); many of those afflicted die young

3 8.1 Overview of Carbohydrate Breakdown Pathways Photoautotrophs make ATP during photosynthesis and use it to synthesize glucose and other carbohydrates Most organisms, including photoautotrophs, make ATP by breaking down glucose and other organic compounds

4 Comparison of the Main Pathways Aerobic respiration Aerobic metabolic pathways (using oxygen) are used by most eukaryotic cells Fermentation Anaerobic metabolic pathways (occur in the absence of oxygen) are used by prokaryotes and protists in anaerobic habitats

5 Comparison of the Main Pathways Aerobic respiration and fermentation both begin with glycolysis, which converts one molecule of glucose into two molecules of pyruvate After glycolysis, the two pathways diverge Fermentation is completed in the cytoplasm, yielding 2 ATP per glucose molecule Aerobic respiration is completed in mitochondria, yielding 36 ATP per glucose molecule

6 Comparison of the Main Pathways

7 Fig. 8-2b, p. 124

8 A Carbohydrate breakdown pathways start in the cytoplasm, with glycolysis. B Fermentation pathways are completed in the semifluid matrix of the cytoplasm. C In eukaryotes, aerobic respiration is completed inside mitochondria. Fig. 8-2b, p. 124

9 Animation: Where pathways start and finish

10 Overview of Aerobic Respiration Three stages Glycolysis Acetyl-CoA formation and Krebs cycle Electron transfer phosphorylation (ATP formation) C 6 H 12 O 6 (glucose) + O 2 (oxygen) CO 2 (carbon dioxide) + H 2 O (water) Coenzymes NADH and FADH 2 carry electrons and hydrogen

11 Overview of Aerobic Respiration

12 glucose 2 ATP ATP Glycolysis 4 ATP (2 net) 2 NADH 2 pyruvate ATP Cytoplasm A The first stage, glycolysis, occurs in the cell s cytoplasm. Enzymes convert a glucose molecule to 2 pyruvate for a net yield of 2 ATP. During the reactions, 2 NAD + pick up electrons and hydrogen ions, so 2 NADH form. oxygen Electron Transfer Phosphorylation Krebs Cycle 6 CO 2 2 ATP ATP 8 NADH, 2 FADH 2 ATP 32 ATP Mitochondrion B The second stage occurs in mitochondria. The 2 pyruvates are converted to acetyl CoA, which enters the Krebs cycle. CO 2 forms and leaves the cell. 2 ATP form. During the reactions, 8 NAD + and 2 FAD pick up electrons and hydrogen ions, so 8 NADH and 2 FADH 2 also form. C The third and final stage, electron transfer phosphorylation, occurs inside mitochondria. 10 NADH and 2 FADH 2 donate electrons and hydrogen ions to electron transfer chains. Electron fl ow through the chains sets up H+ gradients that drive ATP formation. Oxygen accepts electrons at the end of the chains. Fig. 8-3a, p. 125

13 glucose 2 ATP ATP Glycolysis 4 ATP (2 net) oxygen 2 NADH 2 pyruvate Krebs Cycle Electron Transfer Phosphorylation 8 NADH, 2 FADH 2 ATP 6 CO 2 2 ATP ATP 32 ATP ATP Cytoplasm A The first stage, glycolysis, occurs in the cell s cytoplasm. Enzymes convert a glucose molecule to 2 pyruvate for a net yield of 2 ATP. During the reactions, 2 NAD + pick up electrons and hydrogen ions, so 2 NADH form. Mitochondrion B The second stage occurs in mitochondria. The 2 pyruvates are converted to acetyl CoA, which enters the Krebs cycle. CO 2 forms and leaves the cell. 2 ATP form. During the reactions, 8 NAD + and 2 FAD pick up electrons and hydrogen ions, so 8 NADH and 2 FADH 2 also form. C The third and final stage, electron transfer phosphorylation, occurs inside mitochondria. 10 NADH and 2 FADH 2 donate electrons and hydrogen ions to electron transfer chains. Electron fl ow through the chains sets up H+ gradients that drive ATP formation. Oxygen accepts electrons at the end of the chains. Stepped Art Fig. 8-3a, p. 125

14 Fig. 8-3b, p. 125

15 Animation: Overview of aerobic respiration

16 8.1 Key Concepts: Energy From Carbohydrate Breakdown Various degradative pathways convert the chemical energy of glucose and other organic compounds to the chemical energy of ATP Aerobic respiration yields the most ATP from each glucose molecule; in eukaryotes, it is completed inside mitochondria

17 8.2 Glycolysis Glucose Breakdown Starts Glycolysis starts and ends in the cytoplasm of all prokaryotic and eukaryotic cells An energy investment of ATP starts glycolysis

18 Glycolysis Two ATP are used to split glucose and form 2 PGAL, each with one phosphate group Enzymes convert 2 PGAL to 2 PGA, forming 2 NADH Four ATP are formed by substrate-level phosphorylation (net 2 ATP)

19 Glycolysis

20 glycolysis occurs in the cytoplasm Animal Cell (eukaryotic) Plant Cell (eukaryotic) Bacterial Cell (prokaryotic) Fig. 8-4a, p. 126

21 Fig. 8-4a (1), p. 126

22 Fig. 8-4a (2), p. 126

23 Fig. 8-4b, p. 127

24 Fig. 8-4b (1), p. 127

25 Glycolysis glucose ATP ADP glucose-6-phosphate ATP ADP fructose-1,6-bisphosphate ATP-Requiring Steps A An enzyme transfers a phosphate group from ATP to glucose, forming glucose- 6-phosphate. B A phosphate group from a second ATP is transferred to the glucose- 6- phosphate. The resulting molecule is unstable, and it splits into two three- carbon molecules. The molecules are interconvertible, so we will call them both PGAL (phosphoglyceraldehyde). So far, two ATP have been invested in the reactions. Fig. 8-4b (1), p. 127

26 Fig. 8-4b (2), p. 127

27 2 PGAL 2 NAD P i NADH 2 ADP 2 ADP 2 PGA 2 PEP 2 pyruvate to second stage 2 reduced coenzymes ATP 2 ATP produced by substrate-level phosphorylation ATP 2 ATP produced by substrate-level phosphorylation Net 2 ATP + 2 NADH ATP-Generating Steps C Enzymes attach a phosphate to the two PGAL, and transfer two electrons and a hydrogen ion from each PGAL to NAD +. Two PGA (phosphoglycerate) and two NADH are the result. D Enzymes transfer a phosphate group from each PGA to ADP. Thus, two ATP have formed by substratelevel phosphorylation. The original energy investment of two ATP has now been recovered. E Enzymes transfer a phosphate group from each of two intermediates to ADP. Two more ATP have formed by substrate-level phosphorylation. Two molecules of pyruvate form at this last reaction step. F Summing up, glycolysis yields two NADH, two ATP (net), and two pyruvate for each glucose molecule. Fig. 8-4b (2), p. 127

28 Animation: Glycolysis

29 8.2 Key Concepts: Glycolysis Glycolysis is the first stage of aerobic respiration and of anaerobic routes such as fermentation pathways Enzymes of glycolysis convert glucose to pyruvate

30 8.3 Second Stage of Aerobic Respiration The second stage of aerobic respiration finishes breakdown of glucose that began in glycolysis Occurs in mitochondria Includes two stages: acetyl CoA formation and the Krebs cycle (each occurs twice in the breakdown of one glucose molecule)

31 Acetyl CoA Formation In the inner compartment of the mitochondrion, enzymes split pyruvate, forming acetyl CoA and CO 2 (which diffuses out of the cell) NADH is formed

32 The Krebs Cycle Krebs cycle A sequence of enzyme-mediated reactions that break down 1 acetyl CoA into 2 CO 2 Oxaloacetate is used and regenerated 3 NADH and 1 FADH 2 are formed 1 ATP is formed

33 Inside a Mitochondrion

34 Fig. 8-5a, p. 128

35 outer membrane (next to cytoplasm) inner membrane inner mitochondrial compartment outer mitochondrial compartment (in between the two membranes) Fig. 8-5a, p. 128

36 Fig. 8-5b, p. 128

37 glucose 2 pyruvate (glycolysis) OUTER COMPARTMENT 6 CO 2 INNER COMPARTMENT 2 acetyl-coa Krebs Cycle ATP NADH FADH 2 Breakdown of 2 pyruvate 6 CO 2 yields 2 ATP. Also, 10 coenzymes (8 NAD +, 2 FAD) are reduced. The coenzymes carry hydrogen ions and electrons to the third stage of aerobic respiration. Fig. 8-5b, p. 128

38 Animation: Functional zones in mitochondria

39 Acetyl CoA Formation and the Krebs Cycle

40 A An enzyme splits a pyruvate molecule into a two-carbon acetyl group and CO 2. Coenzyme A binds the acetyl group (forming acetyl CoA). NAD + combines with released hydrogen ions and electrons, forming NADH. Acetyl CoA Formation coenzyme A pyruvate NAD + NADH CO 2 B The Krebs cycle starts as one carbon atom is transferred from acetyl CoA to oxaloacetate. Citrate forms, and coenzyme A is regenerated. C A carbon atom is removed from an intermediate and leaves the cell as CO 2. NAD + combines with released hydrogen ions and electrons, forming NADH. D A carbon atom is removed from another intermediate and leaves the cell as CO 2, and another NADH forms. coenzyme A CO 2 NAD + NADH citrate CO 2 NAD + NADH acetyl CoA ADP + P i Krebs Cycle ATP oxaloacetate FAD NADH NAD + FADH 2 H The final steps of the Krebs cycle regenerate oxaloacetate. G NAD + combines with hydrogen ions and electrons, forming NADH. F The coenzyme FAD combines with hydrogen ions and electrons, forming FADH 2. E One ATP forms by substrate-level phosphorylation. Pyruvate s three carbon atoms have now exited the cell, in CO 2. Stepped Art Fig. 8-6, p. 129

41 Animation: The Krebs Cycle - details

42 8.4 Aerobic Respiration s Big Energy Payoff Many ATP are formed during the third and final stage of aerobic respiration Electron transfer phosphorylation Occurs in mitochondria Results in attachment of phosphate to ADP to form ATP

43 Electron Transfer Phosphorylation Coenzymes NADH and FADH 2 donate electrons and H + to electron transfer chains Active transport forms a H + concentration gradient in the outer mitochondrial compartment H + follows its gradient through ATP synthase, which attaches a phosphate to ADP Finally, oxygen accepts electrons and combines with H +, forming water

44 Electron Transfer Phosphorylation

45 Summary: The Energy Harvest Typically, the breakdown of one glucose molecule yields 36 ATP Glycolysis: 2 ATP Acetyl CoA formation and Krebs cycle: 2 ATP Electron transfer phosphorylation: 32 ATP

46 Summary: Aerobic Respiration

47 Animation: Third-stage reactions

48 Key Concepts: How Aerobic Respiration Ends The final stages of aerobic respiration break down pyruvate to CO 2 Many coenzymes that become reduced deliver electrons and hydrogen ions to electron transfer chains; energy released by electrons flowing through the chains is captured in ATP Oxygen accepts electrons at ends of the chains

49 8.5 Anaerobic Energy-Releasing Pathways Fermentation pathways break down carbohydrates without using oxygen The final steps in these pathways regenerate NAD + but do not produce ATP

50 Fermentation Pathways Glycolysis is the first stage of fermentation Forms 2 pyruvate, 2 NADH, and 2 ATP Pyruvate is converted to other molecules, but is not fully broken down to CO 2 and water Regenerates NAD + but doesn t produce ATP Provides enough energy for some single-celled anaerobic species

51 Two Pathways of Fermentation Alcoholic fermentation Pyruvate is split into acetaldehyde and CO 2 Acetaldehyde receives electrons and hydrogen from NADH, forming NAD + and ethanol Lactate fermentation Pyruvate receives electrons and hydrogen from NADH, forming NAD + and lactate

52 Two Pathways of Fermentation

53 Fig. 8-9a, p. 132

54 Glycolysis glucose 2 NAD + 2 ATP 2 NADH 4 ATP pyruvate Alcoholic Fermentation 2 CO 2 acetaldehyde 2 NADH ethanol 2 NAD + Fig. 8-9a, p. 132

55 Fig. 8-9b, p. 132

56 Glycolysis glucose 2 NAD + 2 ATP 2 NADH 4 ATP pyruvate Lactate Fermentation 2 NADH 2 NAD + lactate Fig. 8-9b, p. 132

57 Glycolysis glucose 2 NAD + 2 ATP 2 NADH 4 ATP pyruvate Glycolysis glucose 2 NAD + 2 ATP 2 NADH 4 ATP pyruvate Alcoholic Fermentation 2 CO 2 acetaldehyde 2 NADH Lactate Fermentation 2 NADH 2 NAD + 2 NAD + ethanol lactate Stepped Art Fig. 8-9, p. 132

58 Animation: Fermentation pathways

59 Alcoholic Fermentation

60 Fig. 8-10a, p. 133

61 Fig. 8-10b, p. 133

62 Fig. 8-10c, p. 133

63 8.6 The Twitchers Slow-twitch muscle fibers ( red muscles) make ATP by aerobic respiration Have many mitochondria Dominate in prolonged activity Fast-twitch muscle fibers ( white muscles) make ATP by lactate fermentation Have few mitochondria and no myoglobin Sustain short bursts of activity

64 Sprinters and Lactate Fermentation

65 Fig. 8-11b, p. 133

66 Key Concepts: How Anaerobic Pathways End Fermentation pathways start with glycolysis Substances other than oxygen accept electrons at the end of the pathways Compared with aerobic respiration, the net yield of ATP from fermentation is small

67 8.7 Alternative Energy Sources in the Body Pathways that break down molecules other than carbohydrates also keep organisms alive In humans and other mammals, the entrance of glucose and other organic compounds into an energy-releasing pathway depends on the kinds and proportions of carbohydrates, fats and proteins in the diet

68 The Fate of Glucose at Mealtime and Between Meals When blood glucose concentration rises, the pancreas increases insulin secretion Cells take up glucose faster, more ATP is formed, glycogen and fatty-acid production increases When blood glucose concentration falls, the pancreas increases glucagon secretion Stored glycogen is converted to glucose

69 Energy From Fats About 78% of an adult s energy reserves is stored in fat (mostly triglycerides) Enzymes cleave fats into glycerol and fatty acids Glycerol products enter glycolysis Fatty acid products enter the Krebs cycle Compared to carbohydrates, fatty acid breakdown yields more ATP per carbon atom

70 Energy from Proteins Enzymes split dietary proteins into amino acid subunits, which enter the bloodstream Used to build proteins or other molecules Excess amino acids are broken down into ammonia (NH 3 ) and various products that can enter the Krebs cycle

71 Alternative Energy Sources in the Human Body

72 FOOD FATS COMPLEX CARBOHYDRATES PROTEINS fatty acids glycerol glucose, other simple sugars amino acids acetyl CoA PGAL acetyl CoA Glycolysis NADH pyruvate Krebs Cycle oxaloacetate or another intermediate of the Krebs NADH, FADH 2 Electron Transfer Phosphorylation Fig. 8-12, p. 135

73 FOOD FATS COMPLEX CARBOHYDRATES PROTEINS fatty acids glycerol glucose, other simple sugars amino acids acetyl CoA PGAL acetyl CoA Glycolysis NADH pyruvate Krebs Cycle oxaloacetate or another intermediate of the Krebs NADH, FADH 2 Electron Transfer Phosphorylation Stepped Art Fig. 8-12, p. 135

74 Animation: Alternative energy sources

75 8.7 Key Concepts: Other Metabolic Pathways Molecules other than glucose are common energy sources Different pathways convert lipids and proteins to substances that may enter glycolysis or the Krebs cycle

76 8.8 Reflections on Life s Unity Life s diversity and continuity arise from unity at the level of molecules and energy Energy inputs drive the organization of molecules into cells (one-way flow of energy) Energy from the sun sustains life s organization Photosynthesizers use energy from the sun to feed themselves and other forms of life Aerobic respiration balances photosynthesis

77 Links Between Photosynthesis and Aerobic Respiration

78 energy in (mainly from sunlight) Photosynthesis glucose, oxygen Aerobic Respiration energy out (ATP, heat) carbon dioxide, water energy out (ATP, heat) Fig. 8-13, p. 136

79 Animation: Electron transfer phosphorylation

80 Animation: Electron transfer system and oxidative phosphorylation

81 Animation: Krebs cycle overview

82 Animation: Links with photosynthesis

83 Animation: Recreating the reactions of glycolysis

84 Video: When mitochondria spin their wheels

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