1. LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson © 2011 Pearson Education, Inc. Lectures by Erin Barley Kathleen Fitzpatrick Cellular Respiration and Fermentation Chapter 9

2. Overview: Life Is Work Living cells require energy from outside sources Some animals, such as the chimpanzee, obtain energy by eating plants, and some animals feed on other organisms that eat plants © 2011 Pearson Education, Inc.

3. Figure 9.1

4. Energy flows into an ecosystem as sunlight and leaves as heat Photosynthesis generates O 2 and organic molecules, which are used in cellular respiration Cells use chemical energy stored in organic molecules to regenerate ATP, which powers work © 2011 Pearson Education, Inc.

5. Figure 9.2 Light energy ECOSYSTEM Photosynthesis in chloroplasts Cellular respiration in mitochondria CO 2  H 2 O  O 2 Organic molecules ATP powers most cellular work ATP Heat energy

6. Concept 9.1: Catabolic pathways yield energy by oxidizing organic fuels Several processes are central to cellular respiration and related pathways © 2011 Pearson Education, Inc.

7. Catabolic Pathways and Production of ATP The breakdown of organic molecules is exergonic Fermentation is a partial degradation of sugars that occurs without O 2 Aerobic respiration consumes organic molecules and O 2 and yields ATP Anaerobic respiration is similar to aerobic respiration but consumes compounds other than O 2 © 2011 Pearson Education, Inc.

8. Cellular respiration includes both aerobic and anaerobic respiration but is often used to refer to aerobic respiration Although carbohydrates, fats, and proteins are all consumed as fuel, it is helpful to trace cellular respiration with the sugar glucose C 6 H 12 O 6 + 6 O 2  6 CO 2 + 6 H 2 O + Energy (ATP + heat) © 2011 Pearson Education, Inc.

9. Redox Reactions: Oxidation and Reduction The transfer of electrons during chemical reactions releases energy stored in organic molecules This released energy is ultimately used to synthesize ATP © 2011 Pearson Education, Inc.

10. The Principle of Redox Chemical reactions that transfer electrons between reactants are called oxidation-reduction reactions, or redox reactions In oxidation, a substance loses electrons, or is oxidized In reduction, a substance gains electrons, or is reduced (the amount of positive charge is reduced) © 2011 Pearson Education, Inc.

11. Figure 9.UN01 becomes oxidized (loses electron) becomes reduced (gains electron)

12. Figure 9.UN02 becomes oxidized becomes reduced

13. The electron donor is called the reducing agent The electron receptor is called the oxidizing agent Some redox reactions do not transfer electrons but change the electron sharing in covalent bonds An example is the reaction between methane and O 2 © 2011 Pearson Education, Inc.

14. Figure 9.3 Reactants Products Energy Water Carbon dioxide Methane (reducing agent) Oxygen (oxidizing agent) becomes oxidized becomes reduced

15. Oxidation of Organic Fuel Molecules During Cellular Respiration During cellular respiration, the fuel (such as glucose) is oxidized, and O 2 is reduced © 2011 Pearson Education, Inc.

16. Figure 9.UN03 becomes oxidized becomes reduced

17. Stepwise Energy Harvest via NAD + and the Electron Transport Chain In cellular respiration, glucose and other organic molecules are broken down in a series of steps Electrons from organic compounds are usually first transferred to NAD +, a coenzyme As an electron acceptor, NAD + functions as an oxidizing agent during cellular respiration Each NADH (the reduced form of NAD + ) represents stored energy that is tapped to synthesize ATP © 2011 Pearson Education, Inc.

18. Figure 9.4 Nicotinamide (oxidized form) NAD  (from food) Dehydrogenase Reduction of NAD  Oxidation of NADH Nicotinamide (reduced form) NADH

19. Figure 9.UN04 Dehydrogenase

20. NADH passes the electrons to the electron transport chain Unlike an uncontrolled reaction, the electron transport chain passes electrons in a series of steps instead of one explosive reaction O 2 pulls electrons down the chain in an energy- yielding tumble The energy yielded is used to regenerate ATP © 2011 Pearson Education, Inc.

21. Figure 9.5 (a) Uncontrolled reaction (b) Cellular respiration Explosive release of heat and light energy Controlled release of energy for synthesis of ATP Free energy, G H 2  1 / 2 O 2 2 H  1 / 2 O 2 H2OH2O H2OH2O 2 H +  2 e  2 e  2 H + ATP Electron transport chain (from food via NADH)

22. The Stages of Cellular Respiration: A Preview Harvesting of energy from glucose has three stages –Glycolysis (breaks down glucose into two molecules of pyruvate) –The citric acid cycle (completes the breakdown of glucose) –Oxidative phosphorylation (accounts for most of the ATP synthesis) © 2011 Pearson Education, Inc.

23. Figure 9.UN05 Glycolysis (color-coded teal throughout the chapter) 1. Pyruvate oxidation and the citric acid cycle (color-coded salmon) 2. Oxidative phosphorylation: electron transport and chemiosmosis (color-coded violet) 3.

24. Figure 9.6-1 Electrons carried via NADH Glycolysis Glucose Pyruvate CYTOSOL MITOCHONDRION ATP Substrate-level phosphorylation

25. Figure 9.6-2 Electrons carried via NADH Electrons carried via NADH and FADH 2 Citric acid cycle Pyruvate oxidation Acetyl CoA Glycolysis Glucose Pyruvate CYTOSOL MITOCHONDRION ATP Substrate-level phosphorylation

26. Figure 9.6-3 Electrons carried via NADH Electrons carried via NADH and FADH 2 Citric acid cycle Pyruvate oxidation Acetyl CoA Glycolysis Glucose Pyruvate Oxidative phosphorylation: electron transport and chemiosmosis CYTOSOL MITOCHONDRION ATP Substrate-level phosphorylation Oxidative phosphorylation

27. The process that generates most of the ATP is called oxidative phosphorylation because it is powered by redox reactions © 2011 Pearson Education, Inc.

28. BioFlix: Cellular Respiration

29. Oxidative phosphorylation accounts for almost 90% of the ATP generated by cellular respiration A smaller amount of ATP is formed in glycolysis and the citric acid cycle by substrate-level phosphorylation For each molecule of glucose degraded to CO 2 and water by respiration, the cell makes up to 32 molecules of ATP © 2011 Pearson Education, Inc.

30. Figure 9.7 Substrate Product ADP P ATP Enzyme

31. Concept 9.2: Glycolysis harvests chemical energy by oxidizing glucose to pyruvate Glycolysis (“splitting of sugar”) breaks down glucose into two molecules of pyruvate Glycolysis occurs in the cytoplasm and has two major phases –Energy investment phase –Energy payoff phase Glycolysis occurs whether or not O 2 is present © 2011 Pearson Education, Inc.

32. Figure 9.8 Energy Investment Phase Glucose 2 ADP  2 P 4 ADP  4 P Energy Payoff Phase 2 NAD +  4 e   4 H + 2 Pyruvate  2 H 2 O 2 ATP used 4 ATP formed 2 NADH  2 H + Net Glucose 2 Pyruvate  2 H 2 O 2 ATP 2 NADH  2 H + 2 NAD +  4 e   4 H + 4 ATP formed  2 ATP used

33. Figure 9.9-1 Glycolysis: Energy Investment Phase ATP Glucose Glucose 6-phosphate ADP Hexokinase 1

34. Figure 9.9-2 Glycolysis: Energy Investment Phase ATP Glucose Glucose 6-phosphate Fructose 6-phosphate ADP Hexokinase Phosphogluco- isomerase 12

35. Figure 9.9-3 Glycolysis: Energy Investment Phase ATP Glucose Glucose 6-phosphate Fructose 6-phosphate Fructose 1,6-bisphosphate ADP Hexokinase Phosphogluco- isomerase Phospho- fructokinase 123

36. Figure 9.9-4 Glycolysis: Energy Investment Phase ATP Glucose Glucose 6-phosphate Fructose 6-phosphate Fructose 1,6-bisphosphate Dihydroxyacetone phosphate Glyceraldehyde 3-phosphate To step 6 ADP Hexokinase Phosphogluco- isomerase Phospho- fructokinase Aldolase Isomerase 12345

37. Figure 9.9-5 Glycolysis: Energy Payoff Phase 2 NADH 2 NAD  + 2 H  2 P i 1,3-Bisphospho- glycerate 6 Triose phosphate dehydrogenase

38. Figure 9.9-6 Glycolysis: Energy Payoff Phase 2 ATP 2 NADH 2 NAD  + 2 H  2 P i 2 ADP 1,3-Bisphospho- glycerate 3-Phospho- glycerate 2 Phospho- glycerokinase 6 7 Triose phosphate dehydrogenase

39. Figure 9.9-7 Glycolysis: Energy Payoff Phase 2 ATP 2 NADH 2 NAD  + 2 H  2 P i 2 ADP 1,3-Bisphospho- glycerate 3-Phospho- glycerate 2-Phospho- glycerate 2 2 Phospho- glycerokinase Phospho- glyceromutase 6 7 8 Triose phosphate dehydrogenase

40. Figure 9.9-8 Glycolysis: Energy Payoff Phase 2 ATP 2 NADH 2 NAD  + 2 H  2 P i 2 ADP 1,3-Bisphospho- glycerate 3-Phospho- glycerate 2-Phospho- glycerate Phosphoenol- pyruvate (PEP) 2 22 2 H 2 O Phospho- glycerokinase Phospho- glyceromutase Enolase 6 7 89 Triose phosphate dehydrogenase

41. Figure 9.9-9 Glycolysis: Energy Payoff Phase 2 ATP 2 NADH 2 NAD  + 2 H  2 P i 2 ADP 1,3-Bisphospho- glycerate 3-Phospho- glycerate 2-Phospho- glycerate Phosphoenol- pyruvate (PEP) Pyruvate 2 ADP 2 22 2 H 2 O Phospho- glycerokinase Phospho- glyceromutase EnolasePyruvate kinase 6 7 8910 Triose phosphate dehydrogenase

42. Figure 9.9a Glycolysis: Energy Investment Phase ATP Glucose Glucose 6-phosphate ADP Hexokinase 1 Fructose 6-phosphate Phosphogluco- isomerase 2

43. Figure 9.9b Glycolysis: Energy Investment Phase ATP Fructose 6-phosphate ADP 3 Fructose 1,6-bisphosphate Phospho- fructokinase 45 Aldolase Dihydroxyacetone phosphate Glyceraldehyde 3-phosphate To step 6 Isomerase

44. Figure 9.9c Glycolysis: Energy Payoff Phase 2 NADH 2 ATP 2 ADP 2 2 2 NAD  + 2 H  2 P i 3-Phospho- glycerate 1,3-Bisphospho- glycerate Triose phosphate dehydrogenase Phospho- glycerokinase 67

45. Figure 9.9d Glycolysis: Energy Payoff Phase 2 ATP 2 ADP 2 2 22 2 H 2 O Pyruvate Phosphoenol- pyruvate (PEP) 2-Phospho- glycerate 3-Phospho- glycerate 8 9 10 Phospho- glyceromutase Enolase Pyruvate kinase

46. Concept 9.3: After pyruvate is oxidized, the citric acid cycle completes the energy- yielding oxidation of organic molecules In the presence of O 2, pyruvate enters the mitochondrion (in eukaryotic cells) where the oxidation of glucose is completed © 2011 Pearson Education, Inc.

47. Oxidation of Pyruvate to Acetyl CoA Before the citric acid cycle can begin, pyruvate must be converted to acetyl Coenzyme A (acetyl CoA), which links glycolysis to the citric acid cycle This step is carried out by a multienzyme complex that catalyses three reactions © 2011 Pearson Education, Inc.

48. Figure 9.10 Pyruvate Transport protein CYTOSOL MITOCHONDRION CO 2 Coenzyme A NAD  + H  NADH Acetyl CoA 123

49. The citric acid cycle, also called the Krebs cycle, completes the break down of pyruvate to CO 2 The cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 FADH 2 per turn © 2011 Pearson Education, Inc. The Citric Acid Cycle

50. Figure 9.11 Pyruvate NAD  NADH + H  Acetyl CoA CO 2 CoA 2 CO 2 ADP + P i FADH 2 FAD ATP 3 NADH 3 NAD  Citric acid cycle + 3 H 

51. The citric acid cycle has eight steps, each catalyzed by a specific enzyme The acetyl group of acetyl CoA joins the cycle by combining with oxaloacetate, forming citrate The next seven steps decompose the citrate back to oxaloacetate, making the process a cycle The NADH and FADH 2 produced by the cycle relay electrons extracted from food to the electron transport chain © 2011 Pearson Education, Inc.

52. Figure 9.12-1 1 Acetyl CoA Citrate Citric acid cycle CoA-SH Oxaloacetate

53. Figure 9.12-2 1 Acetyl CoA Citrate Isocitrate Citric acid cycle H2OH2O 2 CoA-SH Oxaloacetate

54. Figure 9.12-3 1 Acetyl CoA Citrate Isocitrate  -Ketoglutarate Citric acid cycle NADH + H  NAD  H2OH2O 32 CoA-SH CO2CO2 Oxaloacetate

55. Figure 9.12-4 1 Acetyl CoA Citrate Isocitrate  -Ketoglutarate Succinyl CoA Citric acid cycle NADH + H  NAD  H2OH2O 324 CoA-SH CO2CO2 CO2CO2 Oxaloacetate

56. Figure 9.12-5 1 Acetyl CoA Citrate Isocitrate  -Ketoglutarate Succinyl CoA Succinate Citric acid cycle NADH ATP + H  NAD  H2OH2O ADP GTPGDP P i 3245 CoA-SH CO2CO2 CO2CO2 Oxaloacetate

57. Figure 9.12-6 1 Acetyl CoA Citrate Isocitrate  -Ketoglutarate Succinyl CoA Succinate Fumarate Citric acid cycle NADH FADH 2 ATP + H  NAD  H2OH2O ADP GTPGDP P i FAD 32456 CoA-SH CO2CO2 CO2CO2 Oxaloacetate

58. Figure 9.12-7 1 Acetyl CoA Citrate Isocitrate  -Ketoglutarate Succinyl CoA Succinate Fumarate Malate Citric acid cycle NADH FADH 2 ATP + H  NAD  H2OH2O H2OH2O ADP GTPGDP P i FAD 324567 CoA-SH CO2CO2 CO2CO2 Oxaloacetate

59. Figure 9.12-8 NADH 1 Acetyl CoA Citrate Isocitrate  -Ketoglutarate Succinyl CoA Succinate Fumarate Malate Citric acid cycle NAD  NADH FADH 2 ATP + H  NAD  H2OH2O H2OH2O ADP GTPGDP P i FAD 3245678 CoA-SH CO2CO2 CO2CO2 Oxaloacetate

60. Figure 9.12a Acetyl CoA Oxaloacetate Citrate Isocitrate H2OH2O CoA-SH 1 2

61. Figure 9.12b Isocitrate  -Ketoglutarate Succinyl CoA NADH NAD  + H  CoA-SH CO2CO2 CO2CO2 34 + H 

62. Figure 9.12c Fumarate FADH 2 CoA-SH 6 Succinate Succinyl CoA FAD ADP GTPGDP P i ATP 5

63. Figure 9.12d Oxaloacetate 8 Malate Fumarate H2OH2O NADH NAD  + H  7

64. Concept 9.4: During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis Following glycolysis and the citric acid cycle, NADH and FADH 2 account for most of the energy extracted from food These two electron carriers donate electrons to the electron transport chain, which powers ATP synthesis via oxidative phosphorylation © 2011 Pearson Education, Inc.

65. The Pathway of Electron Transport The electron transport chain is in the inner membrane (cristae) of the mitochondrion Most of the chain’s components are proteins, which exist in multiprotein complexes The carriers alternate reduced and oxidized states as they accept and donate electrons Electrons drop in free energy as they go down the chain and are finally passed to O 2, forming H 2 O © 2011 Pearson Education, Inc.

66. Figure 9.13 NADH FADH 2 2 H  + 1 / 2 O 2 2 e  H2OH2O NAD  Multiprotein complexes (originally from NADH or FADH 2 ) I II III IV 50 40 30 20 10 0 Free energy (G) relative to O 2 (kcal/mol) FMN Fe  S FAD Q Cyt b Cyt c 1 Cyt c Cyt a Cyt a 3 Fe  S

67. Electrons are transferred from NADH or FADH 2 to the electron transport chain Electrons are passed through a number of proteins including cytochromes (each with an iron atom) to O 2 The electron transport chain generates no ATP directly It breaks the large free-energy drop from food to O 2 into smaller steps that release energy in manageable amounts © 2011 Pearson Education, Inc.

68. Chemiosmosis: The Energy-Coupling Mechanism Electron transfer in the electron transport chain causes proteins to pump H + from the mitochondrial matrix to the intermembrane space H + then moves back across the membrane, passing through the proton, ATP synthase ATP synthase uses the exergonic flow of H + to drive phosphorylation of ATP This is an example of chemiosmosis, the use of energy in a H + gradient to drive cellular work © 2011 Pearson Education, Inc.

69. Figure 9.14 INTERMEMBRANE SPACE Rotor Stator HH Internal rod Catalytic knob ADP + P i ATP MITOCHONDRIAL MATRIX

70. Figure 9.15 Protein complex of electron carriers (carrying electrons from food) Electron transport chain Oxidative phosphorylation Chemiosmosis ATP synth- ase I II III IV Q Cyt c FAD FADH 2 NADH ADP  P i NAD  HH 2 H  + 1 / 2 O 2 HH HH HH 21 HH H2OH2O ATP

71. The energy stored in a H + gradient across a membrane couples the redox reactions of the electron transport chain to ATP synthesis The H + gradient is referred to as a proton- motive force, emphasizing its capacity to do work © 2011 Pearson Education, Inc.

72. An Accounting of ATP Production by Cellular Respiration During cellular respiration, most energy flows in this sequence: glucose  NADH  electron transport chain  proton-motive force  ATP About 34% of the energy in a glucose molecule is transferred to ATP during cellular respiration, making about 32 ATP There are several reasons why the number of ATP is not known exactly © 2011 Pearson Education, Inc.

73. Figure 9.16 Electron shuttles span membrane MITOCHONDRION 2 NADH 6 NADH 2 FADH 2 or  2 ATP  about 26 or 28 ATP Glycolysis Glucose 2 Pyruvate Pyruvate oxidation 2 Acetyl CoA Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis CYTOSOL Maximum per glucose: About 30 or 32 ATP

74. Concept 9.5: Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen Most cellular respiration requires O 2 to produce ATP Without O 2, the electron transport chain will cease to operate In that case, glycolysis couples with fermentation or anaerobic respiration to produce ATP © 2011 Pearson Education, Inc.

75. Anaerobic respiration uses an electron transport chain with a final electron acceptor other than O 2, for example sulfate Fermentation uses substrate-level phosphorylation instead of an electron transport chain to generate ATP © 2011 Pearson Education, Inc.

76. Types of Fermentation Fermentation consists of glycolysis plus reactions that regenerate NAD +, which can be reused by glycolysis Two common types are alcohol fermentation and lactic acid fermentation © 2011 Pearson Education, Inc.

77. In alcohol fermentation, pyruvate is converted to ethanol in two steps, with the first releasing CO 2 Alcohol fermentation by yeast is used in brewing, winemaking, and baking © 2011 Pearson Education, Inc.

78. Animation: Fermentation Overview Right-click slide / select “Play”

79. Figure 9.17 2 ADP 2 ATP Glucose Glycolysis 2 Pyruvate 2 CO 2 2  2 NADH 2 Ethanol 2 Acetaldehyde (a) Alcohol fermentation (b) Lactic acid fermentation 2 Lactate 2 Pyruvate 2 NADH Glucose Glycolysis 2 ATP 2 ADP  2 P i NAD 2 H   2 P i 2 NAD    2 H 

80. 2 ADP  2 P i 2 ATP Glucose Glycolysis 2 Pyruvate 2 CO 2 2 NAD  2 NADH 2 Ethanol 2 Acetaldehyde (a) Alcohol fermentation  2 H  Figure 9.17a

81. In lactic acid fermentation, pyruvate is reduced to NADH, forming lactate as an end product, with no release of CO 2 Lactic acid fermentation by some fungi and bacteria is used to make cheese and yogurt Human muscle cells use lactic acid fermentation to generate ATP when O 2 is scarce © 2011 Pearson Education, Inc.

82. (b) Lactic acid fermentation 2 Lactate 2 Pyruvate 2 NADH Glucose Glycolysis 2 ADP  2 P i 2 ATP 2 NAD   2 H  Figure 9.17b

83. Comparing Fermentation with Anaerobic and Aerobic Respiration All use glycolysis (net ATP = 2) to oxidize glucose and harvest chemical energy of food In all three, NAD + is the oxidizing agent that accepts electrons during glycolysis The processes have different final electron acceptors: an organic molecule (such as pyruvate or acetaldehyde) in fermentation and O 2 in cellular respiration Cellular respiration produces 32 ATP per glucose molecule; fermentation produces 2 ATP per glucose molecule © 2011 Pearson Education, Inc.

84. Obligate anaerobes carry out fermentation or anaerobic respiration and cannot survive in the presence of O 2 Yeast and many bacteria are facultative anaerobes, meaning that they can survive using either fermentation or cellular respiration In a facultative anaerobe, pyruvate is a fork in the metabolic road that leads to two alternative catabolic routes © 2011 Pearson Education, Inc.

85. Figure 9.18 Glucose CYTOSOL Glycolysis Pyruvate No O 2 present: Fermentation O 2 present: Aerobic cellular respiration Ethanol, lactate, or other products Acetyl CoA MITOCHONDRION Citric acid cycle

86. The Evolutionary Significance of Glycolysis Ancient prokaryotes are thought to have used glycolysis long before there was oxygen in the atmosphere Very little O 2 was available in the atmosphere until about 2.7 billion years ago, so early prokaryotes likely used only glycolysis to generate ATP Glycolysis is a very ancient process © 2011 Pearson Education, Inc.

87. Concept 9.6: Glycolysis and the citric acid cycle connect to many other metabolic pathways Gycolysis and the citric acid cycle are major intersections to various catabolic and anabolic pathways © 2011 Pearson Education, Inc.

88. The Versatility of Catabolism Catabolic pathways funnel electrons from many kinds of organic molecules into cellular respiration Glycolysis accepts a wide range of carbohydrates Proteins must be digested to amino acids; amino groups can feed glycolysis or the citric acid cycle © 2011 Pearson Education, Inc.

89. Fats are digested to glycerol (used in glycolysis) and fatty acids (used in generating acetyl CoA) Fatty acids are broken down by beta oxidation and yield acetyl CoA An oxidized gram of fat produces more than twice as much ATP as an oxidized gram of carbohydrate © 2011 Pearson Education, Inc.

90. Figure 9.19 Carbohydrates Proteins Fatty acids Amino acids Sugars Fats Glycerol Glycolysis Glucose Glyceraldehyde 3- P NH 3 Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation

91. Biosynthesis (Anabolic Pathways) The body uses small molecules to build other substances These small molecules may come directly from food, from glycolysis, or from the citric acid cycle © 2011 Pearson Education, Inc.

92. Regulation of Cellular Respiration via Feedback Mechanisms Feedback inhibition is the most common mechanism for control If ATP concentration begins to drop, respiration speeds up; when there is plenty of ATP, respiration slows down Control of catabolism is based mainly on regulating the activity of enzymes at strategic points in the catabolic pathway © 2011 Pearson Education, Inc.

93. Figure 9.20 Phosphofructokinase Glucose Glycolysis AMP Stimulates    Fructose 6-phosphate Fructose 1,6-bisphosphate Pyruvate Inhibits ATPCitrate Citric acid cycle Oxidative phosphorylation Acetyl CoA

94. Figure 9.UN06 Inputs Outputs Glucose Glycolysis 2 Pyruvate  2 ATP  2 NADH

95. Figure 9.UN07 Inputs Outputs 2 Pyruvate2 Acetyl CoA 2 Oxaloacetate Citric acid cycle 2 2 6 8 ATP NADH FADH 2 CO 2

96. Figure 9.UN08 Protein complex of electron carriers (carrying electrons from food) INTERMEMBRANE SPACE MITOCHONDRIAL MATRIX HH HH HH 2 H  + 1 / 2 O 2 H2OH2O NAD  FADH 2 FAD Q NADH I II III IV Cyt c

97. Figure 9.UN09 INTER- MEMBRANE SPACE HH ADP + P i MITO- CHONDRIAL MATRIX ATP synthase HH ATP

98. Figure 9.UN10 Time pH difference across membrane

99. Figure 9.UN11