7 Cellular Respiration and Fermentation

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1 CAMPBELL BIOLOGY IN FOCUS URRY CAIN WASSERMAN MINORSKY REECE 7 Cellular Respiration and Fermentation Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge, Simon Fraser University SECOND EDITION

2 Life Is Work Living cells require energy from outside sources Some animals, such as the giraffe, obtain energy by eating plants, and some animals feed on other organisms that eat plants

3 Figure 7.1 How Do These Leaves Power the Work of Life for This Giraffe?

4 Life Is Work, Continued Energy flows into an ecosystem as sunlight and leaves as heat Photosynthesis generates and organic molecules, which are used as fuel for cellular respiration Cells use chemical energy stored in organic molecules to regenerate ATP, which powers work

5 Animation: Carbon Cycle

6 Figure 7.2 Energy Flow and Chemical Recycling in Ecosystems

7 Concept 7.1: Catabolic pathways yield energy by oxidizing organic fuels Catabolic pathways involving electron transfer are central processes to cellular respiration

8 Catabolic Pathways and Production of ATP The breakdown of organic molecules is exergonic Fermentation is a partial degradation of sugars that occurs without Aerobic respiration consumes organic molecules and and yields ATP Anaerobic respiration is similar to aerobic respiration but consumes compounds other than

9 Catabolic Pathways and Production of ATP, Continued Cellular respiration includes both aerobic and anaerobic processes 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

10 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

11 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)

12 Figure 7.UN01 In-Text Figure, NaCl Redox Reaction, p. 142

13 Figure 7.UN02 In-Text Figure, Generalized Redox Reaction, p. 142

14 The Principle of Redox, Continued The electron donor is called the reducing agent The electron acceptor 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

15 Figure 7.3 Methane Combustion as an Energy- Yielding Redox Reaction

16 The Principle of Redox, Continued-1 Redox reactions that move electrons closer to electronegative atoms, like oxygen, release chemical energy that can be put to work

17 Oxidation of Organic Fuel Molecules During Cellular Respiration During cellular respiration, fuel (such as glucose) is oxidized, and is reduced Organic molecules with an abundance of hydrogen, like carbohydrates and fats, are excellent fuels As hydrogen (with its electron) is transferred to oxygen, energy is released that can be used in ATP synthesis

18 Figure 7.UN03 In-Text Figure, Cellular Respiration Redox Reaction, p. 143

19 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, a coenzyme As an electron acceptor, functions as an oxidizing agent during cellular respiration Each NADH (the reduced form of ) represents stored energy that is tapped to synthesize ATP

20 Stepwise Energy Harvest via NAD + and the Electron Transport Chain, Continued Enzymes called dehydrogenases facilitate the transfer of two electrons and one hydrogen ion to One hydrogen ion is released in this process

21 Figure 7.4 NAD + as an Electron Shuttle

22 Figure NAD + as an Electron Shuttle (Part 1: NAD + Structure)

23 Figure NAD + as an Electron Shuttle (Part 2: NAD + Reaction)

24 Figure 7.UN04 In-Text Figure, Dehydrogenase Reaction, p. 144

25 Stepwise Energy Harvest via NAD + and the Electron Transport Chain, Continued-1 NADH passes the electrons to the electron transport chain Electrons are passed to increasingly electronegative carrier molecules down the chain through a series of redox reactions Electron transfer to oxygen occurs in a series of energy-releasing steps instead of one explosive reaction The energy yielded is used to regenerate ATP

26 Figure 7.5 An Introduction to Electron Transport Chains

27 The Stages of Cellular Respiration: A Preview Harvesting of energy from glucose has three stages Glycolysis breaks down glucose into two molecules of pyruvate in the cytosol Pyruvate oxidation and the citric acid cycle completes the breakdown of glucose in the mitochondrial matrix Oxidative phosphorylation accounts for most of the ATP synthesis and occurs in the inner membrane of the mitochondria

28 Animation: Cellular Respiration

29 Figure 7.UN05 In-Text Figure, Color Code for Stages of Cellular Respiration, p GLYCOLYSIS (color-coded blue throughout the chapter) 2. PYRUVATE OXIDATION and the CITRIC ACID CYCLE (color-coded orange) 3. OXIDATIVE PHOSPHORYLATION: Electron transport and chemiosmosis (color-coded purple)

30 Figure 7.6-s1 An Overview of Cellular Respiration (Step 1)

31 Figure 7.6-s2 An Overview of Cellular Respiration (Step 2)

32 Figure 7.6-s3 An Overview of Cellular Respiration (Step 3)

33 The Stages of Cellular Respiration: A Preview, Continued Oxidative phosphorylation accounts for almost 90% of the ATP generated by cellular respiration This process involves the transfer of inorganic phosphates to ADP

34 The Stages of Cellular Respiration: A Preview, Continued-1 A smaller amount of ATP is formed in glycolysis and the citric acid cycle by substrate-level phosphorylation In this process, an enzyme transfers a phosphate group directly from a substrate molecule to ADP

35 The Stages of Cellular Respiration: A Preview, Continued-2 For each molecule of glucose degraded to water by respiration, the cell makes up to 32 molecules of ATP and

36 Figure 7.7 Substrate-Level Phosphorylation

37 Concept 7.2: Glycolysis harvests chemical energy by oxidizing glucose to pyruvate Glycolysis ( sugar splitting ) breaks down glucose into two molecules of pyruvate Glycolysis occurs in the cytoplasm and has two major phases Energy investment phase Energy payoff phase The net energy yield is 2 ATP plus 2 NADH per glucose molecule Glycolysis occurs whether or not is present

38 Figure 7.UN06 In-Text Figure, Mini-Map, Glycolysis, p. 147

39 Figure 7.8 The Energy Input and Output of Glycolysis

40 Figure A Closer Look at Glycolysis (Part 1: Investment Phase)

41 Figure 7.9-1a-s1 A Closer Look at Glycolysis (Part 1a, Step 1)

42 Figure 7.9-1a-s2 A Closer Look at Glycolysis (Part 1a, Step 2)

43 Figure 7.9-1a-s3 A Closer Look at Glycolysis (Part 1a, Step 3)

44 Figure 7.9-1b-s1 A Closer Look at Glycolysis (Part 1b, Step 1)

45 Figure 7.9-1b-s2 A Closer Look at Glycolysis (Part 1b, Step 2)

46 Figure 7.9-1b-s3 A Closer Look at Glycolysis (Part 1b, Step 3)

47 Figure A Closer Look at Glycolysis (Part 2: Payoff Phase)

48 Figure 7.9-2a-s1 A Closer Look at Glycolysis (Part 2a, Step 1)

49 Figure 7.9-2a-s2 A Closer Look at Glycolysis (Part 2a, Step 2)

50 Figure 7.9-2a-s3 A Closer Look at Glycolysis (Part 2a, Step 3)

51 Figure 7.9-2b-s1 A Closer Look at Glycolysis (Part 2b, Step 1)

52 Figure 7.9-2b-s2 A Closer Look at Glycolysis (Part 2b, Step 2)

53 Figure 7.9-2b-s3 A Closer Look at Glycolysis (Part 2b, Step 3)

54 Concept 7.3: After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic molecules In the presence of, pyruvate enters the mitochondrion (in eukaryotic cells), where the oxidation of glucose is completed 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

55 Figure 7.UN07 In-Text Figure, Mini-Map, Pyruvate Oxidation, p. 148

56 Figure 7.10 An Overview of Pyruvate Oxidation and the Citric Acid Cycle

57 Figure An Overview of Pyruvate Oxidation And the Citric Acid Cycle (Part 1: Pyruvate Oxidation)

58 Figure An Overview of Pyruvate Oxidation and the Citric Acid Cycle (Part 2: Citric Acid Cycle)

59 Concept 7.3: After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic molecules, Continued The citric acid cycle, also called the Krebs cycle, completes the breakdown of pyruvate to The cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 per turn

60 Concept 7.3: After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic molecules, Continued-1 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 produced by the cycle relay electrons extracted from food to the electron transport chain

61 Figure 7.UN08 In-Text Figure, Mini-Map, Citric Acid Cycle, p. 149

62 Figure 7.11-s1 A Closer Look at the Citric Acid Cycle (Step 1)

63 Figure 7.11-s2 A Closer Look at the Citric Acid Cycle (Step 2)

64 Figure 7.11-s3 A Closer Look at the Citric Acid Cycle (Step 3)

65 Figure 7.11-s4 A Closer Look at the Citric Acid Cycle (Step 4)

66 Figure 7.11-s5 A Closer Look at the Citric Acid Cycle (Step 5)

67 Figure 7.11-s6 A Closer Look at the Citric Acid Cycle (Step 6)

68 Figure A Closer Look at the Citric Acid Cycle (Part 1)

69 Figure A Closer Look at the Citric Acid Cycle (Part 2)

70 Figure A Closer Look at the Citric Acid Cycle (Part 3)

71 Figure A Closer Look at the Citric Acid Cycle (Part 4)

72 Concept 7.4: During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis Following glycolysis and the citric acid cycle, NADH and 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

73 The Pathway of Electron Transport The electron transport chain is located 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, forming

74 The Pathway of Electron Transport, Continued Electrons are transferred from NADH or to the electron transport chain Electrons are passed through a number of proteins including cytochromes (each with an iron atom) to The electron transport chain generates no ATP directly It breaks the large free-energy drop from food to into smaller steps that release energy in manageable amounts

75 Figure 7.UN09 In-Text Figure, Mini-Map, Oxidative Phosphorylation, p. 150

76 Figure 7.12 Free-Energy Change During Electron Transport

77 Figure Free-Energy Change During Electron Transport (Part 1)

78 Figure Free-Energy Change During Electron Transport (Part 2)

79 Chemiosmosis: The Energy-Coupling Mechanism Electron transfer in the electron transport chain causes proteins to pump from the mitochondrial matrix to the intermembrane space then moves back across the membrane, passing through the protein complex, ATP synthase ATP synthase uses the exergonic flow of to drive phosphorylation of ATP This is an example of chemiosmosis, the use of energy in a gradient to drive cellular work

80 Video: ATP Synthase 3-D Side View

81 Video: ATP Synthase 3-D Top View

82 Figure 7.13 ATP Synthase, A Molecular Mill

83 Figure ATP Synthase, A Molecular Mill (Part 1: )

84 Figure ATP Synthase, A Molecular Mill (Part 2: )

85 Chemiosmosis: The Energy-Coupling Mechanism, Continued The energy stored in a gradient across a membrane couples the redox reactions of the electron transport chain to ATP synthesis The gradient is referred to as a proton-motive force, emphasizing its capacity to do work

86 Figure 7.UN09 In-Text Figure, Mini-Map, Oxidative Phosphorylation, p. 150, Continued

87 Figure 7.14 Chemiosmosis Couples the Electron Transport Chain to ATP Synthesis

88 Figure Chemiosmosis Couples the Electron Transport Chain to ATP Synthesis (Part 1: Electron Transport Chain)

89 Figure Chemiosmosis Couples the Electron Transport Chain to ATP Synthesis (Part 2: Chemiosmosis)

90 An Accounting of ATP Production by Cellular Respiration During cellular respiration, most energy flows in the following sequence: glucose NADH electron transport chain protonmotive 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 molecules is not known exactly

91 Figure 7.15 ATP Yield Per Molecule of Glucose at Each Stage of Cellular Respiration

92 Figure ATP Yield Per Molecule of Glucose at Each Stage of Cellular Respiration (Part 1: Glycolysis)

93 Figure ATP Yield Per Molecule of Glucose at Each Stage of Cellular Respiration (Part 2: Citric Acid Cycle)

94 Figure ATP Yield Per Molecule of Glucose at Each Stage of Cellular Respiration (Part 3: Oxidative Phosphorylation)

95 Figure ATP Yield Per Molecule of Glucose at Each Stage of Cellular Respiration (Part 4: Maximum ATP Yield Per Glucose)

96 Concept 7.5: Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen Most cellular respiration requires to produce ATP Without, the electron transport chain will cease to operate In that case, glycolysis couples with fermentation or anaerobic respiration to produce ATP

97 Concept 7.5: Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen, Continued Anaerobic respiration uses an electron transport chain with a final electron acceptor other than, for example, sulfate Fermentation uses substrate-level phosphorylation instead of an electron transport chain to generate ATP

98 Types of Fermentation Fermentation consists of glycolysis plus reactions that regenerate, which can be reused by glycolysis Two common types are alcohol fermentation and lactic acid fermentation

99 Types of Fermentation, Continued In alcohol fermentation, pyruvate is converted to ethanol in two steps The first step releases from pyruvate, and the second step reduces the resulting acetaldehyde to ethanol Alcohol fermentation by yeast is used in brewing, winemaking, and baking

100 Animation: Fermentation Overview

101 Figure 7.16 Fermentation (Part 1: Alcohol)

102 Types of Fermentation, Continued-1 In lactic acid fermentation, pyruvate is reduced by NADH, forming lactate as an end product, with no release of 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 is scarce

103 Figure 7.16 Fermentation (Part 2: Lactic Acid)

104 Figure 7.16 Fermentation

105 Comparing Fermentation with Anaerobic and Aerobic Respiration All use glycolysis (net ATP = 2) to oxidize glucose and other organic fuels to pyruvate In all three, is the oxidizing agent that accepts electrons from food during glycolysis The mechanism of NADH oxidation differs In fermentation the final electron acceptor is an organic molecule such as pyruvate or acetaldehyde Cellular respiration transfers electrons from NADH to a carrier molecule in the electron transport chain

106 Comparing Fermentation with Anaerobic and Aerobic Respiration, Continued Cellular respiration produces about 32 ATP per glucose molecule; fermentation produces 2 ATP per glucose molecule

107 Comparing Fermentation with Anaerobic and Aerobic Respiration, Continued-1 Obligate anaerobes carry out only fermentation or anaerobic respiration and cannot survive in the presence of 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

108 Figure 7.17 Pyruvate as a Key Juncture in Catabolism

109 The Evolutionary Significance of Glycolysis Glycolysis is the most common metabolic pathway among organisms on Earth, indicating that it evolved early in the history of life Early prokaryotes may have generated ATP exclusively through glycolysis due to the low oxygen content in the atmosphere The location of glycolysis in the cytosol also indicates its ancient origins; eukaryotic cells with mitochondria evolved much later than prokaryotic cells

110 Concept 7.6: Glycolysis and the citric acid cycle connect to many other metabolic pathways Glycolysis and the citric acid cycle are major intersections to various catabolic and anabolic pathways

111 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 and amino groups must be removed before amino acids can feed glycolysis or the citric acid cycle

112 The Versatility of Catabolism, Continued Fats are digested to glycerol (used in glycolysis) and fatty acids 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

113 Figure 7.18-s1 The Catabolism of Various Molecules From Food (Step 1)

114 Figure 7.18-s2 The Catabolism of Various Molecules From Food (Step 2)

115 Figure 7.18-s3 The Catabolism of Various Molecules From Food (Step 3)

116 Figure 7.18-s4 The Catabolism of Various Molecules From Food (Step 4)

117 Figure 7.18-s5 The Catabolism of Various Molecules From Food (Step 5)

118 Biosynthesis (Anabolic Pathways) The body uses small molecules to build other substances Some of these small molecules come directly from food; others can be produced during glycolysis or the citric acid cycle

119 Figure 7.UN10-1 Skills Exercise: Making a Bar Graph and Evaluating a Hypothesis (Part 1) Thyroid Hormone Level Low Normal Elevated Oxygen Consumption Rate [nmol /(min mg cells)] Data from M.E. Harper and M.D. Brand, the quantitative contributions of mitochondrial proton leak and ATP turnover reactions to the changed respiration rates of hepatocytes from rats of different thyroid status, Journal of Biological Chemistry 268:14850:14860 (1993).

120 Figure 7.UN10-2 Skills Exercise: Making a Bar Graph and Evaluating a Hypothesis (Part 2)

121 Figure 7.UN11 Summary of Key Concepts: Glycolysis

122 Figure 7.UN12 Summary of Key Concepts: Citric Acid Cycle

123 Figure 7.UN13 Summary of Key Concepts: Electron Transport Chain

124 Figure 7.UN14 Summary of Key Concepts: Chemiosmosis

125 Figure 7.UN15 Test Your Understanding, Question 8 (ph vs. Time)

126 Figure 7.UN16 Test Your Understanding, Question 9 (Regulation of Phosphofructokinase)

127 Figure 7.UN17 Test Your Understanding, Question 13 (Coenzyme Q)

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