Cellular Respiration: Harvesting Chemical Energy

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1 Chapter 9 Cellular Respiration: Harvesting Chemical Energy

2 You should be able to: 1. Explain how redox reactions are involved in energy exchanges. Name and describe the three stages of cellular respiration; for each, state the region of the eukaryotic cell where it occurs and the products that result 3. Explain the role of the electron transport chain in cellular respiration Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

3 4. Explain where and how the respiratory electron transport chain creates a proton gradient 5. Distinguish and explain fermentation and anaerobic respiration 6. Distinguish between obligate and facultative anaerobes Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

4 Overview: Life Is Work Living cells require energy from outside sources Some animals, such as the giant panda, obtain energy by eating plants, and some animals feed on other organisms that eat plants Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

5 Fig. 9-1

6 Energy flows into an ecosystem as sunlight and leaves as heat Photosynthesis generates O and organic molecules, which are used in cellular respiration Cells use chemical energy stored in organic molecules to regenerate ATP, which powers work Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

7 Fig. 9- Light energy ECOSYSTEM CO + H O Photosynthesis in chloroplasts Cellular respiration in mitochondria Organic molecules + O ATP ATP powers most cellular work Heat energy

8 Concept 9.1: Catabolic pathways yield energy by oxidizing organic fuels Several processes are central to cellular respiration and related pathways Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

9 Catabolic Pathways and Production of ATP The breakdown of organic molecules is exergonic Fermentation is a partial degradation of sugars that occurs without O Aerobic respiration consumes organic molecules and O and yields ATP Anaerobic respiration is similar to aerobic respiration but consumes compounds other than O Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

10 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 1 O O 6 CO + 6 H O + Energy (ATP + heat) Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

11 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

12 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) Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

13 Fig. 9-UN1 becomes oxidized (loses electron) becomes reduced (gains electron)

14 Fig. 9-UN becomes oxidized becomes reduced

15 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

16 Fig. 9-3 Reactants becomes oxidized Methane (reducing agent) Oxygen (oxidizing agent) Products becomes reduced Carbon dioxide Water

17 Oxidation of Organic Fuel Molecules During Cellular Respiration During cellular respiration, the fuel (such as glucose) is oxidized, and O is reduced: Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

18 Fig. 9-UN3 becomes oxidized becomes reduced

19 Fig. 9-UN4 Dehydrogenase

20 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

21 Fig. 9-4 NAD + + [H] Nicotinamide (oxidized form) e + H + e + H + Dehydrogenase Reduction of NAD + Oxidation of NADH NADH Nicotinamide (reduced form) + H + H +

22 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 pulls electrons down the chain in an energyyielding tumble The energy yielded is used to regenerate ATP Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

23 Fig. 9-5 H + 1 / O Explosive release of heat and light energy (a) Uncontrolled reaction H 1 / O (from food via NADH) H + + e Controlled release of energy for synthesis of ATP 1 / O (b) Cellular respiration

24 The Stages of Cellular Respiration: A Preview Cellular respiration 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) Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

25 Fig Electrons carried via NADH Glycolysis Glucose Pyruvate Cytosol ATP Substrate-level phosphorylation

26 Fig Electrons carried via NADH Glycolysis Glucose Pyruvate Cytosol ATP Substrate-level phosphorylation Electrons carried via NADH and FADH Citric acid cycle Mitochondrion ATP Substrate-level phosphorylation

27 Fig Electrons carried via NADH Glycolysis Glucose Pyruvate Cytosol ATP Substrate-level phosphorylation Electrons carried via NADH and FADH Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis Mitochondrion ATP ATP Substrate-level phosphorylation Oxidative phosphorylation

28 The process that generates most of the ATP is called oxidative phosphorylation because it is powered by redox reactions Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

30 Fig. 9-7 Enzyme ADP P Substrate Enzyme + ATP Product

31 Concept 9.: 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

32 Fig. 9-8 Energy investment phase Glucose ADP + P ATP used Energy payoff phase 4 ADP + 4 P 4 ATP formed NAD e + 4 H + NADH + H + Pyruvate + H O Net Glucose Pyruvate + H O 4 ATP formed ATP used ATP NAD e + 4 H + NADH + H +

33 Fig Glucose ATP 1 Hexokinase ADP Glucose-6-phosphate Glucose ATP 1 Hexokinase ADP Glucose-6-phosphate

34 Fig Glucose ATP 1 Hexokinase ADP Glucose-6-phosphate Phosphoglucoisomerase Fructose-6-phosphate Glucose-6-phosphate Phosphoglucoisomerase Fructose-6-phosphate

35 Fig Glucose ATP 1 Hexokinase ADP Glucose-6-phosphate Phosphoglucoisomerase Fructose-6-phosphate ATP 3 Phosphofructokinase ADP Fructose- 1, 6-bisphosphate Fructose-6-phosphate ATP ADP 3 Phosphofructokinase Fructose- 1, 6-bisphosphate 1

36 Fig Glucose ATP 1 Hexokinase ADP Glucose-6-phosphate Phosphoglucoisomerase Fructose-6-phosphate ATP 3 Phosphofructokinase ADP Fructose- 1, 6-bisphosphate 4 Aldolase 5 Isomerase Dihydroxyacetone phosphate Glyceraldehyde- 3-phosphate Fructose- 1, 6-bisphosphate 4 Aldolase 5 Isomerase Dihydroxyacetone phosphate Glyceraldehyde- 3-phosphate

37 Fig NAD + NADH + H + 6 Triose phosphate dehydrogenase P i 1, 3-Bisphosphoglycerate Glyceraldehyde- 3-phosphate NAD + 6 Triose phosphate dehydrogenase NADH P i + H + 1, 3-Bisphosphoglycerate

38 Fig NAD + 6 NADH P i + H + Triose phosphate dehydrogenase 1, 3-Bisphosphoglycerate ADP ATP 7 Phosphoglycerokinase 3-Phosphoglycerate 1, 3-Bisphosphoglycerate ADP ATP 7 Phosphoglycerokinase 3-Phosphoglycerate

39 Fig NAD + NADH + H + P i 6 Triose phosphate dehydrogenase 1, 3-Bisphosphoglycerate ADP 7 Phosphoglycerokinase ATP 3-Phosphoglycerate 8 Phosphoglyceromutase -Phosphoglycerate 3-Phosphoglycerate 8 Phosphoglyceromutase -Phosphoglycerate

40 Fig NAD + NADH + H + 6 Triose phosphate dehydrogenase P i 1, 3-Bisphosphoglycerate ADP 7 Phosphoglycerokinase ATP 3-Phosphoglycerate 8 Phosphoglyceromutase -Phosphoglycerate H O 9 Enolase Phosphoenolpyruvate -Phosphoglycerate H O 9 Enolase Phosphoenolpyruvate

41 Fig NAD + NADH + H + 6 Triose phosphate dehydrogenase P i 1, 3-Bisphosphoglycerate ADP 7 Phosphoglycerokinase ATP 3-Phosphoglycerate 8 Phosphoglyceromutase -Phosphoglycerate 9 Enolase H O Phosphoenolpyruvate ADP ATP 10 Pyruvate kinase Pyruvate Phosphoenolpyruvate ADP ATP 10 Pyruvate kinase Pyruvate

42 Concept 9.3: The citric acid cycle completes the energy-yielding oxidation of organic molecules In the presence of O, pyruvate enters the mitochondrion Before the citric acid cycle can begin, pyruvate must be converted to acetyl CoA, which links the cycle to glycolysis Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

43 Fig CYTOSOL MITOCHONDRION NAD + NADH + H Acetyl CoA Pyruvate CO Coenzyme A Transport protein

44 The citric acid cycle, also called the Krebs cycle, takes place within the mitochondrial matrix The cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 FADH per turn Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

45 Fig Pyruvate NAD + NADH + H + Acetyl CoA CoA CO CoA CoA Citric acid cycle FADH FAD ADP + P i ATP 3 CO 3 NAD + NADH + 3 H +

46 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 produced by the cycle relay electrons extracted from food to the electron transport chain Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

47 Fig Acetyl CoA CoA SH 1 Oxaloacetate Citrate Citric acid cycle

48 Fig Acetyl CoA CoA SH 1 Oxaloacetate Citrate Citric acid cycle H O Isocitrate

49 Fig Acetyl CoA CoA SH 1 Oxaloacetate Citrate Citric acid cycle H O Isocitrate NAD + 3 NADH + H + CO -Ketoglutarate

50 Fig Acetyl CoA CoA SH 1 H O Oxaloacetate Citric acid cycle Citrate Isocitrate NAD + 3 CO NADH + H + CoA SH 4 -Ketoglutarate NAD + CO Succinyl CoA NADH + H +

51 Fig Acetyl CoA CoA SH 1 H O Oxaloacetate Citric acid cycle Citrate Isocitrate NAD + 3 CO NADH + H + CoA SH CoA SH 4 -Ketoglutarate 5 NAD + CO Succinate GTP GDP P i Succinyl CoA NADH + H + ADP ATP

52 Fig FADH Acetyl CoA CoA SH 1 H O Oxaloacetate Citric acid cycle Citrate Isocitrate NAD + 3 CO NADH + H + Fumarate 6 CoA SH CoA SH 4 -Ketoglutarate FAD Succinate 5 GTP GDP P i Succinyl CoA NAD + NADH + H + CO ADP ATP

53 Fig H O 7 FADH Acetyl CoA CoA SH 1 H O Oxaloacetate Malate Citric acid cycle Citrate Isocitrate NAD + 3 CO NADH + H + Fumarate 6 CoA SH CoA SH 4 -Ketoglutarate FAD Succinate 5 GTP GDP P i Succinyl CoA NAD + NADH + H + CO ADP ATP

54 Fig H O Acetyl CoA CoA SH NADH +H + 1 H O NAD + 8 Oxaloacetate 7 Malate Citric acid cycle Citrate Isocitrate NAD + 3 CO NADH + H + Fumarate 6 CoA SH CoA SH 4 -Ketoglutarate FADH FAD Succinate 5 GTP GDP P i Succinyl CoA NAD + NADH + H + CO ADP ATP

55 Concept 9.4: During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis Following glycolysis and the citric acid cycle, NADH and FADH 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

56 The Pathway of Electron Transport The electron transport chain is in the 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, forming H O Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

57 Fig NADH 50 e NAD + FADH 40 FMN Fe S e FAD Fe S FAD Multiprotein complexes 30 0 Q Cyt b Fe S Cyt c 1 Cyt c Cyt a I V Cyt a 3 10 e (from NADH or FADH ) 0 H / O H O

58 Electrons are transferred from NADH or FADH to the electron transport chain Electrons are passed through a number of proteins including cytochromes (each with an iron atom) to O The electron transport chain generates no ATP The chain s function is to break the large freeenergy drop from food to O into smaller steps that release energy in manageable amounts Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

59 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 channels in 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

60 Fig INTERMEMBRANE SPACE Rotor H + Stator Internal rod Catalytic knob ADP + P ATP i MITOCHONDRIAL MATRIX

61 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 protonmotive force, emphasizing its capacity to do work Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

62 Fig H + Protein complex of electron carriers NADH (carrying electrons from food) H + H + Cyt c Q V FADH FAD H / O H O NAD + 1 Electron transport chain Oxidative phosphorylation H + ATP synthase ADP + P i ATP H + Chemiosmosis

63 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 40% of the energy in a glucose molecule is transferred to ATP during cellular respiration, making about 38 ATP Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

64 Fig CYTOSOL Electron shuttles span membrane NADH or FADH MITOCHONDRION NADH NADH 6 NADH FADH Glycolysis Glucose Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis + ATP + ATP + about 3 or 34 ATP Maximum per glucose: About 36 or 38 ATP

65 Concept 9.5: Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen Most cellular respiration requires O to produce ATP Glycolysis can produce ATP with or without O (in aerobic or anaerobic conditions) In the absence of O, glycolysis couples with fermentation or anaerobic respiration to produce ATP Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

66 Anaerobic respiration uses an electron transport chain with an electron acceptor other than O, for example sulfate Fermentation uses phosphorylation instead of an electron transport chain to generate ATP Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

67 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

68 In alcohol fermentation, pyruvate is converted to ethanol in two steps, with the first releasing CO Alcohol fermentation by yeast is used in brewing, winemaking, and baking Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

69 Fig ADP + P i ATP Glucose Glycolysis Pyruvate NAD + NADH + H + CO Ethanol Acetaldehyde (a) Alcohol fermentation ADP + P i ATP Glucose Glycolysis NAD + NADH + H + Pyruvate Lactate (b) Lactic acid fermentation

70 Fig. 9-18a ADP + P i ATP Glucose Glycolysis Pyruvate NAD + NADH CO + H + Ethanol Acetaldehyde (a) Alcohol fermentation

71 In lactic acid fermentation, pyruvate is reduced to NADH, forming lactate as an end product, with no release of CO 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 is scarce Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

72 Fig. 9-18b ADP + P i ATP Glucose Glycolysis NAD + NADH + H + Pyruvate Lactate (b) Lactic acid fermentation

73 Fermentation and Aerobic Respiration Compared Both processes use glycolysis to oxidize glucose and other organic fuels to pyruvate The processes have different final electron acceptors: an organic molecule (such as pyruvate or acetaldehyde) in fermentation and O in cellular respiration Cellular respiration produces 38 ATP per glucose molecule; fermentation produces ATP per glucose molecule Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

74 Obligate anaerobes carry out fermentation or anaerobic respiration and cannot survive in the presence of O 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

75 Fig CYTOSOL No O present: Fermentation Ethanol or lactate Glucose Glycolysis Pyruvate O present: Aerobic cellular respiration Acetyl CoA MITOCHONDRION Citric acid cycle

76 The Evolutionary Significance of Glycolysis Glycolysis occurs in nearly all organisms Glycolysis probably evolved in ancient prokaryotes before there was oxygen in the atmosphere Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

77 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

78 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

79 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

80 Fig. 9-0 NH 3 Proteins Carbohydrates Amino acids Sugars Glycolysis Glucose Glyceraldehyde-3- P Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation Fats Glycerol Fatty acids

81 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

82 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 Copyright 008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

83 Fig. 9-1 Inhibits ATP Glucose Glycolysis Fructose-6-phosphate Phosphofructokinase Fructose-1,6-bisphosphate Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation AMP Stimulates + Inhibits Citrate

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