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Chapter 9 Cellular Respiration: Harvesting Chemical Energy PowerPoint Lectures for Biology, Seventh Edition Neil Campbell and Jane Reece Lectures by Chris Romero Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Overview: Life Is Work • Living cells require energy from outside sources • Some animals, such as the giant panda, obtain energy by eating plants; others feed on organisms that eat plants Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Energy flows into an ecosystem as sunlight and leaves as heat • Photosynthesis generates oxygen and organic molecules, which are used in cellular respiration • Cells use chemical energy stored in organic molecules to regenerate ATP, which powers work Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-2 Light energy ECOSYSTEM Photosynthesis in chloroplasts Organic + O molecules 2 CO2 + H2O Cellular respiration in mitochondria ATP powers most cellular work Heat energy Concept 9.1: Catabolic pathways yield energy by oxidizing organic fuels • Several processes are central to cellular respiration and related pathways Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Catabolic Pathways and Production of ATP • The breakdown of organic molecules is exergonic • __________ is a partial degradation of sugars that occurs without oxygen • Cellular respiration consumes oxygen and organic molecules and yields ATP • Although carbohydrates, fats, and proteins are all consumed as fuel, it is helpful to trace cellular respiration with the sugar glucose: write balanced equation: Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 LEO • In reduction, a substance gains electrons, or is reduced (the amount of positive charge is reduced) GER becomes oxidized (loses electron) Xe- + Y X + becomes reduced (gains electron) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ye- • The electron donor is called the reducing agent • The electron receptor is called the oxidizing agent Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Some redox reactions do not transfer electrons but change the electron sharing in covalent bonds • An example is the reaction between methane and oxygen Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-3 Products Reactants becomes oxidized CH4 2 O2 + CO2 C Energy 2 H2O + becomes reduced H H + H O O O C O H O H Methane (reducing agent) Oxygen (oxidizing agent) Carbon dioxide Water H Oxidation of Organic Fuel Molecules During Cellular Respiration • During cellular respiration, the fuel (such as glucose) is oxidized and oxygen is reduced: becomes oxidized C6H12O6 + 6O2 6CO2 + 6H2O + Energy becomes reduced Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-4 2 e– + 2 H+ NAD+ 2 e– + H+ H+ NADH Dehydrogenase + 2[H] (from food) Nicotinamide (oxidized form) + Nicotinamide (reduced form) H+ • 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 • Oxygen pulls electrons down the chain in an energy-yielding tumble • The energy yielded is used to regenerate ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-5 H2 + 1/2 O2 + 2H 1 /2 O2 1/2 O2 (from food via NADH) Explosive release of heat and light energy Free energy, G Free energy, G 2 H+ + 2 e– Controlled release of energy for synthesis of ATP ATP ATP ATP 2 e– 2 H+ H2O Uncontrolled reaction H2O Cellular respiration 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) • The process that generates most of the ATP is called oxidative phosphorylation because it is powered by redox reactions [Animation listed on slide following figure] Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-6_1 Glycolysis Pyruvate Glucose Cytosol Mitochondrion ATP Substrate-level phosphorylation LE 9-6_2 Glycolysis Pyruvate Glucose Cytosol Citric acid cycle Mitochondrion ATP ATP Substrate-level phosphorylation Substrate-level phosphorylation LE 9-6_3 Electrons carried via NADH and FADH2 Electrons carried via NADH Glycolysis Pyruvate Glucose Cytosol Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis Mitochondrion ATP ATP ATP Substrate-level phosphorylation Substrate-level phosphorylation Oxidative phosphorylation Animation: Cell Respiration Overview Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Oxidative phosphorylation accounts for almost 90% of the ATP generated by cellular respiration • A small amount of ATP is formed in glycolysis and the citric acid cycle by substrate-level phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-7 Enzyme Enzyme ADP P Substrate + Product ATP Concept 9.2: Glycolysis harvests 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 Animation: Glycolysis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-8 Energy investment phase Glucose 2 ATP used 2 ADP + 2 P Glycolysis Citric acid cycle Oxidative phosphorylation Energy payoff phase ATP ATP ATP 4 ADP + 4 P 2 NAD+ + 4 e– + 4 H+ 4 ATP formed 2 NADH + 2 H+ 2 Pyruvate + 2 H2O Net Glucose 4 ATP formed – 2 ATP used 2 NAD+ + 4 e– + 4 H+ 2 Pyruvate + 2 H2O 2 ATP 2 NADH + 2 H+ LE 9-9a_1 Glucose ATP Hexokinase ADP Glucose-6-phosphate Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation ATP LE 9-9a_2 Glucose ATP Hexokinase ADP Glucose-6-phosphate Phosphoglucoisomerase Fructose-6-phosphate ATP Phosphofructokinase ADP Fructose1, 6-bisphosphate Aldolase Isomerase Dihydroxyacetone phosphate Glyceraldehyde3-phosphate Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation ATP LE 9-9b_1 2 NAD+ Triose phosphate dehydrogenase 2 NADH + 2 H+ 1, 3-Bisphosphoglycerate 2 ADP Phosphoglycerokinase 2 ATP 3-Phosphoglycerate Phosphoglyceromutase 2-Phosphoglycerate LE 9-9b_2 2 NAD+ Triose phosphate dehydrogenase 2 NADH + 2 H+ 1, 3-Bisphosphoglycerate 2 ADP Phosphoglycerokinase 2 ATP 3-Phosphoglycerate Phosphoglyceromutase 2-Phosphoglycerate 2 H2O Enolase Phosphoenolpyruvate 2 ADP Pyruvate kinase 2 ATP Pyruvate Concept 9.3: The citric acid cycle completes the energy-yielding oxidation of organic molecules • Before the citric acid cycle can begin, pyruvate must be converted to acetyl CoA, which links the cycle to glycolysis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-10 MITOCHONDRION CYTOSOL NAD+ NADH + H+ Acetyl Co A Pyruvate Transport protein CO2 Coenzyme A • The citric acid cycle, also called the Krebs cycle, takes place within the mitochondrial matrix • The cycle oxidizes organic fuel derived from pyruvate, generating one ATP, 3 NADH, and 1 FADH2 per turn Animation: Electron Transport Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-11 Pyruvate (from glycolysis, 2 molecules per glucose) CO2 NAD+ Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation CoA NADH + H+ Acetyl CoA CoA CoA Citric acid cycle FADH2 2 CO2 3 NAD+ 3 NADH + 3 H+ FAD ADP + P i ATP ATP • 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 FADH2 produced by the cycle relay electrons extracted from food to the electron transport chain Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-12_1 Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation ATP Acetyl CoA H2O Oxaloacetate Citrate Isocitrate Citric acid cycle LE 9-12_2 Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation ATP Acetyl CoA H2O Oxaloacetate Citrate Isocitrate CO2 Citric acid cycle NAD+ NADH + H+ a-Ketoglutarate NAD+ Succinyl CoA NADH + H+ CO2 LE 9-12_3 Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation ATP Acetyl CoA H2O Oxaloacetate Citrate Isocitrate CO2 Citric acid cycle NAD+ NADH + H+ Fumarate a-Ketoglutarate FADH2 NAD+ FAD Succinate GTP GDP ADP ATP Pi Succinyl CoA NADH + H+ CO2 LE 9-12_4 Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation ATP Acetyl CoA NADH + H+ H2O NAD+ Oxaloacetate Malate Citrate Isocitrate CO2 Citric acid cycle H2O NAD+ NADH + H+ Fumarate a-Ketoglutarate FADH2 NAD+ FAD Succinate GTP GDP ADP ATP Pi Succinyl CoA NADH + H+ CO2 Concept 9.4: During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis • Following glycolysis and the citric acid cycle, NADH and FADH2 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 O2, forming water Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-13 NADH 50 Free energy (G) relative to O2 (kcal/mol) FADH2 40 FMN I Multiprotein complexes FAD Fe•S II Fe•S Q III Cyt b 30 Fe•S Cyt c1 Glycolysis Citric acid cycle ATP ATP Oxidative phosphorylation: electron transport and chemiosmosis IV Cyt c Cyt a Cyt a3 20 10 0 2 H+ + 1/2 O2 H2O ATP • The electron transport chain generates no ATP • The chain’s function is to break the large freeenergy drop from food to O2 into smaller steps that release energy in manageable amounts Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-14 INTERMEMBRANE SPACE H+ H+ H+ H+ H+ H+ A rotor within the membrane spins as shown when H+ flows past it down the H+ gradient. H+ A stator anchored in the membrane holds the knob stationary. A rod (or “stalk”) extending into the knob also spins, activating catalytic sites in the knob. H+ ADP + P ATP i MITOCHONDRAL MATRIX Three catalytic sites in the stationary knob join inorganic phosphate to ADP to make ATP. • 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 Animation: Fermentation Overview Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-15 Inner mitochondrial membrane Glycolysis Citric acid cycle ATP ATP Oxidative phosphorylation: electron transport and chemiosmosis ATP H+ H+ H+ H+ Intermembrane space Cyt c Protein complex of electron carriers Q IV III I ATP synthase II Inner mitochondrial membrane FADH2 NADH + H+ 2H+ + 1/2 O2 H2O FAD NAD+ Mitochondrial matrix ATP ADP + P i (carrying electrons from food) H+ Electron transport chain Electron transport and pumping of protons (H+), Which create an H+ gradient across the membrane Oxidative phosphorylation Chemiosmosis ATP synthesis powered by the flow of H+ back across the membrane 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-16 Electron shuttles span membrane CYTOSOL 2 NADH Glycolysis Glucose 2 Pyruvate MITOCHONDRION 2 NADH or 2 FADH2 2 NADH 2 Acetyl CoA 6 NADH Citric acid cycle + 2 ATP + 2 ATP by substrate-level phosphorylation by substrate-level phosphorylation Maximum per glucose: About 36 or 38 ATP 2 FADH2 Oxidative phosphorylation: electron transport and chemiosmosis + about 32 or 34 ATP by oxidation phosphorylation, depending on which shuttle transports electrons form NADH in cytosol Concept 9.5: Fermentation enables some cells to produce ATP without the use of oxygen • Cellular respiration requires O2 to produce ATP • Glycolysis can produce ATP with or without O2 (in aerobic or anaerobic conditions) • In the absence of O2, glycolysis couples with fermentation to produce ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In alcohol fermentation, pyruvate is converted to ethanol in two steps, with the first releasing CO2 • Alcohol fermentation by yeast is used in brewing, winemaking, and baking Play Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-17a 2 ADP + 2 P i Glucose 2 ATP Glycolysis 2 Pyruvate 2 NAD+ 2 Ethanol Alcohol fermentation 2 NADH + 2 H+ 2 CO2 2 Acetaldehyde • In lactic acid fermentation, pyruvate is reduced to NADH, forming lactate as an end product, with no release of CO2 • 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 O2 is scarce Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-17b 2 ADP + 2 P i Glucose 2 ATP Glycolysis 2 NAD+ 2 NADH + 2 H+ 2 CO2 2 Pyruvate 2 Lactate Lactic acid fermentation Fermentation and Cellular 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) in fermentation and O2 in cellular respiration • Cellular respiration produces much more ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-18 Glucose CYTOSOL Pyruvate No O2 present Fermentation O2 present Cellular respiration MITOCHONDRION Ethanol or lactate Acetyl CoA Citric acid cycle 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 • Fats are digested to glycerol (used in glycolysis) and fatty acids (used in generating acetyl CoA) • An oxidized gram of fat produces more than twice as much ATP as an oxidized gram of carbohydrate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-19 Proteins Carbohydrates Amino acids Sugars Glycerol Fatty acids Glycolysis Glucose Glyceraldehyde-3- P NH3 Fats Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 9-20 Glucose AMP Glycolysis Fructose-6-phosphate – Stimulates + Phosphofructokinase – Fructose-1,6-bisphosphate Inhibits Inhibits Pyruvate ATP Citrate Acetyl CoA Citric acid cycle Oxidative phosphorylation