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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 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 Catabolic Pathways and Production of ATP • The breakdown of organic molecules is exergonic • Fermentation is a partial degradation of sugars that occurs without oxygen • Cellular respiration consumes oxygen and organic molecules and yields ATP • Cellular Respiration Equation: 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 • Oxidation: • Reduction: Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The electron donor is called the reducing agent • The electron receptor is called the oxidizing agent becomes oxidized (loses electron) Xe- + Y X + becomes reduced (gains electron) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ye- 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 • 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 • NADH 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 • 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) NADH and NADH Glycolysis Cytosol ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings FADH2 Citric electron transport acid and cycle Mitochondrion chemiosmosis ATP ATP 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 Enzyme Enzyme ADP P Substrate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings + 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 MITOCHONDRION CYTOSOL NAD+ NADH + H+ Acetyl Co A Pyruvate CO2 Transport protein Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Coenzyme A • The citric acid cycle, also called the Krebs cycle, takes place within the mitochondrial matrix Pyruvate (from glycolysis, 2 molecules per glucose) CO2 NAD+ Glycolysis Citric acid cycle ATP ATP Oxidation phosphorylation CoA NADH + H+ ATP Acetyl CoA CoA CoA Citric acid cycle 2 CO2 FADH2 3 NAD+ 3 NADH + 3 H+ FAD ADP + Pi ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Animation: Electron Transport • The citric acid cycle has eight steps, each catalyzed by a specific enzyme acetyl CoA + oxaloacetate citrate • The next seven steps decompose the citrate back to oxaloacetate, making the process a cycle • Electron Carriers: • NADH and FADH2 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 • 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 Chemiosmosis: The Energy-Coupling Mechanism • Electron transfer in the ETC 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 • 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 9-15 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 LE 9-14 INTERMEMBRANE SPACE H+ H+ H+ H+ H+ H+ H+ A rotor within the membrane spins as shown when H+ flows past it down the H+ gradient. A stator anchored in the membrane holds the knob stationary. H+ ADP + Pi MITOCHONDRAL MATRIX ATP A rod (or “stalk”) extending into the knob also spins, activating catalytic sites in the knob. Three catalytic sites in the stationary knob join inorganic phosphate to ADP to make ATP. The H+ gradient is referred to as a proton-motive force, emphasizing its capacity to do work 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, Play winemaking, and baking 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 by 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 • Yeast and many bacteria are facultative anaerobes, meaning that they can survive using either fermentation or cellular respiration Glucose CYTOSOL Pyruvate No O2 present Fermentation Ethanol or lactate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings O2 present Cellular respiration MITOCHONDRIO Acetyl CoA Citric acid cycle The Versatility of Catabolism • Catabolic pathways funnel electrons from many kinds of organic molecules into cellular respiration Proteins Carbohydrates Amino acids Sugars Glycerol Fatty acids Glycolysis Glucose Glyceraldehyde-3- P NH3 Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fats