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Chapter 9 Cellular Respiration: Harvesting Chemical Energy PowerPoint TextEdit Slides for Biology, Seventh Edition Neil Campbell and Jane Reece Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.1 The giant panda is consuming fuel to power the work of life Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.2 Energy flow and chemical recycling in ecosystems Light energy ECOSYSTEM Photosynthesis in chloroplasts CO2 + H2O Organic + O2 Cellular respiration molecules in mitochondria ATP powers most cellular work Heat energy Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.3 Methane combustion as an energyyielding redox reaction Reactants Products becomes oxidized CH4 + 2 O2 CO2 Energy + 2 H2O becomes reduced O O C O H O O H H H Methane (reducing agent) Oxygen (oxidizing agent) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Carbon dioxide Water H H C + Figure 9.4 NAD+ as an electron shuttle 2 e– + 2 H+ NAD+ Dehydrogenase O NH2 H C CH2 O O– O O P O H O P O– HO O N+ Nicotinamide (oxidized form) H OH HO CH2 N O H HO N N N NAD+ Reduction of + 2[H] (from food) Oxidation of NADH NH2 H 2 e– + H+ H H OH Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings NADH H O C H N H+ NH2 Nicotinamide (reduced form) + H+ Figure 9.5 An introduction to electron transport chains H2 + 1/2 O2 2H /2 O2 1 + (from food via NADH) Explosive release of heat and light energy ATP Free energy, G Free energy, G 2 H+ + 2 e– Controlled release of energy for synthesis of ATP ATP ATP 2 e– /2 O2 1 2 H+ H2O (a) Uncontrolled reaction Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings H2O (b) Cellular respiration Figure 9.6 An overview of cellular respiration Glycolsis Glucose Pyruvate Mitochondrion ATP Substrate-level phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.6 An overview of cellular respiration Citric acid cycle Glycolsis Glucose Pyruvate Mitochondrion ATP ATP Substrate-level phosphorylation Substrate-level phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.6 An overview of cellular respiration Electrons carried via NADH Electrons carried via NADH and FADH2 Citric acid cycle Glycolsis Glucose Pyruvate Oxidative phosphorylation: electron transport and chemiosmosis Mitochondrion ATP Substrate-level phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings ATP Substrate-level phosphorylation ATP Oxidative phosphorylation Figure 9.16 ATP yield per molecule of glucose at each stage of cellular respiration Electron shuttles span membrane CYTOSOL MITOCHONDRION 2 NADH or 2 FADH2 2 NADH 2 NADH Glycolysis Glucose 2 Pyruvate 6 NADH Citric acid cycle 2 Acetyl CoA + 2 ATP by substrate-level phosphorylation Maximum per glucose: 2 FADH2 Oxidative phosphorylation: electron transport and chemiosmosis + 2 ATP + about 32 or 34 ATP by oxidative phosphorylation, depending by substrate-level on which shuttle transports electrons phosphorylation from NADH in cytosol About 36 or 38 ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.7 Substrate-level phosphorylation Enzyme Enzyme ADP P Substrate + Product Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings ATP Figure 9.8 The energy input and output of glycolysis Glycolysis ATP Citric acid cycle Oxidative phosphorylation ATP ATP Energy investment phase Glucose 2 ATP + 2 P 2 ATP used 4 ATP formed Energy payoff phase 4 ADP + 4 P 2 NAD+ + 4 e- + 4 H + 2 NADH + 2 H+ 2 Pyruvate + 2 H2O Net Glucose 4 ATP formed – 2 ATP used 2 NAD+ + 4 e– + 4 H + Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2 Pyruvate + 2 H2O 2 ATP 2 NADH + 2 H+ Figure 9.9 A closer look at glycolysis: energy investment phase CH2OH OH HH OH H HO OH H OH Glucose ATP 1 Hexokinase ADP CH2O P O H H H OH H OH HO H OH Glucose-6-phosphate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Glycolysis Citirc acid cycle ATP ATP Oxidative phosphorylation ATP Figure 9.9 A closer look at glycolysis: energy investment phase CH2OH H H OH OH H HO OH H OH Glucose ATP 1 Hexokinase ADP CH2O P HH O H OH H HO H OH Glucose-6-phosphate 2 Phosphoglucoisomerase CH2O P O CH2OH H HO HO H H HO Fructose-6-phosphate 3 ATP Phosphofructokinase ADP P O CH2 O CH2 O H HO OH H HO H Fructose1, 6-bisphosphate P 4 Aldolase 5 P O CH2 C O CH2OH Dihydroxyacetone phosphate Isomerase H C O CHOH CH2 O P Glyceraldehyde3-phosphate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Glycolysis Citirc acid cycle ATP ATP Oxidative phosphorylation ATP Figure 9.9 A closer look at glycolysis: energy payoff phase 6 2 NAD+ Triose phosphate dehydrogenase 2 Pi 2 NADH + 2 H+ 2 P O C O CHOH CH2 O P 1, 3-Bisphosphoglycerate 2 ADP 7 Phosphoglycerokinase 2 ATP O– 2 C O CHOH CH2 O P 3-Phosphoglycerate 8 Phosphoglyceromutase O– 2 H C O C O P CH2OH 2-Phosphoglycerate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.9 A closer look at glycolysis: energy payoff phase 6 Triose phosphate dehydrogenase 2 NAD+ 2 Pi 2 NADH + 2 H+ 2 P O C O CHOH CH2 O P 1, 3-Bisphosphoglycerate 2 ADP 7 Phosphoglycerokinase 2 ATP O– 2 C CHOH CH2 O P 3-Phosphoglycerate 8 Phosphoglyceromutase 2 O– C O H C O P CH2OH 2-Phosphoglycerate 9 Enolase 2H O 2 2 O– C O C O P CH2 Phosphoenolpyruvate 2 ADP 10 Pyruvate kinase 2 ATP 2 O– C O C O CH3 Pyruvate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.10 Conversion of pyruvate to acetyle CoA, the junction between glycolysis and the citric acid cycle CYTOSOL MITOCHONDRION NAD+ NADH + H+ O– S CoA C O 2 C C O O 1 3 CH3 Pyruvate CH3 Acetyle CoA CO2 Transport protein Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Coenzyme A Figure 9.11 An overview of the citric acid cycle Pyruvate (from glycolysis, 2 molecules per glucose) Glycolysis Citric acid cycle ATP ATP Oxidative phosphorylation ATP CO2 NAD+ CoA NADH + H+ Acetyle CoA CoA CoA Citric acid cycle 2 CO2 3 NAD+ FADH2 FAD 3 NADH + 3 H+ ADP + P i ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.12 Closer look at the citric acid cycle Glycolysis ATP Citirc acid cycle ATP Oxidative phosphorylation ATP S CoA C O CH3 Acetyl CoA CoA SH O C COO– CH2 COO– 1 COO– Oxaloacetate H2O COO– CH2 HO C COO– CH2 COO– Citrate CH2 2 HC COO– HO CH COO– Isocitrate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.12 Closer look at the citric acid cycle Glycolysis ATP Citirc acid cycle ATP Oxidative phosphorylation ATP S CoA C O CH3 Acetyl CoA CoA SH C COO– CH2 H2O COO– 1 1 COO– COO– CH2 Oxaloacetate HO C COO– CH2 2 CH2 COO– COO– HC HO CH Citrate COO– Isocitrate Citric acid cycle CO2 3 NAD+ NADH + H+ COO– CoA SH CH2 a-Ketoglutarate CH2 4 COO– C O COO– CH2 CH2 C O S CoA SUCCINYL CoA Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings NAD+ NADH + H+ CO2 Figure 9.12 Closer look at the citric acid cycle Glycolysis ATP Citirc acid cycle Oxidative phosphorylation ATP ATP S CoA C O CH3 Acetyl CoA CoA SH O C COO– CH2 COO– 1 COO– H2O COO– CH2 Oxaloacetate HO C COO– CH2 CH2 2 HC COO– COO– HO CH Citrate COO– Isocitrate Citric acid cycle COO– CH NAD+ NADH + H+ COO– Fumarate HC CO2 3 CoA SH CH2 a-Ketoglutarate CH2 COO– 6 4 CoA SH COO– FAD CH2 CH2 COO– C O Succinate Pi S CoA GTP GDP Succinyl CoA ADP ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings COO– CH2 5 CH2 FADH2 COO– C O NAD+ NADH + H+ CO2 Figure 9.12 Closer look at the citric acid cycle Glycolysis ATP Citirc acid cycle Oxidative phosphorylation ATP ATP S CoA C O CH3 Acetyl CoA CoA SH C COO– O NADH + H+ COO– 1 CH2 COO– NAD+ 8 Oxaloacetate HO CH COO– C COO– CH2 COO– HO H2O CH2 CH2 2 HC COO– Malate HO Citrate CH2 CH COO– Isocitrate COO– H2 O COO– Citric acid cycle 7 COO– CO2 3 NAD+ COO– CH Fumarate HC CH2 CH2 a-Ketoglutarate CoA SH COO– 6 CoA SH COO– FAD CH2 CH2 COO– C O Succinate Pi S CoA GTP GDP Succinyl CoA ADP ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 4 C O COO– CH2 5 CH2 FADH2 COO– NADH + H+ NAD+ NADH + H+ CO2 Figure 9.13 Free-energy change during electron transport Glycolysis Citirc acid cycle ATP ATP Oxidative phosphorylation ATP NADH 50 Free energy (G) relative to O2 (kcl/mol) FADH2 40 FMN I Fe•S Multiprotein complexes FAD Fe•S II O Cyt b 30 20 III Fe•S Cyt c1 Cyt c IV Cyt a Cyt a3 10 0 2H++ 2 O 2 1 H2O Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.14 ATP synthase, a molecular mill INTERMEMBRANE SPACE H+ H+ H+ H+ H+ H+ H+ A rotor within the membrane spins clockwise when H+ flows past it down the H+ gradient. A stator anchored in the membrane holds the knob stationary. H+ ADP + Pi MITOCHONDRIAL MATRIX Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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. Figure 9.15 Chemiosmosis couples the electron transport chain to ATP synthesis Glycolysis ATP Citirc acid cycle ATP Inner Mitochondrial membrane Oxidative phosphorylation electron transport and chemiosmosis ATP H+ H+ H+ Intermembrane space Protein complex of electron carners Q I Inner mitochondrial membrane IV III ATP synthase II FADH2 NADH+ Mitochondrial matrix H+ Cyt c FAD+ NAD+ 2 H+ + 1/2 O2 H2O ADP + (Carrying electrons from food) ATP Pi H+ Chemiosmosis Electron transport chain Electron transport and pumping of protons (H+), ATP synthesis powered by the flow which create an H+ gradient across the membrane Of H+ back across the membrane Oxidative phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.17 Fermentation 2 ADP + 2 Glucose 2 ATP Pi Glycolysis O– C O C O CH3 2 Pyruvate 2 NADH +2 H+ 2 NAD+ H 2 CO2 H H C OH C CH3 O CH3 2 Acetaldehyde 2 Ethanol (a) Alcohol fermentation 2 ADP + 2 Glucose P i 2 ATP Glycolysis 2 NAD+ O 2 NADH C O H C OH CH3 2 Lactate (b) Lactic acid fermentation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings O– C O C O CH3 2 Pyruvate Figure 9.18 Pyruvate as a key juncture in catabolism Glucose CYTOSOL Pyruvate No O2 present Fermentation O2 present Cellular respiration MITOCHONDRION Ethanol or lactate Acetyl CoA Citric acid cycle Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 9.19 The catabolism of various molecules from food Proteins Carbohydrates Amino acids Sugars Glycolysis Glucose Glyceraldehyde-3- P NH3 Pyruvate Acetyl CoA Citric acid cycle Oxidative phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fats Glycerol Fatty acids Figure 9.20 The control of cellular respiration Glucose AMP Glycolysis Fructose-6-phosphate – Inhibits Stimulates + Phosphofructokinase – Fructose-1,6-bisphosphate Inhibits Pyruvate Citrate ATP Acetyl CoA Citric acid cycle Oxidative phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings