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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 • Energy – Flows into an ecosystem as sunlight and leaves as heat Light energy ECOSYSTEM Photosynthesis in chloroplasts Organic CO2 + H2O + O2 Cellular molecules respiration in mitochondria ATP powers most cellular work Figure 9.2 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Heat energy Glucose CYTOSOL Pyruvate No O2 present Fermentation O2 present Cellular respiration MITOCHONDRION Ethanol or lactate Acetyl CoA Citric acid cycle Figure 9.18 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Electrons carried via NADH and FADH2 Electrons carried via NADH Citric acid cycle Glycolsis Pyruvate Glucose Cytosol Mitochondrion ATP Figure 9.6 Oxidative phosphorylation: electron transport and chemiosmosis Substrate-level phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings ATP Substrate-level phosphorylation ATP Oxidative phosphorylation The Stages of Cellular Respiration: A Preview • Respiration is a cumulative function of three metabolic stages – Glycolysis – The citric acid cycle (Krebs Cycle) – Oxidative phosphorylation (Electron Transport) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Glycolysis – Breaks down glucose into two molecules of pyruvate • The citric acid cycle – Completes the breakdown of glucose Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Oxidative phosphorylation – Is driven by the electron transport chain – Generates ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • An overview of cellular respiration Electrons carried via NADH and FADH2 Electrons carried via NADH Citric acid cycle Glycolsis Pyruvate Glucose Cytosol Mitochondrion ATP Figure 9.6 Oxidative phosphorylation: electron transport and chemiosmosis Substrate-level phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings ATP Substrate-level phosphorylation ATP Oxidative phosphorylation • Both glycolysis and the citric acid cycle – Can generate ATP by substrate-level phosphorylation Enzyme Enzyme ADP P Substrate + Figure 9.7 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Product ATP • Concept 9.2: Glycolysis harvests energy by oxidizing glucose to pyruvate • Glycolysis – Means “splitting of sugar” – Breaks down glucose into pyruvate – Occurs in the cytoplasm of the cell Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Glycolysis consists of two major phases – Energy investment phase – Energy payoff phase Citric acid cycle Glycolysis Oxidative phosphorylation ATP ATP ATP Energy investment phase Glucose 2 ATP + 2 P 2 ATP used Energy payoff phase 4 ADP + 4 P 2 NAD+ + 4 e- + 4 H + 4 ATP formed 2 NADH + 2 H+ 2 Pyruvate + 2 H2O Glucose 4 ATP formed – 2 ATP used Figure 9.8 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2 NAD+ + 4 e– + 4 H + 2 Pyruvate + 2 H2O 2 ATP + 2 H+ 2 NADH CH2OH HH H HO H HO OH H OH Glycolysis Glucose ATP 1 Hexokinase ADP CH2OH P HH OH OH H HO H OH Glucose-6-phosphate 2 Phosphoglucoisomerase CH2O P O CH2OH H HO HO H HO H Fructose-6-phosphate ATP 3 Phosphofructokinase ADP P O CH2 O CH2 O P HO H OH HO H Fructose1, 6-bisphosphate 4 Aldolase 5 H P O CH2 Isomerase C O C O CHOH CH2OH CH2 O P Figure 9.9 A Dihydroxyacetone phosphate Glyceraldehyde3-phosphate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Citric Oxidative acid cycle phosphorylation 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 Figure 9.8 B CH3 Pyruvate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Before the citric acid cycle can begin – Pyruvate must first be converted to acetyl CoA, which links the cycle to glycolysis CYTOSOL MITOCHONDRION NAD+ NADH + H+ O– S CoA C O 2 C C O O 1 3 CH3 Pyruvate Transport protein Figure 9.10 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings CH3 Acetyle CoA CO2 Coenzyme A • An overview of the citric acid cycle Pyruvate (from glycolysis, 2 molecules per glucose) Glycolysis Citric acid cycle ATP ATP Oxidative phosphorylatio n ATP CO2 CoA NADH + 3 H+ Acetyle CoA CoA CoA Citric acid cycle 2 CO2 3 NAD+ FADH2 FAD 3 NADH + 3 H+ ADP + P i ATP Figure 9.11 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Glycolysis Citric Oxidative acid phosphorylation cycle S CoA C O CH3 Acetyl CoA CoA SH O NADH + H+ C COO– COO– 1 CH2 COO– NAD+ 8 Oxaloacetate HO C COO– COO– CH2 COO– HO CH H2O CH2 CH2 2 HC COO– COO– Malate Figure CH2 HO Citrate 9.12 COO– Isocitrate COO– H2O COO– CH CO2 Citric acid cycle 7 3 NAD+ COO– Fumarate HC CH CH2 CoA SH 6 CoA SH COO– FAD CH2 CH2 COO– C O Succinate Pi S CoA GTP GDP Succinyl CoA ADP ATP Figure 9.12 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 4 C O COO– CH2 5 CH2 FADH2 COO– NAD+ NADH + H+ + H+ a-Ketoglutarate CH2 COO– NADH CO2 • At the end of the chain – Electrons are passed to oxygen, forming water NADH 50 Free energy (G) relative to O2 (kcl/mol) FADH2 40 FMN I Fe•S Fe•S II O 30 Multiprotein complexes FAD III Cyt b Fe•S 20 Cyt c1 IV Cyt c Cyt a Cyt a3 10 0 Figure 9.13 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2 H + + 12 O2 H2 O Chemiosmosis: The Energy-Coupling Mechanism • ATP synthase – Is the enzyme that actually makes ATP 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 Figure 9.14 MITOCHONDRIAL MATRIX Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings ATP A rod (for “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. • Chemiosmosis – Is an energy-coupling mechanism that uses energy in the form of a H+ gradient across a membrane to drive cellular work Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Chemiosmosis and the electron transport chain Oxidative phosphorylation. electron transport and chemiosmosis Glycolysis ATP Inner Mitochondrial membrane ATP 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 + ATP synthesis powered by the flow Electron transport and pumping of protons (H ), + + which create an H gradient across the membrane Of H back across the membrane Figure 9.15 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Oxidative phosphorylation • There are three main processes in this metabolic enterprise 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 ATP 2 FADH2 Oxidative phosphorylation: electron transport and chemiosmosis + about 32 or 34 ATP by substrate-level by oxidative phosphorylation, depending on which shuttle transports electrons phosphorylation from NADH in cytosol About 36 or 38 ATP Figure 9.16 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 glyocolysis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In alcohol fermentation – Pyruvate is converted to ethanol in two steps, one of which releases CO2 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • During lactic acid fermentation – Pyruvate is reduced directly to NADH to form lactate as a waste product Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2 ADP + 2 P1 2 ATP O– C O Glucose Glycolysis C O CH3 2 Pyruvate 2 NADH 2 NAD+ H 2 CO2 H H C OH C O CH3 CH3 2 Ethanol 2 Acetaldehyde (a) Alcohol fermentation 2 ADP + 2 Glucose P1 2 ATP Glycolysis O– C O C O O 2 NAD+ 2 NADH C O H C OH CH3 2 Lactate Figure 9.17 (b) Lactic acid fermentation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings CH3 Fermentation and Cellular Respiration Compared • Both fermentation and cellular respiration – Use glycolysis to oxidize glucose and other organic fuels to pyruvate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Pyruvate is a key juncture in catabolism Glucose CYTOSOL Pyruvate No O2 present Fermentation O2 present Cellular respiration MITOCHONDRION Ethanol or lactate Figure 9.18 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Acetyl CoA Citric acid cycle • The catabolism of various molecules from food Proteins Carbohydrates Amino acids Sugars Fats Glycerol Glycolysis Glucose Glyceraldehyde-3- P NH3 Pyruvate Acetyl CoA Citric acid cycle Figure 9.19 Oxidative phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fatty acids • The control of cellular respiration Glucose Glycolysis Fructose-6-phosphate – Inhibits AMP Stimulates + Phosphofructokinase – Fructose-1,6-bisphosphate Inhibits Pyruvate Citrate ATP Acetyl CoA Citric acid cycle Figure 9.20 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Oxidative phosphorylation