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Chapter 9 Cellular Respiration: Harvesting Chemical Energy 高醫 生物醫學暨環境生物學系 助理教授 張學偉 http://genomed.dlearn.kmu.edu.tw 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 • To keep working, cells must regenerate ATP 本圖補充 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 Glycolysis 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 Concept 1: Catabolic pathways yield energy by oxidizing organic fuels 2: Glycolysis harvests energy by oxidizing glucose to pyruvate 3: The citric acid cycle completes the energy-yielding oxidation of organic molecules 4: During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis 5: Fermentation enables some cells to produce ATP without the use of oxygen 6: Glycolysis and the citric acid cycle connect to many other metabolic pathways Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 9.1: Catabolic pathways yield energy by oxidizing organic fuels Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Catabolic Pathways and Production of ATP Catabolic pathways (異化) yield energy by oxidizing organic fuels 同化作用 本圖為補充 exergonic (產能的) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings endergonic 耗能的 • One catabolic process, fermentation (發酵) – Is a partial degradation of sugars that occurs without oxygen 稍後再講 Conecept 5: Fermentation enables some cells to produce ATP without the use of oxygen Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Cellular respiration – Is the most prevalent and efficient catabolic pathway – Consumes oxygen (with O2 as the ultimate electron acceptor) and organic molecules such as glucose – Yields ATP It is called cellular respiration to distinguish it from the respiration of breathing. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Redox Reactions: Oxidation and Reduction • Catabolic pathways yield energy – Due to the transfer of electrons The Principle of Redox • Redox reactions – Transfer electrons from one reactant to another by oxidation and reduction Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Definition of Oxidation & Reduction and its agent • In oxidation – A substance loses electrons, or is oxidized Reducting Agent • In reduction – A substance gains electrons, or is reduced Oxidating Agent becomes oxidized (loses electron) Na + Cl Na+ + becomes reduced (gains electron) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Cl– • Some redox reactions – Do not completely exchange electrons – Change the degree of electron sharing in covalent bonds 注意 : blue ball位置 Products Reactants becomes oxidized + CH4 2O2 2 H2O O O C O H O H H H O 注意 : blue ball位置 H Methane (reducing agent) + Energy becomes reduced H C + CO2 Oxygen (oxidizing agent) Figure 9.3 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Carbon dioxide Water Oxidation of Organic Fuel Molecules During Cellular Respiration • During cellular respiration: - Glucose is oxidized in a series of steps and oxygen is reduced becomes oxidized C6H12O6 + 6O2 6CO2 + 6H2O + Energy △G = - 686 Kcal becomes reduced △G < 0 1. spontaneous reaction (without an input of energy) 2. Liberating energy Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Stepwise Energy Harvest via NAD+ and the Electron Transport Chain NAD+ = Nicotinamide adenine dinucleotide 2 e– + 2 H+ NAD+ H O O– O O P O H – O P O HO NH2 O N+ Nicotinamide (oxidized form) H OH HO CH2 O H HO H OH N H H + 2[H] • Nucleosides = Sugar (from food) Oxidation of NADH phosphate) N H N Reduction of NAD+ NH2 N H+ NADH • Nucleotide = Base + Sugar Dehydrogenase O + Phosphate O C CH2 2 e– + H+ adenine Figure 9.4 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings H C NH2 + Base (no N Nicotinamide (reduced form) + H+ • Electrons from organic compounds – Are usually first transferred to NAD+, a coenzyme electron 2 e– + proton 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 H H HO N H OH Reduction of NAD+ + 2[H] (from food) Oxidation of NADH Hydrogen atom H O C H N H+ NADH NH2 + H+ Nicotinamide (reduced form) NH2 N O 2 e– + H+ N H 2H NAD+ + H+ +2e= NAD+ + 2H Figure 9.4 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2H+ +2e- (oxidation) NADH (reduction) NADH + H+ Ex. A B + 2H+ +2e- (oxidation) (+) FAD + 2H+ +2e- FADH2 (reduction) = A+ FAD <=> B+ FADH2 A+ FAD <=> B+ FADH2 A+ NAD(P) <=> B+ NAD(P)H A is the electron donor (so it is oxidized) FAD & NAD(P) is the electron acceptor (so it is reduced). The products of the reaction are B (oxidized state) FADH2 or NAD(P)H (reduced state) . Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 圖形為補充 • NADH passes the electrons to the electron transport chain Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • If electron transfer is not stepwise – A large release of energy occurs – As in the reaction of hydrogen and oxygen to form water Free energy, G H2 + 1/2 O2 Figure 9.5 A Explosive release of heat and light energy H2O Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (a) Uncontrolled reaction 比較 The electron transport chain 2H • Passes electrons in a series of steps instead of in one explosive reaction 2 O2 1/ O2 (from food via NADH) 2 H+ + 2 e– Controlled release of energy for synthesis of ATP Free energy, G • Uses the energy from the electron transfer to form ATP 1/ + ATP ATP ATP 2 e– 2 H+ H2O Figure 9.5 B Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (b) Cellular respiration 2 • An overview of cellular respiration Electrons carried via NADH and FADH2 Electrons carried via NADH 1 2 Glycolsis Pyruvate Glucose Cytosol Oxidative 3phosphorylation: electron transport and chemiosmosis Mitochondrion ATP Figure 9.6 Citric acid cycle Substrate-level phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings ATP Substrate-level phosphorylation ATP Oxidative phosphorylation • Glycolysis – Breaks down glucose into two molecules of pyruvate • The citric acid cycle – Completes the breakdown of glucose • Oxidative phosphorylation – Is driven by the electron transport chain – Generates ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings substrate-level phosphorylation • generate ATP from glycolysis and TCA cycle 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 ATP Citric acid cycle Oxidative phosphorylation ATP ATP Energy investment phase • Glycolysis consists of two major phases Glucose 2 ATP + 2 – Energy investment phase used 2 ATP Energy payoff phase 4 ADP + 4 – Energy payoff phase P 4 ATP P formed 2 NADH 2 NAD+ + 4 e- + 4 H + + 2 H+ 2 Pyruvate + 2 H2O Figure 9.8 Net Rx Glucose 4 ATP formed – 2 ATP used 2 NAD+ + 4 e– + 4 H + 2 Pyruvate + 2 H2O 2 ATP + 2 H+ 2 NADH Net Rx: Glucose + 2ADP+ 2Pi + NAD+ 2 Pyruvate + 2ATP + 2NADH + 2 H+ Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings investment phase payoff phase Glycolysis Citric acid cycle Oxidative phosphorylation 2 CH2 O P 1, 3-Bisphosphoglycerate 2 ADP 7 Phosphoglycerokinase ADP 2 ATP H OH Glucose-6-phosphate 2 Phosphoglucoisomerase P O CH2OH H HO OH H H HO Fructose-6-phosphate CH2O 3 ATP O C O CHOH 1 Hexokinase CH2O P HH O H OH H OH HO 2 Pi 2 NADH + 2 H+ P ATP 6 Triose phosphate dehydrogenase 2 NAD+ CH2OH OH HH OH H HO OH H OH Glucose Kinase isomerase Mutase Aldolase enolase Dehydrogenase Phosphofructokinase ADP 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 P O CH2 O CH2 O H HO OH H HO H Fructose1, 6-bisphosphate 2 P O– C O C O 4 Aldolase Pyruvate kinase 2 ATP 5 P O CH2 C O CH2OH Dihydroxyacetone phosphate Isomerase H 2 C O Figure 9.9 A Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings O– C O CHOH CH2 O P Glyceraldehyde3-phosphate P CH2 Phosphoenolpyruvate 2 ADP 10 C O Figure 9.8 CH3 Pyruvate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 9.3: The citric acid cycle completes the energy-yielding oxidation of organic molecules Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The citric acid cycle – Takes place in the matrix of the mitochondrion Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Before 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 O Pyruvate Oxidation C O 1 3 CH3 Pyruvate Acetyle CoA CO2 Coenzyme A From Vit B Transport protein Figure 9.10 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings CH3 • An overview of the citric acid cycle = TCA (Tricarboxyl acid) 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 = Krebs Cycle A closer look at the citric acid cycle 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 HO Citrate CH2 COO– Isocitrate COO– H2O CH 7 COO– CH CO2 Citric acid cycle 3 NAD+ COO– Fumarate HC CH2 CoA SH 6 CoA SH COO– FAD CH2 CH2 COO– C O Succinate 4 C O COO– CH2 5 CH2 FADH2 COO– + H+ a-Ketoglutarate CH2 COO– NADH Pi S CoA GTP GDP Succinyl NAD+ CO2 NADH + H+ CoA ADP ATP Figure 9.12 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings A closer look at the citric acid cycle Citric Oxidative acid phosphorylation cycle Glycolysis 2C 3C pyruvate S CoA C O CH3 NAD + CoA NADH + CO2 Acetyl CoA 4C O NADH + H+ CoA SH 6C C COO– COO– 1 CH2 COO– NAD+ 8 Oxaloacetate HO C ½ Glucose 4C COO– COO– CH2 COO– HO CH H2O CH2 CH2 2 HC COO– COO– Malate Figure CH2 HO Citrate 9.12 Isocitrate COO– 4C CH CO2 Citric acid cycle 7 3 NAD+ COO– Fumarate HC CH2 CoA SH CoA SH COO– FAD CH2 CH2 COO– C O Succinate 4C 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– 4C NAD+ NADH + H+ NADH + H+ a-Ketoglutarate CH2 COO– 6 6C COO– COO– H2O CH CO2 5C Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Energetics DHase Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 9.4: During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis 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 ATP Substrate-level phosphorylation ATP Oxidative phosphorylation • NADH and FADH2: Donate electrons to the electron transport chain (ETC), which powers ATP synthesis via oxidative phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Pathway of Electron Transport • In the ETC NADH 50 – Electrons from NADH and FADH2 lose energy in several steps FADH2 Free energy (G) relative to O2 (kcl/mol) 40 • At the end of the chain Multiprotein complexes FAD Fe•S Fe•S II O III Cyt b 30 Fe•S Cyt c1 IV Cyt c Cyt a Cyt a3 20 10 – Electrons are passed to oxygen, forming water 0 Figure 9.13 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings I FMN 2 H + + 12 O2 H2 O Chemiosmosis: The Energy-Coupling Mechanism • ATP synthase is the enzyme that actually makes ATP INTERMEMBRANE SPACE H+ rotor H+ H+ H+ H+ H+ H+ A within the membrane spins clockwise when H+ flows past it down the H+ gradient. stator A anchored in the membrane holds the knob stationary. 注意: H+& ATP的方向 rod H+ A (for “stalk”) extending into the knob also spins, activating catalytic sites in the knob. Three catalytic sites in the ADP + Pi Figure 9.14 MITOCHONDRIAL MATRIX Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings knob ATP stationary join inorganic Phosphate to ADP to make ATP. • At certain steps along the electron transport chain – Electron transfer causes protein complexes to pump H+ from the mitochondrial matrix to the intermembrane space. [逆反應; 與產生ATP時 的H+運送方向相反] 目的: resulting H+ gradient Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The resulting H+ gradient – Stores energy – Is referred to as a proton-motive force – Drives chemiosmosis in ATP synthase • 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 & 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 IV III Inner mitochondrial membrane ATP synthase II FADH2 NADH+ Mitochondrial matrix H+ Cyt c FAD+ NAD+ 2 H+ + 1/2 O2 H2O ADP + ATP Pi (Carrying electrons from, food) 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 Figure 9.15 NADH 3 ATP; FADH2 2 ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings An Accounting of ATP Production by 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 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 Shuttles for transfer of reducing equivalents from cytosol into mitochondria 補充 不考 DHAP/GA3P shuttle Malate/Asp shuttle Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Glycolysis (aerobic) Glucose +2ADP + 2Pi+ 2NAD+ 2 pyruvate +2ATP+2NADH+2H++2H2O aerobic Pyruvate Oxidation 1 Pyruvate + NAD+ + CoASH 1 Acetyl-CoA + 1 NADH + CO2 2 Pyruvate + 2 NAD+ + 2 CoASH 2 Acetyl-CoA + 2 NADH + 2 CO2 TCA 2 Acetyl-CoA + 4 H2O + 6 NAD+ + 2FAD + 2GDP + 2 Pi 2 CO2 + 6 NADH + 2 FADH2 + 2 CoA-SH + 2 GTP Glycolysis 2 ATP 2 NADH (2H) Pyruvate Oxidation TCA 2 (2x1) NADH Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2 (2x1) GTP/ATP 6 (2x3) NADH 2 (2x1) FADH2 Glycolysis (aerobic) Glycolysis 2 ATP 2 NADH (2H) Pyruvate Oxidation TCA 2 (2x1) NADH 2 (2x1) GTP/ATP 6 (2x3) NADH 2 (2x1) FADH2 NADH 3 ATP; FADH2 2 ATP EST & chemiosmosis ATP 8 or (6) shuttle 6 2 + 18 + 4 = 38 = 36 • About 40% of the energy in a glucose molecule – Is transferred to ATP during cellular respiration, making approximately 38 ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 9.5: Fermentation enables some cells to produce ATP without the use of oxygen Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Cellular respiration – Relies on oxygen to produce ATP • In the absence of oxygen [no cellular respiration] – Cells can still produce ATP through fermentation • fermentation is a partial degradation of sugars that occurs without oxygen Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Glycolysis – Can produce ATP with or without oxygen, in aerobic or anaerobic conditions • Fermentation consists of – Glycolysis + reactions that regenerate NAD+, which can be reused by glyocolysis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Types of Fermentation 2 ADP + 2 P1 2 ATP • alcohol fermentation O– C O Glucose Glycolysis C O 3C CH3 2 Pyruvate 2 NADH 2 NAD+ H 2 CO2 – Pyruvate is converted to ethanol in two steps, one of which releases CO2 H H C OH C O CH3 2C CH3 2 Ethanol 2 Acetaldehyde (a) Alcohol fermentation 2 ADP + 2 Glucose P1 • lactic acid fermentation 2 ATP Glycolysis O– C O C O O 2 NAD+ 2 NADH 3C CH3 C O 3C H C OH CH3 2 Lactate Figure 9.17 (b) Lactic acid fermentation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings – Pyruvate is reduced directly to NADH to form lactate as a waste product • Anaerobic metabolism Electrons carried via NADH 2 3x2 2x2 Citric acid cycle Glycolsis Pyruvate Glucose Cytosol lactate Or ethanol Oxidative phosphorylation: electron transport and chemiosmosis Mitochondrion 2 ATP Figure 9.6 Electrons carried via NADH and FADH2 Substrate-level phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 1x2 ATP Substrate-level phosphorylation 32-34 ATP Oxidative phosphorylation • 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 Summaries of Glycolysis 1. anaerobic Glucose -> 2 Lactate + 2 ATP (lactic acid fermentation) Glucose -> 2 Ethanol + 2 ATP (alcoholic fermentation) 2. aerobic Glucose -> 2 Pyruvate + 2ATP + 2NADH (aerobic subtotal) 註: 2 NADH -> 6 ATP (aerobic conversion: 氧化磷酸化) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Fermentation and Cellular Respiration Compared • 相同點 Use glycolysis to oxidize glucose and other organic fuels to pyruvate • 相異點 1.Differ in their final electron acceptor Fermentation by acetylaldehyde (ethanol ferment.); by pyruvate (lactate ferment.) Cellular Respiration by O2 2.Cellular respiration produces more ATP Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Evolutionary Significance of Glycolysis • Glycolysis – Occurs in nearly all organisms – 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 The Versatility of Catabolism • Catabolic pathways – Funnel electrons from many kinds of organic molecules into cellular respiration Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The catabolism of various molecules from food Proteins Carbohydrates Amino acids Sugars Fats Glycerol Fatty acids 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 b-oxidation Biosynthesis (Anabolic Pathways) • The body – Uses small molecules to build other substances • These small molecules – May come directly from food or through glycolysis or the citric acid cycle Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Regulation of Cellular Respiration via Feedback Mechanisms • Cellular respiration is controlled by allosteric enzymes (e.g., Phosphofructokinase) at key points in glycolysis and the citric acid cycle Allosteric regulation (see glossary) The binding of a molecule to a protein that affects the function of the protein at a different site. 補充圖 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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 Oxidative phosphorylation Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • ATP AMP + ppi