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Chapter 9. Cellular Respiration: Harvesting Chemical Energy. Overview: Life Is Work Living cells Require transfusions of energy from outside sources to perform their many tasks. Figure 9.1. The giant panda Obtains energy for its cells by eating plants. Light energy. ECOSYSTEM.
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Chapter 9 Cellular Respiration: Harvesting Chemical Energy
Overview: Life Is Work • Living cells • Require transfusions of energy from outside sources to perform their many tasks
Figure 9.1 • The giant panda • Obtains energy for its cells by eating plants
Light energy ECOSYSTEM Photosynthesisin chloroplasts Organicmolecules CO2 + H2O + O2 Cellular respirationin mitochondria ATP powers most cellular work Heatenergy Figure 9.2 • Energy • Flows into an ecosystem as sunlight and leaves as heat
Concept 9.1: Catabolic pathways yield energy by oxidizing organic fuels
Catabolic Pathways and Production of ATP • The breakdown of organic molecules is exergonic
One catabolic process, fermentation • Is a partial degradation of sugars that occurs without oxygen
Cellular respiration • Is the most prevalent and efficient catabolic pathway • Consumes oxygen and organic molecules such as glucose • Yields ATP
To keep working • Cells must regenerate ATP • Catabolic pathways yield energy • Due to the transfer of electrons • Redox reactions • Transfer electrons from one reactant to another by oxidation and reduction
In oxidation • A substance loses electrons, or is oxidized • In reduction • A substance gains electrons, or is reduced
becomes oxidized(loses electron) Na + Cl Na+ + Cl– becomes reduced(gains electron) • Examples of redox reactions
Products Reactants becomes oxidized + + + Energy 2O2 CO2 2 H2O CH4 becomes reduced H C C O O O O H O H H H H Oxygen(oxidizingagent) Methane(reducingagent) Carbon dioxide Water Figure 9.3 • Some redox reactions • Do not completely exchange electrons • Change the degree of electron sharing in covalent bonds
becomes oxidized C6H12O6 + 6O2 6CO2 + 6H2O + Energy becomes reduced Oxidation of Organic Fuel Molecules During Cellular Respiration • During cellular respiration • Glucose is oxidized and oxygen is reduced
Stepwise Energy Harvest via NAD+ and the Electron Transport Chain • Cellular respiration • Oxidizes glucose in a series of steps
2 e– + 2 H+ 2 e– + H+ NAD+ NADH H Dehydrogenase O O H H Reduction of NAD+ + + 2[H] C NH2 NH2 C (from food) Oxidation of NADH N N+ Nicotinamide(reduced form) Nicotinamide(oxidized form) CH2 O O O O– P O H H OH O O– HO P NH2 HO CH2 O N N H N H N O H H HO OH Figure 9.4 • Electrons from organic compounds • Are usually first transferred to NAD+, a coenzyme
NADH, the reduced form of NAD+ • Passes the electrons to the electron transport chain
H2 + 1/2 O2 Explosiverelease ofheat and lightenergy (a) Uncontrolled reaction Free energy, G Figure 9.5 A H2O • If electron transfer is not stepwise • A large release of energy occurs • As in the reaction of hydrogen and oxygen to form water
The electron transport chain • Passes electrons in a series of steps instead of in one explosive reaction • Uses the energy from the electron transfer to form ATP
2 H + 1/2 O2 (from food via NADH) Controlled release of energy for synthesis ofATP 2 H+ + 2 e– ATP ATP Free energy, G Electron transport chain ATP 2 e– 1/2 O2 2 H+ H2O Figure 9.5 B (b) Cellular respiration
The Stages of Cellular Respiration: A Preview • Respiration is a cumulative function of three metabolic stages • Glycolysis • The citric acid cycle • 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
Electrons carried via NADH and FADH2 Electrons carried via NADH Oxidativephosphorylation:electron transport andchemiosmosis Citric acid cycle Glycolsis Pyruvate Glucose Cytosol Mitochondrion ATP ATP ATP Substrate-level phosphorylation Oxidative phosphorylation Substrate-level phosphorylation Figure 9.6 • An overview of cellular respiration
Enzyme Enzyme ADP P Substrate + ATP Product Figure 9.7 • Both glycolysis and the citric acid cycle • Can generate ATP by substrate-level phosphorylation
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
Glycolysis Oxidativephosphorylation Citricacidcycle ATP ATP ATP Energy investment phase Glucose P 2 ATP + 2 used 2 ATP Energy payoff phase formed P 4 ATP 4 ADP + 4 2 NAD+ + 4 e- + 4 H + + 2 H+ 2 NADH 2 Pyruvate + 2 H2O Glucose 2 Pyruvate + 2 H2O 4 ATP formed – 2 ATP used 2 ATP + 2 H+ 2 NADH 2 NAD+ + 4 e– + 4 H + Figure 9.8 • Glycolysis consists of two major phases • Energy investment phase • Energy payoff phase
CH2OH Citric acid cycle H H Oxidative phosphorylation H Glycolysis H HO HO OH H OH Glucose 1 2 3 5 4 ATP Hexokinase ADP CH2OH P O H H H H OH HO H OH Glucose-6-phosphate Phosphoglucoisomerase CH2O P O CH2OH H HO HO H H HO Fructose-6-phosphate ATP Phosphofructokinase ADP CH2 O O CH2 P P O HO H OH H HO Fructose- 1, 6-bisphosphate Aldolase H O CH2 P Isomerase C O O C CHOH CH2OH O CH2 P Dihydroxyacetone phosphate Glyceraldehyde- 3-phosphate Figure 9.9 A
2 NAD+ Triose phosphate dehydrogenase P i 2 2 NADH + 2 H+ 10 7 9 8 6 2 O C O P CHOH P CH2 O 1, 3-Bisphosphoglycerate 2 ADP Phosphoglycerokinase 2 ATP O– 2 C CHOH O P CH2 3-Phosphoglycerate Phosphoglyceromutase O– 2 C O P C H O CH2OH 2-Phosphoglycerate Enolase 2 H2O O– 2 C O P C O CH2 Phosphoenolpyruvate 2 ADP Pyruvate kinase 2 ATP O– 2 C O C O CH3 Figure 9.8 B Pyruvate
Concept 9.3: The citric acid cycle completes the energy-yielding oxidation of organic molecules • The citric acid cycle • Takes place in the matrix of the mitochondrion
CYTOSOL MITOCHONDRION + H+ NAD+ NADH O– CoA S 2 C O C O C O CH3 1 3 CH3 Acetyle CoA Pyruvate CO2 Coenzyme A Transport protein Figure 9.10 • Before the citric acid cycle can begin • Pyruvate must first be converted to acetyl CoA, which links the cycle to glycolysis
Pyruvate(from glycolysis,2 molecules per glucose) Oxidativephosphorylation Glycolysis Citricacidcycle ATP ATP ATP CO2 CoA NADH + 3 H+ Acetyle CoA CoA CoA Citricacidcycle 2 CO2 3 NAD+ FADH2 FAD 3 NADH + 3 H+ ADP + Pi ATP Figure 9.11 • An overview of the citric acid cycle
Citric acid cycle Oxidative phosphorylation Glycolysis S CoA C O CH3 Acetyl CoA CoA SH H2O O C COO– NADH 1 COO– CH2 + H+ COO– CH2 COO– NAD+ Oxaloacetate 8 C COO– HO CH2 2 CH2 HC COO– COO– COO– HO CH HO CH Malate Citrate COO– CH2 Isocitrate COO– CO2 Citric acid cycle 3 H2O 7 NAD+ COO– NADH COO– CH + H+ Fumarate CH2 CoA SH HC a-Ketoglutarate CH2 COO– C O 4 6 SH CoA COO– COO– COO– CH2 5 CH2 FADH2 CO2 CH2 CH2 NAD+ FAD C O COO– Succinate NADH CoA S P i + H+ Succinyl CoA GDP GTP ADP ATP Figure 9.12 Figure 9.12
Concept 9.4: During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis • NADH and FADH2 • Donate electrons to the electron transport chain, which powers ATP synthesis via oxidative phosphorylation
The Pathway of Electron Transport • In the electron transport chain • Electrons from NADH and FADH2 lose energy in several steps
NADH 50 FADH2 Multiproteincomplexes I 40 FAD FMN II Fe•S Fe•S O III Cyt b 30 Fe•S Cyt c1 IV Free energy (G) relative to O2 (kcl/mol) Cyt c Cyt a Cyt a3 20 10 0 O2 2 H + + 12 Figure 9.13 H2O • At the end of the chain • Electrons are passed to oxygen, forming water
A rotor within the membrane spins clockwise whenH+ flows past it down the H+ gradient. INTERMEMBRANE SPACE H+ H+ H+ H+ H+ H+ H+ A stator anchoredin the membraneholds the knobstationary. A rod (for “stalk”)extending into the knob alsospins, activatingcatalytic sites inthe knob. H+ Three catalytic sites in the stationary knobjoin inorganic Phosphate to ADPto make ATP. ADP + ATP P i MITOCHONDRIAL MATRIX Figure 9.14 Chemiosmosis: The Energy-Coupling Mechanism • ATP synthase • Is the enzyme that actually makes 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
The resulting H+ gradient • Stores energy • Drives chemiosmosis in ATP synthase • Is referred to as a proton-motive force
Chemiosmosis • Is an energy-coupling mechanism that uses energy in the form of a H+ gradient across a membrane to drive cellular work
Inner Mitochondrial membrane Oxidative phosphorylation. electron transport and chemiosmosis Glycolysis ATP ATP ATP H+ H+ H+ H+ Cyt c Protein complex of electron carners Intermembrane space Q IV I III ATP synthase Inner mitochondrial membrane II H2O FADH2 2 H+ + 1/2 O2 FAD+ NADH+ NAD+ ATP ADP + P i (Carrying electrons from, food) H+ Mitochondrial matrix Chemiosmosis ATP synthesis powered by the flow Of H+ back across the membrane Electron transport chain Electron transport and pumping of protons (H+), which create an H+ gradient across the membrane Figure 9.15 Oxidative phosphorylation • Chemiosmosis and the electron transport chain
An Accounting of ATP Production by Cellular Respiration • During respiration, most energy flows in this sequence • Glucose to NADH to electron transport chain to proton-motive force to ATP
Electron shuttles span membrane MITOCHONDRION CYTOSOL 2 NADH or 2 FADH2 2 FADH2 2 NADH 2 NADH 6 NADH Glycolysis Oxidative phosphorylation: electron transport and chemiosmosis Citric acid cycle 2 Acetyl CoA 2 Pyruvate Glucose + 2 ATP + 2 ATP + about 32 or 34 ATP by oxidative phosphorylation, depending on which shuttle transports electrons from NADH in cytosol by substrate-level phosphorylation by substrate-level phosphorylation About 36 or 38 ATP Maximum per glucose: Figure 9.16 • There are three main processes in this metabolic enterprise
About 40% of the energy in a glucose molecule • Is transferred to ATP during cellular respiration, making approximately 38 ATP
Concept 9.5: Fermentation enables some cells to produce ATP without the use of oxygen • Cellular respiration • Relies on oxygen to produce ATP • In the absence of oxygen • Cells can still produce ATP through fermentation
Glycolysis • Can produce ATP with or without oxygen, in aerobic or anaerobic conditions • Couples with fermentation to produce ATP
Types of Fermentation • Fermentation consists of • Glycolysis plus reactions that regenerate NAD+, which can be reused by glyocolysis
In alcohol fermentation • Pyruvate is converted to ethanol in two steps, one of which releases CO2