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Cellular Respiration: Harvesting Chemical Energy ,[object Object],[object Object],Figure 9.1
[object Object],[object Object],Light energy ECOSYSTEM CO 2  + H 2 O Photosynthesis in chloroplasts Cellular respiration in mitochondria Organic molecules + O 2 ATP powers most cellular work  Heat energy Figure 9.2
[object Object],[object Object],[object Object]
[object Object],[object Object],[object Object],[object Object]
The Principle of Redox ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],Na  +  Cl  Na +   +  Cl – becomes oxidized (loses electron) becomes reduced (gains electron)
Oxidation of Organic Fuel Molecules During Cellular Respiration ,[object Object],[object Object],[object Object],[object Object],C 6 H 12 O 6  + 6O 2   6CO 2  + 6H 2 O + Energy becomes oxidized becomes reduced
[object Object],[object Object],Enzyme NAD + H O O O O – O O O – O O O P P CH 2 CH 2 HO OH H H HO OH HO H H N + C NH 2 H N H NH 2 N N Nicotinamide (oxidized form) NH 2 + 2[H] (from food) Dehydrogenase Reduction of NAD + Oxidation of NADH 2 e –  + 2 H + 2 e –  + H + NADH O H H N C + Nicotinamide (reduced form) N Figure 9.4
[object Object],[object Object],[object Object],[object Object],[object Object],(a) Uncontrolled reaction Free energy,  G H 2 O Explosive release of heat and light energy Figure 9.5 A H 2  +  1 / 2  O 2
[object Object],[object Object],[object Object]
2 H 1 / 2  O 2 (from food via NADH) 2 H +  +  2 e – 2 H + 2 e – H 2 O 1 / 2  O 2 Controlled release of energy for  synthesis of ATP ATP ATP ATP Electron transport chain  Free energy, G (b) Cellular respiration + Figure 9.5 B
The Stages of Cellular Respiration:  A Preview ,[object Object],[object Object],[object Object],[object Object]
[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
[object Object],Figure 9.6 Electrons carried via NADH Glycolsis Glucose Pyruvate ATP Substrate-level phosphorylation Electrons carried  via NADH and  FADH 2 Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis  ATP ATP Substrate-level phosphorylation Oxidative phosphorylation Mitochondrion Cytosol
[object Object],[object Object],Figure 9.7 Enzyme Enzyme ATP ADP Product Substrate P +
[object Object],[object Object],[object Object],[object Object],[object Object]
[object Object],[object Object],[object Object],Glycolysis Citric acid cycle Oxidative phosphorylation ATP ATP ATP 2 ATP 4 ATP used formed Glucose 2 ATP + 2  P 4 ADP + 4 P 2 NAD +  + 4 e -  + 4 H  + 2 NADH + 2 H + 2 Pyruvate + 2 H 2 O Energy investment phase Energy payoff phase Glucose 2 Pyruvate + 2 H 2 O 4 ATP formed – 2 ATP used 2 ATP 2 NAD +  + 4 e –  + 4 H  + 2 NADH + 2 H + Figure 9.8
A closer look at the energy investment phase ,[object Object],[object Object],[object Object],[object Object],1 ATP used – phosphorylation  1 ATP used – phosphorylation  Dihydroxyacetone phosphate Glyceraldehyde- 3-phosphate H H H H H OH OH HO HO CH 2 OH H H H H O H OH HO OH P CH 2 O P H O H HO HO H HO CH 2 OH P O CH 2 O CH 2 O P HO H HO H OH O P CH 2 C O CH 2 OH H C CHOH CH 2 O O P ATP ADP Hexokinase Glucose Glucose-6-phosphate Fructose-6-phosphate ATP ADP Phosphoglucoisomerase Phosphofructokinase Fructose- 1, 6-bisphosphate Aldolase Isomerase Glycolysis 1 2 3 4 5 CH 2 OH Oxidative phosphorylation Citric acid cycle Figure 9.9 A
A closer look at the energy payoff phase 6.  G3P  is oxidized, NAD+ is reduced to form NADH & the energy from this causes a P to bind to G3P changing it to  1,3-diphosphoglycerate 7.  1,3-diphosphoglycerate  phosphorylates an ADP to form ATP and becomes  3-phosphoglycerate 8.  3-phosphoglycerate  rearranges into  2-phosphoglycerate 9.  2-phosphoglycerate  goes through a dehydration reaction and becomes  PEP  (phosphoenolpyruvate) 10. another substrate level phosphorylation reaction producing ATP changes  PEP  into  pyruvate Redox reaction  2 ATP produced – phosphorylation  2 ATP produced – phosphorylation  Water lost  2 NAD + NADH 2 + 2 H + Triose phosphate dehydrogenase 2 P  i 2 P C CHOH O P O CH 2 O 2 O – 1, 3-Bisphosphoglycerate 2 ADP 2 ATP Phosphoglycerokinase CH 2 O P 2 C CHOH 3-Phosphoglycerate Phosphoglyceromutase O – C C CH 2 OH H O P 2-Phosphoglycerate 2 H 2 O 2 O – Enolase C C O P O CH 2 Phosphoenolpyruvate 2 ADP 2 ATP Pyruvate kinase O – C C O O CH 3 2 6 8 7 9 10 Pyruvate O Figure 9.8 B
Glycolysis Output ,[object Object],[object Object],Glycolysis Citric acid cycle Oxidative phosphorylation ATP ATP ATP 2 ATP 4 ATP used formed Glucose 2 ATP + 2  P 4 ADP + 4 P 2 NAD +  + 4 e -  + 4 H  + 2 NADH + 2 H + 2 Pyruvate + 2 H 2 O Energy investment phase Energy payoff phase Glucose 2 Pyruvate + 2 H 2 O 4 ATP formed – 2 ATP used 2 ATP 2 NAD +  + 4 e –  + 4 H  + 2 NADH + 2 H + Figure 9.8
[object Object],[object Object],[object Object]
[object Object],[object Object],[object Object],[object Object],[object Object],CYTOSOL MITOCHONDRION NADH +  H + NAD + 2 3 1 CO 2 Coenzyme A Pyruvate Acetyle CoA S CoA C CH 3 O Transport protein O – O O C C CH 3 Figure 9.10
[object Object],ATP 2 CO 2 3 NAD + 3 NADH + 3 H + ADP +  P   i FAD FADH 2 Citric acid cycle CoA CoA Acetyle CoA NADH + 3 H + CoA CO 2 Pyruvate (from glycolysis, 2 molecules per glucose) ATP ATP ATP Glycolysis Citric acid cycle Oxidative phosphorylation Figure 9.11
Figure 9.12 1. acetyl CoA  enters the cycle, CoA is removed and the remaining  2C acetyl group  is bound to  oxaloacetate  (which is 4C), making  citrate  (a 6C molecule) 2. a redox reaction produces an NADH and releases one C as CO 2  changing  citrate  into  alpha-ketoglutarate  (a 5C molecule) 3. another redox reaction and substrate level phosphorylation produces an ATP, another NADH and CO 2 , changing  alpha-ketoglutarate  into  succinate  (a 4C molecule) A closer look at the citric acid cycle = intermediate molecule (short lived)  Acetyl CoA NADH Oxaloacetate Citrate Malate Fumarate Succinate Succinyl CoA  -Ketoglutarate  Isocitrate Citric acid cycle S CoA CoA SH NADH NADH FADH 2 FAD GTP GDP NAD + ADP P  i NAD + CO 2 CO 2 CoA SH CoA SH CoA S H 2 O + H + + H + H 2 O C CH 3 O O C COO – CH 2 COO – COO – CH 2 HO C COO – CH 2 COO – COO – COO – CH 2 HC COO – HO CH COO – CH CH 2 COO – HO COO – CH HC COO – COO – CH 2 CH 2 COO – COO – CH 2 CH 2 C O COO – CH 2 CH 2 C O COO – 1 2 3 Glycolysis Oxidative phosphorylation NAD + + H + ATP Citric acid cycle Figure 9.12
A closer look at the citric acid cycle .. Cont. ,[object Object],[object Object],Acetyl CoA NADH Oxaloacetate Citrate Malate Fumarate Succinate Succinyl CoA  -Ketoglutarate  Isocitrate Citric acid cycle S CoA CoA SH NADH NADH FADH 2 FAD GTP GDP NAD + ADP P  i NAD + CO 2 CO 2 CoA SH CoA SH CoA S H 2 O + H + + H + H 2 O C CH 3 O O C COO – CH 2 COO – COO – CH 2 HO C COO – CH 2 COO – COO – COO – CH 2 HC COO – HO CH COO – CH CH 2 COO – HO COO – CH HC COO – COO – CH 2 CH 2 COO – COO – CH 2 CH 2 C O COO – CH 2 CH 2 C O COO – 4 5 Glycolysis Oxidative phosphorylation NAD + + H + ATP Citric acid cycle Figure 9.12
Total Output Citric Acid Cycle (total = 2 pyruvate) ,[object Object],[object Object],[object Object],[object Object],Acetyl CoA NADH Oxaloacetate Citrate Malate Fumarate Succinate Succinyl CoA  -Ketoglutarate  Isocitrate Citric acid cycle S CoA CoA SH NADH NADH FADH 2 FAD GTP GDP NAD + ADP P  i NAD + CO 2 CO 2 CoA SH CoA SH CoA S H 2 O + H + + H + H 2 O C CH 3 O O C COO – CH 2 COO – COO – CH 2 HO C COO – CH 2 COO – COO – COO – CH 2 HC COO – HO CH COO – CH CH 2 COO – HO COO – CH HC COO – COO – CH 2 CH 2 COO – COO – CH 2 CH 2 C O COO – CH 2 CH 2 C O COO – 4 5 Glycolysis Oxidative phosphorylation NAD + + H + ATP Citric acid cycle Figure 9.12
The Pathway of Electron Transport ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
Anatomy of a mitochondria
Overview ,[object Object],[object Object],[object Object],[object Object],[object Object],Animation: http://vcell.ndsu.nodak.edu/animations/etc/movie.htm H 2 O O 2 NADH FADH2 FMN Fe•S Fe•S Fe•S O FAD Cyt  b Cyt  c 1 Cyt  c Cyt  a Cyt  a 3 2 H  +  +  1  2 I II III IV Multiprotein complexes 0 10 20 30 40 50 Free energy ( G ) relative to O 2  (kcl/mol) Figure 9.13
Electron movement – Inner Membrane  NADH donates 2 electrons Mitochondrial Matrix  Intermembrane   2 H+ Ubiquinone Electrons passed 1 H+ at a time cytochrome  c 1 electron at a time 4 H+ Four electrons must be transferred to the oxidase complex in order for the next major reaction to occur 2 H20  3 H+ to synthesize one ATP from the substrates ADP and Pi (inorganic phosphate). FADH2 donates another electron
Chemiosmosis: The Energy-Coupling Mechanism ,[object Object],[object Object],ATP synthase animation:  http://vcell.ndsu.nodak.edu/animations/atpgradient/movie.htm INTERMEMBRANE SPACE H + H + H + H + H + H + H + H + P  i + ADP ATP 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. 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.   MITOCHONDRIAL MATRIX Figure 9.14
[object Object],[object Object],[object Object],[object Object]
[object Object],[object Object]
[object Object],Oxidative phosphorylation. electron transport and chemiosmosis Glycolysis ATP ATP ATP Inner Mitochondrial membrane H + H + H + H + H + ATP P  i Protein complex of electron  carners Cyt  c I II III IV (Carrying electrons from, food) NADH + FADH 2 NAD + FAD + 2 H +  +  1 / 2  O 2 H 2 O ADP + Electron transport chain Electron transport and pumping of protons (H + ), which create an H +  gradient across the membrane Chemiosmosis ATP synthesis powered by the flow Of H +  back across the membrane ATP synthase Q Oxidative phosphorylation Intermembrane space Inner mitochondrial membrane Mitochondrial matrix Figure 9.15
An Accounting of ATP Production by Cellular Respiration – A summary! ,[object Object],Electron shuttles span membrane CYTOSOL 2 NADH 2 FADH 2 2 NADH 6 NADH  2 FADH 2 2 NADH Glycolysis Glucose 2 Pyruvate 2 Acetyl CoA Citric acid cycle Oxidative phosphorylation: electron transport and chemiosmosis MITOCHONDRION by substrate-level phosphorylation by substrate-level phosphorylation by oxidative phosphorylation, depending on which shuttle transports electrons from NADH in cytosol  Maximum per glucose: About 36 or 38 ATP + 2 ATP + 2 ATP + about 32 or 34 ATP or Figure 9.16
[object Object],[object Object],With the supply of NADH exhausted, the electron transport chain can no longer maintain the proton gradient that powers ATP synthase, and ATP synthesis comes to a stop.
[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
Types of Fermentation ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],2 ADP + 2 P 1 2 ATP Glycolysis Glucose 2 NAD + 2 NADH 2 Pyruvate 2 Acetaldehyde 2 Ethanol (a) Alcohol fermentation 2 ADP + 2 P 1 2 ATP Glycolysis Glucose 2 NAD + 2 NADH 2 Lactate (b) Lactic acid fermentation H H OH CH 3 C O  – O C C O CH 3 H C O CH 3 O – C O C O CH 3 O C O C OH H CH 3 CO 2 2 Figure 9.17
[object Object],[object Object],[object Object],[object Object],Glucose CYTOSOL Pyruvate No O 2  present Fermentation O 2  present Cellular respiration Ethanol or  lactate Acetyl CoA MITOCHONDRION Citric acid cycle Figure 9.18
[object Object],Amino  acids Sugars Glycerol Fatty acids Glycolysis Glucose Glyceraldehyde-3-  P Pyruvate Acetyl CoA NH 3 Citric acid cycle Oxidative phosphorylation Fats Proteins Carbohydrates Figure 9.19
Biosynthesis (Anabolic Pathways) ,[object Object],[object Object],[object Object],[object Object]
Regulation of Cellular Respiration via Feedback Mechanisms ,[object Object],[object Object],Example: excess ATP production serves to inhibit phosphofructokinase in glycolysis, AMP stimulates its production  ,[object Object],[object Object],Glucose Glycolysis Fructose-6-phosphate Phosphofructokinase Fructose-1,6-bisphosphate Inhibits Inhibits Pyruvate ATP Acetyl CoA Citric acid cycle Citrate Oxidative phosphorylation Stimulates AMP + – – Figure 9.20
Dihydroxyacetone phosphate Glyceraldehyde- 3-phosphate H H H H H OH OH HO HO CH 2 OH H H H H O H OH HO OH P CH 2 O P H O H HO HO H HO CH 2 OH P O CH 2 O CH 2 O P HO H HO H OH O P CH 2 C O CH 2 OH H C CHOH CH 2 O O P ATP ADP Hexokinase Glucose Glucose-6-phosphate Fructose-6-phosphate ATP ADP Phosphoglucoisomerase Phosphofructokinase Fructose- 1, 6-bisphosphate Aldolase Isomerase Glycolysis 1 2 3 4 5 CH 2 OH Oxidative phosphorylation Citric acid cycle Figure 9.9 A 2 NAD + NADH 2 + 2 H + Triose phosphate dehydrogenase 2 P  i 2 P C CHOH O P O CH 2 O 2 O – 1, 3-Bisphosphoglycerate 2 ADP 2 ATP Phosphoglycerokinase CH 2 O P 2 C CHOH 3-Phosphoglycerate Phosphoglyceromutase O – C C CH 2 OH H O P 2-Phosphoglycerate 2 H 2 O 2 O – Enolase C C O P O CH 2 Phosphoenolpyruvate 2 ADP 2 ATP Pyruvate kinase O – C C O O CH 3 2 6 8 7 9 10 Pyruvate O Figure 9.8 B
CYTOSOL MITOCHONDRION NADH +  H + NAD + 2 3 1 CO 2 Coenzyme A Pyruvate Acetyle CoA S CoA C CH 3 O Transport protein O – O O C C CH 3 Figure 9.10
Acetyl CoA NADH Oxaloacetate Citrate Malate Fumarate Succinate Succinyl CoA  -Ketoglutarate  Isocitrate Citric acid cycle S CoA CoA SH NADH NADH FADH 2 FAD GTP GDP NAD + ADP P  i NAD + CO 2 CO 2 CoA SH CoA SH CoA S H 2 O + H + + H + H 2 O C CH 3 O O C COO – CH 2 COO – COO – CH 2 HO C COO – CH 2 COO – COO – COO – CH 2 HC COO – HO CH COO – CH CH 2 COO – HO COO – CH HC COO – COO – CH 2 CH 2 COO – COO – CH 2 CH 2 C O COO – CH 2 CH 2 C O COO – 4 5 Glycolysis Oxidative phosphorylation NAD + + H + ATP Citric acid cycle Figure 9.12
Oxidative phosphorylation. electron transport and chemiosmosis Glycolysis ATP ATP ATP Inner Mitochondrial membrane H + H + H + H + H + ATP P  i Protein complex of electron  carners Cyt  c I II III IV (Carrying electrons from, food) NADH + FADH 2 NAD + FAD + 2 H +  +  1 / 2  O 2 H 2 O ADP + Electron transport chain Electron transport and pumping of protons (H + ), which create an H +  gradient across the membrane Chemiosmosis ATP synthesis powered by the flow Of H +  back across the membrane ATP synthase Q Oxidative phosphorylation Intermembrane space Inner mitochondrial membrane Mitochondrial matrix Figure 9.15

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09 cellrespiration text

  • 1.
  • 2.
  • 3.
  • 4.
  • 5.
  • 6.
  • 7.
  • 8.
  • 9.
  • 10. 2 H 1 / 2 O 2 (from food via NADH) 2 H + + 2 e – 2 H + 2 e – H 2 O 1 / 2 O 2 Controlled release of energy for synthesis of ATP ATP ATP ATP Electron transport chain Free energy, G (b) Cellular respiration + Figure 9.5 B
  • 11.
  • 12.
  • 13.
  • 14.
  • 15.
  • 16.
  • 17.
  • 18. A closer look at the energy payoff phase 6. G3P is oxidized, NAD+ is reduced to form NADH & the energy from this causes a P to bind to G3P changing it to 1,3-diphosphoglycerate 7. 1,3-diphosphoglycerate phosphorylates an ADP to form ATP and becomes 3-phosphoglycerate 8. 3-phosphoglycerate rearranges into 2-phosphoglycerate 9. 2-phosphoglycerate goes through a dehydration reaction and becomes PEP (phosphoenolpyruvate) 10. another substrate level phosphorylation reaction producing ATP changes PEP into pyruvate Redox reaction 2 ATP produced – phosphorylation 2 ATP produced – phosphorylation Water lost 2 NAD + NADH 2 + 2 H + Triose phosphate dehydrogenase 2 P i 2 P C CHOH O P O CH 2 O 2 O – 1, 3-Bisphosphoglycerate 2 ADP 2 ATP Phosphoglycerokinase CH 2 O P 2 C CHOH 3-Phosphoglycerate Phosphoglyceromutase O – C C CH 2 OH H O P 2-Phosphoglycerate 2 H 2 O 2 O – Enolase C C O P O CH 2 Phosphoenolpyruvate 2 ADP 2 ATP Pyruvate kinase O – C C O O CH 3 2 6 8 7 9 10 Pyruvate O Figure 9.8 B
  • 19.
  • 20.
  • 21.
  • 22.
  • 23. Figure 9.12 1. acetyl CoA enters the cycle, CoA is removed and the remaining 2C acetyl group is bound to oxaloacetate (which is 4C), making citrate (a 6C molecule) 2. a redox reaction produces an NADH and releases one C as CO 2 changing citrate into alpha-ketoglutarate (a 5C molecule) 3. another redox reaction and substrate level phosphorylation produces an ATP, another NADH and CO 2 , changing alpha-ketoglutarate into succinate (a 4C molecule) A closer look at the citric acid cycle = intermediate molecule (short lived) Acetyl CoA NADH Oxaloacetate Citrate Malate Fumarate Succinate Succinyl CoA  -Ketoglutarate Isocitrate Citric acid cycle S CoA CoA SH NADH NADH FADH 2 FAD GTP GDP NAD + ADP P i NAD + CO 2 CO 2 CoA SH CoA SH CoA S H 2 O + H + + H + H 2 O C CH 3 O O C COO – CH 2 COO – COO – CH 2 HO C COO – CH 2 COO – COO – COO – CH 2 HC COO – HO CH COO – CH CH 2 COO – HO COO – CH HC COO – COO – CH 2 CH 2 COO – COO – CH 2 CH 2 C O COO – CH 2 CH 2 C O COO – 1 2 3 Glycolysis Oxidative phosphorylation NAD + + H + ATP Citric acid cycle Figure 9.12
  • 24.
  • 25.
  • 26.
  • 27. Anatomy of a mitochondria
  • 28.
  • 29. Electron movement – Inner Membrane NADH donates 2 electrons Mitochondrial Matrix Intermembrane 2 H+ Ubiquinone Electrons passed 1 H+ at a time cytochrome c 1 electron at a time 4 H+ Four electrons must be transferred to the oxidase complex in order for the next major reaction to occur 2 H20 3 H+ to synthesize one ATP from the substrates ADP and Pi (inorganic phosphate). FADH2 donates another electron
  • 30.
  • 31.
  • 32.
  • 33.
  • 34.
  • 35.
  • 36.
  • 37.
  • 38.
  • 39.
  • 40.
  • 41.
  • 42. Dihydroxyacetone phosphate Glyceraldehyde- 3-phosphate H H H H H OH OH HO HO CH 2 OH H H H H O H OH HO OH P CH 2 O P H O H HO HO H HO CH 2 OH P O CH 2 O CH 2 O P HO H HO H OH O P CH 2 C O CH 2 OH H C CHOH CH 2 O O P ATP ADP Hexokinase Glucose Glucose-6-phosphate Fructose-6-phosphate ATP ADP Phosphoglucoisomerase Phosphofructokinase Fructose- 1, 6-bisphosphate Aldolase Isomerase Glycolysis 1 2 3 4 5 CH 2 OH Oxidative phosphorylation Citric acid cycle Figure 9.9 A 2 NAD + NADH 2 + 2 H + Triose phosphate dehydrogenase 2 P i 2 P C CHOH O P O CH 2 O 2 O – 1, 3-Bisphosphoglycerate 2 ADP 2 ATP Phosphoglycerokinase CH 2 O P 2 C CHOH 3-Phosphoglycerate Phosphoglyceromutase O – C C CH 2 OH H O P 2-Phosphoglycerate 2 H 2 O 2 O – Enolase C C O P O CH 2 Phosphoenolpyruvate 2 ADP 2 ATP Pyruvate kinase O – C C O O CH 3 2 6 8 7 9 10 Pyruvate O Figure 9.8 B
  • 43. CYTOSOL MITOCHONDRION NADH + H + NAD + 2 3 1 CO 2 Coenzyme A Pyruvate Acetyle CoA S CoA C CH 3 O Transport protein O – O O C C CH 3 Figure 9.10
  • 44. Acetyl CoA NADH Oxaloacetate Citrate Malate Fumarate Succinate Succinyl CoA  -Ketoglutarate Isocitrate Citric acid cycle S CoA CoA SH NADH NADH FADH 2 FAD GTP GDP NAD + ADP P i NAD + CO 2 CO 2 CoA SH CoA SH CoA S H 2 O + H + + H + H 2 O C CH 3 O O C COO – CH 2 COO – COO – CH 2 HO C COO – CH 2 COO – COO – COO – CH 2 HC COO – HO CH COO – CH CH 2 COO – HO COO – CH HC COO – COO – CH 2 CH 2 COO – COO – CH 2 CH 2 C O COO – CH 2 CH 2 C O COO – 4 5 Glycolysis Oxidative phosphorylation NAD + + H + ATP Citric acid cycle Figure 9.12
  • 45. Oxidative phosphorylation. electron transport and chemiosmosis Glycolysis ATP ATP ATP Inner Mitochondrial membrane H + H + H + H + H + ATP P i Protein complex of electron carners Cyt c I II III IV (Carrying electrons from, food) NADH + FADH 2 NAD + FAD + 2 H + + 1 / 2 O 2 H 2 O ADP + Electron transport chain Electron transport and pumping of protons (H + ), which create an H + gradient across the membrane Chemiosmosis ATP synthesis powered by the flow Of H + back across the membrane ATP synthase Q Oxidative phosphorylation Intermembrane space Inner mitochondrial membrane Mitochondrial matrix Figure 9.15