1. Cellular Respiration and Fermentation
Chapter 9
Cellular Respiration and Fermentation

2. PHOTOSYNTHESIS
CELLULAR RESPIRATION

3. 6 CO2 + 12 H2O + Light energy  C6H12O6 + 6 O2 + 6 H2O
PHOTOSYNTHESIS
6 CO H2O + Light energy  C6H12O6 + 6 O2 + 6 H2O
CELLULAR RESPIRATION
C6H12O6 + 6 O2  6 CO2 + 6 H2O + Energy (ATP + heat)

4. Overview: Life Is Work
Living cells require energy from outside sources
Some animals, such as the chimpanzee, obtain energy by eating plants, and some animals feed on other organisms that eat plants
For the Discovery Video Space Plants, go to Animation and Video Files.
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5. Energy flows into an ecosystem as sunlight and leaves as heat
Photosynthesis generates O2 and organic molecules, which are used in cellular respiration
Cells use chemical energy stored in organic molecules to regenerate ATP, which powers work
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6. Photosynthesis in chloroplasts Cellular respiration in mitochondria
Light energy
ECOSYSTEM
Photosynthesis in chloroplasts
 O2
Organic molecules
CO2  H2O
Cellular respiration in mitochondria
Figure 9.2 Energy flow and chemical recycling in ecosystems.
ATP powers most cellular work
ATP
Heat energy

7. Catabolic Pathways and Production of ATP
Fermentation is a partial degradation of sugars that occurs without O2
Aerobic respiration consumes organic molecules and O2 and yields ATP
Anaerobic respiration is similar to aerobic respiration but consumes compounds other than O2
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8. C6H12O6 + 6 O2  6 CO2 + 6 H2O + Energy (ATP + heat)
Cellular respiration includes both aerobic and anaerobic respiration but is often used to refer to aerobic respiration
Although carbohydrates, fats, and proteins are all consumed as fuel, it is helpful to trace cellular respiration with the sugar glucose
C6H12O6 + 6 O2  6 CO2 + 6 H2O + Energy (ATP + heat)
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9. Redox Reactions: Oxidation and Reduction
The transfer of electrons during chemical reactions releases energy stored in organic molecules
This released energy is ultimately used to synthesize ATP
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10. The Principle of Redox
Chemical reactions that transfer electrons between reactants are called oxidation-reduction reactions, or redox reactions
In oxidation, a substance loses electrons, or is oxidized
In reduction, a substance gains electrons, or is reduced (the amount of positive charge is reduced)
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11. becomes oxidized (loses electron) becomes reduced (gains electron)
Figure 9.UN01 In-text figure, p. 164

12. becomes oxidized becomes reduced
Figure 9.UN02 In-text figure, p. 164

13. The electron donor is called the reducing agent
The electron receptor is called the oxidizing agent
Some redox reactions do not transfer electrons but change the electron sharing in covalent bonds
An example is the reaction between methane and O2
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14. Methane (reducing agent) Oxygen (oxidizing agent)
Reactants
Products
becomes oxidized
Energy
becomes reduced
Figure 9.3 Methane combustion as an energy-yielding redox reaction.
Methane (reducing agent)
Oxygen (oxidizing agent)
Carbon dioxide
Water

15. Oxidation of Organic Fuel Molecules During Cellular Respiration
During cellular respiration, the fuel (such as glucose) is oxidized, and O2 is reduced
becomes oxidized
becomes reduced
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16. Stepwise Energy Harvest via NAD+ and the Electron Transport Chain
In cellular respiration, glucose and other organic molecules are broken down in a series of steps
Electrons from organic compounds are usually first transferred to NAD+, a coenzyme
As an electron acceptor, NAD+ functions as an oxidizing agent during cellular respiration
Each NADH (the reduced form of NAD+) represents stored energy that will synthesize ATP
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17. NAD
NADH
Dehydrogenase
Reduction of NAD
(from food)
Oxidation of NADH
Nicotinamide (oxidized form)
Nicotinamide (reduced form)
Figure 9.4 NAD as an electron shuttle.

18. Dehydrogenase
Figure 9.UN04 In-text figure, p. 166

19. NADH passes the electrons to the electron transport chain
Unlike an uncontrolled reaction, the electron transport chain passes electrons in a series of steps instead of one explosive reaction
O2 pulls electrons down the chain in an energy-yielding tumble
The energy yielded is used to regenerate ATP
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20. Controlled release of energy for synthesis of ATP
H2  1/2 O2
2 H 
1/2 O2
(from food via NADH)
Controlled release of energy for synthesis of ATP
2 H+  2 e
ATP
Explosive release of heat and light energy
ATP
Electron transport chain
Free energy, G
Free energy, G
ATP
2 e
Figure 9.5 An introduction to electron transport chains.
1/2 O2
2 H+
H2O
H2O
(a) Uncontrolled reaction
(b) Cellular respiration

21. The Stages of Cellular Respiration: A Preview
Harvesting of energy from glucose has three stages
Glycolysis (breaks down glucose into two molecules of pyruvate)
The citric acid cycle (completes the breakdown of glucose)
Oxidative phosphorylation (accounts for most of the ATP synthesis)
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22. Glycolysis (color-coded teal throughout the chapter) 1.
Pyruvate oxidation and the citric acid cycle (color-coded salmon) 2.
Oxidative phosphorylation: electron transport and chemiosmosis (color-coded violet) 3.
Figure 9.UN05 In-text figure, p. 167

23. Electrons carried via NADH Substrate-level phosphorylation
Glycolysis
Glucose
Pyruvate
CYTOSOL
MITOCHONDRION
Figure 9.6 An overview of cellular respiration.
ATP
Substrate-level phosphorylation

24. Electrons carried via NADH Electrons carried via NADH and FADH2
Pyruvate oxidation
Glycolysis
Citric acid cycle
Glucose
Pyruvate
Acetyl CoA
CYTOSOL
MITOCHONDRION
Figure 9.6 An overview of cellular respiration.
ATP
ATP
Substrate-level phosphorylation
Substrate-level phosphorylation

25. Electrons carried via NADH Electrons carried via NADH and FADH2
Citric acid cycle
Pyruvate oxidation
Acetyl CoA
Glycolysis
Glucose
Pyruvate
Oxidative phosphorylation: electron transport and chemiosmosis
CYTOSOL
MITOCHONDRION
ATP
Substrate-level phosphorylation
Oxidative phosphorylation
Figure 9.6 An overview of cellular respiration.

26. The process that generates most of the ATP is called oxidative phosphorylation because it is powered by redox reactions
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27. Oxidative phosphorylation accounts for almost 90% of the ATP generated by cellular respiration
A smaller amount of ATP is formed in glycolysis and the citric acid cycle by substrate level phosphorylation
For each molecule of glucose degraded to CO2 and water by respiration, the cell makes up to 32 molecules of ATP
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28. Enzyme Enzyme ADP P Substrate ATP Product
Figure 9.7 Substrate-level phosphorylation.
Product

29. Glycolysis harvests chemical energy by oxidizing glucose to pyruvate
Glycolysis (“splitting of sugar”) breaks down glucose into two molecules of pyruvate
Glycolysis occurs in the cytoplasm and has two major phases
Glycolysis occurs whether or not O2 is present
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30. Inputs
Outputs
Glycolysis
Glucose
2 Pyruvate  2
ATP
 NADH

31. After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic molecules
In the presence of O2, pyruvate enters the mitochondrion (in eukaryotic cells) where the oxidation of glucose is completed
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32. Oxidation of Pyruvate to Acetyl CoA
Before the citric acid cycle can begin, pyruvate must be converted to acetyl Coenzyme A (acetyl CoA), which links glycolysis to the citric acid cycle
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33. MITOCHONDRION CYTOSOL CO2 Coenzyme A NAD NADH + H Acetyl CoA1 3 2
NAD
NADH
+ H
Acetyl CoA
Figure 9.10 Oxidation of pyruvate to acetyl CoA, the step before the citric acid cycle.
Pyruvate
Transport protein

34. The Citric Acid Cycle
The citric acid cycle, also called the Krebs cycle, completes the break down of pyrvate to CO2
The cycle oxidizes organic fuel derived from pyruvate, generating 1 ATP, 3 NADH, and 1 FADH2 per turn
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35. Pyruvate CO2 NAD CoA NADH + H Acetyl CoA CoA CoA Citric acid cycle
Figure 9.11 An overview of pyruvate oxidation and the citric acid cycle.
FADH2
3 NAD
FAD
3 NADH
+ 3 H
ADP + P i
ATP

36. The NADH and FADH2 produced by the cycle relay electrons extracted from food to the electron transport chain
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37. Inputs Outputs 2 Pyruvate 2 Acetyl CoA 2 ATP 8 NADH Citric acid cycle
Figure 9.UN07
Inputs
Outputs
2 Pyruvate
2 Acetyl CoA 2
ATP 8
NADH
Citric acid cycle
2 Oxaloacetate 6
CO2 2
FADH2
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38. During oxidative phosphorylation, chemiosmosis couples electron transport to ATP synthesis
Following glycolysis and the citric acid cycle, NADH and FADH2 account for most of the energy extracted from food
These two electron carriers donate electrons to the electron transport chain, which powers ATP synthesis via oxidative phosphorylation
For the Cell Biology Video ATP Synthase 3D Structure — Side View, go to Animation and Video Files.
For the Cell Biology Video ATP Synthase 3D Structure — Top View, go to Animation and Video Files.
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39. The Pathway of Electron Transport
The electron transport chain is in the inner membrane (cristae) of the mitochondrion
Electrons drop in free energy as they go down the chain and are finally passed to O2, forming H2O
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40. Multiprotein complexes I 40 II
NADH 50
2 e
NAD
FADH2
2 e
FAD
Multiprotein complexes I 40
FMN II
FeS
FeS Q
III
Cyt b 30
FeS
Cyt c1 IV
Free energy (G) relative to O2 (kcal/mol)
Cyt c
Cyt a
Cyt a3 20
Figure 9.13 Free-energy change during electron transport. 10
2 e
(originally from NADH or FADH2)
2 H + 1/2 O2
H2O

41. An Accounting of ATP Production by Cellular Respiration
During cellular respiration, most energy flows in this sequence:
glucose  NADH  electron transport chain  ATP
About 34% of the energy in a glucose molecule is transferred to ATP during cellular respiration, making about 32 ATP
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42. Oxidative phosphorylation: electron transport and chemiosmosis
Electron shuttles span membrane
MITOCHONDRION
2 NADH or
2 FADH2
2 NADH
2 NADH
6 NADH
2 FADH2
Oxidative phosphorylation: electron transport and chemiosmosis
Glycolysis
Pyruvate oxidation
Citric acid cycle
Glucose
2 Pyruvate
2 Acetyl CoA
Figure 9.16 ATP yield per molecule of glucose at each stage of cellular respiration.
 2 ATP
 2 ATP
 about 26 or 28 ATP
About 30 or 32 ATP
Maximum per glucose:
CYTOSOL

43. Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen
Most cellular respiration requires O2 to produce ATP
Without O2, the electron transport chain will cease to operate
In that case, glycolysis couples with fermentation or anaerobic respiration to produce ATP
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44. Anaerobic respiration uses an electron transport chain with a final electron acceptor other than O2, for example sulfate
Fermentation uses substrate-level phosphorylation instead of an electron transport chain to generate ATP
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45. Types of Fermentation Two common types are: Alcohol fermentation
Lactic acid fermentation
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46. In alcohol fermentation, pyruvate is converted to ethanol in two steps, with the first releasing CO2
Alcohol fermentation by yeast is used in brewing, wine making, and baking
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47.   2 ADP  2 P i 2 ATP Glucose Glycolysis 2 Pyruvate 2 NAD 2 NADH
Figure 9.17 Fermentation.
2 Acetaldehyde
2 Ethanol
(a) Alcohol fermentation

48. In lactic acid fermentation, pyruvate is reduced to NADH, forming lactate as an end product, with no release of CO2
Lactic acid fermentation by some fungi and bacteria is used to make cheese and yogurt
Human muscle cells use lactic acid fermentation to generate ATP when O2 is scarce
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49.   2 ADP  2 P i 2 ATP Glucose Glycolysis 2 NAD 2 NADH  2 H
2 Pyruvate
Figure 9.17 Fermentation.
2 Lactate
(b) Lactic acid fermentation

50. Comparing Fermentation with Anaerobic and Aerobic Respiration
Cellular respiration produces 32 ATP per glucose molecule; fermentation produces 2 ATP per glucose molecule
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51. Ethanol, lactate, or other products
Glucose
Glycolysis
CYTOSOL
Pyruvate
No O2 present: Fermentation
O2 present: Aerobic cellular respiration
MITOCHONDRION
Ethanol, lactate, or other products
Acetyl CoA
Figure 9.18 Pyruvate as a key juncture in catabolism.
Citric acid cycle

52. The Evolutionary Significance of Glycolysis
Ancient prokaryotes are thought to have used glycolysis long before there was oxygen in the atmosphere
Very little O2 was available in the atmosphere until about 2.7 billion years ago, so early prokaryotes likely used only glycolysis to generate ATP
Glycolysis is a very ancient process
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53. Regulation of Cellular Respiration via Feedback Mechanisms
Feedback inhibition is the most common mechanism for control
If ATP concentration begins to drop, respiration speeds up; when there is plenty of ATP, respiration slows down
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54. Oxidative phosphorylation
Glucose
AMP
Glycolysis
Fructose 6-phosphate
Stimulates 
Phosphofructokinase  
Fructose 1,6-bisphosphate
Inhibits
Inhibits
Pyruvate
ATP
Citrate
Acetyl CoA
Figure 9.20 The control of cellular respiration.
Citric acid cycle
Oxidative phosphorylation

55. Inputs Outputs Glycolysis Glucose 2 Pyruvate  2 ATP  2 NADH
Figure 9.UN06
Inputs
Outputs
Glycolysis
Glucose
2 Pyruvate  2
ATP
 NADH
Figure 9.UN06 Summary figure, Concept 9.2

56. Inputs Outputs 2 Pyruvate 2 Acetyl CoA 2 ATP 8 NADH Citric acid cycle
Figure 9.UN07
Inputs
Outputs
2 Pyruvate
2 Acetyl CoA 2
ATP 8
NADH
Citric acid cycle
2 Oxaloacetate 6
CO2 2
FADH2
Figure 9.UN07 Summary figure, Concept 9.3

57. Protein complex of electron carriers Cyt c
Figure 9.UN08
INTERMEMBRANE SPACE H H H
Protein complex of electron carriers
Cyt c IV Q
III I
Figure 9.UN08 Summary figure, Concept 9.4 (part 1) II
2 H + 1/2 O2
H2O
FADH2
FAD
NAD
NADH
MITOCHONDRIAL MATRIX
(carrying electrons from food)

58. MITO- CHONDRIAL MATRIX ATP synthase
Figure 9.UN09
INTER- MEMBRANE SPACE H
MITO- CHONDRIAL MATRIX
ATP synthase
Figure 9.UN09 Summary figure, Concept 9.4 (part 2)
ADP + P i H
ATP

59. pH difference across membrane
Figure 9.UN10
pH difference across membrane
Figure 9.UN10 Test Your Understanding, question 8
Time

60. Figure 9.UN11
Figure 9.UN11 Appendix A: answer to Test Your Understanding, question 8