1. Cellular Respiration: Harvesting Chemical Energy
Chapter 9
Cellular Respiration: Harvesting Chemical Energy

2. Overview: Life Is Work Living cells
Require transfusions of energy from outside sources to perform their many tasks

3. The giant panda Obtains energy for its cells by eating plants
Figure 9.1

4. powers most cellular work
Energy
Flows into an ecosystem as sunlight and leaves as heat
Light energy
ECOSYSTEM
CO2 + H2O
Photosynthesis in chloroplasts
Cellular respiration in mitochondria
Organic molecules
+ O2
ATP
powers most cellular work
Heat energy
Figure 9.2

5. Concept 9.1: Catabolic pathways yield energy by oxidizing organic fuels

6. Catabolic Pathways and Production of ATP
The breakdown of organic molecules is exergonic

7. One catabolic process, fermentation
Is a partial degradation of sugars that occurs without oxygen

8. Cellular respiration
Is the most prevalent and efficient catabolic pathway
Consumes oxygen and organic molecules such as glucose
Yields ATP

9. To keep working
Cells must regenerate ATP

10. Redox Reactions: Oxidation and Reduction
Catabolic pathways yield energy
Due to the transfer of electrons

11. The Principle of Redox Redox reactions
Transfer electrons from one reactant to another by oxidation and reduction

12. In oxidation In reduction A substance loses electrons, or is oxidized
A substance gains electrons, or is reduced

13. Examples of redox reactions
Na Cl Na Cl–
becomes oxidized (loses electron)
becomes reduced (gains electron)

14. Some redox reactions Do not completely exchange electrons
Change the degree of electron sharing in covalent bonds
CH4 H C O
Methane (reducing agent)
Oxygen (oxidizing agent)
Carbon dioxide
Water +
2O2
CO2
Energy
2 H2O
becomes oxidized
becomes reduced
Reactants
Products
Figure 9.3

15. Oxidation of Organic Fuel Molecules During Cellular Respiration
Glucose is oxidized and oxygen is reduced
C6H12O6 + 6O CO2 + 6H2O + Energy
becomes oxidized
becomes reduced

16. Stepwise Energy Harvest via NAD+ and the Electron Transport Chain
Cellular respiration
Oxidizes glucose in a series of steps

17. Electrons from organic compounds
Are usually first transferred to NAD+, a coenzyme
NAD+ H O O– P
CH2 HO OH N+ C
NH2 N
Nicotinamide (oxidized form) +
2[H]
(from food)
Dehydrogenase
Reduction of NAD+
Oxidation of NADH
2 e– + 2 H+
2 e– + H+
NADH
Nicotinamide (reduced form)
Figure 9.4

18. NADH, the reduced form of NAD+
Passes the electrons to the electron transport chain

19. (a) Uncontrolled reaction
If electron transfer is not stepwise
A large release of energy occurs
As in the reaction of hydrogen and oxygen to form water
(a) Uncontrolled reaction
Free energy, G
H2O
Explosive release of heat and light energy
Figure 9.5 A
H2 + 1/2 O2

20. 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

21. Electron transport chain (b) Cellular respiration
(from food via NADH)
2 H e–
2 H+
2 e–
H2O
Controlled release of energy for synthesis of ATP
ATP
Electron transport chain
Free energy, G
(b) Cellular respiration +
Figure 9.5 B

22. The Stages of Cellular Respiration: A Preview
Respiration is a cumulative function of three metabolic stages
Glycolysis
The citric acid cycle
Oxidative phosphorylation

23. Glycolysis The citric acid cycle
Breaks down glucose into two molecules of pyruvate
The citric acid cycle
Completes the breakdown of glucose

24. Oxidative phosphorylation
Is driven by the electron transport chain
Generates ATP

25. Oxidative phosphorylation: electron transport and chemiosmosis
An overview of cellular respiration
Figure 9.6
Electrons
carried
via NADH
Glycolsis
Glucose
Pyruvate
ATP
Substrate-level
phosphorylation
Electrons carried
via NADH and
FADH2
Citric acid cycle
Oxidative phosphorylation: electron transport and chemiosmosis
Oxidative
Mitochondrion
Cytosol

26. Both glycolysis and the citric acid cycle
Can generate ATP by substrate-level phosphorylation
Figure 9.7
Enzyme
ATP
ADP
Product
Substrate P +

27. Concept 9.2: Glycolysis harvests energy by oxidizing glucose to pyruvate
Means “splitting of sugar”
Breaks down glucose into pyruvate
Occurs in the cytoplasm of the cell

28. Energy investment phase
Glycolysis consists of two major phases
Energy investment phase
Energy payoff phase
Glycolysis
Citric acid cycle
Oxidative phosphorylation
ATP
2 ATP
4 ATP
used
formed
Glucose
2 ATP + 2 P
4 ADP + 4
2 NAD+ + 4 e- + 4 H +
2 NADH
+ 2 H+
2 Pyruvate + 2 H2O
Energy investment phase
Energy payoff phase
4 ATP formed – 2 ATP used
2 NAD+ + 4 e– + 4 H +
Figure 9.8

29. A closer look at the energy investment phase

30. Phosphoglucoisomerase
Dihydroxyacetone
phosphate
Glyceraldehyde-
3-phosphate H OH HO
CH2OH O P
CH2O
CH2 C
CHOH
ATP
ADP
Hexokinase
Glucose
Glucose-6-phosphate
Fructose-6-phosphate
Phosphoglucoisomerase
Phosphofructokinase
Fructose-
1, 6-bisphosphate
Aldolase
Isomerase
Glycolysis 1 2 3 4 5
Oxidative
phosphorylation
Citric
acid
cycle
Figure 9.9 A

31. A closer look at the energy payoff phase

32. 1, 3-Bisphosphoglycerate
2 NAD+
NADH 2
+ 2 H+
Triose phosphate
dehydrogenase
P i P C
CHOH O
CH2 O–
1, 3-Bisphosphoglycerate
2 ADP
2 ATP
Phosphoglycerokinase
3-Phosphoglycerate
Phosphoglyceromutase
CH2OH H
2-Phosphoglycerate
2 H2O
Enolase
Phosphoenolpyruvate
Pyruvate kinase
CH3 6 8 7 9 10
Pyruvate
Figure 9.8 B

33. Concept 9.3: The citric acid cycle completes the energy-yielding oxidation of organic molecules
Takes place in the matrix of the mitochondrion

34. Before the citric acid cycle can begin
Pyruvate must first be converted to acetyl CoA, which links the cycle to glycolysis
CYTOSOL
MITOCHONDRION
NADH
+ H+
NAD+ 2 3 1
CO2
Coenzyme A
Pyruvate
Acetyle CoA S
CoA C
CH3 O
Transport protein O–
Figure 9.10

35. Oxidative phosphorylation
An overview of the citric acid cycle
ATP
2 CO2
3 NAD+
3 NADH
+ 3 H+
ADP + P i
FAD
FADH2
Citric acid cycle
CoA
Acetyle CoA
NADH
CO2
Pyruvate (from glycolysis, 2 molecules per glucose)
Glycolysis
Oxidative phosphorylation
Figure 9.11

36. A closer look at the citric acid cycle

37. Figure 9.12 Figure 9.12 Acetyl CoA Oxaloacetate Malate Citrate
NADH
Oxaloacetate
Citrate
Malate
Fumarate
Succinate
Succinyl
CoA
a-Ketoglutarate
Isocitrate
Citric
acid
cycle S SH
FADH2
FAD
GTP
GDP
NAD+
ADP
P i
CO2
H2O
+ H+ C
CH3 O
COO–
CH2 HO HC CH 1 2 3 4 5 6 7 8
Glycolysis
Oxidative
phosphorylation
ATP
Figure 9.12
Figure 9.12

38. 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

39. The Pathway of Electron Transport
In the electron transport chain
Electrons from NADH and FADH2 lose energy in several steps

40. At the end of the chain Electrons are passed to oxygen, forming water
H2O O2
NADH
FADH2
FMN
Fe•S O
FAD
Cyt b
Cyt c1
Cyt c
Cyt a
Cyt a3
2 H + + 12 I II
III IV
Multiprotein complexes 10 20 30 40 50
Free energy (G) relative to O2 (kcl/mol)
Figure 9.13

41. Chemiosmosis: The Energy-Coupling Mechanism
ATP synthase
Is the enzyme that actually makes ATP
INTERMEMBRANE SPACE 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

42. At certain steps along the electron transport chain
Electron transfer causes protein complexes to pump H+ from the mitochondrial matrix to the intermembrane space

43. The resulting H+ gradient
Stores energy
Drives chemiosmosis in ATP synthase
Is referred to as a proton-motive force

44. Chemiosmosis
Is an energy-coupling mechanism that uses energy in the form of a H+ gradient across a membrane to drive cellular work

45. Electron transport chain Oxidative phosphorylation
Chemiosmosis and the electron transport chain
Oxidative
phosphorylation.
electron transport
and chemiosmosis
Glycolysis
ATP
Inner
Mitochondrial
membrane H+
P i
Protein complex
of electron
carners
Cyt c I II
III IV
(Carrying electrons
from, food)
NADH+
FADH2
NAD+
FAD+
2 H+ + 1/2 O2
H2O
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
synthase Q
Oxidative phosphorylation
Intermembrane
space
mitochondrial
matrix
Figure 9.15

46. 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

47. There are three main processes in this metabolic enterprise
Electron shuttles
span membrane
CYTOSOL
2 NADH
2 FADH2
6 NADH
Glycolysis
Glucose 2
Pyruvate
Acetyl
CoA
Citric
acid
cycle
Oxidative
phosphorylation:
electron transport
and
chemiosmosis
MITOCHONDRION
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
+ about 32 or 34 ATP or
Figure 9.16

48. About 40% of the energy in a glucose molecule
Is transferred to ATP during cellular respiration, making approximately 38 ATP

49. In the absence of oxygen
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

50. Glycolysis
Can produce ATP with or without oxygen, in aerobic or anaerobic conditions
Couples with fermentation to produce ATP

51. Fermentation consists of
Types of Fermentation
Fermentation consists of
Glycolysis plus reactions that regenerate NAD+, which can be reused by glyocolysis

52. In alcohol fermentation
Pyruvate is converted to ethanol in two steps, one of which releases CO2

53. During lactic acid fermentation
Pyruvate is reduced directly to NADH to form lactate as a waste product

54. (a) Alcohol fermentation (b) Lactic acid fermentation
2 ADP + 2 P1
2 ATP
Glycolysis
Glucose
2 NAD+
2 NADH
2 Pyruvate
2 Acetaldehyde
2 Ethanol
(a) Alcohol fermentation
2 Lactate
(b) Lactic acid fermentation H OH
CH3 C
O – O O–
CO2 2
Figure 9.17

55. Fermentation and Cellular Respiration Compared
Both fermentation and cellular respiration
Use glycolysis to oxidize glucose and other organic fuels to pyruvate

56. Fermentation and cellular respiration
Differ in their final electron acceptor
Cellular respiration
Produces more ATP

57. Pyruvate is a key juncture in catabolism
Glucose
CYTOSOL
Pyruvate
No O2 present
Fermentation
O2 present
Cellular respiration
Ethanol or
lactate
Acetyl CoA
MITOCHONDRION
Citric
acid
cycle
Figure 9.18

58. The Evolutionary Significance of Glycolysis
Occurs in nearly all organisms
Probably evolved in ancient prokaryotes before there was oxygen in the atmosphere

59. Concept 9.6: Glycolysis and the citric acid cycle connect to many other metabolic pathways

60. The Versatility of Catabolism
Catabolic pathways
Funnel electrons from many kinds of organic molecules into cellular respiration

61. The catabolism of various molecules from food
Amino
acids
Sugars
Glycerol
Fatty
Glycolysis
Glucose
Glyceraldehyde-3- P
Pyruvate
Acetyl CoA
NH3
Citric
acid
cycle
Oxidative
phosphorylation
Fats
Proteins
Carbohydrates
Figure 9.19

62. 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

63. Regulation of Cellular Respiration via Feedback Mechanisms
Is controlled by allosteric enzymes at key points in glycolysis and the citric acid cycle

64. Fructose-1,6-bisphosphate
The control of cellular respiration
Glucose
Glycolysis
Fructose-6-phosphate
Phosphofructokinase
Fructose-1,6-bisphosphate
Inhibits
Pyruvate
ATP
Acetyl CoA
Citric
acid
cycle
Citrate
Oxidative
phosphorylation
Stimulates
AMP + –
Figure 9.20