Chapter 9

Cellular Respiration: Harvesting Chemical Energy
PowerPoint Lectures for Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero
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• Overview: Life Is Work • Living cells
– Require transfusions of energy from outside sources to perform their many tasks

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• The giant panda
– Obtains energy for its cells by eating plants

Figure 9.1
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• 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
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Heat energy

• Concept 9.1: Catabolic pathways yield energy by oxidizing organic fuels

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

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• One catabolic process, fermentation
– Is a partial degradation of sugars that occurs without oxygen

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

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• To keep working
– Cells must regenerate ATP

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

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

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• In oxidation
– A substance loses electrons, or is oxidized

• In reduction
– A substance gains electrons, or is reduced

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• Examples of redox reactions

becomes oxidized (loses electron)

Na

+

Cl

Na+

+

Cl–

becomes reduced (gains electron)

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• Some redox reactions
– Do not completely exchange electrons – Change the degree of electron sharing in covalent bonds
Reactants
becomes oxidized

Products

CH4 H

+

2O2

CO2

+

Energy

+

2 H2O

becomes reduced

Figure 9.3

Methane (reducing agent)

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H

C
H

O

O

O

C

O

O

Oxygen (oxidizing agent)

Carbon dioxide

Water

H

H

H

Oxidation of Organic Fuel Molecules During Cellular Respiration • During cellular respiration
– Glucose is oxidized and oxygen is reduced
becomes oxidized

C6H12O6 + 6O2

6CO2 + 6H2O + Energy
becomes reduced

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

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• Electrons from organic compounds
– Are usually first transferred to NAD+, a coenzyme 2e +2H
– +

NAD

+

2 e– + H + H N

NADH H O C

O

CH2

O P O O H O P O– HO O

O
H OH N N

CH2HO H

O
H HO

H OH

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H
N+

O
C NH2

Dehydrogenase Reduction of NAD+ + 2[H] (from food) Oxidation of NADH

NH2

+

Nicotinamide (oxidized form)

Nicotinamide (reduced form)

NH2 N N H

Figure 9.4

• NADH, the reduced form of NAD+
– Passes the electrons to the electron transport chain

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• If electron transfer is not stepwise
– A large release of energy occurs – As in the reaction of hydrogen and oxygen to form water
H2 + 1/2 O2

Free energy, G

Explosive release of heat and light energy

(a) Uncontrolled reaction

Figure 9.5 A

H2O

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• 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

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2H (from food via NADH)

+

1

/ 2 O2

2 H + + 2 e–
Elec

Controlled release of energy for synthesis of ATP

Free energy, G

ATP ATP ATP

Figure 9.5 B
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t tron c port rans hain

2 e– 2H
+

1

/ 2 O2

H2O

(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

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• Glycolysis
– Breaks down glucose into two molecules of pyruvate

• The citric acid cycle
– Completes the breakdown of glucose

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• Oxidative phosphorylation
– Is driven by the electron transport chain – Generates ATP

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• An overview of cellular respiration
Electrons carried via NADH Electrons carried via NADH and FADH2

Glucos e

Glycolsis Pyruvate

Citric acid cycle

Oxidative phosphorylation: electron transport and chemiosmosis

Cytosol

Mitochondrion

ATP Substrate-level phosphorylation

ATP Substrate-level phosphorylation

ATP Oxidative phosphorylation

Figure 9.6

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• Both glycolysis and the citric acid cycle
– Can generate ATP by substrate-level phosphorylation

Enzyme

Enzyme

ADP P + Product ATP

Substrate

Figure 9.7

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• 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

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• Glycolysis consists of two major phases
– Energy investment phase – Energy payoff phase
Glycolysis
Citric acid cycle Oxidative phosphorylation

ATP

ATP

ATP

Energy investment phase Glucose

2 ATP + 2 P Energy payoff phase 4 ADP + 4 P

2 ATP

used

4 ATP formed

2 NAD+ + 4 e- + 4 H
+

2 NADH + 2 H+

2 Pyruvate + 2 H2O

Glucose 4 ATP formed – 2 ATP used

2 Pyruvate + 2 H2O 2 ATP + 2 H+ 2 NADH

Figure 9.8
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2 NAD+ + 4 e– + 4 H
+

• A closer look at the energy investment phase

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CH2OH HH H HO H HO OH H OH
Glucose 1

Glycolysis

Citric Oxidative acid cycle phosphorylation

ATP ADP

Hexokinase

Glucose-6-phosphate 2

CH2OH P HH OH OH H HO H OH

Fructose-6-phosphate

Phosphoglucoisomerase CH2O P O CH2OH H HO HO H HO H
3

ATP ADP

Phosphofructokinase

P O CH2 O CH2 O P HO H OH HO H
Fructose1, 6-bisphosphate

Aldolase

4

Figure 9.9 A

Dihydroxyacetone phosphate

5 H P O CH2 Isomerase C O C O CHOH CH2OH CH2 O P

Glyceraldehyde3-phosphate

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• A closer look at the energy payoff phase

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2 NAD+ 2 NADH + 2 H+ 2 P

6 Triose phosphate dehydrogenase 2 Pi

O C O CHOH

CH2 O P 1, 3-Bisphosphoglycerate 2 ADP 7 Phosphoglycerokinase 2 ATP 2 O– C CHOH CH2 O P 3-Phosphoglycerate 8 Phosphoglyceromutase 2 O– C O P H C O

CH2OH 2-Phosphoglycerate 9 Enolase 2 H2O 2 O– C O C O P CH2 Phosphoenolpyruvate 2 ADP 10 Pyruvate kinase 2 ATP 2 O– C O C O CH3 Pyruvate

Figure 9.8 B

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• 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

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• Before the citric acid cycle can begin
– Pyruvate must first be converted to acetyl CoA, which links the cycle to glycolysis
CYTOSOL MITOCHONDRION

NAD+ O– 2 C O

NADH

+ H+ S CoA

C C O 1 CH3 Pyruvate CO2
Coenzyme A

O

3

CH3 Acetyle CoA

Transport protein

Figure 9.10
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• An overview of the citric acid cycle
Pyruvate (from glycolysis, 2 molecules per glucose)
Glycolysis
Citric acid cycle Oxidative phosphorylation

ATP

ATP

ATP

CO2 CoA NADH + 3 H+ Acetyle CoA CoA CoA

FADH2 FAD

Citric acid cycle

2 CO2 3 NAD+ 3 NADH + 3 H+

ADP + P i ATP

Figure 9.11
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• A closer look at the citric acid cycle

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Glycolysis

Citric Oxidative acid phosphorylation cycle

S C

CoA O

CH3

Acetyl CoA
CoA SH NADH + H+ O C COO– CH2 COO–

1

COO– CH2 HO C COO CH2 COO–

H2 O
COO–

NAD

+

8 Oxaloacetate

COO HO CH CH2

2
HO

CH2 HC COO– CH COO–

Malate

Figure

Citrate 9.12

COO–

Isocitrate
3
CO2

H2 O

7
COO– CH HC COO–

Citric acid cycle Fumarate
6
CoA SH

NAD+ NADH
+ H+

COO– CH2 CH2 COO– CH2 CH2 COO

α-Ketoglutarate

CoA SH

COO– CH2 CH2 C O

4

C O COO–

FADH2

5

FAD

NAD+ NADH + H+

CO2

Succinate

GTP GDP Succinyl

Pi

S

CoA

CoA

ADP ATP

Figure 9.12
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• 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

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

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• At the end of the chain
– Electrons are passed to oxygen, forming water

50

NADH

FADH2

Free energy (G) relative to O2 (kcl/mol)

40

FMN

I
Fe•S O

FAD Fe•S II

Multiprotein complexes III

30

Cyt b Fe•S Cyt c1

20

IV Cyt c Cyt a Cyt a3

10

0

2 H + + 1⁄2 O2

Figure 9.13
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H2O

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

H+ H+

H+

H+ H+ H+ H+

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.

H+

ADP + Pi

ATP

Figure 9.14

MITOCHONDRIAL MATRIX

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

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

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

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• Chemiosmosis and the electron transport chain
Glycolysis
Oxidative phosphorylation. electron transport and chemiosmosis

Inner Mitochondrial membrane

ATP

ATP

ATP

H H+ Intermembrane space Protein complex of electron carners I Inner mitochondrial membrane
NADH+

+

H+

Cyt c

H+

Q
III II
FADH2

IV ATP synthase
Pi

FAD+

2 H+ + 1/2 O2

H2O
ADP +

NAD+

ATP H+

Mitochondrial matrix

(Carrying electrons from, food)

Chemiosmosis Electron transport chain + ATP synthesis powered by the flow Electron transport and pumping of protons (H ), which create an H+ gradient across the membrane Of H+ back across the membrane Oxidative phosphorylation

Figure 9.15
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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

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• There are three main processes in this metabolic enterprise
CYTOSOL
Electron shuttles span membrane 2 NADH

MITOCHONDRION

or

2 FADH2 2 NADH 2 NADH 6 NADH 2 FADH2

Glycolysis Glucose 2 Pyruvate

2 Acetyl CoA

Citric acid cycle

Oxidative phosphorylation: electron transport and chemiosmosis

+ 2 ATP by substrate-level phosphorylation

+ 2 ATP

+ 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

Maximum per glucose:

Figure 9.16
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• About 40% of the energy in a glucose molecule
– Is transferred to ATP during cellular respiration, making approximately 38 ATP

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• 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

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

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

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

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

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2 ADP + 2

P1

2 ATP

O– C O C O CH3 2 Pyruvate 2 CO2 H C O CH3 2 Acetaldehyde

Glucose

Glycolysis

2 NAD+ H H C OH CH3 2 Ethanol (a) Alcohol fermentation

2 NADH

2 ADP + 2

P1

2 ATP

Glucose

Glycolysis

O– C O C O CH3

O C O H C OH CH3 2 Lactate

2 NAD+

2 NADH

Figure 9.17

(b) Lactic acid fermentation

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

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• Fermentation and cellular respiration
– Differ in their final electron acceptor

• Cellular respiration
– Produces more ATP

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• Pyruvate is a key juncture in catabolism
Glucose CYTOSOL Pyruvate
No O2 present Fermentation O2 present Cellular respiration

MITOCHONDRION

Ethanol or lactate

Acetyl CoA Citric acid cycle

Figure 9.18
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The Evolutionary Significance of Glycolysis • Glycolysis
– Occurs in nearly all organisms – Probably evolved in ancient prokaryotes before there was oxygen in the atmosphere

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

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

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• The catabolism of various molecules from food
Proteins Carbohydrates Fats Amino acids Sugars Glycerol Fatty acids

Glycolysis

Glucose
Glyceraldehyde-3- P

NH3

Pyruvate
Acetyl CoA

Citric acid cycle

Figure 9.19

Oxidative phosphorylation

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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

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

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• The control of cellular respiration
Glucose
Glycolysis
Fructose-6-phosphate

AMP
Stimulates


Inhibits

Phosphofructokinase Fructose-1,6-bisphosphate

+ –

Inhibits

Pyruvate
ATP
Acetyl CoA

Citrate

Citric acid cycle

Figure 9.20
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Oxidative phosphorylation