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Chapter 9

Cellular Respiration:
Harvesting Chemical Energy

PowerPoint Lectures for


Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero


Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• 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 Cellular + O2
molecules
respiration
in mitochondria

ATP
powers most cellular work

Heat
Figure 9.2 energy

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• 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 Products

becomes oxidized

CH4 + 2O2 CO2 + Energy + 2 H2O

H becomes reduced

H H
C C
H
O O O
H
O O

Methane Oxygen Carbon dioxide Water


(reducing (oxidizing
agent) agent)
Figure 9.3
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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 – +

2 e– + H +
NAD +
NADH
H
O Dehydrogenase
H H O
NH2 + 2[H] Reduction of NAD+ NH2 +
C C
(from food) Oxidation of NADH
N+ N Nicotinamide
Nicotinamide
O CH2 (oxidized form) (reduced form)
O
O P O –

O H H
O P O– HO OH NH2
CH2HO
O N N
H
N N H
O

H H
HO OH Figure 9.4
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• 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 (a) Uncontrolled reaction
heat and light
energy

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 + / 2 O2
1

(from food via NADH)

Controlled
release of
2 H + + 2 e–
energy for
synthesis of

Elec
ATP
ATP
Free energy, G

tron
ATP

t
rans
ATP

port
c
hain

2 e–
1
/ 2 O2
2H +

H2O

Figure 9.5 B (b) Cellular respiration

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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 Electrons carried


carried via NADH and
via NADH FADH2

Oxidative
Citric phosphorylation:
Glycolsis
Glucos acid electron
Pyruvate cycle
e transport and
chemiosmosis

Cytosol
Mitochondrion

ATP ATP ATP

Substrate-level Oxidative
Substrate-level
phosphorylation phosphorylation
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
Substrate
+ ATP

Figure 9.7 Product

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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 Glycolysis Citric Oxidative
acid
cycle phosphorylation

– Energy payoff phase ATP ATP ATP

Energy investment phase

Glucose

2 ATP + 2 P 2 ATP used

Energy payoff phase

4 ADP + 4 P 4 ATP formed

2 NAD+ + 4 e- + 4 H 2 NADH + 2 H+
+

2 Pyruvate + 2 H2O

Glucose 2 Pyruvate + 2 H2O


4 ATP formed – 2 ATP used 2 ATP + 2 H+
2 NAD+ + 4 e– + 4 H 2 NADH
Figure 9.8 +

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• A closer look at the energy investment phase

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CH2OH
Citric
HH H Glycolysis acid Oxidative
HO H cycle phosphorylation
HO OH
H OH
Glucose

ATP 1
Hexokinase
ADP
CH2OH P
HH OH
OH H
HO
H OH
Glucose-6-phosphate
2
Phosphoglucoisomerase
CH2O P
O CH2OH
H HO
H HO
HO H
Fructose-6-phosphate
3
ATP
Phosphofructokinase

ADP
P O CH2 O CH2 O P
HO
H OH
HO H
Fructose-
1, 6-bisphosphate
4
Aldolase

5 H
P O CH2 Isomerase
C O
C O
CHOH
CH2OH
CH2 O P
Dihydroxyacetone Glyceraldehyde-
phosphate 3-phosphate
Figure 9.9 A
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• A closer look at the energy payoff phase

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6
2 NAD+ Triose phosphate
dehydrogenase
2 NADH 2 Pi
+ 2 H+
2
P 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
H C O P
CH2OH
2-Phosphoglycerate
9
2 H2O Enolase
2 O–
C O
C O P
CH2
Phosphoenolpyruvate
2 ADP 10
Pyruvate kinase
2 ATP

2 O–
C O
C O
CH3
Figure 9.8 B Pyruvate

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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+ NADH + H+
O–
S CoA
2
C O
C O

C O
CH3
1 3
CH3
Acetyle CoA
Pyruvate CO2 Coenzyme A

Transport protein

Figure 9.10

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• An overview of the citric acid cycle
Pyruvate Glycolysis Citric Oxidative
acid
(from glycolysis, cycle phosphorylation

2 molecules per glucose)


ATP ATP ATP

CO2

CoA
NADH
+ 3 H+ Acetyle CoA
CoA

CoA

Citric
acid 2 CO2
cycle
FADH2 3 NAD+

FAD 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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Citric Oxidative
Glycolysis
acid phosphorylation
cycle

S CoA
C O
CH3
Acetyl CoA
CoA SH

NADH O C COO–
+ H+ CH2 1 COO– H2 O
COO– CH2 COO–
NAD +

8 Oxaloacetate HO C COO –
CH2
CH2 2
COO – HC COO–
COO– HO CH
HO CH Malate Citrate
CH2 Figure 9.12 COO–
Isocitrate
COO–
Citric CO2
3
7 acid NAD+
H2 O cycle
COO–
COO– NADH
CH + H+
Fumarate CoA SH CH2
HC
CH2 α-Ketoglutarate
COO–
C O
6 4
COO– CoA SH COO– COO–

CH2 5 CH2
FADH2
CH2 CH2 CO2
FAD NAD+
COO – C O
Succinate Pi S CoA NADH
GTP GDP Succinyl + H+
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

NADH
50

FADH2
Free energy (G) relative to O2 (kcl/mol)

I Multiprotein
40 FMN FAD
complexes
Fe•S Fe•S II
O
Cyt b
III
30 Fe•S
Cyt c1
Cyt c IV
Cyt a
Cyt a3
20

10

0 2 H + + 1⁄2 O2

Figure 9.13 H2O

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Chemiosmosis: The Energy-Coupling Mechanism
• ATP synthase
– Is the enzyme that actually makes ATP
INTERMEMBRANE SPACE A rotor within the
H+
membrane spins
H+ H+ clockwise when
H+ H+ flows past
H+
it down the H+
gradient.
H+
H+
A stator anchored
in the membrane
holds the knob
stationary.

A rod (for “stalk”)


extending into
the knob also
spins, activating
catalytic sites in
H+ the knob.
Three catalytic
sites in the
ADP stationary knob
+ join inorganic
Pi ATP
Phosphate to ADP
to make ATP.
MITOCHONDRIAL MATRIX
Figure 9.14
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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
Inner
Mitochondrial
Glycolysis
Oxidative
phosphorylation. membrane
electron transport
and chemiosmosis

ATP ATP ATP

H+
H +

H+
H+
Cyt c
Protein complex
Intermembrane of electron
space carners
Q IV
I III
ATP
Inner II synthase
mitochondrial FADH2 H2O
membrane FAD+ 2 H+ + 1/2 O2
NADH+
NAD+ ADP + Pi ATP
(Carrying electrons
from, food) H+
Mitochondrial Chemiosmosis
Electron transport chain
matrix Electron transport and pumping of protons (H ),
+ ATP synthesis powered by the flow
which create an H gradient across the membrane Of H back across the membrane
+ +

Figure 9.15 Oxidative phosphorylation

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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
Electron shuttles MITOCHONDRION
CYTOSOL 2 NADH
span membrane
or
2 FADH2

2 NADH 2 NADH 6 NADH 2 FADH2

Glycolysis Oxidative
2 Citric phosphorylation:
2 Acetyl acid electron transport
Glucose Pyruvate CoA cycle and
chemiosmosis

+ 2 ATP + 2 ATP + about 32 or 34 ATP


by substrate-level by substrate-level by oxidative phosphorylation, depending
phosphorylation phosphorylation on which shuttle transports electrons
from NADH in cytosol

About
Maximum per glucose: 36 or 38 ATP

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
Glucose Glycolysis
CH3
2 Pyruvate
2 NAD+ 2 NADH 2 CO2

H H
H C OH C O
CH3 CH3
2 Ethanol 2 Acetaldehyde
(a) Alcohol fermentation

2 ADP + 2 P1 2 ATP

Glucose Glycolysis O–
C O
C O
2 NAD+ 2 NADH
O CH3
C O
H C OH
CH3
2 Lactate
(b) Lactic acid fermentation
Figure 9.17
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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 O2 present
Fermentation Cellular respiration

MITOCHONDRION
Ethanol Acetyl CoA
or
lactate
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 Sugars Glycerol Fatty


acids acids

Glycolysis
Glucose

Glyceraldehyde-3- P

NH3 Pyruvate

Acetyl CoA

Citric
acid
cycle

Oxidative
Figure 9.19 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
AMP
Fructose-6-phosphate Stimulates
+
Phosphofructokinase
– –
Fructose-1,6-bisphosphate
Inhibits Inhibits

Pyruvate

ATP Citrate
Acetyl CoA

Citric
acid
cycle

Oxidative
Figure 9.20 phosphorylation

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