Chapter 9

Cellular Respiration and

Fermentation
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

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Figure 9.1
 • 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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Figure 9.2

Light
energy

ECOSYSTEM

Photosynthesis
in chloroplasts
Organic
CO2  H2O
molecules  O2
Cellular respiration
in mitochondria

ATP powers
ATP
most cellular work

Heat
energy
Concept 9.1: Catabolic pathways yield
energy by oxidizing organic fuels
• Several processes are central to cellular
respiration and related pathways

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Catabolic Pathways and Production of ATP
• The breakdown of organic molecules is
exergonic
• 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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 • 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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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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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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Figure 9.UN01

becomes oxidized
(loses electron)

becomes reduced
(gains electron)
 • The electron donor is called the reducing
agent
• The electron receptor is called the oxidizing
agent

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Oxidation of Organic Fuel Molecules During
Cellular Respiration
• During cellular respiration, the fuel (such as
glucose) is oxidized, and O2 is reduced

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Figure 9.UN03

becomes oxidized

becomes reduced
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 is tapped to
synthesize ATP
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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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Figure 9.6-1

Electrons
carried
via NADH

Glycolysis

Glucose Pyruvate

CYTOSOL MITOCHONDRION

ATP

Substrate-level
phosphorylation
Figure 9.6-2

Electrons Electrons carried

carried via NADH and
via NADH FADH2

Pyruvate
Glycolysis Citric
oxidation
acid
Glucose Pyruvate Acetyl CoA cycle

CYTOSOL MITOCHONDRION

ATP ATP

Substrate-level Substrate-level
phosphorylation phosphorylation
Figure 9.6-3

Electrons Electrons carried

carried via NADH and
via NADH FADH2

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

CYTOSOL MITOCHONDRION

ATP ATP ATP

Substrate-level Substrate-level Oxidative

phosphorylation phosphorylation phosphorylation
 • The process that generates most of the ATP is
called oxidative phosphorylation because it is
powered by redox reactions

BioFlix: Cellular Respiration

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 • 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 CO 2
and water by respiration, the cell makes up to
32 molecules of ATP

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Figure 9.7

Enzyme Enzyme

ADP
P
Substrate ATP
Product
Concept 9.2: 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

• Glycolysis occurs whether or not O2 is present

• The product of glycolysis is pyruvate

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Concept 9.3: 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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Figure 9.6-3

Electrons Electrons carried

carried via NADH and
via NADH FADH2

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

CYTOSOL MITOCHONDRION

ATP ATP ATP

Substrate-level Substrate-level Oxidative

phosphorylation phosphorylation phosphorylation
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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Figure 9.11
Pyruvate

CO2
NAD 

CoA
NADH
+ H Acetyl CoA
CoA

CoA

Citric
acid
cycle 2 CO2

FADH2 3 NAD

FAD 3 NADH
+ 3 H
ADP + P i

ATP
Concept 9.4: 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

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The Pathway of Electron Transport
• The electron transport chain is in the inner
membrane (cristae) of the mitochondrion
• Most of the chain’s components are proteins,
which exist in multiprotein complexes
• The carriers alternate reduced and oxidized
states as they accept and donate electrons
• Electrons drop in free energy as they go down
the chain and are finally passed to O 2, forming
H2O

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Figure 9.13
NADH
50
2 e
NAD
FADH2

2 e FAD Multiprotein

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

40 FMN
I complexes
FeS II
FeS
Q
III
Cyt b
FeS
30
Cyt c1
IV
Cyt c
Cyt a
Cyt a3
20

2 e
10
(originally from
NADH or FADH2)

0 2 H + 1/2 O2

H2O
Chemiosmosis: The Energy-Coupling
Mechanism
• Electron transfer in the electron transport chain
causes proteins to pump H+ from the
mitochondrial matrix to the intermembrane space
• H+ then moves back across the membrane,
passing through the proton, ATP synthase
• ATP synthase uses the exergonic flow of H + to
drive phosphorylation of ATP
• This is an example of chemiosmosis, the use of
energy in a H+ gradient to drive cellular work

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Figure 9.15

H
H 

Protein H 

complex H
Cyt c
of electron
carriers
IV
Q
III
I
ATP
II synth-
2 H + 1/2O2 H2O ase
FADH2 FAD

NADH NAD
ADP  P i ATP
(carrying electrons
from food) H

1 Electron transport chain 2 Chemiosmosis

Oxidative phosphorylation
 • The energy stored in a H+ gradient across a
membrane couples the redox reactions of the
electron transport chain to ATP synthesis
• The H+ gradient is referred to as a proton-
motive force, emphasizing its capacity to do
work

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An Accounting of ATP Production by
Cellular Respiration
• During cellular respiration, most energy flows
in this sequence:
glucose  NADH  electron transport chain
 proton-motive force  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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Concept 9.5: 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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 • 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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Types of Fermentation
• Fermentation consists of glycolysis plus
reactions that regenerate NAD+, which can be
reused by glycolysis
• Two common types are alcohol fermentation
and lactic acid fermentation

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 • In alcohol fermentation, pyruvate is converted
to ethanol in two steps, with the first releasing
CO2
• Alcohol fermentation by yeast is used in
brewing, winemaking, and baking

Animation: Fermentation Overview

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Figure 9.17

2 ADP  2 P 2 ATP 2 ADP  2 P 2 ATP

i i

Glucose Glycolysis Glucose Glycolysis

2 Pyruvate

2 NAD  2 NADH 2 CO2 2 NAD  2 NADH

 2 H  2 H
2 Pyruvate

2 Ethanol 2 Acetaldehyde 2 Lactate

(a) Alcohol fermentation (b) Lactic acid fermentation

Figure 9.17a

2 ADP  2 P i 2 ATP

Glucose Glycolysis

2 Pyruvate

2 NAD  2 NADH 2 CO2

 2 H

2 Ethanol 2 Acetaldehyde

(a) Alcohol fermentation

 • 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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Figure 9.17b

2 ADP  2 P i 2 ATP

Glucose Glycolysis

2 NAD  2 NADH
 2 H
2 Pyruvate

2 Lactate

(b) Lactic acid fermentation

Comparing Fermentation with Anaerobic
and Aerobic Respiration
• All use glycolysis (net ATP =2) to oxidize glucose
and harvest chemical energy of food
• In all three, NAD+ is the oxidizing agent that
accepts electrons during glycolysis
• The processes have different final electron
acceptors: an organic molecule (such as pyruvate
or acetaldehyde) in fermentation and O2 in cellular
respiration
• Cellular respiration produces 32 ATP per glucose
molecule; fermentation produces 2 ATP per
glucose molecule
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Figure 9.18
Glucose

Glycolysis
CYTOSOL

Pyruvate
No O2 present: O2 present:
Fermentation Aerobic cellular
respiration

MITOCHONDRION
Ethanol, Acetyl CoA
lactate, or
other products
Citric
acid
cycle
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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The Versatility of Catabolism
• Catabolic pathways funnel electrons from many
kinds of organic molecules into cellular
respiration
• Glycolysis accepts a wide range of
carbohydrates
• Proteins must be digested to amino acids;
amino groups can feed glycolysis or the citric
acid cycle

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 • Fats are digested to glycerol (used in
glycolysis) and fatty acids (used in generating
acetyl CoA)
• Fatty acids are broken down by beta oxidation
and yield acetyl CoA
• An oxidized gram of fat produces more than
twice as much ATP as an oxidized gram of
carbohydrate

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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
• Control of catabolism is based mainly on
regulating the activity of enzymes at
strategic points in the catabolic pathway

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Figure 9.UN06

Inputs Outputs

Glycolysis
Glucose 2 Pyruvate  2 ATP  2 NADH
Figure 9.UN07

Inputs Outputs

2 Pyruvate 2 Acetyl CoA 2 ATP 8 NADH

Citric
2 Oxaloacetate acid
cycle 6 CO2 2 FADH2