Chapter 9
Cellular
Respiration and
Fermentation
Lecture Presentations by
Nicole Tunbridge and
© 2017 Pearson Education, Inc.
Kathleen Fitzpatrick
Figure 9.2
Light
energy
ECOSYSTEM
Photosynthesis
in chloroplasts Organic
CO2 + H2O + O2
molecules
Cellular respiration
in mitochondria
ATP powers
ATP
most cellular work
Heat
energy
© 2017 Pearson Education, Inc.
Concept 9.1: Catabolic pathways yield energy
by oxidizing organic fuels
Catabolic pathways release stored energy by
breaking down complex molecules
Electron transfer plays a major role in these
pathways
These processes are central to cellular respiration
© 2017 Pearson Education, Inc.
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
© 2017 Pearson Education, Inc.
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)
© 2017 Pearson Education, Inc.
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)
© 2017 Pearson Education, Inc.
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
© 2017 Pearson Education, Inc.
Oxidation of Organic Fuel Molecules During
Cellular Respiration
During cellular respiration, the fuel (such as glucose)
is oxidized, and O2 is reduced
Organic molecules with an abundance of hydrogen
are excellent sources of high-energy electrons
Energy is released as the electrons associated with
hydrogen ions are transferred to oxygen, a lower
energy state
© 2017 Pearson Education, Inc.
Figure 9.UN03
becomes oxidized
becomes reduced
© 2017 Pearson Education, Inc.
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
© 2017 Pearson Education, Inc.
Figure 9.4
NAD+ NADH
Dehydrogenase
Reduction of NAD+
2[H]
(from food) Oxidation of NADH
Nicotinamide Nicotinamide
(oxidized form) (reduced form)
© 2017 Pearson Education, Inc.
The Stages of Cellular Respiration: A Preview
Harvesting of energy from glucose has three stages
1. Glycolysis (breaks down glucose into two
molecules of pyruvate)
2. The citric acid cycle (completes the breakdown of
glucose)
3. Oxidative phosphorylation (accounts for most of
the ATP synthesis)
© 2017 Pearson Education, Inc.
Figure 9.UN05
1. GLYCOLYSIS (color-coded blue throughout the chapter)
2. PYRUVATE OXIDATION and the CITRIC ACID CYCLE
(color-coded light orange and dark orange)
3. OXIDATIVE PHOSPHORYLATION: Electron transport and
chemiosmosis (color-coded purple)
© 2017 Pearson Education, Inc.
Figure 9.6_1
Electrons
via NADH
GLYCOLYSIS
Glucose Pyruvate
CYTOSOL MITOCHONDRION
ATP
Substrate-level
© 2017 Pearson Education, Inc.
Figure 9.6_2
Electrons Electrons
via NADH via NADH
and FADH2
GLYCOLYSIS PYRUVATE
OXIDATION CITRIC
ACID
Glucose Pyruvate Acetyl CoA CYCLE
CYTOSOL MITOCHONDRION
ATP ATP
Substrate-level Substrate-level
© 2017 Pearson Education, Inc.
Figure 9.6_3
Electrons Electrons
via NADH via NADH
and FADH2
GLYCOLYSIS PYRUVATE OXIDATIVE
OXIDATION CITRIC PHOSPHORYLATION
ACID
Glucose Pyruvate Acetyl CoA CYCLE (Electron transport
and chemiosmosis)
CYTOSOL MITOCHONDRION
ATP ATP ATP
Substrate-level Substrate-level Oxidative
© 2017 Pearson Education, Inc.
The process that generates almost 90% of the ATP
is called oxidative phosphorylation because it is
powered by redox reactions
A smaller amount of ATP is formed in glycolysis and
the citric acid cycle by substrate-level
phosphorylation
© 2017 Pearson Education, Inc.
For each molecule of glucose degraded to CO2 and
water by respiration, the cell makes up to 32
molecules of ATP
© 2017 Pearson Education, Inc.
Concept 9.2: Glycolysis harvests chemical
energy by oxidizing glucose to pyruvate
Glycolysis (“sugar splitting”) breaks down glucose
into two molecules of pyruvate
Glycolysis occurs in the cytoplasm and has two
major phases
Energy investment phase
Energy payoff phase
Glycolysis occurs whether or not O2 is present
© 2017 Pearson Education, Inc.
Figure 9.UN06
CITRIC OXIDATIVE
PYRUVATE
GLYCOLYSIS ACID PHOSPHORYL-
OXIDATION
CYCLE ATION
ATP
© 2017 Pearson Education, Inc.
Figure 9.8
Energy Investment Phase
Glucose
2 ATP used 2 ADP + 2 P
Energy Payoff Phase
4 ADP + 4 P 4 ATP formed
2 NAD+ + 4 e– + 4 H+ 2 NADH + 2 H+
2 Pyruvate + 2 H2O
Net
Glucose 2 Pyruvate + 2 H2O
4 ATP formed – 2 ATP used 2 ATP
2 NAD+ + 4 e– + 4 H+ 2 NADH + 2 H+
© 2017 Pearson Education, Inc.
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 a
mitochondrion (in eukaryotic cells), where the
oxidation of glucose is completed
© 2017 Pearson Education, Inc.
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
This step is carried out by a multienzyme complex
that catalyzes three reactions
1. Oxidation of pyruvate and release of CO2
2. Reduction of NAD+ to NADH
3. Combination of the remaining two-carbon fragment
and coenzyme A to form acetyl CoA
© 2017 Pearson Education, Inc.
Figure 9.UN07
CITRIC OXIDATIVE
PYRUVATE
GLYCOLYSIS ACID PHOSPHORYL-
OXIDATION
CYCLE ATION
© 2017 Pearson Education, Inc.
Figure 9.10
MITOCHONDRION
CYTOSOL
Coenzyme A
CO2
1 3
NAD+ NADH + H+ Acetyl CoA
Pyruvate
Transport protein
© 2017 Pearson Education, Inc.
The Citric Acid Cycle
The citric acid cycle, also called the Krebs cycle,
completes the breakdown of pyruvate to CO2
The cycle oxidizes organic fuel derived from
pyruvate, generating 1 ATP, 3 NADH, and 1 FADH2
per turn
© 2017 Pearson Education, Inc.
Figure 9.UN08
CITRIC OXIDATIVE
PYRUVATE
GLYCOLYSIS ACID PHOSPHORYL-
OXIDATION
CYCLE ATION
ATP
© 2017 Pearson Education, Inc.
Figure 9.11
PYRUVATE OXIDATION
Pyruvate
(from glycolysis,
2 molecules per glucose)
CO2
NAD+
CoA
NADH
+ H+ Acetyl CoA
CoA
NADH
+ H+ CoA
NAD+
CITRIC
ACID
2 CO2
CYCLE
FADH2 2 NAD+
FAD 2 NADH
+ 2 H+
ADP + P i
ATP
© 2017 Pearson Education, Inc.
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
© 2017 Pearson Education, Inc.
Figure 9.UN09
CITRIC OXIDATIVE
PYRUVATE
GLYCOLYSIS ACID PHOSPHORYL-
OXIDATION
CYCLE ATION
ATP
© 2017 Pearson Education, Inc.
Figure 9.13
NADH (least electronegative)
50
2 e– NAD+
Complexes I-IV
FADH2
each consist of
2 e– FAD multiple proteins
Free energy (G) relative to O2 (kcal/mol)
40 FMN I with electron
Fe•S Fe•S II carriers.
Q
III
Cyt b
30 Fe•S
Cyt c1 IV
Cyt c
Cyt a
Electron transport Cyt a3
20 chain
10 2 e–
2 H+ + ½ O2
0 (most electronegative)
H2O
© 2017 Pearson Education, Inc.
Figure 9.13a
NADH (least electronegative)
50
2 e–
NAD+
Free energy (G) relative to O2 (kcal/mol)
Complexes I-IV
FADH2
each consist of
2 e– FAD multiple proteins
40 I with electron
FMN
II carriers.
Fe•S Fe•S
Q
III
Cyt b
Fe•S
30
Cyt c1 IV
Cyt c
Cyt a
Electron transport chain Cyt a3
20
10 2 e–
© 2017 Pearson Education, Inc.
Figure 9.13b
30
Cyt c1
Free energy (G) relative to O2 (kcal/mol)
IV
Cyt c
Cyt a
Cyt a3
20
10 2 e–
2 H+ + ½ O2
0 (most electronegative)
H2O
© 2017 Pearson Education, Inc.
Chemiosmosis: The Energy-Coupling
Mechanism
The energy released as electrons are passed down
the electron transport chain is used to pump H+ from
the mitochondrial matrix to the intermembrane space
H+ then moves down its concentration gradient back
across the membrane, passing through the protein
complex ATP synthase
© 2017 Pearson Education, Inc.
H+ moves into binding sites on the rotor of ATP
synthase, causing it to spin in a way that catalyzes
phosphorylation of ADP to ATP
This is an example of chemiosmosis, the use of
energy in a H+ gradient to drive cellular work
© 2017 Pearson Education, Inc.
Certain electron carriers in the electron transport
chain accept and release H+ along with the electrons
In this way, 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
© 2017 Pearson Education, Inc.
Figure 9.15
H+ ATP
Protein H+
H+ synthase
complex H+
of electron Cyt c
carriers
IV
Q
I III
II
2 H+ + ½ O2 H2O
FADH2 FAD
NADH NAD+
ADP + P i ATP
(carrying electrons
from food) H+
1 Electron transport chain 2 Chemiosmosis
Oxidative phosphorylation
© 2017 Pearson Education, Inc.
Figure 9.15a
H+
H+
Protein H+
complex
of electron Cyt c
carriers
IV
Q
I III
II
2 H+ + ½ O2 H2O
FADH2 FAD
NADH NAD+
(carrying electrons
from food)
1 Electron transport chain
© 2017 Pearson Education, Inc.
Figure 9.15b
ATP
synthase
H+
ADP + P i ATP
H+
2 Chemiosmosis
© 2017 Pearson Education, Inc.
Figure 9.16
Electron shuttles MITOCHONDRION
span membrane 2 NADH
or
CYTOSOL 2 FADH2
2 NADH 2 NADH 6 NADH 2 FADH2
GLYCOLYSIS PYRUVATE OXIDATIVE
OXIDATION CITRIC
PHOSPHORYLATION
ACID
Glucose 2 Pyruvate 2 Acetyl CoA (Electron transport
CYCLE
and chemiosmosis)
+ 2 ATP + 2 ATP + about 26 or 28 ATP
About
Maximum per glucose:
30 or 32 ATP
© 2017 Pearson Education, Inc.
Figure 9.16a
Electron shuttles
span membrane 2 NADH
or
2 FADH2
2 NADH
GLYCOLYSIS
Glucose 2
Pyruvate
+ 2 ATP
© 2017 Pearson Education, Inc.
Figure 9.16b
2 NADH 6 NADH 2 FADH2
PYRUVATE CITRIC
OXIDATION ACID
2 Acetyl CoA CYCLE
+ 2 ATP
© 2017 Pearson Education, Inc.
Figure 9.16c
2 NADH
or
2 FADH2
2 NADH 6 NADH 2 FADH2
OXIDATIVE
PHOSPHORYLATION
(Electron transport
and chemiosmosis)
+ about 26 or 28 ATP
© 2017 Pearson Education, Inc.
Figure 9.16d
About
Maximum per glucose: 30 or 32 ATP
© 2017 Pearson Education, Inc.
Concept 9.5: Fermentation and anaerobic
respiration enable cells to produce ATP without
the use of oxygen
Most cellular respiration depends on electronegative
oxygen to pull electrons down the transport chain
Without oxygen, the electron transport chain will
cease to operate
In that case, glycolysis couples with anaerobic
respiration or fermentation to produce ATP
© 2017 Pearson Education, Inc.
Anaerobic respiration uses an electron transport
chain with a final electron acceptor other than
oxygen, for example, sulfate
Fermentation uses substrate-level phosphorylation
instead of an electron transport chain to generate
ATP
© 2017 Pearson Education, Inc.
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
© 2017 Pearson Education, Inc.
In alcohol fermentation, pyruvate is converted to
ethanol in two steps
The first step releases CO2 from pyruvate
The second step produces NAD+ and ethanol
Alcohol fermentation by yeast is used in brewing,
winemaking, and baking
© 2017 Pearson Education, Inc.
Figure 9.17
2 ADP + 2 P i 2 ATP 2 ADP + 2 P i 2 ATP
Glucose GLYCOLYSIS Glucose GLYCOLYSIS
2 Pyruvate
2 NAD+ 2 NADH 2 CO2 2 NAD+ 2 NADH
+ 2 H+ + 2 H+
2 Pyruvate
NAD+ REGENERATION NAD+ REGENERATION
2 Ethanol 2 Acetaldehyde 2 Lactate
(a) Alcohol fermentation (b) Lactic acid fermentation
© 2017 Pearson Education, Inc.
In lactic acid fermentation, pyruvate is reduced by
NADH, forming NAD+ and lactate as end products,
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 during strenuous exercise when O2 is
scarce
© 2017 Pearson Education, Inc.
Figure 9.17b
2 ADP + 2 P i 2 ATP
Glucose GLYCOLYSIS
2 NAD+ 2 NADH
+ 2 H+
2 Pyruvate
NAD+ REGENERATION
2 Lactate
(b) Lactic acid fermentation
© 2017 Pearson Education, Inc.
Obligate anaerobes carry out fermentation or
anaerobic respiration and cannot survive in the
presence of O2
Yeast and many bacteria are facultative
anaerobes, meaning that they can survive using
either fermentation or cellular respiration
In a facultative anaerobe, pyruvate is a fork in the
metabolic road that leads to two alternative catabolic
routes
© 2017 Pearson Education, Inc.
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
© 2017 Pearson Education, Inc.