Chapter 6

How Cells Harvest Chemical Energy

Introduction

 In eukaryotes, cellular respiration

– harvests energy from food,
– yields large amounts of ATP, and
– Uses ATP to drive cellular work.

 A similar process takes place in many prokaryotic

organisms.

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Figure 6.0_1
Chapter 6: Big Ideas

Cellular Respiration: Stages of Cellular

Aerobic Harvesting Respiration
of Energy

Fermentation: Anaerobic Connections Between

Harvesting of Energy Metabolic Pathways
 CELLULAR RESPIRATION:
AEROBIC HARVESTING
OF ENERGY

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6.1 Photosynthesis and cellular respiration
provide energy for life
 Life requires energy.
 In almost all ecosystems, energy ultimately comes
from the sun.
 In photosynthesis,
– some of the energy in sunlight is captured by
chloroplasts,
– atoms of carbon dioxide and water are rearranged, and
– glucose and oxygen are produced.

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6.1 Photosynthesis and cellular respiration
provide energy for life
 In cellular respiration
– glucose is broken down to carbon dioxide and water
and
– the cell captures some of the released energy to make
ATP.
– Cellular respiration takes place in the mitochondria of
eukaryotic cells.

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Figure 6.1
Sunlight energy

ECOSYSTEM

Photosynthesis
in chloroplasts

CO2 Glucose

H2O O2

Cellular respiration
in mitochondria

(for cellular
ATP
work)

Heat energy
Figure 6.2

Breathing
O2 CO2

Lungs

CO2 Bloodstream O2

Muscle cells carrying out

Cellular Respiration
Glucose  O2 CO2  H2O  ATP
6.3 Cellular respiration banks energy in ATP
molecules
 Cellular respiration is an exergonic process that
transfers energy from the bonds in glucose to form
ATP.
 Cellular respiration
– produces up to 32 ATP molecules from each glucose
molecule and
– captures only about 34% of the energy originally stored
in glucose.

 Other foods (organic molecules) can also be used

as a source of energy.
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Figure 6.3

C6H12O6 6 O2 6 CO2 6 H2O ATP

Glucose Oxygen Carbon Water  Heat

dioxide
6.4 CONNECTION: The human body uses energy
from ATP for all its activities
 The average adult human needs about 2,200 kcal
of energy per day.
– About 75% of these calories are used to maintain a
healthy body.
– The remaining 25% is used to power physical activities.

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6.4 CONNECTION: The human body uses energy
from ATP for all its activities
 A kilocalorie (kcal) is
– the quantity of heat required to raise the temperature of
1 kilogram (kg) of water by 1oC,
– the same as a food Calorie, and
– used to measure the nutritional values indicated on food
labels.

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Figure 6.4

Activity kcal consumed per hour

by a 67.5-kg (150-lb) person*
Running (8–9 mph) 979
Dancing (fast) 510
Bicycling (10 mph) 490
Swimming (2 mph) 408
Walking (4 mph) 341
Walking (3 mph) 245
Dancing (slow) 204
Driving a car 61
Sitting (writing) 28

*Not including kcal needed for

body maintenance
6.5 Cells tap energy from electrons “falling”
from organic fuels to oxygen
 How do cells extract the energy from glucose?

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6.5 Cells tap energy from electrons “falling”
from organic fuels to oxygen
 When the carbon-hydrogen bonds of glucose are
broken, electrons are transferred to oxygen.
– Oxygen has a strong tendency to attract electrons.
– An electron loses potential energy when it “falls” to
oxygen.

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6.5 Cells tap energy from electrons “falling”
from organic fuels to oxygen
 The movement of electrons from one molecule to
another is an oxidation-reduction reaction, or
redox reaction. In a redox reaction,
– the loss of electrons from one substance is called
oxidation,
– the addition of electrons to another substance is called
reduction,
– a molecule is oxidized when it loses one or more
electrons, and
– reduced when it gains one or more electrons.

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6.5 Cells tap energy from electrons “falling”
from organic fuels to oxygen
 A cellular respiration equation is helpful to show
the changes in hydrogen atom distribution.
 Glucose
– loses its hydrogen atoms and
– becomes oxidized to CO2.

 Oxygen
– gains hydrogen atoms and
– becomes reduced to H2O.

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Figure 6.5A

Loss of hydrogen atoms

(becomes oxidized)

C6H12O6 6 O2 6 CO2 6 H2O ATP

Glucose  Heat
Gain of hydrogen atoms
(becomes reduced)
6.5 Cells tap energy from electrons “falling”
from organic fuels to oxygen
 Energy can be released from glucose by simply
burning it.
 The energy is dissipated as heat and light and is
not available to living organisms.

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6.5 Cells tap energy from electrons “falling”
from organic fuels to oxygen
 On the other hand, cellular respiration is the
controlled breakdown of organic molecules.
 Energy is
– gradually released in small amounts,
– captured by a biological system, and
– stored in ATP.

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6.5 Cells tap energy from electrons “falling”
from organic fuels to oxygen
 Enzymes are necessary to oxidize glucose and
other foods.
 NAD+ (Nicotinamide adenine dinucleotide)
– is an important co-enzyme in oxidizing glucose,
– accepts electrons, and
– becomes reduced to NADH.

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Figure 6.5B

Becomes oxidized
2H

Becomes reduced
NAD 2H NADH H
(carries
2 H 2 2 electrons)
6.5 Cells tap energy from electrons “falling”
from organic fuels to oxygen
 There are other electron “carrier” molecules that
function like NAD+.
– They form a staircase where the electrons pass from
one to the next down the staircase.
– These electron carriers collectively are called the
electron transport chain.
– As electrons are transported down the chain, ATP is
generated.

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Figure 6.5C

NADH

NAD ATP
2 Controlled
H release of
energy for
synthesis
of ATP

2
1 O
2 H 2 2

H 2O
 STAGES OF CELLULAR
RESPIRATION

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 Glycolysis
Glucose Pyruvate x 2

Acetyl-coA
Citric acid cycle CO2 x 2

NAD+, FAD NADH, FADH2

Oxidative phosphorylation
Chemiosmosis
Electron transport chain
ATP synthase
6.6 Overview: Cellular respiration occurs in
three main stages
 Cellular respiration consists of a sequence of steps
that can be divided into three stages.
– Stage 1 – Glycolysis
– Stage 2 – Pyruvate oxidation and citric acid cycle
– Stage 3 – Oxidative phosphorylation

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6.6 Overview: Cellular respiration occurs in
three main stages
 Stage 1: Glycolysis
– occurs in the cytoplasm,
– begins cellular respiration, and
– breaks down glucose into two molecules of a three-
carbon compound called pyruvate.

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6.6 Overview: Cellular respiration occurs in
three main stages
 Stage 2: The citric acid cycle
– takes place in mitochondria,
– oxidizes pyruvate to a two-carbon compound, and
– supplies the third stage with electrons.

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6.6 Overview: Cellular respiration occurs in
three main stages
 Stage 3: Oxidative phosphorylation
– involves electrons carried by NADH and FADH2,
– shuttles these electrons to the electron transport chain
embedded in the inner mitochondrial membrane,
– involves chemiosmosis, and
– generates ATP through oxidative phosphorylation
associated with chemiosmosis.

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Figure 6.6

CYTOPLASM

NADH
Electrons NADH FADH2
carried by NADH

Glycolysis Oxidative
Pyruvate Citric Acid Phosphorylation
Glucose Pyruvate
Oxidation Cycle (electron transport
and chemiosmosis)

Mitochondrion

ATP
Substrate-level Substrate-level Oxidative
ATP ATP
phosphorylation phosphorylation phosphorylation
6.7 Glycolysis harvests chemical energy by
oxidizing glucose to pyruvate
 In glycolysis,
– a single molecule of glucose is enzymatically cut in half
through a series of steps,
– two molecules of pyruvate are produced,
– two molecules of NAD+ are reduced to two molecules of
NADH, and
– a net of two molecules of ATP is produced.

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Figure 6.7A

Glucose

2 ADP
2 NAD
2 P

2 NADH

2 ATP 2 H

2 Pyruvate
6.7 Glycolysis harvests chemical energy by
oxidizing glucose to pyruvate
 ATP is formed in glycolysis by substrate-level
phosphorylation during which
– an enzyme transfers a phosphate group from a
substrate molecule to ADP and
– ATP is formed.

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Figure 6.7B

Enzyme Enzyme

P ADP

ATP

P P
Substrate Product
6.7 Glycolysis harvests chemical energy by
oxidizing glucose to pyruvate
 The steps of glycolysis can be grouped into two
main phases.
– In steps 1–4, the energy investment phase,
– energy is consumed as two ATP molecules are used to
energize a glucose molecule,
– which is then split into two small sugars that are now primed
to release energy.

– In steps 5–9, the energy payoff,

– two NADH molecules are produced for each initial glucose
molecule and
– ATP molecules are generated.
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Figure 6.7Ca_s1
Glucose ENERGY
Steps 1 – 3 A fuel ATP INVESTMENT
molecule is energized, Step PHASE
using ATP. 1
ADP

P Glucose 6-phosphate

P Fructose 6-phosphate
ATP
3
ADP
Fructose
P P
1,6-bisphosphate
Figure 6.7Ca_s2
Glucose ENERGY
Steps 1 – 3 A fuel ATP INVESTMENT
molecule is energized, Step PHASE
using ATP. 1
ADP

P Glucose 6-phosphate

P Fructose 6-phosphate
ATP
3
ADP
Step 4 A six-carbon Fructose
intermediate splits P P
1,6-bisphosphate
into two three-carbon
4
intermediates.

Glyceraldehyde
P P
3-phosphate (G3P)
Figure 6.7Cb_s1
P P ENERGY
PAYOFF
PHASE
Step 5 NAD NAD
5 5
A redox reaction P P
NADH NADH
generates NADH.
H H
P P P P 1,3-Bisphospho-
glycerate
Figure 6.7Cb_s2
P P ENERGY
PAYOFF
PHASE
Step 5 NAD NAD
5 5
A redox reaction P P
NADH NADH
generates NADH.
H H
P P P P 1,3-Bisphospho-
ADP ADP glycerate
6 6
Steps 6 – 9
ATP and pyruvate ATP ATP
are produced.
P P 3-Phospho-
glycerate
7 7

P P

2-Phospho-
glycerate
8 H2O
8
H2O

P P

Phosphoenol-
ADP ADP pyruvate (PEP)
9 9
ATP ATP
Pyruvate
6.8 Pyruvate is oxidized prior to the citric acid
cycle
 The pyruvate formed in glycolysis is transported
from the cytoplasm into a mitochondrion where
– the citric acid cycle and
– oxidative phosphorylation will occur.

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6.8 Pyruvate is oxidized prior to the citric acid
cycle
 Two molecules of pyruvate are produced for each
molecule of glucose that enters glycolysis.
 Pyruvate does not enter the citric acid cycle, but
undergoes some chemical grooming in which
– a carboxyl group is removed and given off as CO2,
– the two-carbon compound remaining is oxidized while a
molecule of NAD+ is reduced to NADH,
– coenzyme A joins with the two-carbon group to form
acetyl coenzyme A, abbreviated as acetyl CoA, and
– acetyl CoA enters the citric acid cycle.
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Figure 6.8

NAD NADH H
2

CoA
Pyruvate 1 Acetyl coenzyme A
3
CO2
Coenzyme A
6.9 The citric acid cycle completes the oxidation of
organic molecules, generating many NADH
and FADH2 molecules
 The citric acid cycle
– is also called the Krebs cycle (after the German-British
researcher Hans Krebs, who worked out much of this
pathway in the 1930s),
– completes the oxidation of organic molecules, and
– generates many NADH and FADH2 molecules.

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Figure 6.9A
Acetyl CoA
CoA
CoA

2 CO2
Citric Acid Cycle

FADH2 3 NAD

FAD 3 NADH

3 H

ATP ADP P
6.9 The citric acid cycle completes the oxidation of
organic molecules, generating many NADH
and FADH2 molecules
 During the citric acid cycle
– the two-carbon group of acetyl CoA is added to a four-
carbon compound, forming citrate,
– citrate is degraded back to the four-carbon compound,
– two CO2 are released, and
– 1 ATP, 3 NADH, and 1 FADH2 are produced.

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Figure 6.9B_s1
Acetyl CoA CoA
CoA

2 carbons enter cycle

Oxaloacetate
1

Citric Acid Cycle

Step 1
Acetyl CoA stokes
the furnace.
Figure 6.9B_s2
Acetyl CoA CoA
CoA

2 carbons enter cycle

Oxaloacetate
1

Citrate
NAD

NADH H
2
Citric Acid Cycle
CO2 leaves cycle

Alpha-ketoglutarate

3
CO2 leaves cycle

NAD

ADP P NADH H

Step 1 Steps 2 – 3 ATP

Acetyl CoA stokes NADH, ATP, and CO2
the furnace. are generated during redox reactions.
Figure 6.9B_s3
Acetyl CoA CoA
CoA

2 carbons enter cycle

Oxaloacetate
1

Citrate
NADH H
NAD
5
NAD
NADH H
2
Citric Acid Cycle
Malate CO2 leaves cycle

FADH2 Alpha-ketoglutarate
4
3
FAD CO2 leaves cycle

NAD
Succinate
ADP P NADH H

Step 1 Steps 2 – 3 ATP Steps 4 – 5

Acetyl CoA stokes NADH, ATP, and CO2 Further redox reactions generate
the furnace. are generated during redox reactions. FADH2 and more NADH.
6.9 The citric acid cycle completes the oxidation of
organic molecules, generating many NADH
and FADH2 molecules
 Remember that the citric acid cycle processes two
molecules of acetyl CoA for each initial glucose.
 Thus, after two turns of the citric acid cycle, the
overall yield per glucose molecule is
– 2 ATP,
– 6 NADH, and
– 2 FADH2.

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6.10 Most ATP production occurs by oxidative
phosphorylation
 Oxidative phosphorylation
– involves electron transport and chemiosmosis and
– requires an adequate supply of oxygen.

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 mitochondria

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6.10 Most ATP production occurs by oxidative
phosphorylation
 Electrons from NADH and FADH2 travel down the
electron transport chain to O2.
 Oxygen picks up H+ to form water.
 Energy released by these redox reactions is used
to pump H+ from the mitochondrial matrix into the
intermembrane space.
 In chemiosmosis, the H+ diffuses back across the
inner membrane through ATP synthase
complexes, driving the synthesis of ATP.

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Figure 6.10

H H H H
H
Intermem- Protein H Mobile H
brane electron H H ATP
complex carriers
space of electron synthase
carriers III
IV
I

Inner mito-
chondrial II
membrane
Electron FADH2 FAD
flow 1
2 H H2O
NADH NAD 2 O2
Mito- H
chondrial
matrix ADP P ATP
H

Electron Transport Chain Chemiosmosis

Oxidative Phosphorylation
 mitochondria

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6.11 CONNECTION: Interrupting cellular respiration
can have both harmful and beneficial effects

 Three categories of cellular poisons obstruct the

process of oxidative phosphorylation. These
poisons
1. block the electron transport chain (for example,
rotenone, cyanide, and carbon monoxide),
2. inhibit ATP synthase (for example, the antibiotic
oligomycin), or
3. make the membrane leaky to hydrogen ions (called
uncouplers, examples include dinitrophenol).

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Figure 6.11

Rotenone Cyanide, Oligomycin

H
H carbon monoxide
H H ATP
synthase
H H H

DNP

FADH2 FAD
1
NAD O 2 H
NADH 2 2

H
H2O ADP P ATP
6.11 CONNECTION: Interrupting cellular respiration
can have both harmful and beneficial effects

 Brown fat is
– a special type of tissue associated with the generation
of heat and
– more abundant in hibernating mammals and newborn
infants.

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6.11 CONNECTION: Interrupting cellular respiration
can have both harmful and beneficial effects

 In brown fat,
– the cells are packed full of mitochondria,
– the inner mitochondrial membrane contains an
uncoupling protein, which allows H+ to flow back down
its concentration gradient without generating ATP, and
– ongoing oxidation of stored fats generates additional
heat.

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6.12 Review: Each molecule of glucose yields
many molecules of ATP
 Recall that the energy payoff of cellular respiration
involves
1. glycolysis,
2. alteration of pyruvate,
3. the citric acid cycle, and
4. oxidative phosphorylation.

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6.12 Review: Each molecule of glucose yields
many molecules of ATP
 The total yield is about 32 ATP molecules per
glucose molecule.
 This is about 34% of the potential energy of a
glucose molecule.
 In addition, water and CO2 are produced.

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Figure 6.12

CYTOPLASM
Electron shuttles Mitochondrion
across membrane 2 NADH
2 NADH or
2 FADH2

2 NADH 6 NADH 2 FADH2

Glycolysis Pyruvate Oxidative

2 Oxidation Phosphorylation
Glucose Citric Acid
Pyruvate 2 Acetyl (electron transport
Cycle
CoA and chemiosmosis)

Maximum
per glucose:

2 2  about
ATP ATP 28 ATP About
32 ATP
by substrate-level by substrate-level by oxidative
phosphorylation phosphorylation phosphorylation
 FERMENTATION: ANAEROBIC
HARVESTING OF ENERGY

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6.13 Fermentation enables cells to produce ATP
without oxygen
 Fermentation is a way of harvesting chemical energy
that does not require oxygen. Fermentation
– takes advantage of glycolysis,
– produces two ATP molecules per glucose, and
– reduces NAD+ to NADH.

 The trick of fermentation is to provide an anaerobic

path for recycling NADH back to NAD+.

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6.13 Fermentation enables cells to produce ATP
without oxygen
 Your muscle cells and certain bacteria can oxidize
NADH through lactic acid fermentation, in which
– NADH is oxidized to NAD+ and
– pyruvate is reduced to lactate.

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Figure 6.13A
Glucose

2 ADP 2 NAD

Glycolysis
2 P

2 ATP 2 NADH

2 Pyruvate

2 NADH

2 NAD

2 Lactate
6.13 Fermentation enables cells to produce ATP
without oxygen
 Lactate is carried by the blood to the liver, where it
is converted back to pyruvate and oxidized in the
mitochondria of liver cells.
 The dairy industry uses lactic acid fermentation by
bacteria to make cheese and yogurt.
 Other types of microbial fermentation turn
– soybeans into soy sauce and
– cabbage into sauerkraut.

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6.13 Fermentation enables cells to produce ATP
without oxygen
 The baking and winemaking industries have used
alcohol fermentation for thousands of years.
 In this process yeasts (single-celled fungi)
– oxidize NADH back to NAD+ and
– convert pyruvate to CO2 and ethanol.

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Figure 6.13B
Glucose

2 ADP 2 NAD

Glycolysis
2 P

2 ATP 2 NADH

2 Pyruvate

2 NADH
2 CO2

2 NAD

2 Ethanol
 CONNECTIONS BETWEEN
METABOLIC PATHWAYS

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6.15 Cells use many kinds of organic molecules
as fuel for cellular respiration
 Although glucose is considered to be the primary
source of sugar for respiration and fermentation,
ATP is generated using
– carbohydrates,
– fats, and
– proteins.

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Figure 6.15
Food, such as
peanuts

Carbohydrates Fats Proteins

Sugars Glycerol Fatty acids Amino acids

Amino
groups

Citric
G3P Pyruvate Pyruvate Oxidative
Glucose Oxidation
Acid
Phosphorylation
Glycolysis Acetyl CoA Cycle

ATP