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Cellular Respiration

and Fermentation
• Chemical evolution was driven by energy from
chemicals, radiation, heat, or other sources.

• In general, a cell contains only enough ATP to

sustain from 30 seconds to a few minutes of
normal activity. Because it has such high potential
energy, ATP is unstable and is not stored. As a
result, most cells are making ATP all the time.

• Much of the ATP our cells produce is made using

the chemical energy from glucose.
How do cells obtain glucose?
Photosynthetic organisms can produce glucose from the
products of photosynthesis, where the energy in sunlight is used
to reduce CO2. These organisms will either use the glucose to
make ATP or store it in other energy-rich molecules like starch.
When photosynthetic organisms are eaten or decompose, their
glucose molecules are obtained by animals, fungi, and many
bacteria and archaea.

To withdraw chemical energy from the Storage carbohydrates to

get ATP, storage carbohydrates are first hydrolyzed into their
glucose monomers. The glucose is then used to produce ATP
through one of two general processes: cellular respiration or
fermentation. The primary difference between these two
processes lies in the degree to which glucose is oxidized.
When glucose undergoes the uncontrolled oxidation reaction
called burning, some of the potential energy stored in its
chemical bonds is converted to kinetic energy in the form of
heat and light:

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O + Heat and light energy


Glucose does not burn in cells, however. Instead, it is oxidized

through a long series of carefully controlled redox reactions.
These reactions are occurring, millions of times per minute,
in our cells right now. Instead of releasing all of this energy as
heat, the released free energy is used to synthesize ATP from
ADP and Pi. We use this ATP to read, think, move and stay
The complete oxidation of glucose via cellular respiration
has a set of four interconnected processes that together
convert the chemical energy in glucose to chemical energy
in ATP.

1. Glycolysis

2. Pyruvate processing

3. Citric acid cycle

4. Electron transport and oxidative

Cellular Respiration Plays a Central Role in

• Sets of reactions that break down molecules are

called catabolic pathways. These reactions often
harvest stored chemical energy to produce ATP.

• Anabolic pathways, on the other hand, are sets

of reactions that synthesize larger molecules from
smaller components. Anabolic reactions often use
energy in the form of ATP.
In the 1890s Hans and Edward Buchner were working out techniques
for breaking open baker’s yeast cells and extracting the contents for
commercial and medicinal use. In one set of experiments, the Buchners
added sucrose to their extracts. At the time, sucrose was commonly used
as a preservative—a substance used to prevent food from decaying.
Instead of preserving the yeast extracts, though, the sucrose was quickly
broken down and alcohol appeared as a by-product. This was a key
finding: It showed that metabolic pathways could be studied in
vitro—outside the organism.

When researchers studied how the sugar was being processed, they
found that the reactions could go on much longer than normal if
inorganic phosphate were added to the mixture. This result implied that
some of the compounds involved were being phosphorylated.

In 1905 researchers found that the processing of sugar by yeast extracts

stopped if they boiled the reaction mix. Because it was known that
enzymes could be inactivated by heat, this discovery suggested that
enzymes were involved in at least some of the processing steps.
In both eukaryotes and
prokaryotes, all 10 reactions of
glycolysis occur in the cytosol.
1. Glycolysis starts by using ATP, not producing it. In
the initial step, glucose is phosphorylated to form

• After the second reaction rearranges the sugar to

form fructose-6-phosphate, the third reaction adds a
second phosphate group, forming the compound
fructose-1,6-bisphosphate observed by early

• Thus, in reactions 1–5, two ATP molecules are used

up before any ATP is produced. This part of glycolysis
is referred to as the energy-investment phase.
2. The energy-payoff phase of glycolysis occurs in
reactions 6–10.

• The first high-energy molecules are produced in the

sixth reaction, where two molecules of
(nicotinamide adenine dinucleotide) NAD+ are
reduced to form two NADH.

• In reactions 7 and 10, enzymes catalyze the transfer

of a phosphate group from a phosphorylated
substrate to ADP, forming ATP. Enzyme catalyzed
reactions that result in ATP production are termed
substrate-level phosphorylation.
3. For each molecule of
glucose processed by
glycolysis, the net yield is two
molecules of NADH, two of
ATP, and two of pyruvate.
Processing Pyruvate to Acetyl CoA

In eukaryotes, the pyruvate

produced by glycolysis is
transported from the cytosol to
Pyruvate moves across the mitochondrial outer
membrane through small pores and is transported
into the matrix through a carrier protein in the inner
membrane. Once it is inside the matrix, a sequence of
reactions occurs inside an enormous and intricate
enzyme complex called pyruvate dehydrogenase.
In eukaryotes, this complex is located in the
mitochondrial matrix.

In bacteria and archaea, pyruvate dehydrogenase is

located in the cytosol.
• As pyruvate is being processed, one of its carbons is
oxidized to CO2 and NAD+ is reduced to NADH. The
remaining two-carbon acetyl unit (-COCH3) reacts with
a compound called coenzyme A (CoA).

• Coenzyme A is sometimes abbreviated as CoA-SH to

call attention to its key sulfhydryl functional group.

• The acetyl is transferred to CoA to produce acetyl

CoA. In this and many other reactions, CoA acts as a
coenzyme by accepting and then later transferring an
acetyl group to another substrate (“A” stands for
Acetyl CoA is the final product of the pyruvate-
processing step in glucose oxidation.

Pyruvate, NAD+, and CoA go in;

CO2, NADH, and acetyl CoA come out.

The Citric Acid Cycle:

Oxidizing Acetyl CoA to CO2

Early researchers identified eight small carboxylic acids
that are rapidly oxidized in sequence, from most reduced
to most oxidized.

Redox reactions that involve carboxylic acids often produce

CO2, which is the endpoint of glucose oxidation via cellular
respiration. When they added one of the eight carboxylic
acids to cells, the rate of glucose oxidation increased,
suggesting that the reactions are somehow connected to
pathways involved in glucose catabolism.

Whichever carboxylic acid they added, it did not appear to

be used up. Instead, virtually all the acids could be
recovered later. How is this possible?
• Hans Krebs solved the mystery when he
proposed that the reaction sequence
occurs in a cycle instead of a linear

• Krebs had another crucial insight when

he suggested that the reaction sequence
was directly tied to the processing of
To test these hypotheses, Krebs set out to
determine if adding pyruvate could link the two
ends of the sequence of eight carboxylic acids. If
pyruvate is the key link in forming a cycle, it would
need to be involved in the conversion of the most
oxidized of the eight carboxylic acids to the most
reduced carboxylic acid.

When Krebs added pyruvate, the series of redox

reactions occurred. The conclusion?
The sequence of eight carboxylic acids is indeed
arranged in a cycle.
Many biologists now refer to the cycle as the
citric acid cycle because it starts with
citrate, which is the salt of citric acid after
the protons are released.

The citric acid cycle is also known as the

tricarboxylic acid (TCA) cycle, because
citrate has three carboxyl groups, or as the
Krebs cycle, after its discoverer.
• In each cycle, the energy released by the
oxidation of one molecule of acetyl CoA is used to
produce three molecules of NADH, one of
FADH2, and one of ATP or guanosine
triphosphate (GTP), through substrate-level

• Whether ATP or GTP is produced depends on the

version of the enzyme used in the fifth reaction.
For example, the enzyme used in muscle cells of
mammals produces ATP, while the enzyme used in
liver cells produces GTP.
• In prokaryotes (bacteria and archaea), the
enzymes responsible for the TCA cycle are located
in the cytosol.

• In eukaryotes, most of the enzymes responsible

for the TCA cycle are located in the mitochondrial

• Because glycolysis produces two molecules of

pyruvate, the cycle turns twice for each molecule
of glucose processed in cellular respiration.
Electron transport and oxidative

Electrons from NADH and FADH2 move through a series

of proteins that together are called an electron
transport chain (ETC). The energy released in this chain
of redox reactions is used to create a proton gradient
across a membrane; the ensuing flow of protons back
across the membrane is used to make ATP. Because this
mode of ATP production links the phosphorylation of
ADP with the oxidation of NADH and FADH2, it is called
oxidative phosphorylation.
Mitchell introduced the term chemiosmosis to describe
the use of a proton gradient to drive the production of ATP.
Similar to osmosis, chemiosmosis involves diffusion across
a membrane, but in this case, protons are diffusing along its
gradient rather than water.

Mitchell’s prediction was correct: ATP production

depended solely on the existence of a proton-motive
force, which is based on a proton electrochemical
gradient. It could occur in the absence of an electron
transport chain. This result, along with many others, has
provided strong support for the hypothesis of
chemiosmosis. Most of the ATP produced by cellular
respiration is made by a flow of protons.
The Proton-Motive Force
Couples Electron Transport
to ATP Synthesis
ATP Synthase
The F0 unit serves as a rotor, whose turning is
conveyed to the F1 unit via the shaft. A flow of
protons through the F0 unit causes the rotor
and shaft to spin. As the shaft spins within the
F1 unit, it is thought to change the conformation
of the F1 subunits in a way that catalyzes the
phosphorylation of ADP to ATP.

When protons pass through the ATP synthase, it

spins and releases energy used to synthesize
• The energy to produce ATP in oxidative
phosphorylation comes from an established
proton gradient.

• ETC transports enough protons to produce

approximately three ATP for each NADH and two
for each FADH2, depending on the type of ATP
synthase used.
The chemical equation that represents the
overall process involved in cellular
respiration is

C6H12O6 + 6 H2O + 6 O2 + 29 ADP + 29 Pi →

6 CO2 + 12 H2O + 29 ATP

During cellular respiration, oxygen is
the electron acceptor used by all
eukaryotes and a wide diversity of

Species that depend on oxygen as an

electron acceptor for the ETC use
aerobic respiration and are called
aerobic organisms.
Cellular respiration can occur without oxygen. Many
thousands of bacterial and archaeal species rely on
electron acceptors other than oxygen, and electron
donors other than glucose.

Cells that depend on electron transport chains with

electron acceptors other than oxygen are said to use
anaerobic (“no air”) respiration. Even though the
starting and ending points of cellular respiration differ,
aerobic and anaerobic respiring cells still use an ETC to
create a proton-motive force that drives the synthesis of

In bacteria and archaea, the ETC and ATP synthase are

located in the plasma membrane.
Aerobic Respiration Is Most Efficient

Even though an array of compounds can serve as the

final electron acceptor in cellular respiration, oxygen
provides the greatest energy yield. Because oxygen is
so highly electronegative, the potential energy in bonds
between an oxygen atom and a non-oxygenic atom, such
as hydrogen, is low.
As a result, there is a large difference between the
potential energy of reduced electron donors, like NADH,
and reduced forms of oxygen, like water. This large
differential in potential energy means that the ETC can
generate a large proton-motive force.
Cells that do not use oxygen as an electron
acceptor cannot generate such a large potential
energy difference. As a result, they make less ATP
per electron donor, such as glucose, than cells that
use aerobic respiration.

If cells that use anaerobic respiration compete

with cells using aerobic respiration, those that
use oxygen as an electron acceptor almost always
grow faster and reproduce more.
What happens when the electron acceptors
in an ETC get used up?
When there is no terminal electron acceptor, the electrons in
each of the complexes of the ETC have no place to go and the
electron transport chain stops.
Without an oxidized complex I, NADH remains reduced.
The concentration of NAD+ drops rapidly as cells continue to
convert NAD+ to NADH.
This situation is life threatening. When there is no
longer any NAD+ to drive glycolysis, pyruvate processing, and
the citric acid cycle, then no ATP can be produced. If NAD+
cannot be regenerated somehow, the cell will die.

How do cells cope?

• Fermentation is another process that oxidizes

How does fermentation differ from cellular


Cellular respiration, like burning, results in the

complete oxidation of glucose into CO2 and water.
Fermentation, on the other hand, does not fully
oxidize glucose. Instead, small, reduced organic
molecules are produced as waste. As a result, cellular
respiration releases more energy from glucose than
• Fermentation is a metabolic pathway that regenerates
NAD+ by oxidizing stockpiles of NADH. The electrons
removed from NADH are transferred to pyruvate, or a
molecule derived from pyruvate, instead of an ETC.

• In respiring cells, fermentation serves as an

emergency backup to produce ATP even when the ETC
and oxidative phosphorylation is shut down.

Fermentation may allow the cell to survive in the

absence of an active electron transport chain by
regenerating NAD+ in the cytosol, where glycolysis can
continue to produce ATP.
How does fermentation regenerate NAD+, and
what is being reduced in this redox reaction?

When we run up a long flight of stairs, our muscles begin

metabolizing glucose so fast that the supply of oxygen is
rapidly used up by their mitochondria. When oxygen runs
out, the electron transport chains shut down and NADH
cannot donate its electrons there.

When fermentation takes place in our cells, the pyruvate

produced by glycolysis then begins to accept electrons
from NADH. This process, called lactic acid
fermentation, regenerates NAD+ by forming lactate: a
deprotonated form of lactic acid.
Alcohol fermentation
It occurs in the eukaryote Saccharomyces cerevisiae, strains of
which are used to make baker’s and brewer’s yeast.

When yeast cells grow in bread dough or a bottle of grape juice,

they quickly use up all the available oxygen. Instead of using
NADH to reduce pyruvate, yeast first convert pyruvate to the
two-carbon compound acetaldehyde.

This reaction gives off carbon dioxide, which causes bread to

rise and produces the bubbles in champagne and beer.
Acetaldehyde then accepts electrons from NADH, forming the
NAD+ required to keep glycolysis going. The addition of
electrons to acetaldehyde forms ethanol as a waste product. The
yeast cells excrete ethanol as waste.
Bacteria and archaea that rely exclusively on fermentation are
called obligate anaerobes.

Fermentation is extremely inefficient compared with cellular

respiration. Fermentation produces just 2 molecules of ATP per
molecule of glucose metabolized, while aerobic cellular
respiration produces about 29—almost 15 times more ATP per
glucose molecule than fermentation. The reason for the
disparity is that the fermentation reactions that follow glycolysis
are not used to generate ATP.

Organisms that can switch between fermentation and aerobic

cellular respiration are called facultative anaerobes. The
adjective “facultative” reflects the ability to use aerobic cellular
respiration when oxygen is present and anaerobic fermentation
when it is absent.
Photosynthesis Harnesses Sunlight to Make

Photosynthetic organisms cannot store the electromagnetic

energy of light unless it is first converted into another form.
In photosynthesis, energy in sunlight is transformed to
chemical energy in the C–C and C–H bonds of carbohydrate.

CO2 + H2O + light energy → (CH2O)n + O2

Photosynthesis is an energy demanding series of redox

reactions that produce sugar and O2 from CO2 and water.
Cellular respiration is an energy-releasing series of redox
reactions that produces CO2 and water from sugar and O2.
• When researchers analyzed the chemical composition of
thylakoid membranes, they found huge quantities of

• Pigments are molecules that absorb only certain

wavelengths of light—other wavelengths are either reflected
or transmitted (pass through).

• The most abundant pigment in the thylakoid membranes of

green plants is chlorophyll (“green-leaf”), which reflects or
transmits green light. As a result, chlorophyll is responsible
for the green color of plants, some algae, and many
photosynthetic bacteria.
Which of these wavelengths drive

Engelmann defined the action spectrum

for photosynthesis—the wavelengths that
drive the light-capturing reactions.

• Engelmann’s data indicate that violet-to-

blue and red photons are the most
effective at driving photosynthesis.
An absorption spectrum measures how the wavelength of photons influences
the amount of light absorbed by a pigment. In the combined graph, peaks
indicate wavelengths where absorbance or photosynthetic activity is high;
troughs indicate wavelengths where absorbance or photosynthetic activity is
Chlorophyll a and chlorophyll b are similar in structure.
Both have two fundamental parts: a long isoprenoid
“tail” and a “head” consisting of a large ring structure
with a magnesium atom in the middle. The tail interacts
with proteins embedded in the thylakoid membrane; the
head is where light is absorbed.
When Light Is Absorbed, Electrons Enter an
Excited State

When a chlorophyll molecule absorbs a photon, the

photon’s energy is transferred to bonds in the
chlorophyll molecule’s head region. In response, an
electron becomes “excited,” or bumped up to a higher
energy state.

If the difference between the possible energy states is

the same as the energy in the photon, the photon can be
absorbed and an electron excited to a higher energy
If a pigment absorbs a photon with the right
amount of energy in the form of electromagnetic
radiation, an electron is excited.

The electron now has high potential energy, but

this excited state is unstable and it will last for
only a short time—on the order of picoseconds (1
× 10-12 seconds).

What happens next?

When pigments in chloroplasts absorb photons,
about 2 percent of the excited electrons produce
fluorescence. The other 98 percent of the energized
pigments use their excited electrons to drive

In the thylakoid membrane, 200–300 chlorophyll

molecules and accessory pigments are organized by
an array of proteins to form structures called the
antenna complex & the reaction center. These
complexes, along with the molecules that capture and
process excited electrons, form a photosystem.
The Reaction Center
When a photon or resonance energy from the antenna
complex reaches the reaction center, the energy is
absorbed by one of two specialized chlorophyll
molecules. When this pigment is energized, an excited
electron is transferred from the pigment to an electron

As the acceptor becomes reduced, its potential energy

increases. This is a key step in the transformation of
light energy: Electromagnetic energy from sunlight
has now been transformed to chemical energy.
There are four possible fates of electrons in
chlorophyll that are excited by photons. The
energy released from these electrons can

1. be emitted in the form of light via

fluorescence, or
2. be given off as heat alone, or
3. excite an electron in a nearby pigment and
induce resonance, or
4. be transferred to an electron acceptor in a
redox reaction.
According to the two-photosystem hypothesis, the
enhancement effect occurs because photosynthesis
is much more efficient when both photosystems
operate together.

Subsequent work has shown that the two-

photosystem hypothesis is correct for cyanobacteria
(“blue-green bacteria”) and the chloroplasts of
eukaryotes, such as algae and plants.

These two photosystems differ in structure and

function, but work together in the light-
capturing reactions.
How Does Photosystem II Work?
In photosystem II, the action begins when the antenna
complex transmits resonance energy to the reaction
center, where the electron acceptor pheophytin
comes into play.

Structurally, pheophytin is identical to chlorophyll

except that pheophytin lacks a magnesium atom in its
head region.

Functionally, the two molecules are extremely

What prevents the electron from being pulled back to
the oxidized pigment?
Since the synthesis of ATP in
chloroplasts is initiated by the energy
from light, it is called

In mitochondria, ATP is exported and

fuels many different cellular processes.
In chloroplasts, however, the ATP
remains within the organelle and is used
for the production of carbohydrate.
To continue this ETC, the electrons removed
from the reaction center pigments need to be

Where do the electrons required by

photosystem II come from?

Photosystem II obtains electrons by

oxidizing water.
To manufacture sugar from oxidized carbons in
CO2, strong reducing agents like NADPH are

If the ETC following photosystem II is responsible

for driving the synthesis of ATP, then what
produces the NADPH for the Calvin cycle?

The answer lies in photosystem I.

1. Pigments in the antenna complex absorb photons and
pass the energy to the photosystem I reaction center.

2. Electrons are excited in reaction center chlorophyll


3. The reaction center pigments are oxidized, and the

excited electrons are passed through a series of carriers
inside the photosystem, then to a molecule called
ferredoxin, and then to the enzyme called NADP+

4. NADP+ reductase transfers two electrons and a proton

to reduce NADP+ and form NADPH.
Where do the electrons from
photosystem II end up?

How does the oxidized reaction

center of photosystem I obtain
electrons so NADPH will continue to
be made?
The Z Scheme:
Photosystems II and I
Work Together
When electrons reach the end of the
cytochrome complex, they are passed to
a small diffusible protein called
plastocyanin (PC).

The reduced PC diffuses through the

lumen of the thylakoid, and donates the
electron to an oxidized reaction center
pigment in photosystem I.
Oxygenic Photosynthesis and the Evolution of
According to the geologic record, oxygen levels in the
atmosphere and oceans began to rise only about 2 billion
years ago, as organisms that performed oxygenic
photosynthesis increased in abundance.

O2 was, in fact, almost nonexistent on Earth before

enzymes evolved that could catalyze the oxidation of

Since ozone is formed from O2 gas, this protective layer

would have arisen in our atmosphere only after the
evolution of oxygenic photosynthesis.
How is CO2 Reduced to Produce Sugars?
The reactions that produce sugar from CO2 are not
triggered directly by light. Instead, they depend on the
ATP and NADPH produced by the light-capturing

Carbon fixation is the addition of CO2 to an organic

The discovery of the Calvin cycle clarified how the
ATP and NADPH produced by light-capturing
reactions allow cells to fix and reduce CO2 gas to
carbohydrate (CH2O)n.

Because sugars store a great deal of potential

energy, producing them takes a great deal of
chemical energy.

In the Calvin cycle, each mole of CO2 requires

the energy from 3 moles of ATP and 2 moles of
NADPH to fix it and reduce it to sugar.
All photosynthetic organisms that use the
Calvin cycle to fix carbon require the CO2 -fixing
enzyme ribulose-1,5-bisphosphate
carboxylase/oxygenase (rubisco).

The rubisco enzyme is roughly cube-shaped and

consists of 16 polypeptides that form eight active
sites where CO2 is fixed.
What Happens to the Sugar
That Is Produced by
The products of the Calvin cycle enter one of several
reaction pathways. The most important of these
reaction sequences uses G3P to produce the
monosaccharide glucose, a process called
gluconeogenesis. This glucose is often combined
with fructose, which is also made from G3P, to form
the disaccharide (“two sugar”) sucrose.

When photosynthesis is taking place slowly, almost all

the G3P that is produced is used to make sucrose.
Sucrose is water soluble and readily transported to
other parts of the plant. If sucrose is delivered to
rapidly growing parts of the plant, it is broken down to
fuel cellular respiration and growth.
An alternative pathway occurs when photosynthesis is
proceeding rapidly and sucrose is abundant. Under these
conditions, the glucose molecules are polymerized to form
starch, which is stored in the cells of leaves and roots.

In photosynthesizing cells, starch acts as a temporary

sugar storage product. At night, the starch that is stored in
leaf cells is broken down to glucose molecules. The glucose
is then fed into cellular respiration or used to manufacture
sucrose for transport to other parts of the plant. In this
way, chloroplasts provide sugars for cells throughout the
plant by day and by night.
If a mouse eats the starch that is stored in the leaves
or roots of a plant, however, the chemical energy in
the reduced carbons of starch fuels the mouse’s
growth and reproduction.

If an owl eats the mouse, the chemical energy in the

mouse’s tissues fuels the predator’s growth and

In this way, virtually all cell activity can be traced

back to the sun’s energy that was originally captured
by photosynthesis. Photosynthesis is the staff of life.