Chapter 7

The first energy-releasing pathways

evolved about 3.8 billion years ago and
could run to completion without oxygen.

Many bacteria and protistans still make

ATP by anaerobic pathways, mainly
fermentation and anaerobic electron
transport.
Some cells in your own body can use an
anaerobic route for short periods, but they do
so only when they don't receive enough
oxygen.

Your cells use mainly aerobic respiration.

With each breath you take, you provide your
actively respiring cells with a fresh supply of
oxygen.

energy_releasing.swf
The main energy-releasing pathways all
start in the cytoplasm with glycolysis, a
pathway in which enzymes cleave and
rearrange each glucose molecule into two
pyruvate molecules. Once this stage is over,
the energy-releasing pathways differ.
When a glucose molecule is the starting
material, aerobic respiration can be
summarized this way:

C H12O6 --------> 6CO2 + 6H2O

6
However, this summary equation tells us
only what the substances are at the start
and finish of the pathway.

In between are three reaction stages. These

stages are briefly described in the clip

aerobic_stages.swf
Each glucose molecule has six carbon atoms,
twelve hydrogen atoms, and six oxygen atoms
covalently bonded to one another.

During glycolysis, glucose or some other

carbohydrate in the cytoplasm partially breaks
down to pyruvate, a molecule with a backbone
of three carbon atoms.
Glycolysis takes place in two major parts.
The first part requires energy input. The
second part releases energy.

The Figure shows a schematic of the

energy changes during glycolysis.

glycolysis_two_stages.swf
Glycolysis begins when ATP molecules each
transfer a phosphate group to glucose and so
donate energy to it.

Such a transfer is known as phosphorylation.

In this case, phosphorylation raises the energy
content of glucose to a level that is high
enough to enable entry into the second part,
the energy-releasing steps of glycolysis.
The first energy-releasing step cleaves the
activated glucose into two molecules,
which we can call PGAL
(phosphoglyceraldehyde).

Each PGAL gives up two electrons and a

hydrogen to the coenzyme NAD+, reducing
it to NADH.
The first energy-releasing step cleaves the
activated glucose into two molecules,
which we can call PGAL
(phosphoglyceraldehyde).

Each PGAL gives up two electrons and a

hydrogen to the coenzyme NAD+, reducing
it to NADH.
As a result of these donations, each PGAL
is converted to an unstable intermediate.

This intermediate enables ATP to form by

giving up a phosphate group to ADP. The
next intermediate in the sequence does the
same thing.
A total of four ATP form by substrate-level
phosphorylation. This metabolic event is defined
as the direct transfer of a phosphate group from
a substrate of a reaction to some other
molecule, such as ADP.

Remember that two ATP were invested to start

the reactions, so the net energy yield from
glycolysis is only two ATP. Two NADH also form.
The animation provides a step-by-step
look at the reactions of glycolysis.

glycolysis.swf
Suppose two pyruvate molecules, formed
by glycolysis, leave the cytoplasm and
enter a mitochondrion.

In this organelle, both the second and

third stages of the aerobic pathway run
to completion. The Figure reviews the
structure of a mitochondrion.

mitochondrion.swf
The second stage occurs in the inner
compartment of the mitochondrion.

It starts with a few preparatory steps in which

an enzyme removes a carbon atom from each
pyruvate molecule.

A coenzyme, known as coenzyme A, becomes

acetyl-CoA when it combines with the remaining
two-carbon fragment.
preparatory_rx.swf
The second-stage reactions continue when
acetyl-CoA transfers the two-carbon
fragment to oxaloacetate, the entry point
of the Krebs cycle.
The reactions of the Krebs
cycle in more detail

The combination of acetyl-CoA and

oxaloacetate forms citrate. The Krebs cycle
breaks down citrate into carbon dioxide and
water in a stepwise fashion. All the carbon
molecules of pyruvate eventually end up in
carbon dioxide.
There are three other important points to
understand about the Krebs cycle:

1. Hydrogen and electrons are transferred

to the coenzymes NAD+ and FAD (flavin
adenine dinucleotide, a different coenzyme)
to produce NADH and FADH2.

2. Substrate-level phosphorylations produce

more ATP.

3. Oxaloacetate regenerates. (That's why

the process is called a cycle.)
The animation shows the reactions and
intermediates of the Krebs cycle in
detail.

krebs_cycle_reactions.swf
In total, the second-stage reactions produce
two ATP, eight NADH, and two FADH2 for each
molecule of glucose. The coenzymes go to the
electron transport system for the final stage of
the aerobic pathway.

The animation will show a final overview of the

second-stage reactions

krebs_telecourse.mov
ATP production goes into high gear in the
third stage of the aerobic pathway, electron
transport phosphorylation.

During the earlier stages, hydrogen and

electrons were stripped from reactants and
loaded onto the coenzymes NAD and FAD,
reducing them to NADH and FADH2.
In the final stage, these coenzymes deliver
hydrogen ions and electrons to an electron
transfer chain in the inner mitochondrial
membrane.

The electrons are transferred from one

molecule of the chain to the next molecule
in line.
When certain molecules accept and then
donate electrons, they also pick up
hydrogen ions in the inner compartment.

Quickly afterward, they release the ions to

the outer compartment.
This shuttling action sets up H+
concentration and electric gradients across
the inner mitochondrial membrane.

Nearby in the membrane, H+ ions follow

the gradients and flow back to the inner
compartment, through the interior of ATP
synthases.
The H+ flow through these transport proteins
drives the formation of ATP from ADP and
unbound phosphate.

Free oxygen keeps ATP production going. When

it withdraws electrons at the end of the
transport systems and then combines with H+,
water is the result.

This process is illustrated in the following

animations
electron_transport.mov

mito_chemiosmosis.swf
Thirty-two ATP typically form during the
third stage of aerobic respiration.

Add these to the net yield from the

preceding stages, and the total harvest is
thirty-six ATP from one glucose molecule.

However, the exact yield varies depending

on the type of cell and prevailing
conditions. See Figure.

energy_harvest.swf
The human body has many alternative sources
of energy.
Complex carbohydrates, fats, and proteins
cannot enter the aerobic pathway directly.
The digestive system and individual cells must
first break apart these molecules into simpler
degradable subunits.
The Figure shows the reaction sites where a
variety of organic compounds can enter the
stage of aerobic respiration.

alt_energy_sources.swf
In common usage, the term fermentation is
used to describe the process by which alcoholic
beverages such as beer and wine are produced.

This is alcoholic fermentation, an energy-

releasing pathway. It is an example of an
anaerobic pathway because, unlike aerobic
respiration, it does not require oxygen.
Many microorganisms rely entirely on
fermentation pathways to meet their modest
energy requirements.

Other microorganisms switch back and forth

between aerobic and anaerobic pathways as
the oxygen levels in their environments
change.
The yeast used to brew alcoholic
beverages is an example.

Human muscle cells also can switch back

and forth. They utilize a fermentation
pathway (lactate fermentation) during
periods of vigorous exercise, when they are
short of oxygen.
Like all the main energy-releasing pathways,
fermentation begins with the breakdown of
glucose by glycolysis.

However, the reactions do not completely

break down glucose to carbon dioxide and
water, and produce no more ATP beyond the
tiny yield from glycolysis.

The final steps serve only to regenerate NAD+,

a coenzyme with central roles in the
breakdown reactions.
Exactly what happens after glycolysis
depends on what type of fermentation is
taking place.

fermentation.swf
Summary of Cellular
Respiration

CellRespiration.svg
The End