Oxidative Phosphorylation - Biology (Article) - Khan Academy

8/29/2017 Oxidative phosphorylation | Biology (article) | Khan Academy
Oxidativephosphorylation
Overview of oxidative phosphorylation. The electron transport
chain forms a proton gradient across the inner mitochondrial
membrane, which drives the synthesis of ATP via chemiosmosis.
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Why do we need oxygen?

You, like many other organisms, need oxygen to live. As you know if youve
ever tried to hold your breath for too long, lack of oxygen can make you feel
dizzy or even black out, and prolonged lack of oxygen can even cause death.
But have you ever wondered why thats the case, or what exactly your body
does with all that oxygen?
As it turns out, the reason you need oxygen is so your cells can use this
molecule during oxidative phosphorylation, the final stage of cellular
respiration. Oxidative phosphorylation is made up of two closely connected
components: the electron transport chain and chemiosmosis. In the electron
transport chain, electrons are passed from one molecule to another, and
energy released in these electron transfers is used to form an
electrochemical gradient. In chemiosmosis, the energy stored in the gradient
is used to make ATP.
So, where does oxygen fit into this picture? Oxygen sits at the end of the
electron transport chain, where it accepts electrons and picks up protons to
form water. If oxygen isnt there to accept electrons (for instance, because a
person is not breathing in enough oxygen), the electron transport chain will
stop running, and ATP will no longer be produced by chemiosmosis. Without
enough ATP, cells cant carry out the reactions they need to function, and,
after a long enough period of time, may even die.
https://www.khanacademy.org/science/biology/cellular-respiration-and-fermentation/oxidative-phosphorylation/a/oxidative-phosphorylation-etc 1/16
In this article, we'll examine oxidative phosphorylation in depth, seeing how it

provides most of the ready chemical energy (ATP) used by the cells in your
body.
Overview: oxidative phosphorylation
Theelectron transport chainis a series of proteins and organic molecules

found in the inner membrane of the mitochondria. Electrons are passed from
one member of the transport chain to another in a series of redox reactions.
Energy released in these reactions is captured as a proton gradient, which is
then used to make ATP in a process calledchemiosmosis. Together, the
electron transport chain and chemiosmosis make upoxidative
phosphorylation. The key steps of this process, shown in simplified form in
the diagram above, include:
Delivery of electrons by NADH and FADH2 .Reduced electron carriers

(NADH and FADH2 ) from other steps of cellular respiration transfer their
electrons to molecules near the beginning of the transport chain. In the
+
process, they turn back into NAD and FAD, which can be reused in other
steps of cellular respiration.
Electron transfer and proton pumping.As electrons are passed down

the chain, they move from a higher to a lower energy level, releasing
+
energy. Some of the energy is used to pump H ions, moving them out of
the matrix and into the intermembrane space. This pumping establishes
an electrochemical gradient.
Splitting of oxygen to form water.At the end of the electron transport

chain, electrons are transferred to molecular oxygen, which splits in half
+
and takes up H to form water.
+
Gradient-driven synthesis of ATP.As H ions flow down their gradient
and back into the matrix, they pass through an enzyme called ATP
synthase, which harnesses the flow of protons to synthesize ATP.
We'll look more closely at both the electron transport chain and
chemiosmosis in the sections below.
The electron transport chain

Theelectron transport chainis a collection of membrane-embedded
proteins and organic molecules, most of them organized into four large
complexes labeled I to IV. In eukaryotes, many copies of these molecules are
found in the inner mitochondrial membrane. In prokaryotes, the electron
transport chain components are found in the plasma membrane.
As the electrons travel through the chain, they go from a higher to a lower
energy level, moving from less electron-hungry to more electron-hungry
molecules. Energy is released in these downhill electron transfers, and
several of the protein complexes use the released energy to pump protons
from the mitochondrial matrix to the intermembrane space, forming a proton
gradient.[Click to see a free energy diagram]
Image modified from "Oxidative phosphorylation: Figure 1", by OpenStax

College, Biology (CC BY 3.0).
All of the electrons that enter the transport chain come from NADH and
FADH2 molecules produced during earlier stages of cellular respiration:
glycolysis, pyruvate oxidation, and the citric acid cycle.
NADHis very good at donating electrons in redox reactions (that is, its
electrons are at a high energy level), so it can transfer its electrons
+
directly to complex I, turning back into NAD . As electrons move through
complex I in a series of redox reactions, energy is released, and the
complex uses this energy to pump protons from the matrix into the
intermembrane space.
FADH2 is not as good at donating electrons as NADH (that is, its

electrons are at a lower energy level), so it cannot transfer its electrons to
complex I. Instead, it feeds them into the transport chain through complex
II, which does not pump protons across the membrane.
Because of this "bypass," each FADH2 molecule causes fewer protons to be

pumped (and contributes less to the proton gradient) than an NADH.
[More about complexes I and II]
Beyond the first two complexes, electrons from NADH and FADH2 travel
exactly the same route. Both complex I and complex II pass their electrons to
a small, mobile electron carrier calledubiquinone(Q), which is reduced to
form QH2 and travels through the membrane, delivering the electrons to
+
complex III. As electrons move through complex III, more H ions are
pumped across the membrane, and the electrons are ultimately delivered to
another mobile carrier calledcytochrome C(cyt C). Cyt C carries the
+
electrons to complex IV, where a final batch of H ions is pumped across the
membrane. Complex IV passes the electrons to O2 , which splits into two
oxygen atoms and accepts protons from the matrix to form water. Four
electrons are required to reduce each molecule of O2 , and two water
molecules are formed in the process.[More about complexes III and IV]
Overall, what does the electron transport chain do for the cell? It has two
important functions:
Regenerates electron carriers.NADH and FADH2 pass their electrons to

+
the electron transport chain, turning back into NAD and FAD. This is
important because the oxidized forms of these electron carriers are used
in glycolysis and the citric acid cycle and must be available to keep these
processes running.
Makes a proton gradient.The transport chain builds a proton gradient

across the inner mitochondrial membrane, with a higher concentration of
+
H in the intermembrane space and a lower concentration in the matrix.
This gradient represents a stored form of energy, and, as well see, it can
be used to make ATP.
Chemiosmosis
Chemiosmosis
Complexes I, III, and IV of the electron transport chain are proton pumps. As
electrons move energetically downhill, the complexes capture the released
+
energy and use it to pump H ions from the matrix to the intermembrane
space. This pumping forms an electrochemical gradient across the inner
mitochondrial membrane. The gradient is sometimes called theproton-
motive force, and you can think of it as a form of stored energy, kind of like a
battery.
Like many other ions, protons can't pass directly through the phospholipid
bilayer of the membrane because its core is too hydrophobic. Instead,
+
H ions can move down their concentration gradient only with the help of
channel proteins that form hydrophilic tunnels across the membrane.
+
In the inner mitochondrial membrane, H ions have just one channel
available: a membrane-spanning protein known asATP synthase.
Conceptually, ATP synthase is a lot like a turbine in a hydroelectric power
+
plant. Instead of being turned by water, its turned by the flow of H ions
moving down their electrochemical gradient. As ATP synthase turns, it
catalyzes the addition of a phosphate to ADP, capturing energy from the
proton gradient as ATP.
Image modified from "Oxidative phosphorylation: Figure 3," by Openstax

College, Biology (CC BY 3.0).
This process, in which energy from a proton gradient is used to make ATP, is
calledchemiosmosis. More broadly, chemiosmosis can refer to any process
in which energy stored in a proton gradient is used to do work. Although
chemiosmosis accounts for over 80% of ATP made during glucose
breakdown in cellular respiration, its not unique to cellular respiration. For
instance, chemiosmosis is also involved in the light reactions
ofphotosynthesis.
What would happen to the energy stored in the proton gradient if it weren't
used to synthesize ATP or do other cellular work? It would be released as
heat, and interestingly enough, some types of cells deliberately use the
proton gradient for heat generation rather than ATP synthesis. This might
seem wasteful, but it's an important strategy for animals that need to keep
warm. For instance, hibernating mammals (such as bears) have specialized
cells known as brown fat cells. In the brown fat cells,uncoupling
proteinsare produced and inserted into the inner mitochondrial membrane.
These proteins are simply channels that allow protons to pass from the
intermembrane space to the matrix without traveling through ATP synthase.

By providing an alternate route for protons to flow back into the matrix, the
uncoupling proteins allow the energy of the gradient to be dissipated as heat.
ATP yield
How many ATP do we get per glucose in cellular respiration? If you look in
different books, or ask different professors, you'll probably get slightly
different answers. However, most current sources estimate that the maximum
2,3,4
ATP yield for a molecule of glucose is around 30-32 ATP . This range is
lower than previous estimates because it accounts for the necessary
transport of ADP into, and ATP out of, the mitochondrion.[More details]
Where does the figure of 30-32 ATP come from? Two net ATP are made in
glycolysis, and another two ATP (or energetically equivalent GTP) are made
in the citric acid cycle. Beyond those four, the remaining ATP all come from
oxidative phosphorylation. Based on a lot of experimental work, it appears
+
that four H ions must flow back into the matrix through ATP synthase to
power the synthesis of one ATP molecule. When electrons from NADH move
+
through the transport chain, about 10 H ions are pumped from the matrix to
the intermembrane space, so each NADH yields about 2.5 ATP. Electrons
from FADH2 , which enter the chain at a later stage, drive pumping of only 6
+
H , leading to production of about 1.5 ATP.
With this information, we can do a little inventory for the breakdown of one
molecule of glucose:
Stage Direct products (net) Ultimate ATP yield (net)

Glycolysis 2 ATP 2 ATP
2 NADH 3-5 ATP
Stage Direct products (net) Ultimate ATP yield (net)

Pyruvate oxidation 2 NADH 5 ATP
Citric acid cycle 2 ATP/GTP 2 ATP
6 NADH 15 ATP
2 FADH2 3 ATP
Total 30-32 ATP
[Click here for a diagram showing ATP production]
One number in this table is still not precise: the ATP yield from NADH made
in glycolysis. This is because glycolysis happens in the cytosol, and NADH
can't cross the inner mitochondrial membrane to deliver its electrons to
complex I. Instead, it must hand its electrons off to a molecular shuttle
system that delivers them, through a series of steps, to the electron transport
chain.
Some cells of your body have a shuttle system that delivers electrons to
the transport chain via FADH2 . In this case, only 3 ATP are produced for
the two NADH of glycolysis.
Other cells of your body have a shuttle system that delivers the electrons
via NADH, resulting in the production of 5 ATP.
In bacteria, both glycolysis and the citric acid cycle happen in the cytosol, so
no shuttle is needed and 5 ATP are produced.
30-32 ATP from the breakdown of one glucose molecule is a high-end

estimate, and the real yield may be lower. For instance, some intermediates
from cellular respiration may be siphoned off by the cell and used in other
biosynthetic pathways, reducing the number of ATP produced. Cellular
respiration is a nexus for many different metabolic pathways in the cell,

forming anetworkthats larger than the glucose breakdown pathways alone.
Self-check questions
1. Cyanide acts as a poison because it inhibits
complex IV, making it unable to transport
electrons.How would cyanide poisoning affect 1)
the electron transport chain and 2) the proton
gradient across the inner mitochondrial
membrane?
Choose 1 answer:
The electron transport chain would speed up, and the

gradient would become stronger
B
The electron transport chain would stop, and the

gradient would decrease
C
Both the electron transport chain and the gradient

would stay the same
D
The electron transport chain would be re-routed

through complex II, and the gradient would become
weaker
[Hint]
2. Dinitrophenol (DNP) is a chemical that acts as an

uncoupling agent, making the inner mitochondrial
membrane leaky to protons. It was used until 1938
as a weight-loss drug.How would DNP affect the
amount of ATP produced in cellular respiration?
Why do you think it is now off the market?*
Choose 1 answer:
It would increase ATP production, but could also

cause dangerously high body temperature
B
It would decrease ATP production, but could also

cause dangerously high body temperature
C
It would decrease ATP production, but could also

cause dangerously low body temperature
D
It would increase ATP production, but could also

cause dangerously low body temperature
[Hint]
Check
[Attribution and references]
Oxidative phosphorylation
Oxidative phosphorylation and the electron transport chain
Oxidative phosphorylation
Practice: Oxidative phosphorylation and the ETC
Next tutorial
Variations on cellular respirat
Ask a question...
Questions Tips & Thanks
how does the nadh from glycolisys gets into the matrix so its
electron could be used?
7votes 1 comment Flag 2 years agoby Maulana Akmal
It's actually called Malate-aspartate shuttle

6votes Comment Flag about a year agoby Eric Ballesteros
Show all 3 answers Answer this question
The new Campbell Biology textbook updated the ATP yield totals
to be 26-28 (instead of 30-32). What does this mean for your table
on the 'breakdown of one molecule of glucose'? Where did the net
yield go down?
4votes Comment Flag 11 months agoby Dallas Huggins
well, seems like scientists have recently discovered that the

old ATP yield is not quite accurate, and the most recent data
shows that it should be around 26-28

2votes Comment Flag 10 months agoby Peony
What does substrate level phosphorylation means?

3votes Comment Flag 10 months agoby Herukm18
Substrate level phosphorylation is when a substrate brings the

phosphate directly to ADP to form ATP. Hope this helps.
2votes Comment Flag 7 months agoby Alex Leung
if the volume of the intermembrane space was increased, what

effect would this have on the function of a mitochondrion?
2votes Comment Flag 4 months agoby na26262
Hm.... A cell stays small to allow easier transport of molecules

and charged particles from organelles. If the intermembrane
space of the mitochondria was increased, I would think that
respiration would be less efficient, because now the electrons
have to cross a larger space and lose much more energy.
So.... That's my guess and it would probably be wrong,
2votes Comment Flag about a month agoby Richard Wu
Answer this question...
I thought it was 38 ATPs from the previous videos

2votes 1 comment Flag 10 months agoby SanteeAlexander
Yes it was, but that is when the ATP production was at it's
highest so to speak. That could be right from what I have
learned.
2votes Comment Flag 9 months agoby MichaelCainBlack
Where do the H+ ions in the electron transport chain come from?

Also, how does the NADH release electrons, and if NADH is not
the thing that releases H+ ions, then why does NAD+ get reduced
to NADH?
1vote Comment Flag 2 years agoby Aiza B.
They come from NADH and FADH(subscript 2)

3votes Comment Flag about a year agoby Chibuzor Nwalor
Why do the elections transfer through all the complexes in such a

specific order. Why not reduce the oxygen directly which seems to
be easier since oxygen is more electron hunger.
2votes 1 comment Flag about a year agoby Lucy Ch
to prevent that energy is lost (cannot be used for work).

1vote Comment Flag about a year agoby mariana rosas
I don't really understand the difference between substrate-level

phosphorylation and oxidative phosphorylation. Maybe someone
can explain? Thanks in advance.
1vote Comment Flag about a year agoby Ugne Steponaviciute
Substrate-level phosphorylation is when a phosphate group is

added forcibly to ADP to form ATP. Oxidative phosphorylation,
however, is specifically when the gradient of H+ across ATP
Synthase sort of "pushes" the reaction of ADP into ATP.
3votes Comment Flag 11 months agoby mariielatham
How much H2O is produced is the electron transport chain?

1vote Comment Flag 10 months agoby richie56rich
For every electron one H2O is produced in the ETC.

2votes 3 comments Flag 9 months agoby MichaelCainBlack
"As the electrons travel through the chain, they go from a higher to
a lower energy level, moving from less electron-hungry to more
electron-hungry molecules."
So is it possible that some electrons move from, lets say, enzyme
1 to enzyme 4 whithout getting to enzyme 3? and thus indirectly
creating less ATP?
1vote Comment Flag 9 months agoby Greacus
From my understanding this occurs because, for the electron

to be passed along the ETC spontaneously each consecutive
acceptor has to have a higher electron affinity than the donor.
(That should explain the statement in quotes) The reason that
this occurs however is because Complexes I, III and IV work
to create a proton gradient that is necessary to power ATP
Synthase. So if the electrons skipped from Complex I (I
assume that is what you mean by enzyme) to Complex IV, per
each electron (thatskips) that results in like 8 protons not
being pumped out into the intermembrane space and thus the
proton motive force wouldn't be as strong. The proton motive
force is the entire reason why ATP synthase even functions

because those protons do not want to remain in the
intermembrane space they want to be in the Matrix so they
escape through the channel in ATP synthase which
coincidently uses that to power oxidative phosphorylation.
Another way to think of the proton is as form of energy that is
released from the electron as it is passed down the chain.
ETC functions to harness every ounce of energy because
energy loss is not favorable to this system. So making less
ATP is redundant to the system that desires to harness
energy. Instead of making less ATP/glucose when there is an
abundance of ATP the process of energy production is slowed
down elsewhere and the energy is not lost but harnessed for
later use. ATP in excess inhibits glycolysis, as well as
pyruvate oxidation to acetyl-CoA and Citric Acid Cycle. This
form of regulation is favorable in terms of energy, because
you never want less ATP, you want the system to function at
max capacity so that energy is not lost.

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Why do we need oxygen?

In this article, we'll examine oxidative phosphorylation in depth, seeing how it

Overview: oxidative phosphorylation

Theelectron transport chainis a series of proteins and organic molecules

Delivery of electrons by NADH and FADH2 .Reduced electron carriers

Electron transfer and proton pumping.As electrons are passed down

Splitting of oxygen to form water.At the end of the electron transport

The electron transport chain

Image modified from "Oxidative phosphorylation: Figure 1", by OpenStax

FADH2 is not as good at donating electrons as NADH (that is, its

Because of this "bypass," each FADH2 molecule causes fewer protons to be

Regenerates electron carriers.NADH and FADH2 pass their electrons to

Makes a proton gradient.The transport chain builds a proton gradient

Image modified from "Oxidative phosphorylation: Figure 3," by Openstax

intermembrane space to the matrix without traveling through ATP synthase.

Stage Direct products (net) Ultimate ATP yield (net)

Stage Direct products (net) Ultimate ATP yield (net)

[Click here for a diagram showing ATP production]

30-32 ATP from the breakdown of one glucose molecule is a high-end

respiration is a nexus for many different metabolic pathways in the cell,

The electron transport chain would speed up, and the

The electron transport chain would stop, and the

Both the electron transport chain and the gradient

The electron transport chain would be re-routed

2. Dinitrophenol (DNP) is a chemical that acts as an

It would increase ATP production, but could also

It would decrease ATP production, but could also

It would decrease ATP production, but could also

It would increase ATP production, but could also

[Attribution and references]

Oxidative phosphorylation and the electron transport chain

Practice: Oxidative phosphorylation and the ETC

Questions Tips & Thanks

It's actually called Malate-aspartate shuttle

Show all 3 answers Answer this question

well, seems like scientists have recently discovered that the

shows that it should be around 26-28

Show all 3 answers Answer this question

What does substrate level phosphorylation means?

Substrate level phosphorylation is when a substrate brings the

Show all 2 answers Answer this question

if the volume of the intermembrane space was increased, what

Hm.... A cell stays small to allow easier transport of molecules

Answer this question...

I thought it was 38 ATPs from the previous videos

Show all 2 answers Answer this question

Where do the H+ ions in the electron transport chain come from?

They come from NADH and FADH(subscript 2)

Show all 3 answers Answer this question

Why do the elections transfer through all the complexes in such a

to prevent that energy is lost (cannot be used for work).

Answer this question...

I don't really understand the difference between substrate-level

Substrate-level phosphorylation is when a phosphate group is

Show all 2 answers Answer this question

How much H2O is produced is the electron transport chain?

For every electron one H2O is produced in the ETC.

Answer this question...

From my understanding this occurs because, for the electron

force is the entire reason why ATP synthase even functions

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