Aerobic Respiration: Chemiosmosis and Electron Transport Chain

Unit 9: Cellular Respiration

Lesson 9.4
Aerobic Respiration: Chemiosmosis and Electron
Transport Chain

Contents
Introduction 1
Learning Objectives 2
Warm Up 3
Learn about It! 5

Electron Transport Chain 5
Complex I 6
Complex II and Ubiquinone 7
Complex III 8
Complex IV 9
Electron Transport Chain and Proton Pumps 11
Chemiosmosis 13
ATP Yield 14
Key Points 15
Check Your Understanding 16
Challenge Yourself 17
Bibliography 17


Lesson 9.4
Aerobic Respiration: Chemiosmosis
and Electron Transport Chain

Introduction
You have probably experienced commuting on your way to school or to your house, and it
must have become a part of your daily routine. We encounter many public utility vehicles,
such as jeepneys, taxis, carpools, and tricycles, as you ride one of them to reach your
destination without getting delayed in your appointments. Despite the differences among
these vehicles, they share one thing in common—they drive you to your destinations by
harnessing the energy that can be generated from burning fuel.

9.4 Aerobic Respiration: Chemiosmosis and Electron Transport Chain 1


The role of vehicles in transporting passengers can be compared to a necessary process
that our bodies need to perform—energy production. In the previous lessons, you have
learned about glycolysis and the Krebs cycle. These stages of cellular respiration generate
energy in the form of ATP, which can be readily used by the cells of our body to drive
chemical reactions for maintenance. During these phases of cellular respiration, electron
carriers have also been reduced. The role of electron carriers can be compared to that of
utility vehicles, where the “passengers” are the electrons. What happens to the electrons
that these electron carriers have? What is their role in aerobic respiration? How can electron
carriers help generate energy?

Learning Objectives DepEd Competencies

In this lesson, you should be able to do the ● Distinguish major features of
glycolysis, Krebs cycle, electron
following:
transport system, and
● Explain the mechanisms of the chemiosmosis
electron transport chain and (STEM_BIO11/12- IIa-j-6).
● Describe reactions that produce
chemiosmosis.
and consume ATP
● Differentiate the protein (STEM_BIO11/12- IIa-j-9).
complexes involved in the ● Describe the role of oxygen in

respiration and describe
electron transport chain.
pathways of electron flow in the
● Determine the ATP yield of the absence of oxygen
electron transport chain and (STEM_BIO11/12- IIa-j-10).

● Compute the number of ATPs
chemiosmosis.
needed or gained in respiration
(STEM_BIO11/12- IIa-j-11).



Warm Up
ETC Simulation 15 minutes
Understanding the complex process of cellular respiration requires patience and several
hours of studying to thoroughly grasp its mechanism. In this activity, you will have a glimpse
of the last stage of cellular respiration through a simulation. This simulation is in the form of
a game that requires speed and alertness.

Materials
● 20 yellow plastic balls
● 50 blue plastic balls
● five small baskets (per group)
● six name tags (with labels NADH, FADH2, C1, C2, C3, C4)

Procedure
1. Divide the class into groups of six.
2. Assign each member of the group to be one of the following items.
a. NADH d. C2
b. FADH2 e. C3
c. C1 f. C4
3. Each group must have four yellow and ten blue plastic balls. The yellow balls
represent electrons, while blue balls represent hydrogen atoms.
4. The members of the group should position themselves, similar to Fig. 9.4.1.
5. One basket should be placed behind the members representing C1, C3, and C4.
Another basket must be placed between NADH and FADH2, and this is where the
balls are placed.



Fig. 9.4.1. The members of each team must position themselves, as shown above.

First Round
1. NADH should throw one yellow and one blue ball to C1. Then, C1 should place the
blue ball in the basket behind him/her. If one of the members fails to catch the ball,
the game will start over again.
2. C1 should pass the yellow ball to C3. When C3 catches the yellow ball, FADH2 should
throw to C3 one blue ball, which should be placed in the basket behind.
3. Then, C3 should pass the yellow ball to C4. When C4 catches the ball, FADH2 should
throw to C4 another blue ball, which should be placed in the basket together with
the yellow ball behind him/her.

Second Round
1. The second round of the game will start with FADH2 throwing one yellow ball to C2.
C2 should pass this yellow ball to C3.
2. If C3 catches the ball, FADH2 should throw to C3 one blue ball, which should be
placed in the basket behind him/her.
3. Then, C3 should pass the yellow ball to C4. If C4 catches the ball, FADH2 should
throw to C4 one blue ball, which should be placed in the basket together with the
yellow ball behind him/her.
4. After the second round, let the game start over from the first round. Let the game
last for two minutes.



5. The group with the most number of blue balls in the basket will be declared as the
winner. The teacher may provide the winning team with a reward.
6. Afterward, answer the guide questions.

Guide Questions
1. What do NADH and FADH2 give off?
2. How do C1, C3, and C4 gain hydrogen balls?
3. What do you think is the specific process that is being simulated when C1, C2, and C4
receive yellow balls that represent electrons?
4. How do you think does the simulation in the game relate to the production of energy
in mitochondria?

Learn about It!

Electron Transport Chain
The first two stages of cellular respiration, which involve glycolysis and Krebs cycle, have
generated energy in the form of ATP. Alongside ATP, the electron carriers NADH and FADH2
are also produced during these processes. These electron carriers transport the electrons to
the inner membrane of mitochondria, where they are used in the electron transport chain
and chemiosmosis (as shown in Fig. 9.4.2). There are four proteins embedded in the
inner membrane of a mitochondrion that forms this transport complex. These protein
complexes receive the electrons from NADH and FADH2. Consequently, hydrogen ions (H+)
from the matrix (the central or innermost region) are pumped (through active transport)
into the intermembrane space. These hydrogen ions play an essential role in the
phosphorylation of ADP to produce ATP through the process of chemiosmosis. Both the
electron transport chain and chemiosmosis constitute the oxidative phosphorylation. This
mechanism of ATP synthesis involves the oxidation of NADH and FADH2 to generate the
energy that will phosphorylate ADP to produce ATP.



Fig. 9.4.2. The protein complexes and ATP synthase of the electron transport chain and
chemiosmosis, respectively, are embedded in the inner mitochondrial membrane.

How do protein complexes differ from each other?

Complex I
The first protein complex that receives electrons from NADH is the complex I. This is
embedded in the inner mitochondrial membrane alongside complexes II to IV. Complex I
has two molecular components that undergo reduction and oxidation or redox reactions
once they receive and release electrons (as shown in Fig. 9.4.3). The first molecule that
accepts the electrons from NADH is a flavoprotein. This molecule contains a prosthetic
group called flavin mononucleotide (FMN). The reduction of FMN happens when it
receives the electrons from the oxidation of NADH to NAD+. These electrons are then
transferred to the second molecule, the iron-sulfur protein (Fe-S). As a result, FMN is



oxidized, while Fe-S is reduced. Also, note that in the figure below, the transfer of electrons
allows complex I to actively pump H+ from the matrix to the intermembrane space.

Fig. 9.4.3. Complex I contains an FMN-containing flavoprotein and an iron-sulfur protein.
These proteins receive and transfer electrons from the oxidation of NADH. Electrons are
then transferred to ubiquinone (labeled as Q in the figure), thus producing QH2.

Complex II and Ubiquinone
Complex II and ubiquinone, although located near each other, differ in function.
Ubiquinone (Q) is a mobile hydrophobic molecule that receives electrons from Fe-S
found in both complexes I and II. Complex II, unlike complex I, does not receive electrons
from the oxidation of NADH. Instead, it accepts electrons from the oxidation of FADH2 to
FAD. Complex II contains succinate dehydrogenase and Fe-S (as shown in Fig. 9.4.4), which
also undergo reduction and oxidation once the complex receives and releases electrons.
Succinate dehydrogenase, the enzyme that is also used in the Krebs cycle, is bound to a FAD
(flavin adenine dinucleotide) molecule. Once FADH2 is oxidized, the complex releases the
electrons (to form FAD) and moves them to Fe-S. Then, the electrons from the Fe-S located
in complex II are transferred to Q, which becomes QH2 (ubiquinol), its reduced form.
Moreover, unlike complex I, complex II does not pump protons into the intermembrane
space.



Fig. 9.4.4. Complex II consists of succinate dehydrogenase (covalently attached to FAD) and
Fe-S that receive electrons from the oxidation of FADH2 to FAD. The succinate
dehydrogenase enzyme also oxidizes succinate to fumarate during the Krebs cycle.

Complex III
The electrons from the Fe-S proteins in Complexes I and II, which are carried by the mobile
carrier QH2, are transferred to Complex III. This complex contains three molecules, namely,
cytochrome b (Cyt b), Rieske center (2Fe-2S center), and cytochrome c1 (Cyt c1).
Cytochromes are proteins that contain a heme group consisting of an iron atom. Iron
atoms in the cytochromes are responsible for receiving and donating electrons.
Cytochromes also differ in the number of heme groups. Cyt b contains two heme groups,
whereas Cyt c1 has one. Electrons from complex I and II are carried by QH2, which can carry
a total of two electrons. After being reduced, QH2 moves and attaches to the complex III and
oxidizes into Q. A series of events then takes place, as shown in Fig. 9.4.5.



Fig. 9.4.5. Q-cycle has two steps that involve the transfer of one electron to cytochrome c
and another electron to Q, reducing it to Q– (step 1). In step 2, another QH2 transfers one
electron to cytochrome c and another electron to Q– to become QH2 again.

The first electron released by QH2 is transferred to Rieske center to Cyt c1. Then, the Cyt c1
transfers the electron to the cytochrome c (Cyt c), which is also considered as a mobile
carrier of electrons similar to ubiquinone. Cyt c transfers electrons from complex III to
complex IV. However, it can only carry one electron at a time. Since QH2 carries two
electrons, one of its electrons passes through another pathway. The second electron is
transferred to Cyt b, then to another molecule of Q located in complex III. Q is converted
into its reduced form semiquinone radical ion (Q–). When another QH2 arrives at the
complex III, it is oxidized again to Q. One electron of QH2 is transferred to the Rieske center
then to Cyt c1 and Cyt c. On the other hand, the second electron is transferred to Cyt b and
then to Q–. Since Q– receives another electron, it becomes QH2. This QH2 molecule is
eventually released to the inner membrane of the mitochondrion. This process is called the
Q-cycle, as shown above in Fig. 9.4.5.

Complex IV
Complex IV, the stepwise reduction of which is shown in Fig. 9.4.6 is the last place where
electrons are transferred before they reduce oxygen to form a water molecule. This
complex consists of cytochrome a (Cyt a), cytochrome a3 (Cyt a3), a pair of copper A
(CuA/CuA), and a copper B (CuB). The mobile cytochrome c that travels from complex III to
complex IV carries one electron. This electron travels to CuA/CuA to Cyt a, and then to Cyt a3
(Step 1). When another Cyt c arrives at the complex IV, the electron it carries follows the
same pathway. However, it does not stop to Cyt a but to CuB (Step 2). Since both Cyt a and



CuB receive electrons, they are now both reduced. Once they are reduced, oxygen (O2)
molecule binds to them, forming a peroxide bridge (Step 3). When two more Cyt c arrives
at the complex IV (Step 4), two more electrons are transferred, which causes the binding of
two hydrogen atoms to break the peroxide bridge forming Cyt a3-OH and CuB-OH (Step
5). Two more hydrogen atoms enter the complex IV to oxidize Cyt a3-OH and CuB-OH (Step
6) into their original forms. As a result, there are two water molecules formed (Step 7).

Fig. 9.4.6. The transfer of electrons from cytochrome leads to the reduction of oxygen
forming water molecules. Thus, in the electron transport chain, oxygen molecules act as the
final electron acceptor during aerobic cellular respiration.



How do heme groups in cytochromes help in the

transport of electrons?

Electron Transport Chain and Proton Pumps
The electron transport chain starts when the electron carriers NADH and FADH2 from
glycolysis and the Krebs cycle are oxidized. When they are oxidized, the electrons they carry
are transferred to the protein complexes of mitochondria. NADH and FADH2 release their
electrons simultaneously. When NADH is oxidized to NAD+, electrons are transferred to
complex I. This causes complex I to pump one hydrogen ion from the matrix to the
intermembrane space. From complex I, electrons travel to the ubiquinone (Q), which
reduces to QH2. Then this molecule moves the electrons to the complex III, which causes it
to pump one hydrogen ion from the matrix to the intermembrane space. The electrons
from complex III are then transferred to complex IV by cytochrome c. When electrons are
transferred to complex IV, another hydrogen ion is pumped to the intermembrane space.
After this, the electrons bind with oxygen (O2), causing it to split and react with hydrogen
ions in the matrix to form water (H2O). Therefore, for every one NADH molecule that is
oxidized, three hydrogen ions are pumped from the matrix into the intermembrane space
(as shown in Fig. 9.4.7).

The electrons released from FADH2 follow the same pathway as the electrons from NADH.
However, the electrons from FADH2 are transferred to complex II instead of complex I.
Unlike complex I, III, and IV, complex II is not a proton pump (which was demonstrated in
the warm-up activity). Thus, when electrons are transferred to complex II, no hydrogen
ions are pumped into the intermembrane space. FADH2 is oxidized to FAD resulting in the
release of electrons. These electrons are transferred to complex II. From complex II,
electrons are transferred to Q, which is again converted to its reduced form QH2. This
molecule transfers the electrons to complex III. One hydrogen ion is pumped by complex III
into the intermembrane space as a result of the transport of electrons. Electrons from
complex III are transported to the complex IV through cytochrome c. The reduction of
complex IV results in the pumping of another hydrogen ion from the matrix to the
intermembrane space. Therefore, for every molecule of FADH2 that is oxidized, two
hydrogen ions are pumped into the intermembrane matrix. The pumping of hydrogen ions



from the matrix to the intermembrane space creates a proton gradient. You can liken a
protein gradient in the intermembrane space as a dam that is building up energy as
complexes I, III, and IV continue pumping protons into it.

Fig. 9.4.7. NADH and FADH2 are oxidized to release electrons into the protein complexes.
These electrons pass through protein complexes and cause the pumping of hydrogen ions
from the matrix to the intermembrane space.

How do hydrogen ions help in the production of

ATP?



Chemiosmosis
The pumping of hydrogen ions into the intermembrane space creates a proton gradient.
This gradient is characterized by a higher concentration of hydrogen ions in the
intermembrane space than in the matrix. The hydrogen ions in the intermembrane space
have to return passively to the matrix to maintain the concentration balance. However,
hydrogen ions cannot just pass through the inner membrane without the help of a
transport protein. The only available channel protein in the inner membrane is ATP
synthase. As hydrogen ions move down their concentration gradient, they power ATP
synthase, which results in the synthesis of ATP molecules. When one hydrogen ion passes
through the ATP synthase, one ATP molecule by adding one phosphate molecule to ADP
(as shown in Fig. 9.4.8). If there are a total of 34 hydrogen ions that pass through ATP
synthase for every electron transport chain, there are also 34 ATP molecules produced.

Fig. 9.4.8. Hydrogen ions in the intermembrane space return to the matrix through ATP
synthase. The movement of the hydrogen ions through the ATP synthase provides energy to
the phosphorylation of ADP to ATP.



ATP Yield
In the electron transport chain, no ATP molecules are produced yet, but the energy that will
drive ATP synthesis is building up through the hydrogen ions. These ions are pumped into
the intermembrane space, where they play a vital role in the production of ATP.
Chemiosmosis is the process by which hydrogen ions are used to power ATP synthase to
phosphorylate ADP into ATP. Because hydrogen ions are used to produce ATP, the number
of NADH and FADH2 produced during glycolysis and Krebs cycle must be accounted for.

As previously discussed, a total of 10 NADH and 2 FADH2 are produced during glycolysis (2
NADH), transition reaction (2 NADH), and the Krebs cycle (6 NADH and 2 FADH2). The
electrons released from NADH pass through complex I, III, and IV. As the electrons pass
through each complex, one hydrogen ion is pumped to the intermembrane space.
Therefore, one NADH molecule is equivalent to three hydrogen ions. Since each hydrogen
ion enables the ATP synthase to produce ATP, it should be noted that for every molecule of
NADH, there are 3 ATP molecules produced. On the other hand, the electrons of FADH2
pass through complex II, III, and IV. However, complex II is not capable of pumping
hydrogen ions to the intermembrane space, while complex III and IV can. Hence, one
molecule of FADH2 is equivalent to 2 hydrogen ions. There are 2 ATP molecules produced
from the oxidation of FADH2. The summary of the products of the electron transport chain
and chemiosmosis is shown in Table 9.4.1.

Table 9.4.1. Summary of the products of the electron transport chain and chemiosmosis
Electron Carriers Number Number of ATP produced
NADH 10 30
FADH2 2 4
Total ATP Yield 34 ATP molecules



Did You Know?

A diet medicine called dinitrophenol (DNP) uncouples ATP
synthesis. DNP affects a person’s ability to obtain energy from the
food he or she eats. However, taking this medicine can cause
hyperthermia because ATP cannot be formed, and the energy is
lost as heat.
Key Points
______________________________________________________________________________________________
● Electron transport chain is a series of four multiprotein complexes embedded
in the inner membrane of the mitochondrion where NADH and FADH2 are
oxidized to release electrons.
● The energy from the released electrons is used to pump hydrogen ions from the
matrix to the intermembrane space. This pumping generates a proton gradient.
● Chemiosmosis involves the downhill transport of hydrogen ions from the
intermembrane space to the matrix. This movement provides energy to ATP
synthase to phosphorylate ADP into ATP.
● In terms of ATP synthesis, NADH is equivalent to three ATP molecules because
its electrons will move through complexes I, III, and IV. By contrast, every FADH2
is equivalent to two ATP molecules because its electrons will only move through
complexes III and IV.
● A total of 34 ATP molecules are produced through electron transport chain and
chemiosmosis. The table below summarizes the net ATP yield for the whole
aerobic respiration process.

Stages Net Direct Products Net ATP Equivalent
2 ATP 2 ATP
Glycolysis
2 NADH 6 ATP
Transition Reaction 2 NADH 6 ATP



2 ATP/GTP 2 ATP
Krebs Cycle 6 NADH 18 ATP
2 FADH2 4 ATP
Maximum ATP Yield 38 ATP molecules

______________________________________________________________________________________________

Check Your Understanding

A. Identify the following terms described in each of the following items.

1. This region of the mitochondrion consists of embedded multiprotein complexes.
2. This mobile protein transfers electrons from complexes I and II to complex III.
3. This enzyme transports hydrogen ions from the intermembrane space to the
matrix.
4. This refers to the core of cytochromes that accept and donate electrons.
5. This protein transfers electrons from complex III to complex IV.
6. This protein receives the first electron from QH2 in complex III.
7. These two molecules serve as the functional components of complex I.
8. This multiprotein complex does not pump hydrogen ions.
9. This is formed when fully reduced Cyt a3 and CuB bind with O2.
10. This is the total number of hydrogen ions pumped into the intermembrane space
in the electron transport chain for every glucose molecule.
11. This molecule is considered as the final electron acceptor in ETC.
12. This is the number of hydrogen ions pumped from the oxidation of one NADH
molecule.
13. This molecule is the reduced form of ubiquinone.
14. This molecule directly transfers electrons to cytochrome c.
15. This is the number of hydrogen ions pumped from the oxidation of one FADH2.



B. Determine the event that immediately follows in each of the

following items.
1. Ubiquinol transfers the electrons to cytochrome b.

2. Hydrogen ions pass through the ATP synthase.
3. Flavin mononucleotide receives electrons from the oxidation of NADH.
4. Cytochrome c transfers the electrons to complex IV.
5. An oxygen molecule is reduced by the electrons that were transported through the
multiprotein complexes.

Challenge Yourself

Provide brief answers and explanations to the following questions.
1. How does the electron transport chain work with chemiosmosis?

2. How do you think the disruption in glycolysis and Krebs cycle will affect the electron
transport chain and chemiosmosis?
3. Barbiturate is a type of drug that inhibits the function of complex I. How do
barbiturates affect the production of ATP?
4. If cyanide only inhibits the function of complex IV, why is cyanide poisoning
considered deadly?
5. What will happen if protons pumped to the intermembrane space leaked into the
matrix without passing through the ATP synthase?

Bibliography
Campbell, Neil A. Biology. Frenchs Forest, N.S.W.: Pearson Benjamin Cummings. 2008.

Ching, Johnny A., Ching, Charmaine E. Biology. Quezon City, Philippines: St. Bernadette
Publishing House Corporation. 2012.



Mader, Sylvia S., Michael Windelspecht, and Sylvia S. Mader. Introductory Biology. United
States: McGraw-Hill Create. 2014.

Miller, Kenneth R., and Joseph S. Levine. Prentice-Hall Biology. Upper Saddle River, NJ:
Pearson Prentice Hall. 2006.

Sabile, Mary Jane G., General Biology 1. Quezon City, Philippines: Phoenix Publishing House,
Inc. 2018.


Aerobic Respiration: Chemiosmosis and Electron Transport Chain

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Unit 9: Cellular Respiration

Learn about It! 5

Key Points 15

Check Your Understanding 16

Challenge Yourself 17

Learning Objectives DepEd Competencies

complexes involved in the ● Describe the role of oxygen in

electron transport chain and (STEM_BIO11/12- IIa-j-10).

Learn about It!

How do protein complexes differ from each other?

How do heme groups in cytochromes help in the

How do hydrogen ions help in the production of

Electron Carriers Number Number of ATP produced

NADH 10 30

Total ATP Yield 34 ATP molecules

Did You Know?

Stages Net Direct Products Net ATP Equivalent

Transition Reaction 2 NADH 6 ATP

Krebs Cycle 6 NADH 18 ATP

Maximum ATP Yield 38 ATP molecules

Check Your Understanding

A. Identify the following terms described in each of the following items.

B. Determine the event that immediately follows in each of the

1. Ubiquinol transfers the electrons to cytochrome b.

Provide brief answers and explanations to the following questions.

1. How does the electron transport chain work with chemiosmosis?

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