1 Oxidative Phosphorylation

B.
Sc Life Science
Semester 4
Core Paper 4 - Plant Physiology and Metabolism
E- lesson on Respiration: Oxidative phosphorylation
By P. Chitralekha
Associate professor
Department of Botany
Dyal Singh College
At the end of TCA cycle,
• Glucose has been oxidized to carbon dioxide, and the energy of oxidation is trapped mostly in NADH
and FADH2, and a few ATP molecules. Most of the reactions during glucose respiration are redox
reactions.
Glycolysis: Glucose + 2NAD+ + 2ADP + 2Pi → 2Pyruvate + 2NADH + 2H++ 2ATP + 2H2O
2Pyruvate + 2NAD+ → 2 Acetyl-CoA + 2NADH + 2H+ + 2CO2
TCA Cycle: 2Acetyl-CoA+ 6NAD+ + 2FAD + 2ADP + 2Pi + 4H2O→ 4CO2 + 6NADH + 6H+ + 2FADH2 + 2ATP
Glucose + 10NAD+ + 2FAD + 4ADP + 4Pi + 2H2O → 6CO2 + 10NADH + 10H+ + 2FADH2 + 4ATP
• The above reactions are balanced only if Pi represents the H2PO4− ion, ADP and GDP, the ADP2− and
GDP2− ions, respectively, and ATP and GTP the ATP3− and GTP3− ions, respectively. So the 4Pi
account for the remaining 4 substrate H2O molecules.  
[ C6H12O6 + 6O2 + 6H2O 6CO2 + 12H2O + Energy]
• Oxygen has not been utilised till now.

Oxidative Phosphorylation
• It is the last step of cellular respiration process where ATP is synthesised from the reduced
products, NADH and FADH2, of glycolysis and TCA cycle, and oxygen is utilised.
• Though NADH is sometimes used directly as reducing molecule in biosynthetic pathways
such as in gluconeogenesis, both NADH and FADH2 are mostly used to reduce molecular
oxygen to water. The energy released in the process drives the synthesis of ATP, the
energy currency of the cell.
• Process occurs in the presence of oxygen.
• NADH and FADH2 get oxidised by donating two electrons each to oxygen and reducing it to
water.
• Since the energy for synthesis of ATP molecules is derived from oxidation of NADH or
FADH2, which in turn are formed by oxidation of energy-rich molecules such as
carbohydrates and fats, this type of phosphorylation or ATP synthesis is called oxidative
phosphorylation.
• Electrons donated by NADH and FADH2 are transported through components of electron
transport chain (ETC) sequentially from one having a more negative electron/reduction
potential to the next with less negative potential, till they are accepted by molecular
oxygen.
• As the electrons move through components of ETC with increasing redox potentials, their
free energy is reduced and is released.
• According to the chemiosmotic theory put forth by Peter Mitchell in 1961, energy released
during the movement of electrons through ETC is used in pumping protons across the
inner mitochondria membrane and generating a proton concentration gradient. It is the
electrochemical energy of this gradient that drives the synthesis of ATP.
• The components of ETC are electron carriers which are single molecules or multi-subunit
protein complexes or enzymes containing prosthetic groups capable of accepting and
donating electrons. These complexes are associated with the inner mitochondrial
membrane. (In prokaryotes, which lack mitochondria, the complexes are present
associated with the plasmamembrane)
• There are five complexes/enzymes of mitochondrial ETC: 

 
Complex I (also called NADH dehydrogenase or NADH:ubiquinone oxidoreductase)
containing FMN (flavin mononucleotide) and Fe-S ( iron-sulfur) centers catalyses the transfer of
2 electrons from NADH to ubiquinone. During this transfer a large amount of energy is released
which is used to pump protons (H+) from the matrix to the inter-membrane space between the
two mitochondrial membranes. Approximately 4 protons are pumped for every pair of electrons
transferred.  
Complex II ( also called Succinate dehydrogenase complex, an enzyme of TCA cycle) with
FAD and Fe-S centers catalyzes the transfer of electrons from FADH2 to ubiquinone. However,
during this electron transfer energy released is not sufficient to transfer any protons. 
Complex III ( also called cytochrome bc1 complex or ubiquinol: cytochrome c
oxidoreductase) has heme groups of cytochrome b and cytochrome c1 as electron carriers
along with Fe-S centers and transfers electrons from ubiquinol (reduced ubiquinone or QH2) ,
one at a time, to cytochrome c which is present attached on the side of the inner membrane
facing the inter-membrane space. The transfer results in a drop in the free energy of electrons
which is used to pump 4 protons from matrix to the inter-membrane space.
Respiratory Electron Transport Chain in Plants

Complex IV (also called cytochrome c oxidase or cytochrome aa3 complex) has heme a  
and a3 along with copper and iron-copper centers and transfers four electrons, one at a time, 
from cytochrome c to molecular oxygen, reducing it to water.  
Enough energy is released to pump 2 protons from matrix to inter-membrane space.  
Complex V or ATP synthase is a multi-subunit protein complex consisting of two 
major components, F1 – the headpiece which projects into the matrix and  
contains the site for synthesis of ATP, and F0 – the stalk-like integral membrane  
protein complex forming a pore or channel through which protons pass from 
the inter-membrane space to the matrix. It is very similar to ATP synthase enzyme  
found in chloroplasts.
Flow of electrons through components of ETC
NADH+H+ ! Complex I ! Q ! Complex III ! cytochrome c ! Complex IV ! H 2O

↑
Complex II
↑
Succinate
• Plant mitochondria have alternative pathways for oxidising NADH and even NADPH through
alternative oxidase and dehydrogenases. These pathways are protective mechanisms to
dissipate excess energy, and to prevent over-reduction of ETC and formation of severely
damaging reactive oxygen species. 
i) An alternative NADH dehydrogenase present on the matrix side of the inner  
membrane transfers electrons from NADH directly to ubiquinone bypassing 
the rotenone sensitive NADH dehydrogenase complex I and proton pumping at the site. 
As a result fewer protons are pumped. This pathway proceeds only when the ratio of 
NADH/ NAD+ is exceptionally high, especially when photorespiration is occurring at a very high 
rate. 
ii) Similarly, an alternative NADPH dehydrogenase transfers electrons from NADPH in the 
matrix to ubiquinone directly.  
iii) External NADH dehydrogenase and NADPH dehydrogenase situated on the outer side of 
inner membrane toward the inter-membrane space oxidise cytosolic NADH and NADPH,  
respectively. Here, too, the pathway becomes active only when the cytosolic NAD(P)H/NAD(P)+  
ratio is high. These enzymes too are rotenone resistant.
iv) Alternative oxidase, another special enzyme found only in plant mitochondria, catalyses the  
transfers of electrons from ubiquinol to oxygen directly. Thus, electron flow through complex  
III and the cyanide sensitive complex IV, and their proton pumps is bypassed. The unused 
energy is released as heat. This process occurs when the pool of ubiquinone is highly reduced  
and is also called the cyanide resistant respiration. Some plants like the voodoo lily, skunk 
cabbage, and the arum lilies use the heat to volatilise compounds and emit strong odour to 
attract pollinators, and therefore, adapt the alternative oxidase pathway during flowering 
period.
• Apart from these complexes, the ETC has two mobile elements, i) ubiquinone, also called
coenzyme Q or Q, a lipid soluble compound which diffuses freely within the lipid bilayer of
inner mitochondrial membrane and, ii) cytochrome c, a small peripheral protein present on
the side of the inner mitochondrial membrane facing the inter-membrane space.
• The ETC ensures that energy of oxidation of NADH and FADH2 is released in small packets at
three stages (complex I, complex III, and complex IV) rather than in one large burst, and
therefore, is conserved more efficiently.
Since membranes have a dynamic fluid structure, these complexes are randomly arranged and
can move within the membranes; their position is neither fixed nor sequentially arranged.
However, recent evidence suggests that many of the complexes assemble into supramolecular
or super molecular complexes called repirasomes. The functional significance of such
aggregation is not clear but is believed to reduce oxidative damage and increase efficiency of
electron transfer.
Mechanism of ATP synthesis
• Electron transfer through ETC is coupled to proton pumping.
• A total of 10 protons are pumped out for each pair of electrons transferred to O2, 4 protons
by Complex I, 4 by Complex III, and 2 by Complex IV.
• The pumping out of protons from the matrix to the inter membrane space as electrons are
transferred through the ETC, builds up an electrochemical proton concentration gradient
across the inner mitochondrial membrane (the inner mitochondrial membrane is
impermeable to protons).
• The electrochemical energy stored in the proton gradient is called the proton-motive force.
• The release of protons by the spontaneous and passive flow down the electrochemical
gradient through the channel provided by F0 of ATP synthase makes available the proton
motive force.
• This force powers the rotational movement and conformational changes in the F1 subunits
of ATP synthase, resulting in the synthesis of ATP from ADP and Pi.
• Approximately, transfer of 3 or 4 protons are required to drive synthesis of one ATP
molecule.
• Hence, for every NADH oxidised, approximately 3 or 2.5 ATP molecules are formed, and for
every FADH2 oxidised, appx. 2 or 1.5 ATP molecules are formed.  
So, the total ATP molecules produced by oxidation of one glucose molecule is  
10 (NADH) x 3 (2.5) = 30 (25) 
+ 2 (FADH2) x 2 (1.5) = 4 (3) 
+ 4 ATP = 4  
38 (32)
• However, in eukaryotes, NADH produced in cytoplasm during glycolysis, is transferred into
the mitochondrion for oxidation and uses up some energy in the process. As a result
cytoplasmic NADH produces only 2(1.5) ATP molecules. So, in plants, the total number of
ATP molecules produced by respiration of one molecule of glucose is 36 (30).
Go through the e-lesson on oxidative phosphorylation and discuss and clarify any
doubts, on the whatsApp group.
Try to find answers to the following questions and discuss with me.
1. Discuss the significance of ETC.
2. Compare and contrast oxidative phosphorylation and photophosphorylation.
3. Explain the mechanism of oxidative phosphorylation.
4. Write briefly about the components of ETC.
5. Elaborate on the alternative pathways/mechanisms present in plant mitochondria
to oxidise NADH/NADPH?
6. In oxidative phosphorylation, electron transfer through ETC is an absolute
requirement for ATP synthesis but vice versa is not true. Comment.

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B.

Core Paper 4 - Plant Physiology and Metabolism

E- lesson on Respiration: Oxidative phosphorylation

• Oxygen has not been utilised till now.

• There are five complexes/enzymes of mitochondrial ETC:

Respiratory Electron Transport Chain in Plants

Flow of electrons through components of ETC

NADH+H+ ! Complex I ! Q ! Complex III ! cytochrome c ! Complex IV ! H 2O

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