Transport Chain

Electron transport chain
&
Oxidative Phosphorylation
 During aerobic catabolism;
 Glucose is completely degraded leading to the releasing CO2
 Reducing equivalent are generated from glucose as well as proteins and fat i.e NADH and FADH2.
?
 Aerobic degradation of carbohydrate, lipid and amino acid converge at the transfer of electron to O 2
 By product of glucose oxidation
 Electrons generated are NOT transfer directly to O2 BUT rather via to
coenzymes i.e.
 NAD+ forming NADH

Review: Generation of reducing equivalents;
 Generation of the reducing coenzymes i.e.
 NADH from NAD+
 FADH2 from FAD

 The transfer of electrons to O2 occurs in the
mitochondria.
 Mitochondria structure
 Two membrane:
1. Outer membrane
2. Inner membrane
 Matrix contains
1. TCA enzymes
2. Amino acid oxidation enzymes
3. Fatty acid β-oxidation enzymes e.t.c
 Reducing equivalents (NADH & FADH2 ) are used to drive ATP
synthesis via oxidative phosphorylation
 Oxidative phosphorylation takes place in the mitochondria;
i. Outer membrane – permeable membrane.
ii. Inner membrane – impermeable membrane
iii. Matrix- -contains various enzymes
 Source of reducing equivalents
 NADH – glycolysis (cytosol) and TCA cycle (mitochondrial
matrix)
 NADH freely passes through the outer membrane to the
intermembrane space.
 However, NADH cannot pass through the impermeable inner
membrane due to lack NADH transporters.
 As a result a NADH transporter system is needed, namely Malate-
Aspartate shuttle.
Malate-Aspartate shuttle
 By malate-aspartate shuttle.
What is the fate of NADH and FADH2 in the
mitochondrial matrix?
 Electrons carried by reducing equivalents are transfer to a terminal electron acceptor
(Oxygen)
 Transfer of electrons is sequential via a series of inner membrane proteins.
 The sequential transfer of electrons is known as electron transport chain
 The by-products of E.T.C are;

1. Water
2. proton gradient (aka. proton motive force or chemiosmotic gradient)
Reactions of the electron transport chain
STEP1: Entry of electron from NADH and FADH2 into ETC
 It is via electron carriers (complexes) in the inner membrane
A. Electron transfer from NADH to Ubiquinone (Q):
− Facilitated by complex I (aka. NADH dehydrogenase).
− NADH dehydrogenase comprises:
1. FMN-containing flavoprotein
2. at least six iron sulfur (Fe-S) centers
− Protons are simultaneously pumped to intermembrane
space.
− Thus, complex I is an electron pump

FMN : Flavin mononucleotide
Reactions of electron transport chain
STEP1: Entry of electrons from NADH and FADH2 into ETC
B. Electron transfer from FADH2 to Ubiquinone:
− Facilitated by complex II (aka. succinate dehydrogenase).
− Complex II is the only membrane enzyme of the TCA
cycle.
− Complex II (FAD) receive electron from TCA cycle
(succinate )
− Electron are then transferred to electron Fe-S centres
which in turn transfer electrons to ubiquinone.

STEP Entry of electrons from NADH and FADH2 into ETC
C. Alternatively, ubiquinone receive electrons from fat (triglyceride)
hydrolysis, :
− That is electron carried by FAD (i.e. FADH2) from by-products of
triglyceride hydrolysis, namely Glycerol and Fatty acid oxidation.
NB: Complex II does not pump electrons into the mitochondrial intermembrane space
STEP 2 : Transfer of electrons from reduced ubiquinone (QH2) to cytochrome C
(cyt− C)Electron transfer is facilitated by complex III also known
as:
− Cytochrome bc1 complex or
− Ubiquinone:cytochrome c oxidoreductase
− Electrons transfer to Cyt C is coupled with transport of
protons from the matrix to the intermembrane space.
NB: Electron transfer involve a switch from a two-electron

STEP 3 : Electron transfer cytochrome C (cyt C) to O2
− Final step of ETC.
− Electrons are transferred to O2.
− O2 is subsequently reduced to form H2O.
− Catalysed by Complex IV aka cytochrome oxidase, Has three subunits
− Subunit I = bind two cytochromes (a and a3) & copper ion (CuB).
− Subunit II= bind CuA centres
− Subunit III
STEP 3 : Electron transfer cytochrome C (Cyt C) to O2
 Cyt C.transfer electron CuA centres
 CuA then transfer electron to Cyt a which transfer it to Cyt a3
 Cyt a3 transfer electron to CuB
 Finally, CuB transfer electron to O2
 For every 4 electrons transfered to O2 , 4 H+ are consumed and
2H2O formed.

STEP 4: Establishment of proton gradient (proton motive
 Transfer of electrons to O2 and pumping of protons into intermembrane space from
force)
matrix create a proton gradient i.e;
 Matrix is negatively charged.
 Intermembrane is positively charged.

STEP 4: Establishment of proton gradient (proton motive
 Proton motive force comprise two components

force)
 Chemical potential energy (difference in concentration of a chemical species i.e H+
and OH-)
 Electrical potential energy (separation of charge)

How does proton gradient drive ATP synthesis?
(Oxidative phosphorylation)
STEP 4: Proton gradient (proton motive force) drive ATP
 This involves spontaneous flow of electrons down

synthesis
electrochemical gradient (back to matrix) provide suficient
energy to drive ATP synthesis from ADP and Pi
 This phenomenon is known as Oxidative phosphorylation,
 It is driven by ATP synthase

STEP 4: proton gradient (proton motive force) and ATP
synthesis
ATP synthase comprise two distinct components:
1. F1, a peripheral membrane protein:
 Comprises α3, β3, Ɣ, δ and ε
 Knoblike portion consist alternating α and β subunits
2. Fo (o denoting oligomycin-sensitive), an integral to the
membrane.
 Consist of subunits a, b and C

STEP 4: proton gradient (proton motive force) drive ATP
synthesis
 ATP synthesis takes please in the β–subunit of F1
 At any given moment :
1. one β–subunit is empty.
2. one β–subunit tightly binds ATP.
3. one β–subunit binds ADP and Pi
 The energy from the flow of protons from intermembrane backto
matrix cause confirmation change of ATP synthase.
 This drives ATP synthesis via rotational catalysis;

STEP 4: proton gradient (proton motive force) and ATP
 When 3 protons (H+) flow back into matrix causes β–subunit
synthesis
undergoes conformational change,
 The consequence conformational change;
1. β–subunit holding ADP and Pi syntheses ATP & held it
with high affinity
2. β–subunit already with tightly bound ATP releases
ATP, as affinity for ATP reduces
3. Empty β–subunit takeup ADP and Pi for another ATP
synthesis cycles
 One complete turn of ATPase yields, 3 ATP molecules
Electron transport chain & Oxidative
Phosphorylation
(Chemiosmotic model)
Inhibition of Electron transport chain & Oxidative Phosphorylation
Cyanide inhibits Cyt A3 of complex IV:

 Inhibits both electron transport chain and oxidative phorphorylation
 Promote anaerobin respiration (↑ lactic acid)
Overview: aerobic glucose
catabolism

Transport Chain

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Electron transport chain

 Electrons generated are NOT transfer directly to O2 BUT rather via to

 NAD+ forming NADH

 Generation of the reducing coenzymes i.e.

 NADH from NAD+

 FADH2 from FAD

 Oxidative phosphorylation takes place in the mitochondria;

i. Outer membrane – permeable membrane.

ii. Inner membrane – impermeable membrane

iii. Matrix- -contains various enzymes

 Source of reducing equivalents

 NADH – glycolysis (cytosol) and TCA cycle (mitochondrial

 However, NADH cannot pass through the impermeable inner

membrane due to lack NADH transporters.

 As a result a NADH transporter system is needed, namely Malate-

 Transfer of electrons is sequential via a series of inner membrane proteins.

 The sequential transfer of electrons is known as electron transport chain

 The by-products of E.T.C are;

 It is via electron carriers (complexes) in the inner membrane

A. Electron transfer from NADH to Ubiquinone (Q):

− Facilitated by complex I (aka. NADH dehydrogenase).

− NADH dehydrogenase comprises:

2. at least six iron sulfur (Fe-S) centers

− Protons are simultaneously pumped to intermembrane

− Thus, complex I is an electron pump

B. Electron transfer from FADH2 to Ubiquinone:

− Facilitated by complex II (aka. succinate dehydrogenase).

− Complex II is the only membrane enzyme of the TCA

− Complex II (FAD) receive electron from TCA cycle

− Electron are then transferred to electron Fe-S centres

which in turn transfer electrons to ubiquinone.

C. Alternatively, ubiquinone receive electrons from fat (triglyceride)

− That is electron carried by FAD (i.e. FADH2) from by-products of

triglyceride hydrolysis, namely Glycerol and Fatty acid oxidation.

(cyt− C)Electron transfer is facilitated by complex III also known

− Cytochrome bc1 complex or

− Electrons transfer to Cyt C is coupled with transport of

protons from the matrix to the intermembrane space.

NB: Electron transfer involve a switch from a two-electron

− Final step of ETC.

− Electrons are transferred to O2.

− O2 is subsequently reduced to form H2O.

− Catalysed by Complex IV aka cytochrome oxidase, Has three subunits

− Subunit II= bind CuA centres

 Cyt C.transfer electron CuA centres

 CuA then transfer electron to Cyt a which transfer it to Cyt a3

 Cyt a3 transfer electron to CuB

 Finally, CuB transfer electron to O2

 For every 4 electrons transfered to O2 , 4 H+ are consumed and

 Matrix is negatively charged.

 Intermembrane is positively charged.

 Proton motive force comprise two components

 Chemical potential energy (difference in concentration of a chemical species i.e H+

 Electrical potential energy (separation of charge)

 This involves spontaneous flow of electrons down

electrochemical gradient (back to matrix) provide suficient

energy to drive ATP synthesis from ADP and Pi