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Oxidative Phosphorylation
Conventional view of
mitochondrial
matrix
structure is at right.
Respiratory chain is
intermembrane
cristae
in cristae of the inner
space
membrane.
inner
outer
Spontaneous electron
membrane mitochondrion membrane
transfer through
respiratory chain complexes I, III & IV is coupled to
H+
ejection from the matrix to the intermembrane space.
Because the outer membrane contains large channels,
these protons may equilibrate with the cytosol.
Respiration-linked H+ pumping out of the matrix conserves some of the
free energy of spontaneous e transfers as potential energy of an
electrochemical H+ gradient.
matrix
cristae
intermembrane
space
inner
membrane mitochondrion
outer
membrane
Matrix
Spontaneous
H+ + NADH NAD+ + 2H+
2H+ + O2 H2O
electron flow
2
e
through each of
I
Q
III
IV
complexes I,
III, & IV is
++
coupled to H+
cyt c
+
+
4H
4H
2H+
ejection from
Intermembrane Space
the matrix.
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
++
4H+
4H+
cyt c
2H+
Intermembrane Space
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
++
4H
4H
cyt c
2H+
Intermembrane Space
O
CH3O
CH3
CH3
CH3O
(CH2 CH
O
CH2)nH
CH3O
CH3
CH3
CH3O
(CH2 CH
O
coenzyme Q
CH2)nH
coenzyme Q
e + 2 H+
OH
CH3O
CH3
CH3
CH3O
(CH2 CH
OH
CH2)nH
coenzyme QH2
matrix
Q
2 H+
Q.
QH2
QH2
cyt bH
cyt bL e
One version
of Q Cycle:
Complex III
Q
2 H+
intermembrane space
Fe-S
cyt c1
cyt c
matrix
QH2 gives up 1e
to the Rieske
iron-sulfur center,
Fe-S.
2 H+
Q.
cyt bH
QH2
QH2
Complex III
Fe-S is reoxidized
cyt bL e
e
by transfer of the
Q
Q
Fe-S
cyt c1
cyt c
matrix
2 H+
A 2nd e is
transferred from
Q
Q.
QH2
QH2
the semiquinone to
cyt bH
cyt bL (heme bL)
Complex III
which passes it via
cyt bH across the
cyt bL e
e
membrane to
Q
Q Fe-S
cyt c1
another CoQ
+
2
H
bound at a site on
cyt c
intermembrane space
the matrix side.
The fully oxidized CoQ, generated as the 2nd e is passed to the b
cytochromes, may then dissociate from its binding site adjacent
to the intermembrane space.
Accompanying the two-electron oxidation of bound QH 2, 2H+
are released to the intermembrane space.
matrix
Q
2 H+
Q.
QH2
QH2
cyt bH
cyt bL e
Q
Complex III
Q
2 H+
intermembrane space
Fe-S
cyt c1
cyt c
matrix
Q
2 H+
Q.
QH2
QH2
cyt bH
cyt bL e
Q
Complex III
Q
2 H+
intermembrane space
Fe-S
cyt c1
cyt c
It takes 2 cycles for CoQ bound at the site hear the matrix
to be reduced to QH2, as it accepts 2e from the b hemes,
and 2H+ are extracted from the matrix compartment.
In 2 cycles, 2 QH2 enter the pathway & one is regenerated.
matrix
Animation
2 H+
Q.
QH2
QH2
cyt bH
Overall reaction
catalyzed by
complex III,
including net
inputs & outputs
of the Q cycle :
cyt bL e
Q
Complex III
Q
2 H+
intermembrane space
Fe-S
cyt c1
cyt c
matrix
Q
2 H+
Q.
QH2
QH2
cyt bH
cyt bL e
Q
Complex III
Q
2 H+
intermembrane space
Fe-S
cyt c1
cyt c
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
++
4H+
4H+
cyt c
2H+
Intermembrane Space
Complex III:
Half of the homodimeric
structure is shown.
PDB
1BE3
Complex III
(bc1 Complex)
membrane
Approximate location of
the membrane bilayer is
indicated.
heme bH
heme bL
Fe-S
heme c1
membrane
PDB
1BE3
Complex III
(bc1 Complex)
heme bH
heme bL
Fe-S
heme c1
membrane
PDB
1BE3
Complex III
(bc1 Complex)
heme bH
heme bL
Fe-S
heme c1
Complex III
homo-dimer
PDB-1BGY
Fe-S
heme c1
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
Complex IV
(Cytochrome
Oxidase):
2H+ + O2 H2O
III
IV
++
4H+
4H+
cyt c
2H+
Intermembrane Space
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
++
4H+
4H+
cyt c
2H+
Intermembrane Space
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
Simplified
animation
depicting:
2H+ + O2 H2O
III
IV
++
4H+
4H+
cyt c
2H+
Intermembrane Space
ADP + Pi
ATP
F1
3 H+
matrix
Fo
intermembrane
space
ADP + Pi
ATP
F1
3 H+
matrix
Fo
intermembrane
space
ADP + Pi ATP
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
Fo
++
4H+
F1
4H+
cyt c
2H+
3H+
Intermembrane Space
ADP + Pi ATP
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
Fo
++
4H+
F1
4H+
cyt c
2H+
3H+
Intermembrane Space
ADP + Pi ATP
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
Fo
++
4H+
F1
4H+
cyt c
2H+
3H+
Intermembrane Space
ADP + Pi
ATP
ATP4
matrix
lower [H+]
__
++
3 H+
energy
requiring
reactions
ADP + Pi
higher [H+]
cytosol
ADP + Pi
ATP
ATP4
matrix
lower [H+]
__
++
Animation
3 H+
energy
requiring
reactions
ADP + Pi
higher [H+]
cytosol
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
++
4H+
4H+
cyt c
2H+
Intermembrane Space
Matrix
H+ + NADH NAD+ + 2H+
For, summing up
synthesis of ~P
bonds via ox
phos, assume:
2 e
Q
2H+ + O2 H2O
III
IV
++
4H+
4H+
cyt c
2H+
Intermembrane Space
An oxygen electrode
may be used to record
[O2] in a closed vessel. [O2]
Electron transfer, e.g.,
NADH O2, is
monitored by the rate
of O2 disappearance.
a
c
ADP added
ADP all
converted
to ATP
time
[O2]
ADP added
ADP all
converted
to ATP
time
G
negative value*
positive; depends on H+
gradient**
algebraic sum of above
ADP + Pi ATP
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
Fo
++
4H
F1
4H
cyt c
2H
3H+
Intermembrane Space
ADP + Pi ATP
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
2H+ + O2 H2O
III
IV
Fo
++
4H+
F1
4H+
cyt c
2H+
3H+
Intermembrane Space
OH
NO2
NO2
2,4-dinitrophenol
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
4H
2H+ + O2 H2O
III
4H
IV
cyt c
2H+
uncoupler
H+
Intermembrane Space
Matrix
H+ + NADH NAD+ + 2H+
2 e
Q
4H
2H+ + O2 H2O
III
4H
IV
cyt c
2H+
uncoupler
H+
Intermembrane Space
ADP + Pi
ATP
F1
matrix
3H
Fo
intermembrane
space
Uncoupling Protein
An uncoupling protein (thermogenin) is produced in
brown adipose tissue of newborn mammals and
hibernating mammals.
This protein of the inner mitochondrial membrane
functions as a H+carrier.
The uncoupling protein blocks development of a H+
electrochemical gradient, thereby stimulating
respiration. G of respiration is dissipated as heat.
This "non-shivering thermogenesis" is costly in terms
of respiratory energy unavailable for ATP synthesis, but
provides valuable warming of the organism.