Post on 22-Feb-2018
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Discovery of Oxidative Phosphorylation and the
Mechanism of ATP SynthaseDr. Peter Mitchell was awarded the 1978Nobel Prize in Chemistry for formulating
the chemiosmotic hypothesis that
describes the process by which oxidative
phosphorylation occurs.
In 1997, Drs. Paul Boyer and John
Walker jointly received the NobelPrize in Chemistry for elucidating
the mechanism of ATP Synthase.
Boyer Walker
Mitchell
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NADH + 1/2 O2 + H+ => H2O + NAD+ !G = - 53 kcal/mol
ADP + Pi + H+ => ATP + H2O !G = + 7.3 kcal/mol
The pH in the inter-membrane space is 1.4 units lower than in the matrix
The potential over the inter-membrane due to the charge separation is 0.14 V
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Reaction Catalyzed by ATP Synthase
• ATP is synthesized through the condensation of ADP and Pi, the reverse of hyolysis
•
Mg2+ complexes of ADP and ATP are the actual substrates (similar to NMP kinase).
8
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Matrix
Intermembrane
+," $," 8+- F:/$,4<" >< 4A./? $," H.&=5< <A4=="<$ A.="#%=4& A.$.&<
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R #./7.&A41./< .7
L = Loose State: Binds ADP +
Pi tightly
T = Tight State: Binds ATP
more tightly than ADP + Pi; this
favors ATP formation
O = Open State: Binds ADP or
ATP loosely, permitting their
dissociation
"
subunits - Can bind ATP,but do not participate
chemically in catalysis.
subunit - Rotation
interconverts the T, L, and O
states of the 3 #
subunits.
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"
#
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subunits, catalyzing the
formation and release of
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Entry of ADP and Exit of ATP Is Tightly Controlled
by the ATP-ADP Translocase
•
ATP and ADP are membrane impermeable - ATP generated in the mitochondrialmatrix cannot diffuse into the cytoplasm.
•
The ATP-ADP translocase catalyzes the exchange of ADP from the cytoplasm intothe mitochondrial matrix and also transfers a molecule of ATP from the
mitochondrion into the cytoplasm.
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• The synthesis of ATP from ADP and Pi regulates the flow of electrons throughthe electron transport chain.
• The availability of NAD+ and FAD regulate the rate of the TCA cycle. 22
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Uncoupled e – transport:Energy is released, but no
ATP is generated
DNP is in some Herbicides
and fungicides
Inhibition of oxidative phosphorylation
DCCD
Thermogenin
An uncoupler
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Other Transporters of
Mitochondria
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Summary
• Oxidative Phosphorylation occurs in mitochondria
• Ox-Phos depends on electron transfer
•
The respiratory chain consists of four complexes:three are proton pumps
• A proton gradient powers the synthesis of ATP
• Shuttles allow movement across membranes
• Ox-Phos is regulated by the need for ATP