Electron Transport

Introduction:
The majority of the energy conserved during catabolism reactions occurs near the end of the
metabolic series of reactions in the electron transport chain. The electron transport or
respiratory chain gets its name from the fact electrons are transported to meet up with oxygen
from respiration at the end of the chain. The overall electron chain transport reaction is:
2 H
+
+ 2 e
+
+ 1/2 O
2
---> H
2
O + energy
Notice that 2 hydrogen ions, 2 electrons, and an oxygen molecule react to form as a product
water with energy released in an exothermic reaction. This relatively straight forward reaction
actually requires eight or more steps. The energy released is coupled with the formation of three
ATP molecules per every use of the electron transport chain.
Pre-Initiation of Electron Transport Chain:
The electron transport chain is initiated by the reaction of an organic metabolite (intermediate in
metabolic reactions) with the coenzyme NAD
+
(nicotinamide adenine dinucleotide). This is an
oxidation reaction where 2 hydrogen atoms (or 2 hydrogen ions and 2 electrons) are removed
from the organic metabolite. (The organic metabolites are usually from the citric acid cycle and
the oxidation of fatty acids--details in following pages.) The reaction can be represented simply
where M = any metabolite.

MH
2
+ NAD
+
-----> NADH + H
+
+ M: + energy
One hydrogen is removed with 2 electrons as a hydride ion (H
-
) while the other is removed as the
positive ion (H
+
). Usually the metabolite is some type of alcohol which is oxidized to a ketone.
NAD
+
is a coenzyme containing the B-vitamin, nicotinamide, shown on a previous page.
The purpose of the other seven steps in the electron transport chain is threefold:
1) to pass along 2H
+
ions and 2e
-
to eventually react with oxygen;
2) to conserve energy by forming three ATP's; and
3) to regenerate the coenzymes back to their original form as oxidizing agents.



Initiation of Electron Transport Chain:
Once the NADH has been made from a metabolite in the citric acid cycle inside of the
mitochondria, it interacts with the first complex 1 enzyme, known as NADH
reductase. This complex 1 contains a coenzyme flavin mononucleotide (FMN) which
is similar to FAD.
The sequence of events is that the NADH, plus another hydrogen ion enter the
enzyme complex and pass along the 2 hydrogen ions, ultimately to an interspace in
the mitochondria. These hydrogen ions, acting as a pump, are utilized by ATP
synthetase to produce an ATP for every two hydrogen ions produced. Three
complexes (1, 3, 4) act in this manner to produce 2 hydrogen ions each, and thus will
produce 3 ATP for every use of the complete electron transport chain.
In addition, NADH passes along 2 electrons to first FMN, then to an iron-sulfur
protein (FeS), and finally to coenzyme Q. The net effect of these reactions are to
regenerate coenzyme NAD
+
. This regeneration of reactants occurs in many of the
reactions so that a cycling effect occurs. The NAD
+
is ready to react further with
metabolites in the citric acid cycle.
Coenzyme Q, which also picks up an additional 2 hydrogen ions to make CoQH
2
, is
soluble in the lipid membrane and can move through the membrane to come into
contact with enzyme complex 3.
In summary, the very first enzyme complex in the electron transport chain is coupled
with the formation of ATP. The coupled reaction may be written as:

a) MH
2
+ NAD
+
---> NADH + H
+
+ M + energy
b) ADP + P + energy ---> ATP + H
2
O







Electron Transport - Enzyme Complex 3:
Coenzyme QH
2
carrying an extra 2 electrons and 2 hydrogen ions now starts a
cascade of events through enzyme complex 3, also known as cytochrome reductase
bc.
Cytochromes are very similar to the structure of myoglobin or hemoglobin. The
significant feature is the heme structure containing the iron ions, initially in the +3
state and changed to the +2 state by the addition of an electron. The
CoQH
2
(yellow)passes along the 2 electrons first to cytochrome (blue) b1 heme
(magenta), then b2 heme , then to an iron-sulfur protein (green), then to cytochrome
c1 (red with black heme), and finally to cytochrome c (not shown). Co Q is
represented by the inhibitor antimycin (yellow) in the graphic.
In the meantime the 2 hydrogen ions are channeled to the interspace of the
mitochondria for ultimate conversion into ATP.
Complex 4: Refer to the middle graphic: Cytochrome c is a small molecule which is
also able to move in the lipid membrane layer and diffuses toward cytochrome a
complex 4.
Cytochrome c - Chime in new window
At this time it continues the transport of the electrons, and provides the third and final
time that 2 hydrogen ions are channeled to the interspace of the mitochondria for
ultimate conversion into ATP.
ATP synthetase is also found at numerous locations in the bilayer membrane of the
mitochondria. The function of this ATP enzyme is found in an earlier page. Three
ATP are produced by the pumping action of the re-entry of the hydrogen ions
through the ATP synthetase.
Finally, oxygen has diffused into the cell and the mitochondria for the finally reaction
of metabolism. Oxygen atom reacts with the 2 electrons and 2 hydrogens to produce
a water molecule.