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Shuttle Involved in
Cytosol-Matrix
Exchange of NADH
and NADPH
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Reduced coenzymes such as NADH and

NADPH do not permeate the inner
membrane of the mitochondrion to any
significant extent.

However, reduced pyridine nucleotides

are known to be produced in a number
of reactions in the cytosol (the reduction
of NAD+ at the glyceraldehyde step in
glycolysis is an important example) and
the re-oxidation of NADH occurs via the
mitochondrion.

The mechanism involves a set of

reactions called a shuttle (Fig. 16-32).
1. Glycerol Phosphate Shuttle:

The glycerol phosphate shuttle (Fig. 16-

32) involves:

(1) Glycerol phosphate dehydrogenase in

the cytosol,

(2) Glycerol phosphate dehydrogenase on

the outer surface of the inner
mitochondrial membrane, and

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(3) The reduction of Q in the electron

transport chain.

Dihydroxyacetone phosphate, NADH,

and H+ react in the cytosol to form
glycerol phosphate, which diffuses
through the outer mitochondrial
membrane to the outer surface of the
inner membrane. There the glycerol
phosphate reacts with the membrane
dehydrogenase to form
dihydroxyacetone phosphate, which
returns to the cytosol. The membrane-
bound dehydrogenase employs a
flavoprotein as a coenzyme, and the FP
becomes reduced during the reaction.

Subsequently, the electrons from the

reduced FP are passed directly into the
electron transport system at the Q step.
Because the NAD+ →FP step of electron
transport is skipped, only two ATP are
generated for each pair of electrons that
enters in this fashion from the cytosol.

2. Malate-Aspartate Shuttle:

Another shuttle, the malate-aspartate

shuttle, can also transport the hydrogens
accepted during the reduction of NAD +
in the cytosol across the inner membrane
(Fig. 16-32). H+ transported in this
manner into the matrix, enters the
electron transport chain via NADH
dehydrogenase, and as a result three ATP
are generated for each pair of electrons.

3. Total ATP Production from the

Catabolism of Glucose:

The complete oxidation of a molecule of

glucose by glycolysis, the TCA cycle, and
electron transport is accompanied by a
gross production of 38 or 40 ATP or a net
production of 36 or 38 ATP. The ATPs
result from substrate-level
phosphorylations and electron transport
coupled to oxidative phosphorylation. An
accounting of ATP production is
described in Figure 16- 33.

The different totals for ATP production

(i.e., gross production of either 38 or 40)
depend on which shuttle is used to
transport H+ from NADH into the mito-
chondrion. H + from the glycerol
phosphate shuttle is accepted by FP and
as a result one coupling step is bypassed.
Hydrogen from the malate-aspartate
shuttle enters the electron transport
chain earlier, so that all three coupling
sites are utilized.

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in Aerobic Respiration