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The mitochondrial shuttles are systems used to transport reducing agents across the inner mitochondrial
membrane. NADH cannot cross the membrane, but it can reduce another molecule that can cross the
membrane, so that its electrons can reach the electron transport chain.

The two main systems in humans are:

Name In To ETC Out

To mitochondrion To cytosol

Glycerol phosphate shuttle Glycerol 3-phosphate QH2 (~1.5 ATP) Dihydroxyacetone phosphate

Malate-aspartate shuttle Malate NADH (~3 ATP) Oxaloacetate aspartate

Malate Aspartate Shuttle

 Mitochondrial membrane is impermeable to
 The NADH equivalents generated in glycolysis are
therefore to be transported from cytoplasm to
mitochondria for oxidation. This is achieved by
malate-aspartate shuttle or malate shuttle, which
operates mainly in liver, kidney and heart.
 The cycle is operated with the help of enzymes
malate dehydrogenase (MDH) and aspartate
amino transferase.
 From one molecule of NADH in the mitochondria 2½
ATP molecules are generated.

Malate Aspartate Shuttle

Glycerol-3-phosphate Shuttle
 In skeletal muscle and brain, the reducing
equivalents from cytoplasmic NADH are transported
to mitochondria asFADH2 through glycerol-3-
phosphate shuttle.
 Hence only 1½ ATPs are generated when this
system is operating.
 In this shuttle, the enzyme called
cytoplasmic glycerol-3-phosphate dehydrogenase 1
(GPDH-C) converts dihydroxyacetone phosphate (2)
to glycerol 3-phosphate (1) by oxidizing one molecule
of NADH to NAD+ .
Glycerol-3-phosphate Shuttle