In Complex I (NADH ubiquinone oxidoreductase, Type I NADH dehydrogenase, or mitochondrial complex I; ), two electrons are removed from NADH and transferred to a lipid-soluble carrier, ubiquinone (Q). The reduced product, ubiquinol (QH), freely diffuses within the membrane, and Complex I translocates four protons (H) across the membrane, thus producing a proton gradient. Complex I is one of the main sites at which premature electron leakage to oxygen occurs, thus being one of the main sites of production of superoxide.
The pathway of electrons is as follows:
NADH is oxidized to NAD, by reducing flavin mononucleotide to FMNH in one two-electron step. FMNH is then oxidized in two one-electron steps, through a semiquinone intermediate. Each electron thus transfers from the FMNH to an Fe–S cluster, from the Fe-S cluster to ubiquinone (Q). Transfer of the first electron results in the free-radical (semiquinone) form of Q, and transfer of the second electron reduces the semiquinone form to the ubiquinol form, QH. During this process, four protons are translocated from the mitochondrial matrix to the intermembrane space. As the electrons move through the complex an electron current is produced along the 180 Angstrom width of the complex within the membrane. This current powers the active transport of four protons to the intermembrane space per two electrons from NADH.