Structure
The complex is a large integral membrane protein composed of several metal prosthetic sites and 13 protein subunits in mammals. In mammals, ten subunits are nuclear in origin, and three are synthesized in the mitochondria. The complex contains two hemes, a cytochrome a and cytochrome a3, and two copper centers, the CuA and CuB centers. In fact, the cytochrome a3 and CuB form a binuclear center that is the site of oxygen reduction. Cytochrome c which is reduced by the preceding component of the respiratory chain, (cytochrome bc1 complex, complex III) and docks near the CuA binuclear center, passing an electron to it and being oxidized back to cytochrome c containing Fe3+. The reduced CuA binuclear center now passes an electron on to cytochrome a, which in turn passes an electron on to the cytochrome a3- CuB binuclear center. The two metal ions in this binuclear center are 4.5 Å apart and coordinate a hydroxide ion in the fully oxidized state.
Crystallographic studies of cytochrome c oxidase show an unusual post-translational modification, linking C6 of Tyr(244) and the ε-N of His(240) (bovine enzyme numbering). It plays a vital role in enabling the cytochrome a3- CuB binuclear center to accept four electrons in reducing molecular oxygen to water. The mechanism of reduction was formerly thought to involve a peroxide intermediate, which was believed to lead to superoxide production. However, the currently accepted mechanism involves a rapid four-electron reduction involving immediate oxygen-oxygen bond cleavage, avoiding any intermediate likely to form superoxide.
Read more about this topic: Cytochrome C Oxidase
Famous quotes containing the word structure:
“The structure was designed by an old sea captain who believed that the world would end in a flood. He built a home in the traditional shape of the Ark, inverted, with the roof forming the hull of the proposed vessel. The builder expected that the deluge would cause the house to topple and then reverse itself, floating away on its roof until it should land on some new Ararat.”
—For the State of New Jersey, U.S. public relief program (1935-1943)
“The verbal poetical texture of Shakespeare is the greatest the world has known, and is immensely superior to the structure of his plays as plays. With Shakespeare it is the metaphor that is the thing, not the play.”
—Vladimir Nabokov (1899–1977)
“There is no such thing as a language, not if a language is anything like what many philosophers and linguists have supposed. There is therefore no such thing to be learned, mastered, or born with. We must give up the idea of a clearly defined shared structure which language-users acquire and then apply to cases.”
—Donald Davidson (b. 1917)