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Berg JM, Tymoczko JL, Stryer L. Biochemistry. 5th edition. New York: W H Freeman; 2002.

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Biochemistry. 5th edition.

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Chapter 18Oxidative Phosphorylation

Mitochondria, Stained Green, Form a Network Inside a Fibroblast Cell (Left).


Mitochondria, Stained Green, Form a Network Inside a Fibroblast Cell (Left). Mitochondria oxidize carbon fuels to form cellular energy. This transformation requires electron transfer through several large protein complexes (above), some of which pump (more...)

The NADH and FADH2 formed in glycolysis, fatty acid oxidation, and the citric acid cycle are energy-rich molecules because each contains a pair of electrons having a high transfer potential. When these electrons are used to reduce molecular oxygen to water, a large amount of free energy is liberated, which can be used to generate ATP. Oxidative phosphorylation is the process in which ATP is formed as a result of the transfer of electrons from NADH or FADH 2 to O 2 by a series of electron carriers. This process, which takes place in mitochondria, is the major source of ATP in aerobic organisms (Figure 18.1). For example, oxidative phosphorylation generates 26 of the 30 molecules of ATP that are formed when glucose is completely oxidized to CO2 and H2O.

Figure 18.1. Electron Micrograph of a Mitochondrion.

Figure 18.1

Electron Micrograph of a Mitochondrion. [Courtesy of Dr. George Palade.]

Oxidative phosphorylation is conceptually simple and mechanistically complex. Indeed, the unraveling of the mechanism of oxidative phosphorylation has been one of the most challenging problems of biochemistry. The flow of electrons from NADH or FADH2 to O2 through protein complexes located in the mitochondrial inner membrane leads to the pumping of protons out of the mitochondrial matrix. The resulting uneven distribution of protons generates a pH gradient and a transmembrane electrical potential that creates a proton-motive force. ATP is synthesized when protons flow back to the mitochondrial matrix through an enzyme complex. Thus, the oxidation of fuels and the phosphorylation of ADP are coupled by a proton gradient across the inner mitochondrial membrane (Figure 18.2).

Figure 18.2. Essence of Oxidative Phosphorylation.

Figure 18.2

Essence of Oxidative Phosphorylation. Oxidation and ATP synthesis are coupled by transmembrane proton fluxes.

Oxidative phosphorylation is the culmination of a series of energy transformations that are called cellular respiration or simply respiration in their entirety. First, carbon fuels are oxidized in the citric acid cycle to yield electrons with high transfer potential. Then, this electron-motive force is converted into a proton-motive force and, finally, the proton-motive force is converted into phosphoryl transfer potential. The conversion of electron-motive force into proton-motive force is carried out by three electron-driven proton pumps—NADH-Q oxidoreductase, Q-cytochrome c oxidoreductase, and cytochrome c oxidase. These large transmembrane complexes contain multiple oxidation-reduction centers, including quinones, flavins, iron-sulfur clusters, hemes, and copper ions. The final phase of oxidative phosphorylation is carried out by ATP synthase, an ATP-synthesizing assembly that is driven by the flow of protons back into the mitochondrial matrix. Components of this remarkable enzyme rotate as part of its catalytic mechanism. Oxidative phosphorylation vividly shows that proton gradients are an interconvertible currency of free energy in biological systems.


An ATP-generating process in which an inorganic compound (such as molecular oxygen) serves as the ultimate electron acceptor. The electron donor can be either an organic compound or an inorganic one.

  • 18.1. Oxidative Phosphorylation in Eukaryotes Takes Place in Mitochondria
  • 18.2. Oxidative Phosphorylation Depends on Electron Transfer
  • 18.3. The Respiratory Chain Consists of Four Complexes: Three Proton Pumps and a Physical Link to the Citric Acid Cycle
  • 18.4. A Proton Gradient Powers the Synthesis of ATP
  • 18.5. Many Shuttles Allow Movement Across the Mitochondrial Membranes
  • 18.6. The Regulation of Cellular Respiration Is Governed Primarily by the Need for ATP
  • Summary
  • Problems
  • Selected Readings

By agreement with the publisher, this book is accessible by the search feature, but cannot be browsed.

Copyright © 2002, W. H. Freeman and Company.
Bookshelf ID: NBK21208


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