All photosynthetic membranes contain a cytochrome bc 1 or b 6 f complex that catalyzes the oxidation of quinols and the reduction of a high-potential electron carrier, such as cytochrome c 2 or plastocyanin. The cytochrome complex also functions in the translocation of protons across the membrane and as a consequence, establishes the proton motive force that is used for the synthesis of ATP. The structure and function of the cytochrome complexes are first reviewed in this chapter. Amino acid sequence information for almost all of the protein subunits of these complexes is now available, and these allow for a detailed consideration of functional domains in the protein subunits and for a further discussion of the evolution of the cytochrome complex in photosynthetic organisms.