Cilia and Eukaryotic Flagella

 Cilia and eukaryotic flagella are long (10 µm to 40 µm), narrow, membrane-bounded structures that contain a highly ordered, stable microtubule array, consisting of nine fused outer doublet microtubules surrounding a central pair of singlet microtubules. These microtubules are firmly anchored at the base of the cilium or flagellum in basal bodies. The microtubules of cilia and flagella are extremely stable, and their movement is not complicated by dynamics. In addition to the tubulin backbone, these microtubules contain a large and diverse array of microtubule-associated proteins) MAPs), most of which have not been characterized, and several forms of dynein, a large multicomponent ATP-transducing mechanochemical motor protein. The various MAPs create and maintain the stable axoneme structure, and the dynein is responsible for creating the movement. The movements of cilia and flagella involve bending of these long organelles, which is caused by the sliding of stable double-microtubule pairs past each other. Sliding is effected by the dynein motors; the base of the motor is attached permanently to the A microtubule of one outer doublet pair, and the head, which contains the motor domain, transiently attaches and detaches to the B subfiber of an adjacent outer doublet pair. The dynein motor “walks” along the B subfiber toward the minus end, creating the movement. The dynein-mediated sliding of outer double-microtubules occurs in a fashion similar to filament sliding in muscle contraction, in which actin filaments and myosin filaments slide past each other through the action of the ATP-transducing myosin motor “heads.”