Structure of cilia/flagella.
(A) Cilia/flagella have a cylindrical
architecture composed of axoneme, plasma membrane, and basal body.
(B) Axoneme has a microtubule-based structure formed of nine microtubule
doublets at the periphery and two microtubule singlets at the center,
which are associated with protein complexes such as radial spokes,
inner dynein arm, outer dynein arm, and nexin linkers. There is a
mechanical feedback from the bending on the regulation of dynein activity,
which switches the activity of dyneins on the opposite side of the
central pair microtubules.15 (C) In the
presence of nexin proteins which cross-link the neighboring doublets,
microtubules are not free to slide and active forces generated by
dynein molecular motors bend the cilium/flagellum. Nexin linkers play
an important role to convert the action of dynein motors in microtubule
bending,16,17 and radial spokes ensure that dyneins work
together in a highly coordinated manner to generate a regular wave
pattern.18