
Figure 2. Simplified schematic representation of the mechanism by which axonemal
dynein arms generate the flagellar beat. The parallel lines in the region of
the flagellum represent adjacent outer microtubule doublets of the axoneme
(OMDA). To simplify the figure, only 2 outer microtubule doublets are
represented (rather than the actual 9). The central pair of microtubule
doublets is not shown, nor are the any of the accessory structures. Dynein
arms between the pair of OMDAs are numbered. Each symbol represents a pair of
inner and outer dynein arms originating from the microtubule on the right and
extending toward the microtubule on the left. (A) None of the dynein
arms are active; as such, the adjacent microtubule doublets are straight and
there is no flagellar bend. (B) The first dynein arm has engaged the
adjacent microtubule doublet. (C) The first dynein arm has generated a
downward stroke, resulting in the adjacent microtubules sliding past one
another (large arrows). Because both microtubule doublets are anchored to the
sperm head, this sliding force is translated into a bend in the
axonemethe start of a flagellar beat. Also, the second dynein arm has
engaged the adjacent microtubule doublet. (D) The first dynein arm has
released the adjacent microtubule doublet, whereas the second dynein arm has
generated a downward stroke. This results in the propagation of the flagellar
bend down the length of the microtubules. The third dynein arm has engaged the
adjacent microtubule. (E) The flagellar beat is being propagated by the
downward stroke of the third dynein arm, whereas the second dynein arm has
released the adjacent microtubule. The first dynein arm has returned to its
original position in preparation for another stroke. This sequence will be
repeated by the fourth dynein arm, and so on along the length of the
microtubules. One must imagine a similar sequence occurring for all 9 dynein
arm pairs and their associated 9 outer microtubule doublets. Additionally, one
must imagine this sequence occurring in an asynchronous but coordinated
fashion around the circumference and along the entire length of the flagellum
(ie, in 3 dimensions). It is this coordinated use of dynein arms and the
associated bending of the outer microtubule doublets that result in a normal
flagellar beat. The ODFs and the FS add structural support. In Chlamydomonas,
it has been shown that the central pair of microtubules, together with the
radial spokes emanating from the 9 outer microtubule doublets, coordinate and
regulate the actions of the dynein arms and so control the size and shape of
the axonemal bend.