A novel motility pattern in quail spermatozoa with implications for the mechanism of flagellar beating

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Abstract
Background information. The spermatozoon of the quail (Coturnix coturnix L., var japonica) has a ‘9+2’ flagellum that is unusually long. When it moves in a viscous medium, near to the coverslip, it develops a meander waveform. Because of the high viscosity, the meander bends are static in relation to the field of view; bend propagation is therefore manifest as the forward movement of the flagellum through the meander shape. At the same time, the origin of the oscillation typically shifts proximally in a stepwise fashion. These movements have been analysed in the hope of contributing to the resolution of problems in flagellar mechanics.Results. (1) Meander waves originate from spontaneous sigmoid bend complexes. (2) On a given flagellum, fully developed meander bends are uniform in their large angle, curvature and propagation speed; interbends can vary in length and shape. (3) No intra‐axonemal sliding is transmitted through formed bends; sliding related to new bends is accommodated proximally. (4) Sliding reversal is initiated at a threshold shear angle of approx. 1 rad. (5) The arc wavespeed is the product of the arc wavelength and the beat frequency. (6) Physical obstruction to bend development causes a pause in the oscillation. (7) New bend initiation can thus be dissociated from bend propagation on the distal flagellum. (8) The steps in the forward advance of the oscillation site occur during the early phase of bend growth.Conclusions. (1) The main conclusion is that, in meander waves, the mechanical basis of the oscillation appears to be that the propulsive thrust arising from bend propagation acts as a bending stress to trigger sliding reversal, thus perpetuating the rhythmic beating. (2) Oscillations can originate at any position, provided the position is distal to a location where doublet sliding is restrained. (3) Meander waves are an example of new bend development without ‘paradoxical’ classes of sliding.