The sperm connecting piece is a complex structure that, from a mechanical perspective, appears to play a role in stabilizing the proximal part of the sperm tail. We report the three-dimensional structure of the intact bovine sperm connecting piece, revealing an intricate, asymmetrical architecture with the segmented columns held together by filamentous linkages. The columns fuse, at the proximal end, with each other into structures that form the centriolar vault, and at the distal end, with the outer dense fibers (ODFs). The grouping of the fibers into these structures is consistent with bending only in the plane of the head. Structures reminiscent of the proximal centriole were observed in the vault, while the association of a novel bar structure with ODFs 3 and 8 organizes the distal centriolar vault. It has been proposed that the elastic compliance of the connecting piece provides the underlying mechanism behind initiation of the sperm beat cycle and bend propagation. According to the basal sliding theory of sperm movement, distortion of the connecting piece may store energy that initiates a new beat. The intersegment linkers could serve as mechanosensitive elements that regulate alternation of the sperm tail's bending direction in the beat cycle in addition to providing structural stabilization for the connecting piece segmented structures. On the other hand, our video recordings of the bull sperm movement show little bending of the head with respect to the tail, so it appears that there may be normally little strain within the connecting piece.

Electron cryotomography of the structure of the sperm connecting piece yields novel insights into the motility of spermatozoa.