Unusual axonemes of hexapod spermatozoa

Hexapod spermatozoa exhibit a great variation in their axoneme structure. The 9+2 pattern organization is present in a few basal taxa and in some derived groups. In most hexapods, a crown of nine accessory microtubules surrounds the 9+2 array, giving rise to the so-called 9+9+2 pattern. This general organization, however, displays a number of modifications in several taxa. In this review, the main variations concerning the number and localization of the accessory tubules, microtubular doublets, central microtubules, dynein arms, and axonemal length are summarized. We discuss the phylogenetic significance of all this structural information as well as the current hypotheses relating the sperm size and sperm polymorphism with reproductive success of some hexapod species. Also described are the biochemical data and the motility patterns which are currently known on some peculiar aberrant axonemes, in light of the contribution these models may give to the comprehension of the general functioning of the conventional 9+2 axoneme. Finally, we summarize methodological developments for the study of axoneme ultrastructure and the new opportunities for the molecular analysis of hexapod axonemes.