Ultrastructure of spermiogenesis in the gastropodHeliacus variegatus (Architectonicidae), with description of a banded periaxonemal helix

Abstract

Spermiogenesis and spermatozoa of the marine gastropodHeliacus variegatus Gmelin were examined using transmission electron microscopy (TEM). Mature spermatozoa are composed of an acrosomal vesicle, helical nucleus, elongate midpiece, annulus, glycogen piece and short end piece. The midpiece consists of a 9+2 axoneme, nine coarse fibres, and a banded helix, all enclosed by a continuous mitochondrial sheath (with multiple, helically coiled grooves). Anterior extensions of the mitochondrial sheath and banded helix form a double sheath around the basal half of the nucleus—an arrangement possibly unique in the Mollusca. During spermiogenesis, dense plaques delineating the anterior and posterior poles of the spermatid nucleus become attachment sites for the acrosomal vesicle and the axial complex (respectively). As the nucleus condenses and elongates, midpiece formation involves fusion of numerous, oblong mitochondria along the length of the axoneme. The coarse fibres and banded helix of the midpiece probably are derived through centriolar activity. Results of the study support inclusion of the Architectonicidae within the Heterobranchia, but in view of midpiece specializations, do not clarify the precise relationship of the family within this subclass.