Male reproductive system of the deep-sea acorn worm Quatuoralisia malakhovi (Hemichordata, Enteropneusta, Torquaratoridae) from the Bering Sea

The deep-sea acorn worm Quatuoralisia malakhovi belongs to the phylum Hemichordata, class Enteropneusta, family Torquaratoridae, which was described in 2005. Owing to their epibenthic lifestyle and deep-sea habitat features, torquaratorids differ anatomically from shallow-water acorn worms; however, their morphology and fine structure are poorly studied. We have the opportunity to make three complete detailed series of histological sections of Q. malakhovi and to study the microscopic anatomy, histology and fine structure of the reproductive system of this acorn worm using scanning and transmission electron microscopy. The sexes of Q. malakhovi are separate and indistinguishable externally. The lobed testes occupy the dorsal side of the genital wings and distinctly bulge into the peribranchial cavity by their mature lobes. The central part of the testis is always submerged into the genital wing and opens via a single gonad pore. The monociliary muscle cells stretch along the external wall of the testis and surround the gonad pore, probably taking part in the contraction of the testis lobes for spawning. The germinative epithelium of the testis contains spermatogenic cells at different stages of development and interstitial cells. Yolk cells are not found. Interstitial cells embrace the spermatogonia and spermatogenic columns, providing horizontal compartmentalization of the germinative epithelium, and contain numerous phagosomes with remnants of degenerating spermatogenic cells. The testis wall contains haemal lacunae, which are usually located on the side opposite the gonad pore. We describe the fine structure of spermatogonia, spermatocytes clustered in spermatogenic columns, spermatids, and spermatozoa. Spermatozoa are of the ectaquasperm type and consist of an acorn-shaped head and a flagellum 18–25 µm long. The sperm head includes a beak-shaped acrosomal part, a spherical nucleus and a midpiece containing a ring of 5 or rarely 6 mitochondria. The male reproductive system and sperm structure of Q. malakhovi, a representative of the family Torquaratoridae, have a number of differences from shallow-water acorn worms; however, the spermatogenesis and sperm structure of Q. malakhovi generally follow the pattern of the other three enteropneust families, and the phylogenetic significance of these deviations should be the subject of further research.