Anastassya S. Maiorova

Reproduction and development of common species of peanut worms (Sipuncula) from the Sea of Japan

This study deals with the reproduction and development of the most common species of peanut worms from the Sea of Japan: Thysanocardia nigra, Themiste pyroides, and Phascolosoma agassizii. Data on the time of reproduction and larval settlement and the distribution of these species in Peter the Great Bay are provided. The peculiarities of gametogenesis, spawning, and embryonic, larval, and postlarval development are described. The reproductive biology of representatives of these species from the western and eastern Pacific is examined in a comparative aspect.

A new species of Aspidiophorus (Gastrotricha, Chaetonotida) from the Russian Far East with a key to marine species of the genus

Abstract The Russian gastrotrich fauna is virtually unknown, particularly the marine fauna. In the ocean, investigations have been restricted to the White Sea, from where only three fully described species have been reported so far. In this study we describe a new species of Aspidiophorus found in a sandy sample collected from off of Vladivostok (Peter the Great Bay, Sea of Japan). Aspidiophorus oculatus n. sp. is the first marine chaetonotid gastrotrich described from Russia and the first representative of the order Chaetonotida reported from the Sea of Japan. Specimens of the new species are characterized by a body up to 147.5 µm in total length; enveloped by 57 alternating columns of 65–70 keeled, elongate scales. Pharyngeo-intestinal junction at U27; the head bears cephalion, hypostomion, pleuria and a pair of eye-spots; the furca is 20 µm long including the 12-µm long adhesive tube; the ventral interciliary field is naked, with the exception of a pair of elliptical, keeled scales occurring near the anus. Morphological differences between the new species and the other 13 marine con-generic taxa are discussed. A key to the known marine species of the world based on easily identifiable traits, visible in both living and formalin-fixed specimens is provided.

Microscopic Anatomy and Ultrastructure of Nephridium in the Sipunculan Thysanocardia nigra Ikeda, 1904 from the Sea of Japan

The microscopic anatomy and ultrastructure of nephridium have been studied in the sipunculan Thysanocardia nigra Ikeda, 1904 (Sipuncula, Sipunculidea) from the Sea of Japan using histological and electron microscopic techniques (SEM and TEM). This paper describes ultrastructural features of nephridial epithelium, muscle grid, and coelomic epithelium on the surface of the nephridium, the area of the ciliary funnel, and the “tongue.” Several types of cells were distinguished in the excretory tube of the nephridium: (1) a columnar epithelium of the excretory bunches; (2) a cubical or flattened epithelium of flask-shaped infoldings; and (3) granulocytes that migrate from the coelom to the extracellular matrix of the nephridial wall. The system of podocytes and multiciliary cells were described in the nephridial coelothelium. Two types of secretion of nephridial epithelium have been discovered: a merocrine secretion of columnar cells and an apocrine secretion of cells of the flask-shaped infoldings. Using ultrastructural data, two zones of filtration through the wall of excretory tube have been found, namely (1) the tips of flask-shaped infoldings (via the extracellular matrix and microvillary canals between the epithelial cells) and (2) areas between the flask-shaped infoldings (via the contacts of podocytes, extracellular matrix, and the basal labyrinth of the columnar cells). Unlike previously studied representatives of the genus Phascolosoma, no coelomic epithelium is present on the tips of the flask-shaped infoldings in Th. nigra. This data on the anatomy and histology allow us to conclude that the funnel only works like a gonoduct.

Spermatozeugma and sperm ultrastructure of Thysanocardia nigra, Ikeda, 1904 (Sipuncula; Sipunculidea)

Summary This study presents details of the ultrastructure of spermatozeugmata and spermatozoa of the peanut worm, Thysanocardia nigra, from the Sea of Japan. Spermatozoa are tightly packaged into unencapsulated conical bundles, spermatozeugmata, with as many as 300–350 cells comprising a single packet. Within an individual spermatozeugma, cells are aligned side by side such that head of the bundle is comprised of the heads of spermatozoa and the internal cavity of the bundle harbors numerous flagella that together form a tail for the spermatozeugma. The integrity of the packet appears to be maintained by specialized junctions between adjacent cells. The spermatozeugmata break up when discharged into the sea water. Each spermatozoon is characterized by a rounded head, consisting of the acrosome, nucleus, and midpiece. The midpiece contains mitochondria, centrioles, and some residual cytoplasm that contains dark granules. The method of “sperm packaging” and morphology of spermatozeugmata are unique to the Sipuncula. The ultrastructural modifications of spermatozeugmata are discussed as possible adaptations for the reproductive mode and unique excretory system of sipunculans.

Embryonic and larval development of the peanut worm Themiste pyroides (Sipuncula: Sipunculoidea) from the Sea of Japan

Summary Embryonic and larval development are described for the sipunculan Themiste pyroides from the Peter the Great Bay, the Sea of Japan. The eggs are 120 μm in diameter and coated by a 2 μm thick envelope, with a dense concentration of yolk. Egg cleavage is complete, spiral, and unequal. T. pyroides from the Sea of Japan is characterized by an indirect development with two pelagic larval stages—a lecithotrophic trochophore and a lecithotrophic pelagosphera that is short lived. At the blastula stage, the embryos begin to move in the near-bottom layer of water. At this stage, the embryo resembles a dense sphere with a wide ciliary band extending along its equator. The actively swimming trochophore after 3 days of development metamorphoses to a lecithotrophic pelagic larva, the pelagosphera. The pelagosphera of T. pyroides does not feed but relies on stored yolk, whose large granules are visible in the gut. At day 14, larvae finally settle on the bottom and creep peristaltically. Results of our study of the development of T. pyroides from the Sea of Japan substantially differ from the earlier published data (Rice, 1967) on development of the same species in British Columbia. The species from the East Pacific develops directly within the jelly coat, the embryo hatching from the jelly as a small, crawling worm (see Rice, 1967). These differences correlate with the differences in size of the eggs, time of breeding season and water temperature in the localities of this amphi-pacific species in the East and West Pacific.

Ultrastructural observations on spermiogenesis in the peanut worm, Themiste pyroides (Chamberlin, 1920) (Sipuncula; Sipunculidea)

Summary The process of spermiogenesis and the ultrastructure of the spermatozoa in the peanut worm, Themiste pyroides, from the Sea of Japan were observed with electron microscopy (SEM and TEM). The testes are composed of groups of spermatogonia and are covered by peritoneal cells. Clusters of spermatocytes are released from the testes into the coelomic fluid. Connected by intercellular bridges, the spermatocytes within a given cluster develop asynchronously. Proacrosomal vesicles and a flagellum appear in spermatocytes. Spermatids in the clusters retain the intercellular connections. During spermiogenesis, the acrosomal vesicle, formed by coalescence of small proacrosomal vesicles in the basal part of the spermatid, migrates to the apical part of the cell to form a conical-shaped acrosome. The basal concavity lying above the nucleus is filled with subacrosomal substance. The midpiece contains four mitochondria, two centrioles, and some residual cytoplasm with dark glycogen-like granules. A peculiar annulus structure develops around the base of the flagellum. The distal centriole has a pericentriolar complex consisting of radially oriented elements. Before the spawning process, the spermatozoa are filtered throughout the ciliary nephrostomal funnel into the excretory sac of paired nephridia where they are stored for a short time. The sperm are released into the sea water via nephridiopores. Spermatozoa remaining in the coelomic fluid after spawning are resorbed by amoebocytes. This species from Vostok Bay is characterized by a prolonged spawning period from June to early October. The reproductive strategy of T. pyroides is discussed in comparison with that of Thysanocardia nigra, the latter having a unique pattern of packaging of the spermatozoa, resulting in the formation of spermatozeugmata, as a reproductive adaptation to the very short spawning period.

Ultrastructure of the Coelomocytes in the Tentacular Coelom of Thysanocardia nigra Ikeda, 1904 (Sipuncula)

Free-floating coelomocytes in the tentacular coelomic cavity of the sipunculan Thysanocardia nigra Ikeda, 1904, were studied using light interference contrast microscopy and scanning and transmission electron microscopy. The following coelomocyte types were distinguished: hemerythrocytes, amoebocytes, and two morphological types of granular cells. No clusters of specialized cells that had been reported to occur in the trunk coelom of Th. nigra were found in the tentacular coelom. The corresponding types of coelomocytes from the tentacular and trunk coelomic cavities were shown to differ in size. These two coeloms are completely separated in sipunculans.

Embryonic and larval development of the peanut worm Phascolosoma agassizii (Keferstein 1867) from the Sea of Japan (Sipuncula: Phascolosomatidea)

All stages of the embryonic and larval development of Phascolosoma agassizii from Peter the Great Bay (Sea of Japan) were studied and illustrated using light and electron microscopy. The eggs of P. agassizii have the form of an ellipsoid (long and short axes about 100 and 70 µm, respectively). Egg cleavage is typical, spiral, and unequal. Gastrulation occurs by epiboly. This species possesses two pelagic larval stages, a lecithotrophic trochophore and a planktotrophic pelagosphera. The transformation of trochophore into pelagosphera occurs 80–90 h after fertilization. After 120–180 h, the larva has developed all systems of organs characteristic of the pelagosphera and is capable of feeding. At day 10, pelagospheras can settle, for some time, on the aquarium bottom and move on a ciliated lip, collecting food with the aid of a buccal organ. In addition, the larvae periodically attach themselves to the aquarium bottom or to the surface film of the water by means of a terminal organ. The trunk of the larva elongates by enlargement of the region behind the dorsal anal opening, which is located almost in the middle of the trunk region in the 15-day old larva. In the laboratory, 1-month old larvae spend the greater part of time in the attached state. Being attached by a glandular terminal organ to the aquarium bottom, they characteristically bend the body, actively feeding on microalgae from the substratum surface. The differences in the development of P. agassizii in the isolated West-Pacific and East-Pacific populations are shown and discussed.

Ultrastructure of tentacles in the sipunculid worm Thysanocardia nigra Ikeda, 1904 (Sipuncula)

The ultrastructure of the tentacles was studied in the sipunculid worm Thysanocardia nigra. Flexible digitate tentacles are arranged into the dorsal and ventral tentacular crowns at the anterior end of the introvert of Th. nigra. The tentacle bears oral, lateral, and aboral rows of cilia; on the oral side, there is a longitudinal groove. Each tentacle contains two oral tentacular canals and an aboral tentacular canal. The oral side of the tentacle is covered by a simple columnar epithelium, which contains large glandular cells that secrete their products onto the apical surface of the epithelium. The lateral and aboral epithelia are composed of cuboidal and flattened cells. The tentacular canals are lined with a flattened coelomic epithelium that consists of podocytes with their processes and multiciliated cells. The tentacular canals are continuous with the radial coelomic canals of the head and constitute the terminal parts of the tentacular coelom, which shows a highly complex morphology. Five tentacular nerves and circular and longitudinal muscle bands lie in the connective tissue of the tentacle wall. Similarities and differences in the tentacle morphology between Th. nigra and other sipunculan species are discussed.

Microscopic Anatomy and Ultrastructure of a Polian Vessel in the Sipunculan Thysanocardia nigra Ikeda, 1904 from the Sea of Japan

The microscopic anatomy and ultrastructure of a Polian vessel have been studied in the sipunculan Thysanocardia nigra Ikeda, 1904 from the Sea of Japan using the methods of histology and electron microscopy. We describe ultrastructural features of the inner and outer coelothelium, which is constructed of podocytes and multiciliary cells. Between the processes of the podocyte cells, we found double diaphragms that are considered characteristic macromolecular filters. We conclude from an analysis of the ultrastructural features of the vessel wall that coelomic fluid may be filtered from the tentacular coelom to the trunk coelom via the wall of the Polian vessel.