Development of the enteropneust Ptychodera flava: ciliary bands and nervous system

Ripe specimens of Ptychodera flava were collected at Paiko Peninsula, Oahu, Hawaii, USA, and the development from egg to tornaria larva was followed in the laboratory. To complete the series, large tornaria larvae were collected from the plankton off the nearby Ala Moana Beach, and followed through metamorphosis to a juvenile stage with four pairs of gill slits. Ciliary band development was examined by scanning electron microscopy, and the development of the serotonergic nervous system was followed by means of immunostaining. The development of the apical tuft and neotroch (circumoral/perioral ciliary band) and their subsequent degeneration accorded fully with previous descriptions. A perianal ciliary ring of separate cilia develops just after hatching. This later develops a midventral extension, the neurotroch, extending to the neotroch posterior to the mouth. The cilia of this ring apparently beat diaplectically, with the effective stroke in the clockwise direction when seen from behind. An additional ring of cilia develops several days later anterior to the perianal ring. This opisthotroch (called telotroch by previous authors) consists at first of separate cilia, but later they became organized as large compound cilia. The apical tuft disappears after about a week, the neotroch degenerates at the transition to the Agassiz stage, and the opisthotroch degenerates just after metamorphosis. The serotonergic nervous system of the fully grown tornaria consists of an apical ganglion with many perikarya, a paired lateral group of perikarya on the postoral ciliary band, and scattered perikarya along the opisthotroch. Serotonergic processes are found along the ciliary bands except for the ventral and perianal ciliary bands and are scattered along the epidermis. At the Spengel stage and at metamorphosis (Agassiz stage), the processes along the ciliary bands are concentrated in the three ciliated food grooves so as to form three separate nerves, and are retained on the proboscis at least until 2-3 gill slit stage. No serotonergic processes were found to extend from the proboscis to the collar region, and no serotonergic neurons were observed in the collar cord or in the ventral nerve cord. Our results therefore do not provide any clues as to the origin of the chordate neural tube relative to the dorsal-ventral orientation of the enteropneusts.