Search

The adult ascidian neural complex forms from a thin tube called the neurohypophyseal duct and from the primordium of the cerebral ganglion from the sensory vesicle in metamorphosing larvae. Neurohypophyseal duct cells, located in the anterior left side of the sensory vesicle of swimming larvae, are derived from the anterior embryonic neural plate, which expresses common transcription factors in vertebrates and urochordates. The cerebral ganglion primordium is probably derived from the posterior sensory vesicle during metamorphosis. After metamorphosis begins, the duct elongates anteriorly and fuses with the stomodeal ectoderm, where the dorsal tubercle, a large ciliated structure that opens into the upper part of the pharynx, later develops. The rudiment of the cerebral ganglion and the duct elongate posteriorly. The duct also differentiates into the neural gland. The dorsal wall of the neural gland in adult ascidians has a thick epithelium (placode), the central part of which forms the dorsal strand by repeated invaginations along the visceral nerve. Both gonadotropin-releasing hormone (GnRH) neurons and prolactin-like (non-GnRH) neurons are generated in the dorsal strand and migrate to the cerebral ganglion along the visceral nerve throughout adulthood. Thus, the epithelium derived from the neurohypophyseal duct possesses neurogenic potential to generate neural stem cells of the central (cerebral ganglion) and peripheral (dorsal strand) nervous systems. The generation of prolactin-like neurons and their migration into the brain with GnRH neurons suggest that the ascidian dorsal strand is homologous to the craniate olfactory placode, and provide unequivocal support for the existence of the clade Olfactores.

Electron-microscopic studies of the neural complex (neural gland, dorsal strand, and cerebral ganglion) of an ascidian, Halocynthia roretzi, were performed, paying particular attention to the secretory systems. We found that cells scattered along the dorsal strand and neural cells in the cerebral ganglion contained electron-dense secretory granules of variable size. Immunoelectron-microscopic studies with an antiserum to bullfrog prolactin revealed that the secretory granules (100-250 nm in diameter) of some granulated cells contained a prolactin-like substance. Cells belonging to the neural gland and dorsal strand neither contained electron-dense granules nor showed immunoreactivity. The possibility that cells in the cerebral ganglion and those along the dorsal strand are phylogenetic progenitors of vertebrate adenohypophyseal cells is discussed.