Placode formation and generation of gonadotropin-releasing hormone (GnRH) neurons in ascidians.

Neurogenic placodes, a chordate innovation, generate several neuronal populations, including gonadotropin-releasing hormone (GnRH) neurons which are crucial for vertebrate and solitary ascidian urochordate reproduction. The dorsal strand placode of ascidians Is derived from the anterior ridge of the embryonic neural plate and thus shares a common developmental origin and expression of various transcription factors with vertebrate placodes. Despite their importance for understanding vertebrate origins, the evolutionary and developmental origins of the neurogenic placode remain obscure. Here I demonstrate the formation of an elaborate neurogenic placode, which forms the dorsal strand, on part of the neural gland epithelium in a solitary ascidian urochordate, Halocynthia roretzi. Two modes of GnRH neurogenesis in the dorsal strand (a peripheral organ) and the migration of GnRH neurons into the brain along the visceral nerve are also described. Ontogenetically, GnRH neurons are first detected in the dorsal strand and cerebral ganglion of very young Juveniles at almost the same time, demonstrating that ascidians possess morphological and developmental features in common with vertebrates. These results further indicate that the onset of peripheral GnRH neurogenesis and the ability of neurons to migrate into the brain predate the divergence of ascidians and vertebrates. Thus, based on the generation of GnRH neurons, the dorsal strand in ascidians may be homologous to the vertebrate olfactory placode. These organs are derived from the anterior region of the embryonic neural plate, which expresses several transcription factors that invertebrate chordates and vertebrates have in common. These results provide unequivocal support for the clade Olfactories (tunicates + vertebrates).