Annelid Brain and Nerve Cord in Siboglinid Riftia pachyptila

Abstract

Vestimentifera is a peculiar group of marine gutless siboglinids which has uncertain position in annelid tree. The detailed study of the fragmentary explored central nervous system of vestimentiferans and other siboglinids is requested to trace the evolution of the siboglinid group. Among all siboglinids the vestimentiferans preserve the gut rudiment what makes them a key group to homologize main cerebral structures with the ones of typical annelids, such as supra- and subesophageal commissures, cirsumesophageal connectives etc. Histologically we revealed main annelid brain structures in the compact large brain of Riftia pachyptila: circumesophageal connectives (longitudinal nerve tracts) and commissures (dorsal, supra- and subenteral commissures). Innervation of tentacles makes them homologous to peristomial palps of the rest annelids. The single nerve cord is represented by paired intraepidermal longitudinal strands associated with the ventral ciliary field in vestimentum and bearing giant axons originating from at least four pairs of perikarya. The absence of regularly positioned ganglia and lateral nerves in the nerve cord in vestimentum and trunk and presence of them in the opisthosome segments. Among siboglinids, the vestimentiferans distinguished by a large and significatly differentiated brain which is reflection of the high development of the palp apparatus. Osedax, frenulates and Sclerolinum have less developped brain. Frenulates and Sclerolinum have good ganglionization in the opisthosome, which probably indicates its high mobility. Comparative neuroanatomical analysis of the siboglinids and annelid sister clades allows us to hypothesize that the last common ancestor of siboglinids might had brain with a dorsal commissure giving rise neurite bundles to palps and paired ventral nerve cord.