Evolution of neuronal patterning in free-living rhabditid nematodes I: Sex-specific serotonin-containing neurons.

As a first step toward understanding the evolution of neuronal patterning and function in a group of simple animals, we have examined serotonin-containing neurons in 17 species of free-living rhabditid nematodes and compared them with identified neurons of Caenorhabditis elegans. We found many serotonin-immunoreactive (serotonin-IR) neurons that are likely homologs of those in C. elegans; this paper focuses on sex-specific neurons such as the egg laying hermaphrodite-specific neurons (HSNs), VCs, and male CAs, CPs, and ray sensory neurons known to function in mating. These cells vary in number and position in the species examined but are consistent with a current molecularly based phylogeny. Two groups (Oscheius and Pristionchus) appear independently to have lost a serotonin-IR HSN. Oscheius furthermore has no serotonin-IR innervation of the vulval region, in contrast to every other species we examined. We also saw variation in the location of somas of putative HSN, consistent with evolutionary changes in HSN migration. In C. elegans, the HSN soma migrates during embryogenesis from the tail to the central body, where it innervates its major postsynaptic targets, the vulval muscles. For other species, we observed putative HSN homologs along the anterior-posterior axis from the head to the tail, but typically HSNs were located near the vulva, which also varies in anterior-posterior position among the species we examined. The varying positions of the HSN somas in other species are reminiscent of phenotypes seen in various C. elegans mutants with altered HSN migration, suggesting possible mechanisms for the evolutionary differences we observed.