Steps towards a centralized nervous system in basal bilaterians: insights from neurogenesis of the acoel Symsagittifera roscoffensis.

Due to its proposed basal position in the bilaterian Tree of Life, Acoela may hold the key to our understanding of the evolution of a number of bodyplan features including the central nervous system. In order to contribute novel data to this discussion we investigated the distribution of α-tubulin and the neurotransmitters serotonin and RFamide in juveniles and adults of the sagittiferid Symsagittifera roscoffensis. In addition, we present the expression pattern of the neuropatterning gene SoxB1. Adults and juveniles exhibit six serotonergic longitudinal neurite bundles and an anterior concentration of serotonergic sensory cells. While juveniles show an "orthogon-like" arrangement of longitudinal neurite bundles along the anterior-posterior axis, it appears more diffuse in the posterior region of adults. Commissures between the six neurite bundles are present only in the anterior body region of adults, while irregularly distributed individual neurites, often interconnected by serotonergic nerve cells, are found in the posterior region. Anti-RFamide staining shows numerous individual neurites around the statocyst. The orthogon-like nervous system of S. roscoffensis is confirmed by α-tubulin immunoreactivity. In the region of highest neurotransmitter density (i.e., anterior), the HMG-box gene SrSoxB1, a transcription factor known to be involved in neurogenesis in other bilaterians, is expressed in juvenile specimens. Accordingly, SoxB1 expression in S. roscoffensis follows the typical pattern of higher bilaterians that have a brain. Thus, our data support the notion that Urbilateria already had the genetic toolkit required to form brain-like neural structures, but that its morphological degree of neural concentration was still low.