Comparison of the serotonergic nervous system among Tunicata: implications for its evolution within Chordata

Using immunohistochemistry in combination with confocal laser scanning microscopy, the serotonergic nervous systems of major tunicate taxa were studied in three-dimensional detail. Organisms analyzed included aplousobranchiate, phlebobranchiate, and stolidobranchiate ascidian larvae, appendicularian juveniles and adults, and doliolid oozooids. Outgroup comparisons to notochordates showed that the serotonergic nervous system of the last common ancestor of Chordata consisted of two elements: (i) an anterior concentration of serotonergic cell bodies, and (ii) a fiber network that followed posteriorly and gave rise to fiber tracts that descended towards the effective somatic lateral musculature. Within Tunicata, the nervous systems of Appendicularia and Aplousobranchiata appear serotonin-reduced or negative. This could be interpreted as a heterochronic reduction and a synapomorphy between Appendicularia and Aplousobranchiata. In this hypothesis, free-living Appendicularia are derived within Tunicata, and a biphasic life cycle with a free-swimming larva and a sessile, ascidian-like adult is most parsimoniously reconstructed for the last common ancestor of Tunicata. The close spatial relation of the serotonergic cell cluster with the statocyte complex suggests a function as an integrative control center for the coordination of locomotion. A similar anterior concentration of serotonergic nerve cells is known from tornaria larvae.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-02.htm