Urochordate Nervous Systems

Urochordates are chordate siblings that comprise the following marine invertebrates: the sessile Ascidiaceae, or sea squirts; planktonic Larvacea; and the pelagic salps, doliolids, and pyrosomes (collectively the Thaliacea), each more beautiful than the next. Tadpole larvae of ascidians and adult larvaceans both have a body plan that is chordate, with a notochord and dorsal, tubular nervous system that forms from a neural plate. Thalaciacea have a ganglion developed from a tubular structure, which has been compared to the vertebrate mes-metencephalic region, and while salps have well developed eyes, other anterior brain components are absent, and the connections within their central nervous system, as well as the neurobiology of other Thaliacea are all little reported. The ascidian tadpole larva is extensively reported, especially in the model species Ciona intestinalis, as is the caudal nerve cord in the larvacean Oikopleura dioica. Chordate features that share proposed homology with vertebrate features include ciliary photoreceptors that hyperpolarize to light, descending decussating motor pathways that resemble Mauthner cell pathways, coronet cells in the ascidian larva and saccus vasculosus of fishes, the neural canal’s Reissner’s fiber; secondary mechanoreceptors that resemble hair cells; and ascidian bipolar cells that are like dorsal root ganglion cells.

Evolution and the origin of the visual retinoid cycle in vertebrates

Absorption of a photon by visual pigments induces isomerization of 11- cis -retinaldehyde (RAL) chromophore to all- trans -RAL. Since the opsins lacking 11- cis -RAL lose light sensitivity, sustained vision requires continuous regeneration of 11- cis -RAL via the process called ‘visual cycle’. Protostomes and vertebrates use essentially different machinery of visual pigment regeneration, and the origin and early evolution of the vertebrate visual cycle is an unsolved mystery. Here we compare visual retinoid cycles between different photoreceptors of vertebrates, including rods, cones and non-visual photoreceptors, as well as between vertebrates and invertebrates. The visual cycle systems in ascidians, the closest living relatives of vertebrates, show an intermediate state between vertebrates and non-chordate invertebrates. The ascidian larva may use retinochrome-like opsin as the major isomerase. The entire process of the visual cycle can occur inside the photoreceptor cells with distinct subcellular compartmentalization, although the visual cycle components are also present in surrounding non-photoreceptor cells. The adult ascidian probably uses RPE65 isomerase, and trans -to- cis isomerization may occur in distinct cellular compartments, which is similar to the vertebrate situation. The complete transition to the sophisticated retinoid cycle of vertebrates may have required acquisition of new genes, such as interphotoreceptor retinoid-binding protein, and functional evolution of the visual cycle genes.