Ontogeny of the head of the Pacific hagfish ( Eptatretus stouti , Myxinoidea): development of the lateral line system

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Abstract
The head of adult hagfishes (jawless craniates, Myxinoidea) of the family Eptatretidae displays a number of skin grooves of uncertain origin. These grooves have been homologized to the neuromast lines of other craniates, and they are innervated by two ganglionated cranial nerves that have been interpreted as lateral line nerves. The grooves do not, however, contain the compound receptors that are typical of a lateral line (i.e. neuromasts or electroreceptors), and both their development and function have remained enigmatic. To elucidate the embryonic origin of the grooves (which should develop from placodes if they are homologues of the lateral line system), embryos of Pacific hagfish were examined by means of threedimensional reconstructions from serial sections. Because of the scarcity of specimens of embryonic hagfishes, only two embryos were reconstructed, but these reconstructions clearly show that a number of placodes and placodal derivatives (i.e. sensory ridges, receptor primordia, and cranial ganglia) occur in the head of embryonic eptatretid hagfishes. Some of these placodes correspond to the lens and epibranchial placodes of other craniates, but there are also three other placodes which represent possible homologues of lateral line placodes. The topology of the placodes in this latter group corresponds to the topology of the grooves of adult hagfishes, and we therefore reach three conclusions: (i) that an embryonic lateral line system is present in hagfishes; (ii) that the grooves of adult hagfishes in all probability derive from lateral line placodes; and (iii) that the presence of lateral line placodes is a primitive character of craniates.