A layer of nerve fibres is present almost everywhere at the base of the epidermis. It consists of a very thin basal layer of irregularly arranged fibres, and generally a thicker, more superficial layer of orientated fibres, which forms the main nerve cords and subsidiary systems of smaller through-conducting bundles. In the proboscis there are numerous longitudinal bundles, an anterior nerve ring round the basal periphery and a nerve loop under the pre-oral ciliary organ. The neurocord appears to be a simple conducting tract. In the collar epidermis numerous bundles are formed posteriorly, connecting with the prebranchial nerve ring. In the trunk the size of the longitudinal cords and the distribution of the general plexus is related to the degree of development of the muscles and cilia. The gut is well equipped with nerve fibres anteriorly, where it is particularly muscular. Practically all the nerve-cell nuclei lie outside the plexus of nerve fibres. They are very numerous and widely distributed. A concentration of bipolar neurones (Hess’s cells) occurs at the proboscis tip. Cells regarded as sensory on histological grounds are abundant round the base of the proboscis and in the groove of the ciliary organ. Large unipolar neurones are concentrated in the neurocord, some possessing ‘giant’ axons, which run posteriorly or anteriorly. The cilia of the epidermis are the chief agents of locomotion, those of the trunk being capable of synchronized reversal. They are aided by peristaltic contractions of the longitudinal muscles, which are controlled by the main longitudinal nerve cords. Burrowing peristalsis is controlled by the dorsal nerve cord of the proboscis. Some reactions to light, to the presence of fine particles and to adrenaline are described. The proboscis is necessary for spontaneous and varied activity, but the considerable degree of co-ordination shown is not due to any localized centre but to a longitudinal reflex path involving the main nerve cords. Rapid contractions of the anterior end are probably due to the giant axons. The peculiarities of the neurocord are difficult to interpret, except as a result of degeneration and paedomorphosis. The greater part of the richly nervous epidermis may be compared with the vertebrate neural plate.
Read full abstract