Fig. 114.—Distribution of Trigeminal Nerve in Ammocœtes.

ps. br., pseudo-branchial groove; met., nerve to lower lip, or metastomal nerve; t., nerve to tongue; tent., nerve to tentacles. The mandibular and internal maxillary nerves are coloured red; the purely sensory nerves to the external surface are coloured black.

The Trigeminal Nerve of Ammocœtes.

Striking is the answer. In Fig. [114], Miss Alcock has drawn the distribution of the trigeminal nerve as traced by her through a series of sections. It arises, as is well known, from two separate ganglia, of which the foremost gives rise to a purely cutaneous nerve, the ophthalmic nerve, and the hindmost to three nerves, the most posterior of which is purely cutaneous and passes tailwards over the ventral branchial region, as shown in the figure; the other two nerves, both of which contain motor fibres, are called by Hatschek the mandibular and maxillary nerves. Of these the mandibular or velar nerve (met.) is a large, conspicuous nerve, which arises so separately from the rest of the trigeminal as almost to deserve the title of a separate nerve. When it leaves the large posterior ganglion, it passes into the anterior part of the velum, runs along with the tubular muscles, which it supplies, to the ventral surface as far as the junction of the lower lip with the thyroid plate, and has not been followed further by Hatschek. Miss Alcock, however, by means of serial sections, has traced it further, and shown that at this point it turns abruptly headwards to terminate in the muscles of the lower lip. If, then, as suggested, the lower lip represents the metastoma—the last pair of prosomatic appendages—then this mandibular or velar nerve represents that segmental nerve.

The other nerve—the maxillary nerve of Hatschek—which constitutes the larger part of the trigeminal, passes forwards from the ganglion, and at a point somewhere about the anterior region of the eyeball, divides into two, an external (black in Fig. [114]) and an internal (red in Fig. [114]) nerve. The external branch is apparently entirely sensory, and supplies the external surfaces of the upper and lower lips. The internal branch is mainly motor, and supplies the muscles of the upper lip; it contains also the nerves of the tentacles.

The nerve to the median ventral tentacle (t.) or tongue leaves the internal division of the maxillary immediately after its separation from the external; it runs ventralwards, and at the same time passes internally until it reaches a position between the muco-cartilage and the epithelium lining the cavity of the throat. It then turns, and passing posteriorly (towards the tail) to the point where the median ventral tentacle is attached to the lower lip, it supplies some very rudimentary-looking muscles which run from the tentacle to the adjoining surface, and no doubt serve to move the tentacle from side to side. A portion of the nerve still continues to run along the side of the median ventral ridge, as far back as the point where the muscles of the hyoid segment pass round to the ventral side between the velum and the thyroid; in fact, this small nerve passes along the whole length of the median ventral ridge.

This description shows that the trigeminal nerve divides itself into two groups: the one represented black in the figure, which is purely cutaneous and sensory, corresponding, in the main, according to my theory, to the epimeral nerves of Limulus; the other coloured red, which supplies muscles belonging to the visceral or splanchnic muscle-group, and contains also the nerves to the tentacles.

This latter group, which is formed by two distinct well-defined nerves, viz. the mandibular and the internal branch of the maxillary, corresponds, according to my theory, to the amalgamated nerves of the prosomatic appendages, and is clearly divisible into three distinct nerves—

1. The lower lip-nerve or the metastomal nerve (met.).