The Origin of Vertebrates

by Walter Holbrook Gaskell

THE EVIDENCE OF THE AUDITORY APPARATUS AND THE ORGANS OF THE LATERAL LINE

Lateral line organs.—Function of this group of organs.—Poriferous sense-organs on the appendages in Limulus.—Branchial sense-organs.—Prosomatic sense organs.—Flabellum.—Its structure and position.—Sense-organs of mandibles.—Auditory organs of insects and arachnids.—Poriferous chordotonal organs.—Balancers of Diptera.—Resemblance to organs of flabellum.—Racquet-organs of Galeodes.—Pectens of scorpions.—Large size of nerve to all these special sense-organs.—Origin of parachordals and auditory capsule.—Reason why VIIth nerve passes in and out of capsule.—Evidence of Ammocœtes.—Intrusion of glandular mass round brain into auditory capsule.—Intrusion of generative and hepatic mass round brain into base of flabellum.—Summary.

When speaking of the tripartite arrangement of the cranial nerves, an arrangement which gave the clue to the meaning of the cranial segments, I spoke of the trigeminal as supplying the sensory nerves to the skin in the head-region, and I compared this dorsal system of afferent nerves to the system of epimeral nerves in Limulus which supply the prosomatic and mesosomatic carapaces of Limulus with sensory fibres. I compared the ventral system of eye-muscle nerves with the system of nerves supplying the segmental dorso-ventral somatic muscles of the prosomatic region, and I compared the lateral system of mixed nerves with the nerves supplying the prosomatic and mesosomatic appendages of Limulus. I compared, also, the optic nerves and the olfactory nerves with the corresponding nerves in the same invertebrate group. My readers will see at once that one well-marked group of nerves—the auditory and lateral line system—has been entirely omitted up to the present, it has not even been mentioned in the scheme of the cranial segments; I have purposely reserved its consideration until now, because the organs these nerves supply, though situated in the skin, are of such a special character as to form a category by themselves. These nerves cannot be classed among the afferent nerves of the skin any more than the nerves of the optic and olfactory apparatus; they require separate consideration. A very extensive literature has grown up on the subject of this system of lateral line sense-organs and their innervation, the outcome of which is decisively in favour of this system being classed with the sense-organs supplied by the auditory nerve, so that in endeavouring to understand the position of the auditory nerve, we must always bear in mind that any theory as to its origin must apply to the system of lateral line nerves as well.

Now, although the auditory apparatus is common to all vertebrates, the lateral line system is not found in any land-dwelling animals; it belongs essentially to the fishes, and is, therefore, an old system so far as concerns the vertebrate group. Its sense-organs are arranged along the lateral line of the fish, and, in addition, on the head-region in three well-marked lines known as the supra-orbital, infra-orbital, and mandibular line systems. These sense-organs lie in the skin in a system of canals, and are innervated by a special nervous system different to that innervating adjacent skin-areas. The great peculiarity of their innervation consists in the fact that their nerves all belong to the branchial system of nerves; no fibres arise in connection with the trigeminal, but all of them in connection with the facial, glossopharyngeal and vagus nerves. In other words, although organs in the skin, their nerve-supply belongs to the lateral nervous system which supplies splanchnic and not somatic segments, a system which, according to the theory advanced in this book, originated in the nerves supplying appendages. The conclusion, therefore, is that in order to obtain some clue as to the origin of the sense-organs of this system in the assumed palæostracan ancestor, we must examine the mesosomatic appendages and see whether they possess any special sense-organs of similar function.

Further, considering that the auditory organ is to be regarded as a specially developed member of this system, we must especially look for an exceptionally developed organ in the region supplied by the auditory nerve.

The question of the origin of this system of lateral line sense-organs possesses a special interest for all those who attempt to obtain a solution of the origin of vertebrates, for the upholders of the view that the vertebrates have descended from annelids have always found its strongest support in the similarity of two sets of segmental organs found in annelids and vertebrates. On the one hand, great stress was laid upon the similarity of the segmental excretory organs in the two groups of animals, as will be discussed later; on the other, of the similarity of the segmentally arranged lateral sense-organs.

These lateral sense-organs of the annelids have been specially described by Eisig in the Capitellidæ, and, according to Lang, "there are many reasons for considering these lateral organs to be homologous with the dorsal cirri of the ventral parapodia of other Polychæta, and in the family of the Glyceridæ we can follow, almost step by step, the transformation of the cirri into lateral organs." Eisig describes them in the thoracic prebranchial region as slightly different from those in the abdominal branchial region; in the latter region, the ventral parapodia are gill-bearing, so that these lateral organs are in the branchial region closely connected with the branchiæ, just as is also the case in the vertebrates. It is but a small step from the gill-bearing ventral parapodia of the annelid to the gill-bearing appendages of the phyllopod-like protostracan; so that if we assume that this is the correct line along which to search for the origin of the vertebrate auditory apparatus, then, on my theory of the origin of the vertebrates from a group resembling the Protostraca, it follows that special sense-organs must have existed either on or in close connection with the branchial and prebranchial appendages of the protostracan ancestor of the vertebrates, which would form an intermediate link between the lateral organs of the annelids and the lateral and auditory organs of the vertebrates.

Further, these special sense-organs could not have been mere tactile hairs, but must have possessed some special function, and their structure must have been compatible with that function. Can we obtain any clear conception of the original function of this whole system of sense-organs?

A large amount of experimental work has been done to determine the function of the lateral line organs in fishes, and they have been thought at one time or another to be supplementary organs for equilibration, organs for estimating pressure, etc. The latest experimental work done by Parker points directly to their being organs for estimating slow vibrations in water in contradistinction to the quicker vibrations constituting sound. He concludes that surface wave-movements, whether produced by air moving on the water or solid bodies falling into the water, are accompanied by disturbances which are stimuli for the lateral line organs.

One of these segmental organs has become especially important and exists throughout the whole vertebrate group, whether the animal lives on land or in water—this is the auditory organ. Throughout, the auditory organ has a double function—the function of hearing and the function of equilibration. If, then, this is, as is generally supposed, a specialized member of the group, it follows that the less specialized members must possess the commencement of both these functions, just as the experimental evidence suggests.