The Origin of Vertebrates

by Walter Holbrook Gaskell

In our search, then, for the origin of the auditory organ of vertebrates, we must look for special organs for the estimation of vibrations and for the maintenance of the equilibrium of the animal, situated on the appendages, especially the branchial or mesosomatic appendages; and, further, we must specially look for an exceptional development of such segmental organs at the junction of the prosomatic and mesosomatic regions.

Throughout this book the evidence which I have put forward has in all cases pointed to the same conclusion, viz. that the vertebrate arose by way of the Cephalaspidæ from some arthropod, either belonging to, or closely allied to, the group called Palæostraca, of which the only living representative is Limulus. If, then, my argument so far is sound, the appendages of Limulus, both prosomatic and mesosomatic, ought to possess special sense-organs which are concerned in equilibration or the appreciation of the depth of the water, or in some modification of such function, and among these we might expect to find that somewhere at the junction of the prosoma and mesosoma such sense-organs were specially developed to form the beginning of the auditory organ.

Now, it is a striking fact that the appendages of Limulus do possess special sense-organs of a remarkable character, which are clearly not simply tactile. Thus Gegenbaur, as already stated, has drawn attention to the remarkable branchial sense-organs of Limulus; and Patten has pointed out that special organs, which he considers to be gustatory in function, are present on the mandibles of the prosomatic appendages. I myself, as mentioned in my address to the British Association at Liverpool in 1896, searched for some special sense-organ at the junction of the prosoma and mesosoma, and was rewarded by finding that that extraordinary adjunct to the last locomotor appendage, known as the flabellum, was an elaborate sense-organ. I now propose to show that all these special sense-organs are constructed on a somewhat similar plan; that the structure of the branchial sense-organs suggests that they are organs for the estimation of water pressures; that among air-breathing arthropods sense-organs, built up on a somewhat similar plan, are universally found, and are considered to be of the nature of auditory and equilibration organs; and, what is especially of importance, in view of the fact that the most prominent members of the Palæostraca were the sea-scorpions, that the remarkable sense-organs of the scorpions known as the pectens belong apparently to the same group.

The Poriferous Sense-Organs of the Appendages in Limulus.

On all the branchial appendages in Limulus, special sense-organs are found of a most conspicuous character. They form in the living animal bluish convex circular patches, the situation of which on the appendages is shown in Fig. [58]. These organs are not found on the non-branchial operculum. Gegenbaur, who was the first to describe them, has pointed out how the surface of the organ is closely set with chitinous goblets shaped as seen in Fig. [144], A, which do not necessarily project free on the surface, but are extruded on the slightest pressure. Each goblet fits into a socket in the chitinous covering, and is apparently easily protruded by variations of pressure from within. The whole surface of the organ on the appendage is slightly bulged in the living condition, and the chitin is markedly softer here than in the surrounding part of the limb. Each of these organs is surrounded by a thick protection of strongly branching spines. On the surface of the organ itself no spines are found, only these goblets, so that the surface-view presents an appearance as in Fig. [144], B. Each goblet possesses a central pore, which is the termination of a very fine, very tortuous, very brittle chitinous tubule (ch.t.), which passes from the goblet through the layers of the chitin into the subjacent tissue. The goblets vary considerably in size, a few very large ones being scattered here and there. The fine chitinous tubule is especially conspicuous in connection with these largest goblets. In the smaller ones there is the same appearance of a pore and a commencing tube, but I have not been able to trace the tube through the chitinous layers, as in the case of the larger goblets.

Fig. 144.—A, A Goblet from one of the Branchial Sense-organs of Limulus (ch.t., chitinous tubule); B, Surface View of a Portion of a Branchial Sense-Organ.	Fig. 145.—The Endognaths of Limulus pushed out of the way on one side in order to show the position of the Flabellum (fl.) projecting towards the crack between the Prosomatic and Mesosomatic Carapaces.

Gegenbaur, in his picture, draws a straight tubule passing from every goblet among the fine canaliculi of the chitin. He says they are difficult to see, except in the case of the larger goblets. The tubule from the larger goblets is most conspicuous, and is in my sections always tortuous, never straight, as represented by Gegenbaur. A special branch of the appendage-nerve passes to these organs, and upon the fine branches of this nerve groups of ganglion-cells are seen, very similar in appearance to the groups described by Patten on the terminal branches of the nerves which supply the mandibular organs. At present I can see no mechanism by which the goblets are extruded or returned into place. In the case of the Capitellidæ, Eisig describes retractor muscles by means of which the lateral sense-organs are brought below the level of the surface, and he imagines that the protrusion is effected by hydraulic means, by the aid of the vascular system. In the branchial sense-organs of Limulus there are no retractor muscles, and it seems to me that both retraction and protrusion must be brought about by alterations of pressure in the vascular fluids. Certainly the cavity of the organ is very vascular. If this be so, it seems likely enough that such an organ should be a very delicate organ for estimating changes in the pressure of the external medium, for the position of the goblets would depend on the relation between the pressure of the fluid inside the organ and that on the surface of the appendage. Whether the chitinous tubule contains a nerve-terminal or not I am unable to decide from my specimens, but, judging from Patten's description of the similar chitinous tubules belonging to the mandibular organs, it is most highly probable that these tubules also contain a fine terminal nerve-fibre.

These organs, then, represent segmental branchial sense-organs, of which it can be said their structure suggests that they may be pressure-organs; but the experimental evidence is at present wanting.

Passing now from the branchial to the prosomatic region, the first thing that struck me was the presence of that most conspicuous projection at the base of the last locomotor appendage, which is usually called the flabellum, and has been described by Lankester as an exopodite of this appendage. It is jointed on to the most basal portion of the limb (cf. Fig. [155]), and projects dorsally from the limb into the open slit between the prosomatic and mesosomatic carapace, as is seen in Fig. [145] (fl.). Of its two surfaces, the undermost is very convex and the uppermost nearly flat from side to side, the whole organ being bent, so that when the animal is lying half buried in sand, entirely covered over by the prosomatic and mesosomatic carapaces except along this slit between the two, the upper flat or slightly convex surface of the flabellum is exposed to any movement of water through this slit, and owing to its possessing a joint, the direction of the whole organ can be altered to a limited extent. The whole of this flat upper surface is one large sense-organ of a striking character, thus forming a great contrast to the convex under surface, which is remarkably free from tactile spines or special sense-organs.

The nerve going to the flabellum is very large, almost as large as the nerve to the rest of the appendage, and the very large majority of the nerve-fibres turn towards the flat, uppermost side, where the sense-organ is situated. Between the nerve-fibres (n.) and the chitinous surface containing the special sense-tubes masses of cells (gl.) are seen, as in Fig. [146], apparently nerve-cells, which form a broad border between the nerve-fibres and the pigmented chitinogenous layer (p.). On the opposite side, nothing of the sort intervenes between the pigmented layer and the blood-spaces and nerve-fibres which constitute the central mass of the flabellum.