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To give a full account of the morphology, development, and classification of the Medusæ would be both unnecessary for our present purposes and impracticable within the space which is allotted to the present work.[2] But, for the sake of clearness in what follows, I shall begin by briefly describing such features in the anatomy of the jelly-fish as will afterwards be found especially to concern us.

Fig. 1.

Sarsia (natural size).

In size, the different species of Medusæ vary from that of a small pea to that of a large umbrella having streamers a hundred feet long. The general form of these animals varies in different species from that of a thimble (Fig. 1) to that of a bowl, a parasol, or a saucer (see figures in subsequent chapters). Or we may say that the form of the animals always resembles that of a mushroom, and that the resemblance extends to a tolerably close imitation by different species of the various forms which are characteristic of different species of mushrooms, from the thimble-like kinds to the saucer-like kinds. Moreover, this accidental resemblance to a mushroom is increased by the presence of a central organ, occupying the position of, and more or leas resembling in form, the stalk of a mushroom. This organ is called the "manubrium," on account of its looking like the "handle" of an umbrella, and the term "umbrella" is applied to the other portion of the animal. The manubrium, like the umbrella, varies much in size and shape in different species, as a glance at any figures of these animals will show. Both the manubrium and umbrella are almost entirely composed of a thick, transparent, and non-contractile jelly; but the whole surface of the manubrium and the whole concave surface of the umbrella are overlayed by a thin layer or sheet of contractile tissue. This tissue constitutes the earliest appearance in the animal kingdom of true muscular fibres, and its thickness, which is pretty uniform, is nowhere greater than that of very thin paper.

The manubrium is the mouth and stomach of the animal, and at the point where it is attached to or suspended from the umbrella its central cavity opens into a tube-system, which radiates through the lower or concave aspect of the umbrella. This tube-system, which serves to convey digested material and may therefore be regarded as intestinal in function, presents two different forms in the two main groups into which the Medusæ are divided. In the "naked-eyed" group, the tubes are unbranched and run in a straight course to the margin of the umbrella, where they open into a common circular tube which runs all the way round the margin (see Figs. 1 and 22). In the "covered-eyed" group, on the other hand, the tubes are strongly branched (see Fig. 8), although they likewise all eventually terminate in a single circular tube. This circular or marginal tube in both cases communicates by minute apertures with the external medium.

The margin of the umbrella, both in the naked and covered eyed Medusæ, supports a series of contractile tentacles, which vary greatly in size and number in different species (see Figs. 1 and 8). The margin also supports another series of bodies which will presently be found to be of much importance for us. These are the so-called "marginal bodies," which vary in number, size, and structure in different species. In all the covered-eyed species these marginal bodies occur in the form of little bags of crystals (therefore they are called "lithocysts"), which are protected by curiously formed "hoods" or "covers" of gelatinous tissue; and it is on this account that the group is called "covered-eyed," in contradistinction to the "naked-eyed," where these little hoods or coverings are invariably absent (compare Fig. 1 with Fig. 22), and the crystals frequently so. In nearly all cases these marginal bodies contain more or less brightly coloured pigments.

The question whether any nervous tissue is present in the Medusæ is one which has long occupied the more or less arduous labours of many naturalists. The question attracted so much investigation on account of its being one of unusual interest in biology. Nerve-tissue had been clearly shown to occur in all animals higher in the zoological scale than the Medusæ, so that it was of much importance to ascertain whether or not the first occurrence of this tissue was to be met with in this class. But, notwithstanding the diligent application of so much skilled labour, up to the time when my own researches began there had been so little agreement in the results obtained by the numerous investigators, that Professor Huxley—himself one of the greatest authorities upon the group—thus defined the position of the matter in his "Classification of Animals" (p. 22): "No nervous system has yet been discovered in any of these animals."

The following is a list of the more important researches on this topic up to the time which I have just named:—Ehrenberg, "Die Acalephen des rothen Meeres und der Organismvs der Medusen der Ostsee," Berlin, 1836; Kölliker, "Ueber die Randkörper der Quallen, Polypen und Strahlthiere," Froriep's neue Notizen, bd. xxv., 1843; Von Beneden, "Mémoire sur les Campanulaires de la côte d'Ostende," "Mémoires de l'Académie de Bruxelles," vol. xvii., 1843; Desor, "Sur la Génération Medusipare des Polypes hydraires," "Annales d. Scienc. Natur. Zool.," ser. iii. t. xii. p. 204; Krohn, "Ueber Podocoryna carnea," "Archiv. f. Naturgeschichte," 1851, b. i.; McCrady, "Descriptions of Oceania, etc.," "Proceedings of the Elliot Society of Natural History," vol. i., 1859; L. Agassiz, "Contributions to the Acaliphæ of North America," "Memoirs of the American Academy of Arts and Sciences," vol. iii., 1860, vol. iv., 1862; Leuckart, "Archiv. f. Naturgeschichte," Jahrg. 38, b. ii., 1872; Hensen, "Studien über das Gehörorgan der Decapoden," "Zeitchr. f. wiss. Zool.," bd. xiii., 1863; Semper, "Reisebericht," "Zeitschr. f. wiss. Zool.," bd. xiii. vol. xiv.; Claus, "Bemerkungen über Clenophoren und Medusen," "Zeitschr. f. wiss. Zool.," bd. xiv., 1864; Allman, "Note on the Structure of Certain Hydroid Medusæ," "Brit. Assoc. Rep.," 1867; Fritz Müller, "Polypen und Quallen von S. Catherina," "Archiv. f. Naturgesch.," Jahrg. 25, bd. i., 1859; also "Ueber die Randbläschen der Hydroidquallen," "Archiv. f. Anatomie und Physiologie," 1852; Haeckel, "Beiträge zur Naturgesch. der Hydromedusen," 1865; Eimer, "Zoologische Untersuchungen," Würzburg, "Verhandlungen der Phys.-med. Gesellschaft," N.F. vi. bd., 1874.

The most important of these memoirs for us to consider are the two last. I shall subsequently consider the work of Dr. Eimer, which up to this date was of a purely physiological character. Professor Haeckel, who made his microscopical observations chiefly upon the Geryonidæ, described the nervous elements as forming a continuous circle all round the margin of the umbrella, following the course of the radial or nutrient tubes throughout their entire length, and proceeding also to the tentacles and marginal bodies. At the base of each tentacle there is a ganglionic swelling, and it is from these ganglionic swellings that the nerves just mentioned take their origin. The most conspicuous of these nerves are those that proceed to the radial canals and marginal bodies, while the least conspicuous are those that proceed to the tentacles. Cells, as a rule, can only be observed in the ganglionic swellings, where they appear as fusiform and distinctly nucleated bodies of great transparency and high refractive power. On the other hand, the nerves that emanate from the ganglia are composed of a delicate and transparent tissue, in which no cellular elements can be distinguished, but which is longitudinally striated in a manner very suggestive of fibrillation. Treatment with acetic acid, however, brings out distinct nuclei in the case of the nerves that are situated in the marginal vesicles, while in those that accompany the radial canals ganglion-cells are sometimes met with.

A brief sketch of the contents of these and other memoirs on the histology of the Medusæ is given by Drs. Hertwig in their more recently published work on the nervous system and sense-organs of the Medusæ, and these authors point to the important fact that before the appearance of Haeckel's memoir, Leuckart was the only observer who spoke for the fibrillar character of the so-called marginal ring-nerve; so that in Haeckel's researches on Geryonia, whereby both true ganglion-cells and true nerve-fibres were first demonstrated as occurring in the Medusæ, we have a most important step in the histology of these animals. Haeckel's results in these respects have since been confirmed by Claus, "Grundzüge der Zoologie," 1872; Allman, "A Monograph of the Gymnoblastic or Tubularian Hydroids," 1871; Harting, "Notices Zoologiques," Niedlandisches "Archiv. f. Zool.," bd. ii., Heft 3, 1873; F. E. Schulze, "Ueber den Bau von Syncorzne Sarsii"; O. and R. Hertwig, "Das Nervensystem und die Sinnesorgane der Medusen."

The last-named monograph is much the most important that has appeared upon the histology of the Medusæ. I shall, therefore, give a condensed epitome of the leading results which it has established.

There is so great a difference between the nervous system of the naked and of the covered eyed Medusæ, that a simultaneous description of the nervous system in both groups is not by these authors considered practicable. Beginning, therefore, with the naked-eyed division, they describe the nervous system as here consisting of two parts, a central and a peripheral. The central part is localized in the margin of the swimming-bell, and there forms a "nerve-ring," which is divided by the insertion of the "veil"[3] into an upper and a lower nerve-ring. In many species the upper nerve-ring is spread out in the form of a flattish layer, which is somewhat thickened where it is in contact with the veil. In these species the nerve-ring is only indistinctly marked off from the surrounding tissues. But in other species the crowding together of the nerve-fibres at the insertion of the veil gives rise to a considerable concentration of nervous structures; while in others, again, this concentration proceeds to the extent of causing a well-defined swelling of nervous tissue against the epithelium of the veil and umbrella. In the Geryonidæ this swelling is still further strengthened by a peculiar modification of the other tissues in the neighbourhood, which had been previously described by Professor Haeckel. In all species the upper nerve-ring lies entirely in the ectoderm. Its principal mass is composed of nerve-fibres of wonderful tenuity, among which are to be found sparsely scattered ganglion-cells. The latter are for the most part bi-polar, more seldom multi-polar. The fibres which emanate from them are very delicate, and, becoming mixed with others, do not admit of being further traced. Where the nervous tissue meets the enveloping epithelium it is connected with the latter from within, but differs widely from it; for the nerve-cells contain a longitudinally striated cylindrical or thread-like nucleus which carries on its peripheral end a delicate hair, while its central end is prolonged into a fine nerve-fibre. There are, besides these, two other kinds of cells which form a transition between the ganglion and the epithelium cells. The first kind are of a long and cylindrical form, the free ends of which reach as far as the upper surface of the epithelium The second kind lie for the most part under the upper surface. They are of a large size, and present, coursing towards the upper surface, a long continuation, which at its free extremity supports a hair. In some cases this continuation is smaller, and stops short before reaching the outer surface. Drs. Hertwig observe that in these peculiar cells we have tissue elements which become more and more like the ordinary ganglion-cells of the nerve-ring the more that their long continuation towards the surface epithelium is shortened or lost, and these authors are thus led to conclude that the upper nerve-ring was originally constituted only by such prolongations of the epithelium-cells, and that afterwards these prolongations gradually disappeared, leaving only their remnants to develop into the ordinary ganglion-cells already described.

Beneath the upper nerve-ring lies the lower nerve-ring. It is inserted between the muscle-tissue of the veil and umbrella, in the midst of a broad strand wherein muscle-fibres are entirely absent. It here constitutes a thin though broad layer which, like the upper nerve-ring, belongs to the ectoderm. It also consists of the same elements as the upper nerve-ring, viz. of nerve-fibres and ganglion-cells. Yet there is so distinct a difference of character between the elements composing the two nerve-rings, that even in an isolated portion it is easy to tell from which ring the portion has been taken. That is to say, in the lower nerve-ring there are numerous nerve-fibres of considerable thickness, which contrast in a striking manner with the almost immeasurably slender fibres of the upper nerve-ring. A second point of difference consists in the surprising wealth of ganglion-cells in the one ring as compared with the other. Thus, on the whole, there is no doubt that the lower nerve-ring presents a higher grade of structure than does the upper, as shown not only by the greater multiplicity of nerve-cells and fibres, but also by the relation in which these elements stand to the epithelium. For in the case of the lower nerve-ring, the presumably primitive connections of the nervous elements with the epithelium is well-nigh dissolved—this nerve-ring having thus separated itself from its parent structure, and formed for itself an independent layer beneath the epithelium. The two nerve-rings are separated from one another by a very thin membrane, which, in some species at all events, is bored through by strands of nerve-fibres which serve to connect the two nerve-rings with one another.

The peripheral nervous system is also situated in the ectoderm, and springs from the central nervous system, not by any observable nerve-trunks, but directly as a nervous plexus composed both of cells and fibres. Such a nervous plexus admits of being detected in the sub-umbrella of all Medusæ, and in some species may be traced also into the tentacles. It invariably lies between the layer of muscle-fibre and that of the epithelium. The processes of neighbouring ganglion-cells in the plexus either coalesce or dwindle in their course to small fibres: at the margin of the umbrella these unite themselves with the elements of the nerve-rings. There are also described several peculiar tissue elements, such as, in the umbrella, nerve-fibres which probably stand in connection with epithelium-cells; nerve-cells which pass into muscle-fibres, similar to those which Kleinenberg has called neuro-muscular cells; and, in the tentacles, neuro-muscular cells joined with cells of special sensation (Sinneszellen).

No nervous elements could be detected in the convex surface of the umbrella, and it is doubtful whether they occur in the veil.

In some species the nerve-fibres become aggregated in the region of the generative organs, and in that of the radial canals, thus giving rise in these localities to what may be called nerve-trunks. But in other species no such aggregations are apparent, the nervous plexus spreading out in the form of an even trellis-work.

In the covered-eyed Medusæ the central nervous system consists of a series of separate centres which are not connected by any commissures. These nerve-centres are situated in the margin of the umbrella, and are generally eight in number, more rarely twelve, and in some species sixteen. They are thickenings of the ectoderm, which either enclose the bases of the sense-organs, or only cover the ventral side of the same. Histologically they consist of cells of special sensation, together with a thick layer of slender nerve-fibres. Ganglion-cells, however, are absent, so that the nerve-fibres are merely processes of epithelium-cells.

Drs. Hertwig made no observations on the peripheral nervous system of the covered-eyed Medusæ; but they do not doubt that such a system would admit of being demonstrated, and in this connection they cite the observations of Claus, who describes numerous ganglion-cells as occurring in the sub-umbrella of Chrysaora. Here I may appropriately state that before Drs. Hertwig had published their results, Professor Schäfer, F.R.S., conducted in my laboratory a careful research upon the histology of the Medusæ, and succeeded in showing an intricate plexus of cells and fibres overspreading the sub-umbrella tissue of another covered-eyed Medusa (Aurelia aurita).[4] He also found that the marginal bodies present a peculiar modification of epithelium tissue, which is on its way, so to speak, towards becoming fully differentiated into ganglionic cells.

Lastly, returning to the researches of Drs. Hertwig, these authors compare the nervous system of the naked-eyed with that of the covered-eyed Medusæ, with the view of indicating the points which show the latter to be less developed than the former. These points are, that in the nerve-centres of the covered-eyed Medusæ there are no true ganglion-cells, or only very few; that the mass of the central nervous system is very small; and that the centralization of the nervous system is less complete in the one group than in the other. In their memoir these authors further supply much interesting information touching the structure of the sense-organs in various species of Medusæ; but it seems scarcely necessary to extend the present résumé of their work by entering into this division of their subject.

In a later publication, entitled "Der Organismus der Medusen und seine Stellung zur Keimblättertheorie," Drs. Hertwig treat of sundry features in the morphology of the Medusæ which are of great theoretical importance; but here again it would unduly extend the limits of the present treatise if I were to include all the ground which has been so ably cultivated by these industrious workers.

It will presently be seen in how striking a manner all the microscopical observations to which I have now briefly alluded are confirmed by the physiological observations—or, more correctly, I might say that the microscopical observations, in so far as they were concerned with demonstrating the existence of nerve-tissue in the Medusæ, were forestalled by these physiological experiments; for, with the exception of Professor Haeckel's work on Geryonidæ, they were all of later publication. But in matters of scientific inquiry mere priority is not of so much importance as it is too often supposed to be. Thus, in the present instance, no one of the workers was in any way assisted by the publications of another. In each case the work was independent and almost simultaneous.

The remark just made applies also to the only research which still remains to be mentioned. This is the investigation undertaken and published by Professor Eimer.[5] He began, like myself, by what in the next chapter I call the "fundamental observation" on the effects of excising the nerve-centres, and from this basis he worked both at the physiology and the morphology of the neuro-muscular tissues. In point of time, I was the first to make the fundamental observation, and he was the first to publish it. The sundry features in which our subsequent investigations agreed, and those in which they differed, I shall mention throughout the course of the following pages.

I shall now conclude this chapter by giving a brief account of those general principles of the physiology of nerve and muscle with which it is necessary to be fully acquainted, in order to understand the course of the following experiments.

Nerve-tissue, then, universally consists of two elementary structures, viz. very minute nerve-cells and very minute nerve-fibres. The fibres proceed to and from the cells, so in some cases serving to unite the cells with one another, and in other cases with distant parts of the animal body. Nerve-cells are usually found collected together in aggregates, which are called nerve-centres or ganglia, to and from which large bundles of nerve-fibres come and go.

To explain the function of nerve-tissue, it is necessary to begin by explaining what physiologists mean by the term "excitability." Suppose that a muscle has been cut from the body of a freshly killed animal; so long as it is not interfered with in any way, so long will it remain quite passive. But every time a stimulus is supplied to it, either by means of a pinch, a burn, an electrical shock, or a chemical irritant, the muscle will give a single contraction in response to every stimulation. And it is this readiness of organic tissues to respond to a suitable stimulus that physiologists designate by the term "excitability."

Nerves, no less than muscles, present the property of being excitable. If, together with the excised muscle, there had been removed from the animal's body an attached nerve, every time any part of this nerve is stimulated the attached muscle will contract as before. But it must be carefully observed that there is this great difference between these two cases of response on the part of the muscle—that while in the former case the muscle responded to a stimulus applied directly to its own substance, in the latter case the muscle responded to a stimulus applied at a distance from its own substance, which stimulus was then conducted to the muscle by the nerve. And in this we perceive the characteristic function of nerve-fibres, viz. that of conducting stimuli to a distance. The function of nerve-cells is different, viz. that of accumulating nervous energy, and, at fitting times, of discharging this energy into the attached nerve-fibres. The nervous energy, when thus discharged, acts as a stimulus to the nerve-fibre; so that if a muscle is attached to the end of a fibre, it contracts on receiving this stimulus. I may add that when nerve-cells are collected into ganglia, they often appear to discharge their energy spontaneously; so that in all but the very lowest animals, whenever we see apparently spontaneous action, we infer that ganglia are probably present. Lastly, another important distinction must be borne in mind—the distinction, namely, which is to be drawn between muscle and nerve. A stimulus applied to a nerveless muscle can only course through the muscle by giving rise to a visible wave of contraction, which spreads in all directions from the seat of disturbance as from a centre. A nerve, on the other hand, conducts the stimulus without sensibly moving or undergoing any change of shape. Now, in order not to forget this distinction, I shall always speak of muscle-fibres as conveying a visible wave of contraction, and of nerve-fibres as conveying an invisible, or molecular, wave of stimulation. Nerve-fibres, then, are functionally distinguished from muscle-fibres—and also from protoplasm—by displaying the property of conducting invisible, or molecular, waves of stimulation from one part of an organism to another, so establishing physiological continuity between such parts, without the necessary passage of waves of contraction.