Jellyfish, Starfish, and Sea-Urchins: Being a Research on Primitive Nervous Systems

To give a full account of the morphology, development, and classification of the Medus? 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 Medus? 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 Medus? 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 Medus?, 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 Medus? 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 Medus?, 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; K?lliker, "Ueber die Randk?rper der Quallen, Polypen und Strahlthiere," Froriep's neue Notizen, bd. xxv., 1843; Von Beneden, "Mémoire sur les Campanulaires de la c?te 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 Acaliph? 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 Geh?rorgan 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 Medus?," "Brit. Assoc. Rep.," 1867; Fritz Müller, "Polypen und Quallen von S. Catherina," "Archiv. f. Naturgesch.," Jahrg. 25, bd. i., 1859; also "Ueber die Randbl?schen der Hydroidquallen," "Archiv. f. Anatomie und Physiologie," 1852; Haeckel, "Beitr?ge 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 Geryonid?, 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 Medus? is given by Drs. Hertwig in their more recently published work on the nervous system and sense-organs of the Medus?, 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 Medus?, 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 Medus?. 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 Medus?, 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 Geryonid? 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 Medus?, 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 Medus? 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 Medus?; 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 Sch?fer, F.R.S., conducted in my laboratory a careful research upon the histology of the Medus?, 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 Medus?, 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 Medus? 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 Medus?; 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 Keimbl?ttertheorie," Drs. Hertwig treat of sundry features in the morphology of the Medus? 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 Medus?, were forestalled by these physiological experiments; for, with the exception of Professor Haeckel's work on Geryonid?, 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.

The naked-eyed Medus? are very much smaller in size than the covered-eyed, and as we shall find that the distribution of their nervous elements is somewhat different, it will be convenient to use different names for the large umbrella-shaped part of a covered-eyed Medusa, and the much smaller though corresponding part of a naked-eyed Medusa. The former, therefore, I shall call the umbrella, and the latter the swimming-bell, or nectocalyx. In each case alike this portion of the animal performs the office of locomotion, and it does so in the same way.

I have already said that this mushroom-like organ, which constitutes the main bulk of the animal, is itself mainly constituted of thick transparent and non-contractile jelly, but that the whole of its concave surface is lined with a thin sheet of muscular tissue. Such being the structure of the organ, the mechanism whereby it effects locomotion is very simple, consisting merely of an alternate contraction and relaxation of the entire muscular sheet which lines the cavity of the bell. At each contraction of this muscular sheet the gelatinous walls of the bell are drawn together; the capacity of the bell being thus diminished, water is ejected from the open mouth of the bell backwards, and the consequent reaction propels the animal forwards. In these swimming movements, systole and diastole follow one another with as perfect a rhythm as they do in the beating of a heart.

Effects of excising the entire Margins of Nectocalyces.

Confining our attention under this heading to the naked-eyed Medus?, I find that the following proposition applies to every species of the group which I have as yet had the opportunity of examining: Excision of the extreme margin of a nectocalyx causes immediate, total, and permanent paralysis of the entire organ. Nothing can possibly be more definite than in this highly remarkable effect. I have made hundreds of observations upon various species of the naked-eyed Medus?, of all ages and conditions of freshness, vigour, etc.; and I have constantly found that if the experiment be made with ordinary care, so as to avoid certain sources of error presently to be named, the result is as striking and decided as it is possible to desire.[6] Indeed, I do not know of any case in the animal kingdom where the removal of a centre of spontaneity causes so sudden and so complete a paralysis of the muscular system, there being no subsequent movements or twitchings of a reflex kind to disturb the absolute quiescence of the mutilated organism. The experiment is particularly beautiful if performed on Sarsia; for the members of this genus being remarkably active, the death-like stillness which results from the loss of so minute a portion of their substance is rendered by contrast the more surprising.

From this experiment, therefore, I conclude that in the margin of all the species of naked-eyed Medus? which I have as yet had the opportunity of examining, there is situated an intensely localized system of centres of spontaneity, having at least for one of its functions the origination of impulses, to which the contractions of the nectocalyx, under ordinary circumstances, are exclusively due. And this obvious deduction is confirmed (if it can be conceived to require confirmation) by the behaviour of the severed margin. This continues its rhythmical contractions with a vigour and a pertinacity not in the least impaired by its severance from the main organism, so that the contrast between the perfectly motionless swimming-bell and the active contractions of the thread-like portion which has just been removed from its margin is as striking a contrast as it is possible to conceive. Hence it is not surprising that if the margin be left in situ, while other portions of the swimming-bell are mutilated to any extent, the spontaneity of the animal is not at all interfered with. For instance, if the equator of any individual belonging to the genus Sarsia (Fig. 1) be cut completely through, so that the swimming-bell instead of being closed at the top is converted into an open tube, this open tube continues its rhythmical contractions for an indefinitely long time, notwithstanding that the organism so mutilated is, of course, unable to progress. Thus it is a matter of no consequence how small or how large a portion of contractile tissue is left adhering to the severed margin of the swimming-bell; for whether this portion be large or small, the locomotor centres contained in the margin are alike sufficient to supply the stimulus to contraction. Indeed, if only the tiniest piece of contractile tissue be left adhering to a single marginal body cut out of the bell of Sarsia, this tiny piece of tissue, in this isolated state, will continue its contractions for hours, or even for days.

Effects of excising the entire Margins of Umbrellas.

Turning now to the covered-eyed division of the Medus?, I find, in all the species I have come across, that excision of the margins of umbrellas produces an effect analogous to that which is produced by excision of the margins of swimming-bells. There is an important difference, however, between the two cases, in that the paralyzing effect of the operation on umbrellas is neither so certain nor so complete as it is on swimming-bells. That is to say, although in the majority of experiments such mutilation of umbrellas is followed by immediate paralysis, this is not invariably the case; so that one cannot here, as with the naked-eyed Medus?, predict with any great confidence what will be the immediate result of any particular experiment. Further, although such mutilation of an umbrella is usually followed by a paralysis as sudden and marked as that which follows such mutilation of a swimming-bell, the paralysis of the former differs from the paralysis of the latter, in that it is very seldom permanent. After periods varying from a few seconds to half an hour or more, occasional weak and unrhythmical contractions begin to manifest themselves, or the contractions may even be resumed with but little apparent change in their character and frequency. The condition of the animal before the operation, as to general vigour, etc., appears to be one factor in determining the effect of the operation; but this is very far from being the only factor.

Upon the whole, then, although in the species of covered-eyed Medus? which I have as yet had the opportunity of examining, the effects which result from excising the margins of umbrellas are such as to warrant me in saying that the main supply of locomotor centres appears to be usually situated in that part of these organs, these effects are nevertheless such as to compel me at the same time to conclude that the locomotor centres of the covered-eyed Medus? are more diffused or segregated than are those of the naked-eyed Medus?. Lastly, it should be stated that all the species of covered-eyed Medus? resemble all the species of naked-eyed Medus?, in that their members will endure any amount of section it is possible to make upon any of their parts other than their margins without their spontaneity being in the smallest degree affected.

Effects of excising Certain Portions of the Margins of Nectocalyces.

The next question which naturally presents itself is as to whether the locomotor centres are equally distributed all round the margin of a swimming organ, or situated only, or chiefly, in the so-called marginal bodies. To take the case of the naked-eyed Medus? first, it is evident that in most of the genera, in consequence of the intertentacular spaces being so small, it is impossible to cut out the marginal bodies (which are situated at the bases of the tentacles) without at the same time cutting out the intervening portions of the margin. The genus Sarsia, however, is admirably adapted (as a glance at Fig. 1 will show) for trying the effects of removing the marginal bodies without injuring the rest of the margin, and vice versa. The results of such experiments upon members of this genus are as follow.

Whatever be the condition of the individual operated upon as to freshness, vigour, etc., it endures excision of three of its marginal bodies without suffering any apparent detriment; but in most cases, as soon as the last marginal body is cut out, the animal falls to the bottom of the water quite motionless. If the subject of the experiment happens to be a weakly specimen, it will, perhaps, never move again: it has been killed by something very much resembling nervous shock. On the other hand, if the specimen operated upon be one which is in a fresh and vigorous state, its period of quiescence will probably be but short; the nervous shock, if we may so term it, although evidently considerable at the time, soon passes away, and the animal resumes its motions as before. In the great majority of cases, however, the activity of these motions is conspicuously diminished.

The effect of excising all the marginal tissue from between the marginal bodies and leaving the latter untouched, is not so definite as is the effect of the converse experiment just described. Moreover, allowance must here be made for the fact that in this experiment the principal portion of the "veil"[7] is of necessity removed, so that it becomes impossible to decide how much of the enfeebling effect of the section is due to the removal of locomotor centres from the swimming-bell, and how much to a change in the merely mechanical conditions of the organ. From the fact, however, that excision of the entire margin of Sarsia produces total paralysis, while excision of the marginal bodies alone produces merely partial paralysis, there can be no doubt that both causes are combined. Indeed, it has been a matter of the greatest surprise to me how very minute a portion of the intertentacular marginal tissue is sufficient, in case of this genus, to animate the entire swimming-bell. Choosing vigorous specimens of Sarsia, I have tried, by cutting out all the margin besides, to ascertain how minute a portion of intertentacular tissue is sufficient to perform this function, and I find that this portion may be so small as to be quite invisible without the aid of a lens.

From numerous observations, then, upon Sarsia, I conclude that in this genus (and so, from analogy, probably in all the other genera of the true Medus?) locomotor centres are situated in every part of the extreme margin of a nectocalyx, but that there is a greater supply of such centres in the marginal bodies than elsewhere.

Effects of excising Certain Portions of the Margin of Umbrellas.

Coming now to the covered-eyed Medus?, I find that the concentration of the locomotor centres of the margin into the marginal bodies, or lithocysts, is still more decided than it is in the case of Sarsia. Taking Aurelia aurita as a type of the group, I cannot say that, either by excising the lithocysts alone or by leaving the lithocysts in situ and excising all the rest of the marginal tissue, I have ever detected the slightest indications of locomotor centres being present in any part of the margin of the umbrella other than the eight lithocysts; so that all the remarks previously made upon this species, while we were dealing with the effects of excising the entire margin of umbrellas, are equally applicable to the experiment we are now considering, viz. that of excising the lithocysts alone. In other words, but for the sake of symmetry, I might as well have stated at the first that in the case of the covered-eyed Medus? all the remarkable paralyzing effects which are obtained by excising the entire margin of an umbrella are obtained in exactly the same degree by excising the eight lithocysts alone; the intermediate marginal tissue, in the case of these Medus?, is totally destitute of locomotor centres.

Effects upon the Manubrium of excising the Margin of a Nectocalyx or Umbrella.

Lastly, it must now be stated, and always borne in mind, that neither in the case of naked nor covered-eyed Medus? does excision of the margin of a swimming organ produce the smallest effect upon the manubrium. For hours and days after the former, in consequence of this operation, has ceased to move, the latter continues to perform whatever movements are characteristic of it in the unmutilated organism-indeed, these movements are not at all interfered with even by a complete severance of the manubrium from the rest of the animal. In many of the experiments subsequently to be detailed, therefore, I began by removing the manubrium, in order to afford better facilities for manipulation.

Summary of Chapter.

With a single exception to hundreds of observations upon six widely divergent genera of naked-eyed Medus?, I find it to be uniformly true that removal of the extreme periphery of the animal causes instantaneous, complete, and permanent paralysis of the locomotor system. In the genus Sarsia, my observations point very decidedly to the conclusion that the principal locomotor centres are the marginal bodies, but that, nevertheless, every microscopical portion of the intertentacular spaces of the margin is likewise endowed with the property of originating locomotor impulses.

In the covered-eyed division of the Medus?, I find that the principal seat of spontaneity is the margin, but that the latter is not, as in the naked-eyed Medus?, the exclusive seat of spontaneity. Although in the vast majority of cases I have found that excision of the margin impairs or destroys the spontaneity of the animal for a time, I have also found that the paralysis so produced is very seldom of a permanent nature. After a variable period occasional contractions are usually given, or, in some cases, the contractions may be resumed with but little apparent detriment. Considerable differences, however, in these respects are manifested by different species, and also by different individuals of the same species. Hence, in comparing the covered-eyed group as a whole with the naked-eyed group as a whole, so far as my observations extend, I should say that the former resembles the latter in that its representatives usually have their main supply of locomotor centres situated in their margins, but that it differs from the latter in that its representatives usually have a greater or less supply of their locomotor centres scattered through the general contractile tissue of their swimming organs. But although the locomotor centres of a covered-eyed Medusa are thus, generally speaking, more diffused than are those of a naked-eyed Medusa, if we consider the organism as a whole, the locomotor centres in the margin of a covered-eyed Medusa are less diffused than are those in the margin of a naked-eyed Medusa. In no case does the excision of the margin of a swimming organ produce any effect upon the movements of the manubrium.

Mechanical, Chemical, and Thermal Stimulation.

So far as my observations extend, I find that all Medus?, after removal of their locomotor centres, invariably respond to every kind of stimulation. To take the case of Sarsia as a type, nothing can possibly be more definite than is the single sharp contraction of the mutilated nectocalyx in response to every nip with the forceps. The contraction is precisely similar to the ordinary ones that are performed by the unmutilated animal; so that by repeating the stimulus a number of times, the nectocalyx, with its centres of spontaneity removed, may be made to progress by a succession of contractions round and round the vessel in which it is contained, just as a frog, with its cerebral hemispheres removed, may be made to hop along the table in response to a succession of stimulations.[8]

Different species of Medus? exhibit different degrees of irritability in responding to stimuli; but in all the cases I have met with the degree of irritability is remarkably high. Thus, I have seen responsive contractions of the whole umbrella follow upon the exceedingly slight stimulus caused by a single drop of sea-water let fall upon the irritable surface from the height of one inch. As regards chemical stimulation, dilute spirit or other irritant, when dropped on the paralyzed swimming organ of Aurelia aurita, often gives rise to a whole series of rhythmical pulsations, the systoles and diastoles following one another at about the same rate as is observable in the normal swimming motions of the unmutilated animal.

It is somewhat difficult, in the case of paralyzed swimming organs, to prove the occurrence of a contraction in response to thermal stimulation, from the fact that while these tissues are not nearly so sensitive to this mode of excitation as might be anticipated, they are, as just observed, extraordinarily sensitive to mechanical excitation. It therefore becomes difficult to administer the appropriate thermal stimulus without at the same time causing a sufficient mechanical disturbance to render it doubtful to which of the stimuli the response is due. This may be done, however, by allowing a few drops of heated sea-water to run over the excitable surface while it is exposed to the air. In this and in other ways I have satisfied myself that the paralyzed tissues of swimming organs respond to sudden elevations of temperature.

Luminous Stimulation.

It is interesting to note that, in the case of some of the naked-eyed Medus?, the action of light as a stimulus is most marked and unfailing. In the case of Sarsia, for instance, a flash of light let fall upon a living specimen almost invariably causes it to respond with one or more contractions. If the animal is vigorous and swimming freely in water, the effect of a momentary flash thrown upon it during one of the natural pauses is immediately to originate a bout of swimming. But if the animal is non-vigorous, or if it be removed from the water and spread flat upon an object-glass, it usually gives only one contraction in response to every flash. There can thus be no doubt that a sudden transition from darkness to light acts upon Sarsia as a stimulus, and this even though the transition be but of momentary duration. The question therefore arises as to whether the stimulus consists in the presence of light, or in the occurrence of the sudden transition from darkness to light and from light to darkness. To answer this question, I tried the converse experiment of placing a vigorous specimen in sunlight, waiting till the middle of one of the quiescent stages in the swimming motions had come on, and then suddenly darkening. In no case, however, under these circumstances, did I obtain any response; so that I cannot doubt it is the light per se, and not the sudden nature of the transition from darkness to light, which in the former experiment acted as the stimulus. Indeed, the effect of the converse experiment just described is rather that of inhibiting contractions; for, if the sunlight be suddenly shut off during the occurrence of a swimming bout, it frequently happens that the quiescent stage immediately sets in. Again, in a general way, it is observable that Sarsi? are more active in the light than they are in the dark, the comparative duration of the quiescent stages being less in the former than in the latter case. Light thus appears to act towards these animals as a constant stimulus. Lastly, it may be stated that when the marginal bodies of Sarsia are removed, the swimming-bell, although still able to contract spontaneously, no longer responds to luminous stimulation of any kind or degree. But if only one body be left in situ, or if the severed margin alone be experimented upon, the same unfailing response may be obtained to luminous stimulation as that which is obtained from the entire animal.

The fact last mentioned indicates that the marginal bodies are organs of special sense, adapted to respond to luminous stimulation; or, in more simple words, that they perform the functions of sight. Now it has long been thought more or less probable that these marginal bodies are rudimentary or incipient "eyes," but hitherto the supposition has not been tested by experiment, and was therefore of no more value than a guess.[9] The guess in this instance, however, happens to have been correct, as the results of the following experiments will show.

Having put two or three hundred Sarsi? into a large bell-jar, I completely shut out the daylight from the room in which the jar was placed. By means of a dark lantern and a concentrating lens, I then cast a beam of light through the water in which the Sarsi? were swimming. The effect upon the latter was most decided. From all parts of the bell-jar they crowded into the path of the beam, and were most numerous at that side of the jar which was nearest to the light. Indeed, close against the glass they formed an almost solid mass, which followed the light wherever it was moved. The individuals composing this mass dashed themselves against the glass nearest the light with a vigour and determination closely resembling the behaviour of moths under similar circumstances. There can thus be no doubt about Sarsia possessing a visual sense.

The method of ascertaining whether this sense is lodged in the marginal bodies was, of course, extremely simple. Choosing a dozen of the most vigorous specimens, I removed all the marginal bodies from nine, and placed these, together with the three unmutilated ones, in another bell-jar. After a few minutes the mutilated animals recovered from their nervous shock, and began to swim about with tolerable vigour. I now darkened the room, and threw the concentrated beam of light into the water as before. The difference in the behaviour of the mutilated and of the unmutilated specimens was very marked. The three individuals which still had their marginal bodies sought the light as before, while the nine without their marginal bodies swam hither and thither, without paying it any regard.

A further question, however, still remained to be determined. The pigment spot of the marginal body in Medus? is, as L. Agassiz observed, placed in front of the presumably nervous tissue, and for this reason he naturally enough suggested that if the marginal body has a visual function to perform, the probability is that the rays by which the organ is affected are the heat-rays lying beyond the range of the visible spectrum. Accordingly I brought a heated iron, just ceasing to be red, close against the large bell-jar which contained the numerous specimens of Sarsia; but not one of the latter approached the heated metal.

From these observations, therefore, I conclude that in Sarsia the faculty of appreciating luminous rays is present, and that this faculty is lodged exclusively in the marginal bodies; while from observations conducted on the covered-eyed Medus?, I have come to the same conclusion respecting them. But although I have tested many species of naked-eyed Medus? besides Sarsia, I have obtained indications of response to luminous stimulation only in the case of one other. This is a species which I have called Tiaropsis polydiademata, and the response which it gives to luminous stimulation is even more marked and decided than that which is given by Sarsia; for a sudden exposure to sunlight causes this animal to go into a kind of tonic spasm, the whole of the nectocalyx being drawn together in a manner resembling cramp. Now, in one remarkable particular this response to luminous stimulation on the part of Tiaropsis polydiademata differs from that given by Sarsia tubulosa; and the difference consists in the fact that, while with Sarsia the period of latency (i.e. the time between the fall of the stimulus and the occurrence of the response) is, so far as the eye can judge, as instantaneous in the case of response to luminous stimulation as it is in the case of response to any other kind of stimulation, such is far from being true with Tiaropsis polydiademata. The period of latency in the last-named species is, so far as the eye can judge, quite as instantaneous as it is in the case of Sarsia, when the stimulus employed is other than luminous; but in response to light, the characteristic spasm does not take place till slightly more than a second has elapsed after the first occurrence of the stimulus. As this extraordinary difference in the latent period exhibited by the same animal towards different kinds of stimuli appeared to me a matter of considerable interest, I was led to reflect upon the probable cause of the difference. It occurred to me that the only respect in which luminous stimulation of the Medus? differed from all the other modes of stimulation I had employed consisted in this-that, as proved by my previous experiments on Sarsia, which I repeated on Tiaropsis, luminous stimulation directly affected the ganglionic tissues. Now, as in Tiaropsis polydiademata luminous stimulation differed from all the other modes of stimulation in giving rise to an immensely longer period of latency, I seemed here to have an index of the difference between the rapidity of the response to stimuli by the contractile and by the ganglionic tissues respectively. The next question, therefore, which presented itself was as to whether the enormous length of time occupied by the process of stimulation in the ganglia was due to any necessity on the part of the latter to accumulate the stimulating influence prior to originating a discharge, or to an immensely lengthened period of latent stimulation manifested by the ganglia under the influence of light.[10] This is an interesting question, because if such a lengthened period of latent stimulation occurs in this case, it would stand in curious antithesis to the very short period of latent stimulation manifested by the contractile tissues of the same animal under other modes of irritation. To test these alternative hypotheses, I employed the very simple method of first allowing a continuous flood of light to fall suddenly on the Medusa, and then noting the time at which the responsive spasm first began. This time, as already stated, was slightly more than one second. I next allowed the animal to remain for a few minutes in the dark to recover shock, and, lastly, proceeded to throw in single flashes of light of measured duration. I found that unless the flash of light was of slightly more than one second in its duration, no response was given; that is to say, the minimal duration of a flash required to produce a responsive spasm was just the same as the time during which a continuous flood of light required to operate in order to produce a similar spasm. From this, therefore, I conclude that the enormously long period of latent excitation in response to luminous stimuli was not, properly speaking, a period of latent excitation at all; but that it represented the time during which a certain summation of stimulating influence was taking place in the ganglia, which required somewhat more than a second to accumulate, and which then caused the ganglia to originate an abnormally powerful discharge. So that in the action of light upon the ganglionic matter of this Medusa we have some analogy to its action on certain chemical compounds in this respect, that, just as in the case of those compounds which light is able to split up, a more or less lengthened exposure to its influence is necessary in order to admit of the summating influence of its vibrations on the molecules, so in the ease of this ganglionic material, the decomposition which is effected in it by light, and which terminates in an explosion of nervous energy, can only be effected by a prolonged exposure of the unstable material to the summating influence of the luminous vibrations. Probably, therefore, we have here the most rudimentary type of a visual organ that is possible; for it is evident that if the ganglionic matter were a very little more stable than it is, it would altogether fail to be thrown down by the luminous vibrations, or would occupy so long a time in the process that the visual sense would be of no use to its possessor. How great is the contrast between the excitability of such a sense-organ and that of a fully evolved eye, which is able to effect the needful molecular changes in response to a flash as instantaneous as that of lightning.

With regard to luminous stimulation, it is only necessary further to observe that responses were given equally well to direct sunlight, diffused daylight, and to light reflected from a mirror inclined at the polarizing angle. It must also be stated that responses are given to any of the luminous rays of the spectrum when these are employed separately; but that neither the non-luminous rays beyond the red, nor those beyond the violet, appear to exert the smallest degree of stimulating influence.

Electrical Stimulation.

All the excitable parts of all the Medus? which I have examined are highly sensitive to electrical stimulation, both of the constant and of the induced current.

Exploration with needle-point terminals and induction shocks of graduated strength showed that certain parts or tracts of the nectocalyx are more sensitive than others. The most sensitive parts are those which correspond with the distribution of the main nerve-trunks, i.e. round the margin of the nectocalyx and along the course of the radial tubes. The external or convex surface of a nectocalyx or umbrella is totally insensitive to stimulation, and the same statement applies to the whole thickness of the gelatinous substance to which the neuro-muscular sheet is attached.

In all other respects the excitable tissues of the Medus? in their behaviour towards electrical stimulation conform to the rules which are followed by excitable tissues of other animals. Thus, closure of the constant current acts as a stronger stimulus than does opening of the same, while the reverse is true of the induction shock; and exhaustion supervenes under the influence of prolonged excitation. Moreover, I have obtained evidence of that polarization of nerve-tissues under the influence of the constant current, which is known to physiologists by the term "electrotonus;" but it would be somewhat tedious to detail the evidence on this head which I have already published elsewhere.[11] Tetanus produced by faradaic electricity is not of the nature of an apparently single and prolonged contraction, but that of a number of contractions rapidly succeeding one another, as in the case of the heart under similar excitation. This at least applies to Sarsia. In the case of Aurelia, tolerably strong faradization does cause a more or less well-pronounced tetanus. The continuity of the spasm is, indeed, often interrupted by momentary and partial relaxations. These interruptions are the more frequent the weaker the current; so that, at a certain strength of the latter, the tetanus is of a wild and tumultuous nature; but with strong currents the spasm is tolerably uniform. That in all cases the tetanus is due to summation of contractions may be very prettily shown by the following experiment. An Aurelia is cut into a spiral strip, and all its lithocysts are removed; single induction-shocks are then thrown in with a key at one end of the strip-every shock, of course, giving rise to a contraction wave. If these shocks are thrown in at a somewhat fast rate, two contraction waves may be made at the same time to course along the spiral strip, one behind the other; but if the shocks are thrown in at a still faster rate, so as to diminish the distance between any two successive waves, a point soon arrives at which every wave mounts upon its predecessor; and if several waves be thus made to coalesce, the whole strip becomes thrown into a state of persistent contraction.

In this way sustained tetanus, or single contraction waves, or any intermediate phase, may be instantly produced at pleasure. In such experiments, moreover, it is interesting to observe that, no matter how long the strip be, whatever disturbances are set up at one end are faithfully transmitted to the other. For instance, if an Aurelia be cut into the longest possible strip with a remnant of the disk left attached at one end, as represented in Fig. 11 (p. 70), then all the peculiar time relations between successive contractions which are intentionally caused by the experimenter at one end of the strip, are afterwards accurately reproduced at the other end of the strip by the remainder of the disk. Now, as this fact is observable however complex these time relations may be, and however rapidly the successive stimuli are thrown in, I think it is a point of some interest that these complicated relations among rapidly succeeding stimuli do not become blended during their passage along the thirty or forty inches of contractile tissue. The fact, of course, shows that the rate of transmission is so identical in the case of all the stimuli originated, that the sum of the effects of any series of stimuli is delivered at the distal end of the strip, with all its constituent parts as distinct from one another as they were at starting from the proximal end of the strip.

Period of Latency, and Summation of Stimuli.

I shall now give an account of my experiments in the period of latency and the summation of stimuli. To do this, I must first describe the method which I adopted in order to obtain a graphic record of the movements which were given in response to the stimuli supplied. As Aurelia aurita is the only species on which I have experimented in this connection, my remarks under this heading will be confined to it alone.

The method by which I determined the latent period in the case of this species was as follows. A basin containing the Medusa was filled to its brim with sea-water, and placed close beside a smoked cylinder, which, while it lay in a horizontal position, could be rotated at a known rate. The Aurelia[12] was placed with its concave aspect uppermost, and an inch or two below the surface of the water. The animal was held firmly in this position by means of a pair of compasses thrust through it and forced into a piece of wood, which was fastened to the bottom of the basin. The legs of the compasses were provided with india-rubber sliders, so that by placing these under the Medusa, the latter might be kept at any elevation in the water which might be desired. The manubrium and lithocysts were now removed, and also a segment of the umbrella. A light straw was then forced through the gelatinous substance of the umbrella in a radial direction, and close to the gap caused by the missing segment. The other, or free, end of this straw was firmly joined to a capillary glass rod, which was suitably bent to avoid contact with the rim of the basin, and also to write on the smoked cylinder. If the straw was not itself sufficient to support the weight of the capillary rod, a small cross-piece of cork might easily be tied to it, so as to add to the flotation power. A part of the excitable tissue was now raised above the surface of the water by means of a disk of cork placed beneath it, and on the part of the tissue thus raised there were placed a pair of platinum electrodes. These electrodes proceeded from an electro-magnetic apparatus, which was arranged in such a way that every time the current in it was opened or closed, it gave an induction shock and moved a lever at the same instant of time. This lever was therefore placed upon the cylinder immediately above the capillary glass-writer which proceeded from the Medusa, care being taken to place the two writers in the same line, parallel to the axis of the cylinder. Such being the arrangement, the cylinder was rotated, and thus two parallel lines were marked upon it by the two writers. If the current was now closed, an induction shock was thrown into the tissue at the same instant that the electro-magnet writer recorded the fact, by altering its position on the cylinder. Again, as soon as the paralyzed Medusa responded to the induction shock, the radii of the vacant segment were drawn apart, and in this way a curve was obtained by the other writer on the rotating cylinder. Now, by afterwards dropping a perpendicular line from the point at which the electro-magnet writer changed its position, to the parallel line made by the other writer, and then measuring the distance between the point of contact and the point on the last-mentioned line on which the curve began, the period of latent stimulation was determined. A glance at Figs. 3 and 4 (p. 55) will render this description clear to any one who is not already acquainted with the method, when it is stated that the upper line is a record of the movements of the electro-magnet writer, and the lower line that of the movements of the other writer. It will be observed that the point a in the upper line marks the point at which the induction shock was thrown in; so that by first producing the perpendicular till it meets the lower line at b, and then measuring the distance between the point b and the point c, at which the curve in the lower line first begins, the latent period (b c) is determined-the time occupied by the rotation of the cylinder from b to c being known.

Summation of Stimuli.-In this way I have been able to ascertain the period of latent stimulation in Aurelia aurita with accuracy. It must be stated at the outset, however, that the period is subject to great variations under certain varying conditions, so that we can only arrive at a just estimation of it by understanding the nature of the modifying causes. To take the simplest cause first, suppose that the paralyzed Aurelia has been left quiet for several minutes in sea-water at forty-five degrees, and that it is then stimulated by means of a single induction shock; the responsive contraction will be comparatively feeble with a very long period of latency, viz. five-eighths of a second. If another shock of the same intensity be thrown in as soon as the tissue has relaxed, a somewhat stronger contraction, with a somewhat shorter latent period, will be given. If the process is again repeated, the response will be still more powerful, with a still shorter period of latency; and so on, perhaps, for eight or ten stages, when the maximum force of contraction of which the tissue is capable will have been attained, while the period of latency will have been reduced to its minimum. This period is three-eighths of a second, or, in some cases, slightly less.

Now, we have here a remarkable series of phenomena, and as it is a series which never fails to occur under the conditions named, I append tracings to give a better idea of the very marked and striking character of the results. The first tracing (Fig. 2) is a record of the successive increments of the responses to successive induction shocks of the same intensity, thrown in at three seconds' intervals-the cylinder being stationary during each response, and rotated a short distance with the hand during each interval of repose.

Fig. 2.

The second tracing (Figs. 3 and 4) is a record of the difference between the lengths of the latent period, and also between the strengths of the contraction, in the case (a) of the first of such a series of responses (Fig. 3), and (b) of the last of such a series (Fig. 4). From these tracings it will be manifest, without further comment, how surprising is the effect of a series of stimuli; first, in arousing the tissue, as it were, to increased activity, and, second, in developing a state of expectancy.

In accordance with the now customary terminology, I shall call such a series of responses as are given in Fig. 2 a "staircase." Such a staircase has a greater number of steps in it if caused by a weak current (compare Figs. 2 and 5); and if the strength of the current be suddenly increased after the maximum level of a staircase has been reached by using a feeble current, this level admits of being slightly raised (see Fig. 5). Moreover, I find that a stimulus, which at the bottom of a staircase is of less than minimal intensity, is able, at the top of a staircase, to give rise to a contraction of very nearly maximum intensity. That is to say, by employing an induction stimulus of slightly less than minimal intensity in relation to the original irritability of the tissue, no response is given to the first two or three shocks of a series; but at the third or fourth shock a slight response is given, and from that point onward the staircase is built up as usual. This was the case in the experiment of which Fig. 2 is a record, no response having been given to the first two shocks.

Fig. 3.

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Fig. 4.

Fig. 5.

With regard to this interesting staircase action, two questions naturally present themselves. In the first place, we are anxious to know whether the arousing effect which is so conspicuous in a staircase series is due to the occurrence of the previous stimulations, or to that of previous contractions; and, in the next place, we should like to know whether, during the natural rhythm of the tissue, each contraction exerts a beneficial influence on its successor, analogous to that which occurs in the case of contractions which are due to artificial stimuli. To answer the first of these questions, therefore, I built up a staircase in the ordinary way, and then suddenly transferred the electrodes to the opposite side of the umbrella from that on which they rested while constructing the staircase. On now throwing in another shock at this part of the contractile tissue so remote from the part previously stimulated, the response was a maximum response. Similarly, if the electrodes were transferred in the way just described, not after the maximum effect had been attained, but at any point during the process of constructing a staircase, the response given to the next shock was of an intensity to make it rank as the next step in the staircase. Hence, shifting the position of the electrodes in no wise modifies the peculiar effect we are considering; and this fact conclusively proves that the effect is a general one, pervading the whole mass of the contractile tissue, and not confined to the locality which is the immediate seat of stimulation. Nevertheless, this fact does not tend to prove that the staircase effect depends on the process of contraction as distinguished from the process of stimulation, because the wave of the former process must always precede that of the latter. But, on the other hand, in this connection it is of the first importance to remember the fact already stated, viz. that a current which at the beginning of a series of stimulations is of slightly less than minimal intensity presently becomes minimal, and eventually of much more than minimal intensity-a staircase being thus built up of which the first observable step (or contraction) only occurs in response to the second, third, or even fourth shock of the series. This fact conclusively proves that the staircase effect, at any rate at its commencement, depends on the process of stimulation as distinguished from that of contraction; for it is obvious that the latter process cannot play any part in thus constructing what we may term the invisible steps of a staircase.

To answer the second of the above questions, I placed an Aurelia with its concave surface uppermost, and removed seven of its lithocysts; I then observed the spontaneous discharge of the remaining one, and found it to be conspicuous enough that, after the occurrence of one of the natural pauses (if this were of sufficient duration), the first contraction was feeble, the next stronger, the next still stronger, and so on, till the maximum was attained. This natural staircase action admits of being very prettily shown in another way. If a tolerably large Aurelia is cut into a spiral strip of small width and great length, and if all the lithocysts are removed except one at one end of the strip, it may be observed that, after the occurrence of a natural pause, the first discharge only penetrates perhaps about a quarter of the length of the strip, the next discharge penetrates a little further, the next further, and so on, till finally the contraction waves pass from end to end. On now removing the ganglion, waiting a few minutes, and then stimulating with successive induction shocks, the same progressive penetration is observable as that which previously took place with the ganglionic stimulation. Lastly, the identity of natural and artificial staircase action may be placed beyond all doubt by an experiment in which the effects of induction shocks and of ganglionic discharges are combined. To accomplish this, all the lithocysts save one are removed, and a staircase is then built up in the ordinary way by successive induction shocks. It will now occasionally happen that the ganglion originates a discharge during the process of constructing the staircase, which is being built up by the artificial stimuli; when this happens the resulting contraction takes its proper rank in the series, and this at whatever point the natural contraction happens to come in.

Thus, then, to summarize and conclude these observations, we have seen that if a single stimulation, whether of a natural or artificial kind, is supplied to the excitable tissues of a jelly-fish, a short period, called the period of latency, will elapse, and then the jelly-fish will give a single weak contraction. If, as soon as the tissue has relaxed, the stimulation is again repeated, the period of latency will be somewhat shorter, and will be followed by a somewhat stronger contraction. Similarly, if the stimulation is repeated a third time, the period of latency will be still shorter, and the ensuing contraction still stronger. And so on up to nine or ten times, when the period of latency will be reduced to its minimum, while the force of the contraction will be raised to its maximum; so that in the jelly-fish, the effect of a series of excitations supplied at short intervals from one another is that of both arousing the tissue into a state of increased activity, and also of producing in it a state of greater expectancy. We have, moreover, seen that this effect depends upon the repetition of the process of stimulation, and not upon that of the process of contraction.

Now, effects very similar to these have been found to occur in the case of the excitable plants by Dr. Burdon-Sanderson; in the case of the frog's heart by Dr. Bowditch; and in the case of reflex action of the spinal cord by Dr. Stirling. Indeed, the only difference in this respect between these four tissues, so widely separated from one another in the biological scale, consists in the time which may be allowed to elapse between the occurrence of the successive stimuli, in order to produce this so-called summating effect of one stimulus upon its successor: the memory, so to speak, of the heart-tissue for the occurrence of a former stimulus being longer than the memory of the jelly-fish tissue; while the memory of the latter is longer than that of the plant tissue. And I may here add that even in our own organization we may often observe the action of this principle of the summation of stimuli. For instance, we can tolerate for a time the irritation caused by a crumb in the larynx, but very rapidly the sense of irritation accumulates to a point at which it becomes impossible to avoid coughing. And similarly with tickling generally, the convulsive reflex movements to which it gives rise become more and more incontrollable the longer the stimulation is continued, until they reach a maximum point, where, in persons susceptible to this kind of stimulation, the muscular action passes completely beyond the power of the will. Lastly, I may further observe, what I do not think has ever been observed before, that even in the domain of psychology the action of this principle admits of being clearly traced. We find it, for instance, in the rhythmical waves of emotion characteristic of grief, and at the other extreme we find it in the case of the ludicrous. We can endure for a short time, without giving any visible response, the psychological stimulation which is supplied by a comical spectacle; but if the latter continues sufficiently long in a sufficiently ludicrous manner, our appropriate emotion rapidly runs up to a point at which it becomes uncontrollable, and we burst into an explosion of ill-timed laughter. But in this case of psychological tickling, as in the previous case of physiological tickling, some persons are much more susceptible than others. Nevertheless, there can be no doubt that from the excitable tissues of a plant, through those of a jelly-fish and a frog, up even to the most complex of our psychological processes, we have in this recently discovered principle of the summation of stimuli a very remarkable uniformity of occurrence.

Effects of Temperature on Excitability.

I shall now conclude this chapter with a brief statement of the effects of temperature on the excitability of the Medus?; and before stating my results, I may observe that in all my experiments in this connection I changed the temperature of the Medus? by drawing off the water in which they floated with a siphon, while at the same time I substituted water of a different temperature from that which I thus abstracted. In this way, without modifying any of the other conditions to which the animals were exposed, I was able to observe the effects of changing the temperature alone.

With regard to the effect of temperature on the latent period of stimulation, the following table, setting forth the results of one among several experiments, explains itself.

Period of latent stimulation of the deganglionated tissues of Aurelia aurita as affected by temperature:-

Temperature of water (Fahr.). Period of latent stimulation.

70° 1/5 second

50° 1/3 second

35° 2/5 second

20° 1/2 second

In the case of each observation, several shocks were administered before the latent period was taken, in order to decrease this period to its minimum by the staircase action. When this is not done, the latent period at 20° may be as long as 1-1/5 seconds; but soon after this irritability disappears.

The extraordinary sluggishness of the latent period at very low temperatures is fully equalled by the no less extraordinary sluggishness of the contraction.

In order to render apparent the degree in which both these effects are produced, I here append a pair of tracings which were procured from the same piece of tissue when exposed to the different temperatures named. (N.B.-The seconds are wrongly marked in Fig. 7; they ought to be the same as in Fig. 6.)

Fig. 6.

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Fig. 7.

I may as well state here that in water at all temperatures, within the limits where responses to stimuli are given at all, the staircase action admits of being equally well produced; but in order to procure the maximum effect for any given temperature, the rate at which the successive stimuli are thrown in must be quicker in warm than in cold water.