The naked-eyed MedusÆ 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.

Confining our attention under this heading to the naked-eyed MedusÆ, 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 MedusÆ, 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 MedusÆ 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.

Turning now to the covered-eyed division of the MedusÆ, 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 MedusÆ, 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 MedusÆ 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 MedusÆ are more diffused or segregated than are those of the naked-eyed MedusÆ. Lastly, it should be stated that all the species of covered-eyed MedusÆ resemble all the species of naked-eyed MedusÆ, 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.

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 MedusÆ 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 versÂ. 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 MedusÆ) 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.

Coming now to the covered-eyed MedusÆ, 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 MedusÆ 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 MedusÆ, is totally destitute of locomotor centres.

Lastly, it must now be stated, and always borne in mind, that neither in the case of naked nor covered-eyed MedusÆ 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.

With a single exception to hundreds of observations upon six widely divergent genera of naked-eyed MedusÆ, 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 MedusÆ, I find that the principal seat of spontaneity is the margin, but that the latter is not, as in the naked-eyed MedusÆ, 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.