The above experiments led me to try the effects of cutting out a single lithocyst of Aurelia, and, after the rhythm of the detached segment had become regular, progressively paring down the contractile tissues around the ganglion. I found that this process had no very marked effect on the rhythm, until the paring reached within an inch or two of the ganglion: then, however, the effect began to show itself, and with every successive paring it became more marked. This effect consisted in slowing the rate of the rhythm, but more especially in giving rise to prolonged pauses: indeed, if only a very little contractile tissue was left adhering to the ganglion, the pauses often became immensely prolonged, so that one might almost suppose the ganglion to have entirely ceased discharging. But if a stimulus of any kind were then applied, the rhythmic discharges at once recommenced. These generally continued for some little time at a slower rate than that which they had manifested before they were affected by the paring down of the contractile tissue.

The effects of temperature on the rhythm of Medusæ are very decided. For instance, a specimen of Sarsia which in successive minutes gave the following number of pulsations, 16, 26, 0, 0, 26, gave sixty pulsations during the next minute, while a spirit-lamp was held under the water in which the Medusa was swimming. If hot water be added to that in which Sarsia are contained until the whole is about milk-warm, their swimming motions become frantic. If the same experiment be performed after the margins of the Sarsiæ have been removed, the paralyzed bells remain quite passive, while the severed margins exhibit the frantic motions just alluded to.

In the case of Aurelia aurita, the characteristic effects of temperature on rhythm may be better studied than in that of Sarsia, from the fact that the natural motions are more rhythmical and sustained in the former than in the latter genus. I have, therefore, in this connection made more observations on Aurelia than on Sarsia. The following may be taken as a typical experiment.

A small and active specimen of Aurelia contracted with the greatest regularity 33 times per minute in water kept at 34°; but on transference to water kept at 49°, the contractions always became irregular, in respect (a) of not having a perfectly constant rhythm, and (b) of exhibiting frequent pauses, which was never the case in colder water. The rate of rhythm in the warmer water varied from 37 to 49; and as in these observations no allowance was made for the occurrence of the pauses, the actual rate of rhythm during the swimming motions was about 60 per minute. The following are some sample observations in the case of this specimen:—

This rate continued quite regularly for a quarter of an hour, when the observation terminated.

It might naturally be supposed that when the alterations of temperature between 34° and 49° produce such marked effects on the rhythm, still greater alterations would be attended with still greater effects. Such, however, is not the case. Water at 70° or 80°, for instance, has the effect of permanently diminishing the rate of the rhythm, after having temporarily raised it for a few seconds. The following experiment will serve to convey a just estimation of these facts.

An Aurelia whose rhythm in water at 40° was very regular at eighteen per minute, was suddenly transferred to water at 80°. In the immediately succeeding minutes the rhythm was 22, 20, 14. The latter rate continued for nearly half an hour, when the observation terminated.

The effect of very warm water, therefore, is to slow the rhythm, as well, I may add, as to enfeeble the vigour of the contractions. The case of Medusæ thus differs, in the former respect, from that of the heart; and I think the reason of the difference is to be found in the following considerations. Even slight elevations of temperature are quickly fatal to the Medusæ, so it becomes presumable that considerable elevations act very destructively on the neuro-muscular tissues of those animals. This destructive effect of high temperatures may, therefore, very probably counteract the stimulating effect which such temperatures would otherwise exert on the natural rhythm, and hence a point would somewhere be reached at which the destructive effect would so far overcome the stimulating effect as to slow the rhythm. That this is probably the true, as it certainly is the only explanation to be rendered, will, I think, be conceded when I further state that if an Aurelia be left for some little time in water at 80°, and then again transferred to water at 30° or 40°, its original rate of rhythm at the latter temperature does not again return, but the rhythm remains permanently slowed. And, in favour of the explanation just offered, it may be further pointed out that the first effect of sudden immersion in heated water is to quicken the rhythm, it not being for a few seconds, or for even a minute or two after the immersion, that the rhythm becomes slowed. Lastly, the slowing takes place gradually; and this is what we should expect if, as is probable, the destructive effect takes somewhat more time to become fully developed than does the stimulating effect.

Before leaving the subject of temperature in relation to rhythm, I must say a few words on the effects of cold. The following may be regarded as typical experiments.

An Aurelia presenting a regular rhythm of twenty per minute in water at 45° was placed in water at 19°. Soon after the transference the rhythm began to slow, and the strength of the contractions to diminish. Both these phenomena rapidly became more and more pronounced, till the rhythm fell to ten per minute (still quite regular), and the contractions ceased to penetrate the muscular tissue further than an inch or so from the marginal ganglia. Shortly after this stage pauses became frequent, but mechanical or other irritation always originated a fresh swimming bout. Next, only one very feeble contraction was given at long and irregular intervals, a contraction so feeble that it was restricted to the immediate vicinity of the lithocyst in which it originated. Soon after this stage irritability towards all kinds of stimuli entirely ceased, including even strong spirit dropped on the under surface of the animal when taken momentarily out of the water. All these stages thus described were passed through rapidly, the whole series occupying rather less than five minutes. On now leaving the specimen for ten minutes and then restoring it to its original water at 45°, all the above-mentioned stages were passed through in reverse order. The first faint marginal contraction was confined to the immediate vicinity of the prepotent lithocyst, and all subsequent contractions continued to be so for the next three minutes. Rhythm very slow. Contractions now began to penetrate round the margin, and in eight minutes from the restoration had gone all the way round, the rate of their rhythm meanwhile increasing. In two minutes more all the umbrella was contracting at the rate of fifteen per minute.

In another specimen, subjected to the same conditions, the rate of recovery was even more rapid, occupying only two minutes altogether; but in every case the process of recovery is a gradual one, and differs only in the time it occupies in passing through the various stages.

In conclusion, I will describe some rather interesting experiments that consisted in freezing some specimens of Aurelia into a solid block of ice. Of course, as sea-water had to be employed, the cold required was very considerable; but I succeeded in turning out the Medusæ encased on all sides in a continuous block of sea-water. By now immersing this block in warm water, I was able to release the contained specimens, which then presented a very extraordinary appearance. The thick and massive gelatinous bell of a Medusa is, as every one knows, chiefly composed of sea-water, which everywhere enters very intimately into the structure of the tissue. Now, all this sea-water was, of course, frozen in situ, so that the animals were everywhere and in all directions pierced through by an innumerable multitude of ice-crystals, which formed a very beautiful meshwork, pervading the whole substance of their transparent tissues.

These experiments were made in order to ascertain whether the Medusæ, after having been thus completely frozen, would survive on being again thawed out, and, if so, whether the freezing process would exert any permanent influence on the rate of their rhythm. Now in all the cases the Medusæ, after having been thawed out, presented a ragged appearance, which was due to the disintegrating effect exerted by the ice-crystals while forming in the tissues; yet notwithstanding this mechanical injury superimposed on the physiological effects of such extreme cold, all the Medusæ recovered on being restored to sea-water of the normal temperature. The time occupied by the process of recovery varied in different individuals from a few minutes to half an hour or more, and it was observable that those specimens which recovered soonest had the rate of their rhythm least affected by the freezing. In no case, however, that I observed did the rate of the rhythm after the freezing return fully to that which had been manifested before the freezing.

Oxygen.—I will now conclude my remarks on rhythm by very briefly describing the effects of certain gases. Oxygen forced under pressure into sea-water containing Sarsiæ has the effect of greatly accelerating the rate of their rhythm. The following observation on a single specimen will serve to render this apparent.

Number of pulsations given by Sarsia in successive five-minute intervals.

It will be seen from this observation that the acceleration of the rhythm due to the oxygenation was most marked; indeed, the pulsations followed one another so rapidly that it was no easy matter to count them. It must also be stated that while the animal was under the influence of oxygen, the duration of the natural pauses between the swimming bouts was greatly curtailed, the swimming motions, in fact, being almost quite continuous throughout the five minutes that the Medusa was exposed to such influence. Lastly, it will be observed from the above table that the unnatural amount of activity displayed by the organism while in the oxygenated water entailed on it a considerable degree of exhaustion, as shown by the fact that even a quarter of an hour after its restoration to normal water its original degree of energy had not quite returned.

Carbonic acid.—As might be expected, this gas has the opposite effects to those of oxygen. It is therefore needless to say more about this agent, except that if administered in large doses it destroys both spontaneity and irritability. Nevertheless, if its action is not allowed to last too long, the Medusæ will fully recover on being again restored to normal sea-water.

Nitrous oxide.—This gas at first accelerates the motions of Sarsia, but eventually retards them. I omitted, however, to push the experiment to the stage of complete anæsthesia, which would doubtless have supervened had the pressure of the gas been sufficiently great.

Deficient aëration.—It may now be stated that the Medusæ are exceedingly sensitive to such slight carbonization of the water in which they are contained as results from their being confined in a limited body of it for a few hours. The rhythm becomes slowed and the contractions feeble, while the pauses between the swimming bouts become more frequent and prolonged. If the water is not changed, all these symptoms become more marked, and, in addition, the rhythm becomes very irregular. Eventually the swimming motions entirely cease; but almost immediately after the animals are restored to normal sea-water, they recover themselves completely, the rate and regularity of their rhythm being then quite natural. The suddenness with which this return to the normal state of things is effected cannot but strike the observer as very remarkable, and I may mention that it takes place with equal suddenness at whatever stage in the above-described process of asphyxiation the transference to normal sea-water is accomplished.

If the umbrella of Aurelia aurita has been paralyzed by the removal of its lithocysts, and if it is then subjected to faradaic stimulation of minimal intensity, the response which it gives is not tetanic, but rhythmic. The rate of this artificial rhythm varies in different specimens, but the limits of variation are always within those which are observed by the natural rhythm of different specimens. The artificial rhythm is not in every case strictly regular; but by carefully adjusting the strength of the current, and by shifting the electrodes from one part of the tissue to another until the most appropriate part is ascertained, the artificial rhythm admits in most cases of being rendered tolerably regular, and in many cases as strictly regular as is the natural rhythm of the animal. To show this, I append a tracing of the artificial rhythm (Fig. 25), which may be taken as a fair sample of the most perfect regularity that can be obtained by minimal faradaic stimulation.[24]

This artificial rhythm may be obtained with a portion of irritable tissue of any size, and whether a large or small piece of the tissue employed be included between the electrodes.

As the fact of this wonderfully rhythmic response to faradaic irritation was quite unexpected by me, and as it seemed to be a fact of great significance, I was led to investigate it in as many of its bearings as time permitted. First, I tried the effect on the rhythm of progressively intensifying the strength of the faradaic current. I found that with each increment of the current the rate of the rhythm was increased, and this up to the point at which the rhythm began to pass into tetanus due to summation of the successive contractions. But between the slowest rhythm obtainable by minimal stimulation and the most rapid rhythm obtainable before the appearance of tetanus, there were numerous degrees of rate to be observed. I here append another tracing, to show the effect on the rate of the rhythm of alterations in the strength of the current (Fig. 26).

It will also be observed from this tracing that, in consequence of the current having been strengthened slightly beyond the limit within which strictly rhythmic response was attainable, the curves in the middle part of the tracing, where the current was strengthened, are slightly irregular. This irregularity is, of course, due to the first appearance of tumultuous tetanus. If the faradaic stimulation had in this case been progressively made still stronger, the irregularity would have become still more pronounced up to a certain point, when it would gradually have begun to pass into more persistent tetanus. But as in this case, instead of strengthening the current still further, I again weakened it to its original intensity, the rhythm immediately returned to its original rate and regularity.

Such being the facts, the question arises as to their interpretation. At first I was naturally inclined to suppose that the artificial rhythm was due to a periodic variation in the strength of the stimulus, caused by some slight breach of contact between the terminals and the tissue on each contraction of the latter. This supposition, of course, would divest the phenomena in question of all physiological meaning, and I therefore took pains in the first instance to exclude it. This I did in two ways: first, by observing that in many cases (and especially in Cyanæa capillata) the rate of the rhythm is so slow that the contractions do not follow one another till a considerable interval of total relaxation has intervened; and second, by placing the terminals close together, so as to include only a small piece of tissue between, and then firmly pinning the tissue all round the electrodes to a piece of wood placed beneath the Medusa. In this way the small portion of tissue which served as the seat of stimulation was itself prevented from moving, and therefore the rhythmic motions which the rest of the Medusa presented cannot have been due to any variations in the quality of the contact between the electrodes and this stationary seat of stimulation.

Any such merely mechanical source of fallacy being thus, I think, excluded, we are compelled to regard the facts of artificial rhythm as of a purely physiological kind. The question, therefore, as to the explanation of these facts becomes one of the highest interest, and the hypothesis which I have framed to answer it is as follows. Every time the tissue contracts it must as a consequence suffer a certain amount of exhaustion, and therefore must become slightly less sensitive to stimulation than it was before. After a time, however, the exhaustion will pass away, and the original degree of sensitiveness will thereupon return. Now, the intensity of faradaic stimulation which is alone capable of producing rhythmic response, is either minimal or but slightly more than minimal in relation to the sensitiveness of the tissue when fresh; consequently, when this sensitiveness is somewhat lowered by temporary exhaustion, the intensity of the stimulation becomes somewhat less than minimal in relation to this lower degree of sensitiveness. The tissue, therefore, fails to perceive the presence of the stimulus, and consequently fails to respond. But so soon as the exhaustion is completely recovered from, so soon will the tissue again perceive the presence of the stimulus; it will therefore again respond, again become temporarily exhausted, again fail to perceive the presence of the stimulus, and again become temporarily quiescent. Now it is obvious that if this process occurs once, it may occur an indefinite number of times; and as the conditions of nutrition, as well as those of stimulation, remain constant, it is manifest that the responses may thus become periodic.

In order to test the truth of this hypothesis, I made the following experiments. Having first noted the rate of the rhythm under faradaic stimulation of minimal intensity, without shifting the electrodes or altering the intensity of the current, I discarded the faradaic stimulation, and substituted for it single induction shocks thrown in with a key. I found, as I had hoped, that the maximum number of these single shocks which I could thus throw in in a given time so as to procure a response to every shock, corresponded with the number of contractions which the tissue had previously given during a similar interval of time when under the influence of the faradaic current of similar intensity. To make this quite clear, I shall describe the whole course of one such experiment. The deganglionated tissue under the influence of minimal faradaic stimulation manifested a perfectly regular rhythm of thirty contractions per minute, or one contraction in every two seconds. While the position of the platinum electrodes and the intensity of the current remained unchanged, single induction shocks were now administered with a key at any intervals which might be desired. It was found that if these single induction stimuli were administered at regular intervals of two seconds or more, the tissue responded to every stimulus; while if the stimuli were thrown in more rapidly than this, the tissue did not respond to every stimulus, but only to those that were separated from one another by an interval of at least two seconds' duration. Thus, for instance, if the shocks were thrown in at the rate of one a second, the tissue only, but always, responded to every alternate shock. And similarly, as just stated, if any number of shocks were thrown in, the tissue only responded once in every two seconds. Now, as this rate of response precisely coincided with the rate of rhythm previously shown by the same tissue under the influence of faradaic stimulation of the same intensity, the experiment tended to verify the hypothesis which it was designed to test.

I may give one other experiment having the same object and tendency. Employing single induction shocks of slightly more than minimal intensity, and throwing them in at twice the rate that was required to produce a strong response to every shock, I found that midway between every two strong responses there was a weak response. In other words, a stimulus of uniform intensity gives rise alternately to a strong and to a weak contraction, as shown in the appended tracing (Fig. 27). It will be observed that in this tracing each large curve represents the whole time occupied by the strong contraction, the latter beginning at the highest point of the curve on the left-hand side in each case. The effect of the weak contraction is that of momentarily interrupting the even sweep of diastole after the strong contraction, and therefore the result on the tracing is a slight depression in the otherwise even curve of ascent. Lest any doubt should arise from the smallness of the curves representing the weak contractions that the former are in some way accidental, I may draw attention to the fact that the period of latent stimulation is the same in the case of all the curves. To render this apparent, I have placed crosses below the smaller curves, which show in each case the exact point where the depressing effect of these smaller curves on the ascending sweeps of the larger curves first become apparent—i.e. the point at which the feeble contraction begins. Now, what I wish to be gathered from the whole tracing is this. If the strength of the induction shocks had been much greater than it was, all the contractions would have become strong contractions, and tetanus would have been the result. But, as the strength of the induction shocks was only slightly more than minimal, the exhaustion consequent on every strong contraction so far diminished the irritability of the tissue that when, during the process of relaxation, another shock of the same intensity was thrown in, the stimulus was only strong enough, in relation to the diminished irritability of the partly recovered tissue, to cause a feeble contraction. And these facts tend still further to substantiate the hypothesis whereby I have sought to explain the phenomena of artificial rhythm.

Now, I think that the strictly rhythmic action of the paralyzed swimming-bell of Aurelia in answer to constant stimulation is a fact of the highest significance; for here we have a tissue wholly, or almost wholly, deprived of its centres of spontaneity, yet pulsating as rhythmically in answer to artificial stimulation as it previously did in answer to ganglionic stimulation.[25] Does not this tend to show that for the production of the natural rhythm the presence of the ganglionic element is non-essential; that if we merely suppose the function of this element to be that of supplying a constant stimulus of a low intensity, without in addition supposing the presence of any special resistance-mechanism to regulate the discharges, the periodic sequence of systole and diastole would assuredly result; and, therefore, that the rhythmical character of the natural swimming motions is dependent, not on the peculiar relations of the ganglionic, but on the primary qualities of the contractile tissue? Or, if we do not go so far as this (and, as I may parenthetically observe, I am not myself inclined to go so far), must we not at least conclude that the natural rhythm of these tissues is not exclusively due to any mechanism whereby the discharges of the ganglia are interrupted at regular intervals; but that whether these discharges are supposed to be interrupted or continuous, the natural rhythm is probably in a large measure due to the same cause as the artificial rhythm, viz. in accordance with our previous hypothesis, to the alternate exhaustion and recovery of the excitable tissues? This much, at least, must be allowed even by the most cautious of critics, viz. that if, as current views respecting the theory of rhythm would suppose, it is exclusively the ganglionic element which in the unmutilated Aurelia causes the rhythm of the swimming motions by intermittent stimulation, surely it becomes a most unexpected and unaccountable fact, that after the removal of this element the contractile tissues should still persist in their display of rhythm under the influence of constant stimulation. At any rate no one, I think, will dispute that the facts which I have adduced justify us in reconsidering the whole theory of rhythm as due to ganglia.

As I have already said, I am not inclined to deny that there is probably some truth in the current theory of rhythm as due to ganglia; I merely wish to point out distinctly that this theory is inadequate, and that in order to cover all the facts it will require to be supplemented by the theory which I now propose. The current theory of rhythm as due to ganglia attributes the whole of the effect to the ganglionic element, and thus fails to meet the fact of a rhythm which is artificially produced after the ganglionic element has been removed. It also fails to meet a number of other facts of the first importance; for it is beyond all doubt that rhythmic action of the strictest kind occurs in an innumerable multitude of cases where it is quite impossible to suppose anything resembling ganglia to be present. Not to mention such cases as the Snail's heart, where the most careful scrutiny has failed to detect the least vestige of ganglia, but to descend at once to the lowest forms of animal and vegetable life, rhythmic action may here be said to be the rule rather than the exception. The beautifully regular motions observable in some Algæ, Diatomaceæ, and Ocillatoriæ, in countless numbers of Infusoria, Antherozoids, and Spermatozoa, in ciliary action, and even in the petioles of Hedysarum gyrano, are all instances (to which many others might be added) of rhythmical action where the presence of ganglia is out of the question. Again, in a general way, is it not just as we recede from these primitive forms of contractile tissue that we find rhythmic action to become less usual? And, if this is so, may it not be that those contractile tissues which in the higher animals manifest rhythmic action are the contractile tissues which have longest retained their primitive endowment of rhythmicality? To my mind it seems hard to decide in what respect the beating of a Snail's heart differs from that of the pulsatile vesicles of the Infusoria; and I do not think it would be much easier to decide in what essential respect it differs from the beating of the Mammalian heart. The mere fact that the presence of ganglia can be proved in the one case and not in the other, seems to me scarcely to justify the conclusion that the rhythm is in the one case wholly dependent, and in the other as wholly independent, of the ganglia. At any rate, this fact, if it is a fact, is not of so self-evident a character as to recommend to us the current theory of ganglionic action on à priori grounds.

Coming, then, to experimental tests, we have already seen that in the deganglionated swimming organ of Aurelia aurita, rhythmic response is yielded to constant faradaic stimulation of low intensity. The next question, therefore, which presents itself in relation to our subject is as to whether other modes of constant stimulation elicit a similar response. Now, in a general way, I may say that such is the case, although I have chosen faradaic stimulation for special mention, because, in the first place, its effect in producing rhythmic action is the most certain and precise; and, in the next place, the effects of administering instantaneous shocks at given intervals admit of being compared with the effects of constant faradaic stimulation better than with any other kind of constant stimulation. Nevertheless, as just stated, other modes of constant stimulation certainly have a more or less marked effect in producing rhythmic response. The constant current, during the whole time of its passage, frequently has this effect in the case of the paralyzed nectocalyx of Sarsia; and 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.

From this it will be seen that, both in the case of mechanical and of chemical stimulation, the same tendency to the production of rhythmic response on the part of the paralyzed tissues of Aurelia may be observed as in the case of electrical stimulation. The principal differences consist in the rhythm being much less sustained in the former than in the latter case. But, by experimenting on other species of Medusæ, I have been able to obtain, in response to mechanical and chemical stimulation, artificial rhythm of a much more sustained character than that which, under such modes of stimulation, occurs in Aurelia. I have no explanation to offer why it is that some species, or some tissues, present so much more readiness to manifest sustained rhythm under certain modes of stimulation, and less readiness to manifest it under other modes, than do other species or tissues. Probably these differences depend on some peculiarities in the irritability of the tissues which it is hopeless to ascertain; but, in any case, the facts remain, that while Aurelia, Cyanæa, and the covered-eyed Medusæ generally are the best species for obtaining artificial rhythm under the influence of faradaic stimulation, some of the naked-eyed Medusæ are the best species for obtaining it under the influence of the constant current, and also under that of mechanical and chemical stimulation. I have already spoken of this effect of the constant current in the case of Sarsia; I shall now proceed to describe the effects of mechanical and chemical stimulation on the same species.

It is but rarely that artificial rhythm can be produced in the paralyzed nectocalyx of Sarsia by means of mechanical stimulation, but in the case of the manubrium, a very decided, peculiar, and persistent rhythm admits of being produced by this means. In this particular species, the manubrium never exhibits any spontaneous motion after the ganglia of the nectocalyx have been removed. But if it be nipped with the forceps, or otherwise irritated, it contracts strongly and suddenly; it then very slowly and gradually relaxes until it has regained its original length. After a considerable interval, and without the application of any additional stimulus, it gives another single, sudden, though slight contraction, to be again followed by gradual relaxation and a prolonged interval of repose, which is followed in turn by another contraction, and so on. These sudden and well-marked contractions occur at intervals of many seconds, and show a decided tendency to rhythmic periodicity, though the rhythm is not always perfectly exact. This intensely slow rhythm, as the result of injury, may continue for a long time, particularly if the injury has been of a severe character. There can be no doubt, therefore, that the mechanical (or other) injury in this case acts as a source of constant irritation; so that here again we have evidence of rhythmic action independent of ganglia, and caused by the alternate exhaustion and recovery of contractile tissues.[26]

With regard to artificial rhythm caused by chemical stimuli, by far the most conspicuous instance that I have observed is that of the paralyzed nectocalyx of Sarsia. This consists in a highly peculiar motion of a flurried, shivering character, which is manifested by this organ when its marginal ganglia have been removed and it is exposed to the influence of faintly acidulated water. Now, when read in the light of the foregoing facts, there can be no doubt that the present one falls into its place very satisfactorily: it is an additional and very valuable instance of the display of artificial rhythm under the influence of a constant stimulus of low intensity; for the shivering motions of the mutilated nectocalyx under these circumstances are most unmistakably of a rhythmic nature. Viewed from a little distance, indeed, these motions are not distinguishable from the natural swimming motions of the unmutilated animal, except that, not being of quite such a powerful character, they are not so effective for locomotion. Viewed more closely, however, it may frequently be seen that the whole bell does not contract simultaneously, but that, as it were, clouds of contraction pass now over one part and now over another. Still, whether the contractions are partial or universal, they are more or less rhythmical. As this was the only case that had ever been observed of rhythm as due to a constant chemical stimulus, I studied it with much care, and shall now give a full description of what appears to me an important body of physiological facts.

Ten to twenty drops of acetic acid having been added to one thousand cubic centimetres of sea-water, and the paralyzed bell of Sarsia having been placed in the mixture, an interval of about half a minute will elapse before any movement begins. Sooner or later, however, the artificial rhythm is sure to be induced, and it will then continue for a variable time—occasionally as long as an hour, and generally for a considerable number of minutes. After it ceases it may often be made to recommence, either by adding a few more drops of acid to the sea-water, or by supplying an additional stimulus to the bell by nipping it with the forceps. Eventually, however, all movement ceases, owing to the destruction of irritability by the action of the acid. By this time the whole inner surface of the bell has become strongly opalescent, owing to the destructive influence of the acid on the epithelial cells which overspread the irritable tissues. The latter fact is worth mentioning, because in no case does the artificial rhythm set in until this opalescence has begun to show itself; and as this opalescence is but an optical expression of the damage which the epithelial coat is undergoing, the explanation of the time which elapses after the first immersion of the bell in the acidulated water and the commencement of the artificial rhythm no doubt is, that during this time the acid has not obtained sufficient access to the excitable tissues to serve as an adequate stimulus.

During the soaking stage of the experiment—i.e. before the artificial rhythm begins—the excitability of the tissues may be observed progressively and abnormally to increase; for soon after the soaking stage begins, in response to a single nip with the forceps the bell may give two or three locomotor contractions, instead of a single one, as is invariably the case with a paralyzed bell of Sarsia in normal water. Later on during the soaking stage, four or five successive contractions may be yielded in response to a single mechanical stimulus, and shortly after this a whole bout of rhythmic contractions may be started by the same means. Indeed, in some cases the artificial rhythm in acidulated water requires such a single additional stimulus for its inauguration, the shivering movements failing to begin spontaneously, but beginning immediately upon the application of the additional stimulus. Similarly, after the shivering movements have ceased, a fresh bout may very often be started by again giving the motionless nectocalyx a single stimulation. The interpretation of these facts would seem to be that the general irritability of the excitable tissues is exalted by the universal and constant stimulus supplied by the acid to an extent that is just bordering on that which gives rise to rhythmic movement, so that when the violent contraction is given in response to the mechanical stimulus, the disturbance serves to start the rhythmic movement.

If a paralyzed nectocalyx, while manifesting its artificial rhythm in acidulated sea-water, be suddenly transferred to normal sea-water, the movements do not cease immediately, but continue for a considerable time. This fact can easily be explained by the very probable, and indeed almost necessary, supposition that it takes some time after the transference to the normal sea-water for the acid to be washed out from contact with the excitable tissues. Sooner or later, however, as we should expect, in the normal sea-water the rhythmic movements entirely cease, and the bell becomes quiescent, with a normal irritability as regards single stimuli. If it be now again transferred to the acidulated water, after a short interval the rhythmic movements will again commence, and so on during several repetitions of this experiment, until the irritability of the tissues has finally become destroyed by the influence of the acid.

Other chemical irritants which I have tried produce substantially similar effects on the paralyzed bell of Sarsia. I shall, therefore, only wait to describe the influence of one of these irritants, the action of which in some respects differs from that of acids, and which I have found to be one of the most unfailing in its power to produce the rhythmic movements in question. This irritant is glycerine, and in order to produce its full effect it requires to be added to the sea-water in about the proportion of five per cent. The manifestation of artificial rhythm in solutions of this kind is quite unfailing. It begins after an exposure of from fifteen to thirty seconds, and continues for a variable number of seconds. It generally begins with powerful contractions, of a less shivering character than those which are produced by acids, and therefore still more closely resembling the normal swimming motions of the unmutilated animal. Sometimes, however, the first manifestation of the artificial rhythm is in the form of a few gentle rhythmic contractions, to be followed by a few seconds of quiescence, and then by the commencement of the sustained bout of strong contractions. In either case, when the bout of strong contractions sets in, the rate of the rhythm becomes progressively and rapidly increased, until it runs up into incipient tetanus. The rate of the rhythm still quickening, the tetanus rapidly becomes more and more pronounced, till at last the bell becomes quiescent in tonic spasm.[27]

If the bell is still left in the glycerine solution nothing further happens; the tissues die in this condition of strong systole. But if the bell be transferred to normal sea-water immediately after, or, still better, slightly before the tonic spasm has become complete, an interesting series of phenomena is presented. The spasm persists for a long time after the transference without undergoing any change, the length of this time depending on the stage in the severity and the spasm at which the transference is made. After this time is passed, the spasm becomes less pronounced than it was at the moment of transference, and a reversion takes place to the rhythmic contractions. The spasm may next pass off entirely, leaving only the rhythmic contractions behind. Eventually these too fade away into quiescence, but it is remarkable that they leave behind them a much more persistent exaltation of irritability than is the case with acid. For in the case of glycerine, the paralyzed bell which has been exposed to the influence of the irritant and afterwards become quiescent in normal sea-water, will often continue for hours to respond to single stimuli with a bout of rhythmic contractions. This effect of glycerine in producing an extreme condition of exalted irritability is also rendered apparent in another way; for if, during the soaking stage of the experiment—i.e. before the first of the rhythmic contractions has occurred—the bell be nipped with the forceps, the effect may be that of so precipitating events that the whole of the rhythmic stages are omitted, and the previously quiescent bell enters at once into a state of rigid tonic spasm. This effect is particularly liable to occur after prolonged soaking in weak solutions of glycerine.

As in the case of stimulation by acid, so in that of stimulation by glycerine, the artificial rhythm never begins in any strength of solution until the epithelial surface has become opalescent to a considerable degree.

In the case of stimulation by glycerine, the behaviour of the manubrium is more unequivocal than it is in the case of stimulation by acid. I have therefore reserved till now my description of the behaviour of this organ under the influence of constant chemical stimulation. This behaviour is of a very marked though simple character. The manubrium is always the first to respond to the stimulation, its retraction preceding the first movements of the bell by an interval of several seconds, so that by the time the bell begins its rhythmic response the manubrium is usually retracted to its utmost. The initial response of the manubrium is also rhythmic, and the rhythm which it manifests—especially if the glycerine solution be not over-strong—is of the same slow character which has already been described as manifested by this organ when under the influence of mechanical stimulation. The rhythm, however, is decidedly quicker in the former than in the latter case.

Lastly, with regard to the marginal ganglia, it is to be observed that in the case of all the chemical irritants I have tried, if unmutilated specimens of Sarsia be immersed in a sea-water solution of the irritant which is of a sufficient strength to evoke artificial rhythm in paralyzed specimens, the spontaneity of the ganglia is destroyed in a few seconds after the immersion of the animals, i.e. in a shorter time than is required for the first appearance of artificial rhythm. Consequently, whether the specimens experimented upon be entire or paralyzed by removal of their margins, the phenomena of artificial rhythm under the influence of chemical stimulation are the same. But although the spontaneity of the ganglia disappears before the artificial rhythm sets in, such is not the case with the reflex activity of the ganglia; for on nipping a tentacle of the quiescent bell before the artificial rhythm has set in, the bell will give a single normal response to the stimulation.

Hence, in historical order, on dropping an unmutilated specimen of Sarsia into a solution of glycerine of the strength named, the usual succession of events to be observed is an follows. First, increased activity of the normal swimming motions, to be quickly followed by a rapid and progressive decrease of such activity, till in about fifteen seconds after the immersion total quiescence supervenes. Four or five seconds later the manubrium begins to retract by rhythmical twitches, the rate of this rhythm rapidly increasing until it ends in tonic contraction. When the manubrium has just become fully retracted—or very often a little earlier—the bell suddenly begins its forcible and well-pronounced rhythmic contractions, which rapidly increase in their rate of rhythm until they coalesce into a vigorous and persistent spasm. If the animal be now restored to normal sea-water, spontaneity will return in a feeble manner; but there is always afterwards a great tendency displayed by the bell to exhibit shivering spasms instead of normal swimming movements in response to natural or ganglionic stimulation. And, as already observed, this peculiarity of the excitable tissues is also well marked in the case of the artificial stimulation of deganglionated specimens under otherwise similar conditions.

One further experiment may here be mentioned. Having split open the paralyzed bell of Sarsia along the whole of one side from base to apex of the cone, I suspended the now sheet-like mass of tissue by one corner in the air, leaving the rest of the sheet to hang vertically downwards. By means of a rack-work support I now lowered the sheet of tissue, till one portion of it dipped into a beaker filled with a solution of glycerine of appropriate strength. After allowing this portion to soak in the solution of glycerine until it became slightly opalescent, I dropped the entire mutilated bell, or sheet of tissue, into another beaker containing sea-water. If the exposure to the glycerine solution had been of sufficient duration, I invariably found that in the normal sea-water the rhythmic movements were performed by the whole tissue-mass quite as efficiently as was the case in my other experiments, where the whole tissue-mass, and not merely a portion, had been submitted to the influence of the irritant. But on now suddenly snipping off the opalescent portion of the tissue-mass, i.e. the portion which had been previously alone submitted to the influence of the irritant, all movement in the remainder of the tissue-mass instantly ceased. This experiment I performed repeatedly, sometimes exposing a large and sometimes a small portion of the tissue to the influence of the irritant. As I invariably obtained the same result, there can be no doubt that in the case of chemical stimulation the artificial rhythm depends for its manifestation on the presence of a constant stimulus, and is not merely some kind of obscure fluttering motion which, having been started by a stimulus, is afterwards kept up independently of any stimulus.

Such being the case, I naturally expected that if I were to supply a constant stimulus of a thermal kind, I should also obtain the phenomena of artificial rhythm. In this, however, my expectations have not been realized. With no species of Medusa on of artificial rhythm by immersing the paralyzed animals in heated water. I can only explain this fact by supposing that the stimulus which is supplied by the heated medium is of too uniform a character over the whole extent of the excitable tissues; it would seem that in order to produce artificial rhythm there must be a differential intensity of stimulation in different parts of the responding tissue, for no doubt even the excitatory influence of acidulated water is not of nearly so uniform an intensity over the whole of the tissue-area as is that of heated water.

In now quitting the subject of artificial rhythm as it is manifested by the paralyzed bell of Sarsia, it is desirable again to observe that sustained artificial rhythm cannot be produced by means of chemical irritation in the case of any one of the species of covered-eyed Medusæ that I have met with. In order to evoke any response at all, stronger solutions of the irritants require to be employed in the case of the covered than in that of the naked-eyed Medusæ, and when the responses do occur they are not of so suggestive a character as those which I thought it worth while so fully to describe. Nevertheless, even in the covered-eyed Medusæ well marked, though comparatively brief, displays of artificial rhythm may often be observed as the result of constant chemical stimulation. Thus, for instance, in the case of Aurelia, if the paralyzed umbrella be immersed in a solution of glycerine (ten to twenty per cent.), a few rhythmic pulsations of normal rate are usually given; but shortly after these pulsations occur, the tissue begins to go into a tetanus, which progressively and rapidly becomes more and more pronounced until it ends in violent tonic spasm. So that the history of events really resembles that of Sarsia under similar circumstances, except that the stage of artificial rhythm which inaugurates the spasm is of a character comparatively less pronounced.

Thus far, then, I have detailed all the facts which I have been able to collect with reference to the phenomena of artificial rhythm, as produced by different kinds of constant stimulation. It will not be forgotten that the interest attaching to these facts arises from the bearing which they have on the theory of natural rhythm. My belief is that hitherto the theory of rhythm as due to ganglia has attributed far too much importance to the ganglionic as distinguished from the contractile tissues, and I have founded this belief principally on the facts which have now been stated, and which certainly prove at least this much: that after the removal of the centres of spontaneity, the contractile tissues of the Medusæ display a marked and persistent tendency to break into rhythmic action whenever they are supplied with a constant stimulus of feeble intensity. Without waiting again to indicate how this fact tends to suggest that the natural rhythm of the unmutilated organisms is probably in large part due to that alternate process of exhaustion and restoration of excitability on the part of the contractile tissues, whereby alone the phenomena of artificial rhythm can be explained,[28] I shall go on to describe some further experiments which were designed to test the question whether the influences which affect the character of the natural rhythm likewise, and in the same manner, affect the character of the artificial rhythm. I took the trouble to perform these experiments, because I felt that if they should result in answering this question in the affirmative, they would tend still further to substantiate the view I am endeavouring to uphold, viz. that the natural rhythm may be a function of the contractile as distinguished from the ganglionic tissue. Of the modifying causes in question, the first that I tried was temperature.

Having already treated of the effects of temperature on the natural rhythm, it will now be sufficient to say that we have seen these effects to be similar to those which temperature exerts on the rhythm of ganglionic tissues in general. Now, I find that temperature exerts precisely the same influence on the artificial rhythm of deganglionated tissue as it does on the natural rhythm of the unmutilated animal. To economize space, I shall only quote one of my observations in a table which explains itself. I also append tracings of another observation, to render the difference in the rate of the artificial rhythm more apparent to the eye (Fig. 28).

During the whole progress of such experiments the faradaic stimulation was, of course, kept of uniform intensity; so that the progressive acceleration is undoubtedly due to the increase of temperature alone. With each increment of temperature the rate of the artificial rhythm increases suddenly, just as it does in the case of the natural rhythm. Moreover, there seems to be a sort of rough correspondence between the amount of influence that any given degree of temperature exerts on the rate of the natural and of the artificial rhythm respectively. Further, it will be remembered that in warm water the natural rhythm, besides being quicker, is not so regular as it is in cold water; thus also it is with the artificial rhythm. Again, water below 20° or above 85° suspends the natural rhythm, i.e. stops the contractions; and the artificial rhythm is suspended at about the same degrees. Lastly, just as there are considerable individual variations in the extent to which the natural rhythm is affected by temperature, so the artificial rhythm is in some cases more influenced by this cause than in others, though in all cases it further resembles the natural rhythm in showing some considerable degree of modification under such influence.