Effects of Segmentation on the Rhythm.
We have next to consider Dr. Eimer's observations concerning the effects on the rhythm of Aurelia which result on cutting the animal into segments; and here, again, I much regret to say that I cannot wholly agree with this author. He says he found evidence of a very remarkable fact, viz. that by first counting the natural rhythm of an unmutilated Aurelia, and then dividing the animal into two halves, one of these halves into two quarters, and one of these quarters into two eighths; the sum of the contractions performed by these four segments in a given time was equal to the number which had previously been performed in a similar time by the unmutilated animal. And not only so, but the number of contractions which each segment contributed to this sum was a number that stood in direct proportion to the size of the segment; so that the half contracted half as many times, the quarter a quarter as many times, and the eighth parts one-eighth part the number of times that the unmutilated Aurelia had previously contracted in a period of equal duration. I am glad to observe that Dr. Eimer does not regard this rule otherwise than as liable to frequent exception; for, as already observed, I cannot say that my experiments have tended to confirm it. I am only able to say that there is general tendency for the smaller segments of an Aurelia divided in this way to contract less frequently than the larger segments.
It would be tedious and unnecessary to quote any observations in this connection; but as these observations brought out very clearly a fact which I had previously suspected, I may detail one experiment to illustrate this point. The fact in question is, that the potency of the lithocysts in any given segment of a divided Aurelia has more to do with the frequency of its pulsations than has the size of the segment. As previously mentioned, one or more lithocysts may often be observed to be permanently prepotent over the others; and I may here observe that the segmentation experiments just described have shown the converse to be true, viz. that one or more lithocysts are often permanently feebler than the others. Well, if a specimen of Aurelia exhibiting decided prepotency in one or more of its lithocysts be watched for a considerable length of time, so as to be sure that the prepotency is not of a merely temporary character, and if the animal be then divided into segments in such a way that the prepotent lithocysts shall occupy the smaller segments, it may be observed, provided time be left for the tissues to recover, that the segments containing the prepotent lithocysts, notwithstanding their smaller size, contract more frequently than do the larger segments. Conversely, if the larger segments happen to contain feeble lithocysts, their contractions will be but few. I have, indeed, seen cases in which the lithocysts appeared to be quite functionless, so far as the origination of stimuli was concerned.
The following observations were made on a healthy specimen of Aurelia having all its lithocysts in good condition, but prepotency being well marked in the case of one of them, and also, though in a lesser degree, in the case of another. I divided the animal so as to leave one of these two prepotent lithocysts in each of the eighth-part segments, and the next most powerful lithocysts in the quadrant segment. In the following description, I shall call the two eighth-part segments A and B, the former letter designating the segment containing the most powerful lithocyst. The Aurelia before being divided manifested for several hours a very regular and sustained rhythm of thirty-two per minute. After its division, the various segments contracted at the following rates, in one-minute intervals:—
| Time after operation. | Segment 1/2. | Segment 1/4. | Segment 1/8 A. | Segment 1/8 B. |
| 1/2 hour. | 20 | 25 | 27 | 15 |
| 1 " | 20 | 25 | 27 | 15 |
| 2 hours. | 29 | 25 | 27 | 16 |
| 4 " | 19 | 16 | 27 | 12 |
Next morning, the water which contained the segments was somewhat foul, and this, as is always the case, gave rise to abnormally long pauses. This effect was much more marked in the case of some of the segments than in that of others. I therefore observed the segments over five-minute intervals, instead of one-minute intervals as on the previous day. The following is a sample of several observations, all yielding the same general result.
| Segment 1/2. | Segment 1/4. | Segment 1/8 A. | Segment 1/8 B. | |
| Number of pulsations. | Seconds of rest. | No motion during the hour of observation. | Continued persistently to contract with a nearly perfect rhythm of 78 in the 5 minutes during the hour of observation. | Rhythm tolerably perfect at 78 in the 5 minutes; but this was occasionally interrupted by long pauses of 4 or 5 minutes' duration. |
| 12 | 120 | |||
| 3 | 10 | |||
| 2 | 20 | |||
| 44 | 130 | |||
| 12 | 20 | |||
| 73 | 5 min. | |||
| Average rate 14-3/5 per minute. | No motion. | Continuous rhythm at the rate of 15-3/5 per minute. | Interrupted rhythm at the rate of 15-3/5 per minute. | |
I now transferred all the segments to fresh sea-water, with the following results:—
Next day all the segments were dead except the largest one, in which a single lithocyst still continued to discharge at the rate of twenty-four in five minutes.