Part 4

These locomotive appendages are intimately connected with the circulating tubes, as we shall see when we examine the structural details of these animals, so that in them also breathing and moving are in direct relation to each other. To those unaccustomed to the comparison of functions in animals, the use of the word breathing, as applied to the introduction of water into the body, may seem inappropriate, but it is by the absorption of aerated water that these lower animals receive that amount of oxygen into the system, as necessary to the maintenance of life in them, as a greater supply is to the higher animals. The name of Ctenophoræ or comb-bearers, is derived from these rows of tiny paddles which have been called combs by some naturalists, because they are set upon horizontal bands of muscles, see Fig. 29, reminding one of the base of a comb, while the fringes are compared to its teeth. These flappers add greatly to the beauty of these animals, for a variety of brilliant hues is produced along each row by the decomposition of the rays of light upon them when in motion. They give off all the prismatic colors, and as the combs are exceedingly small, so that at first sight one hardly distinguishes them from the disk itself, the exquisite play of color, rippling in regular lines over the surface of the animal, seems at first to have no external cause.

_Pleurobrachia_. (_Pleurobrachia rhododactyla Ag_.)

Among the most graceful and attractive of these animals are the Pleurobrachia (Fig. 29), and, though not first in order, we will give it the precedence in our description, because it will serve to illustrate some features of the other two groups. The body of the Pleurobrachia consists of a transparent sphere, varying, however, from the perfect sphere in being somewhat oblong, and also by a slight compression on two opposite sides (Figs. 27 and 28), so as to render its horizontal diameter longer in one direction than in the other (Fig. 30). This divergence from the globular form, so slight in Pleurobrachia as to be hardly perceptible to the casual observer, establishing two diameters of different lengths at right angles with each other, is equally true of the other genera. It is interesting and important, as showing the tendency in this highest group of Acalephs to assume a bilateral character. This bilaterality becomes still more marked in the highest class of Radiates, the Echinoderms. Such structural tendencies in the lower animals, hinting at laws to be more fully developed in the higher forms, are always significant, as showing the intimate relation between all parts of the plan of creation. This inequality of the diameters is connected with the disposition of parts in the whole structure, the locomotive fringes and the vertical tubes connected with them being arranged in sets of four on either side of a plane passing through the longer diameter, showing thus a tendency toward the establishment of a right and left side of the body, instead of the perfectly equal disposition of parts around a common centre, as in the lower Radiates.

The Pleurobrachia are so transparent, that, with some preparatory explanation of their structure, the most unscientific observer may trace the relation of parts in them. At one end of the sphere is the transverse split (Fig. 27), that serves them as a mouth; at the opposite pole is a small circumscribed area, in the centre of which is a dark eye-speck. The eight rows of locomotive fringes run from pole to pole, dividing the whole surface of the body like the ribs on a melon. (Figs. 27, 28.) Hanging from either side of the body, a little above the area in which the eye-speck is placed, are two most extraordinary appendages in the shape of long tentacles, possessing such wonderful power of extension and contraction that, while at one moment they may be knotted into a little compact mass no bigger than a pin's head, drawn up close against the side of the body, or hidden within it, the next instant they may be floating behind it in various positions to a distance of half a yard and more, putting out at the same time soft plumy fringes (Fig. 29) along one side, like the beard of a feather. One who has never seen these animals may well be pardoned for doubting even the most literal and matter-of-fact account of these singular tentacles. There is no variety of curve or spiral that does not seem to be represented in their evolutions. Sometimes they unfold gradually, creeping out softly and slowly from a state of contraction, or again the little ball, hardly perceptible against the side of the body, drops suddenly to the bottom of the tank in which the animal floating, and one thinks for a moment, so slight is the thread-like attachment, that it has actually fallen from the body; but watch a little longer, and all the filaments spread out along the side of the thread, it expands to its full length and breadth, and resumes all its graceful evolutions.

One word of the internal structure of these animals, to explain its relation to the external appendages. The mouth opens into a wide digestive cavity (Figs. 27, 28), enclosed between two vertical tubes. Toward the opposite end of the body these tubes terminate or unite in a single funnel-like canal, which is a reservoir as it were for the circulating fluid poured into it through an opening in the bottom of the digestive cavity. The food in the digestive cavity becomes liquefied by mingling with the water entering with it at the mouth, and, thus prepared, it passes into this canal, from which, as we shall presently see, all the circulating tubes ramifying throughout the body are fed. Two of these circulating tubes, or, as they are called from the nature of the liquid they contain, chymiferous tubes, are very large, starting horizontally and at right angles with the digestive cavity from the point of junction between the vertical tubes (Fig. 30) and the canal. Presently they give off two branches, those again ramifying in two directions as they approach the periphery, so that each one of the first main tubes has multiplied to four, before its ramifications reach the surface, thus making in all eight radiating tubes. So far, these eight tubes are horizontal, all diverging on the same level; but as they reach the periphery each one gives rise to a vertical tube, running along the surface of the body from pole to pole, just within the rows of locomotive fringes on the outer surface, and immediately connected with them (Figs. 27, 28). As in all the Ctenophoræ, these fringes keep up a constant play of color by their rapid vibrations. In Pleurobrachia the prevailing tint is a yellowish pink, though it varies to green, red, and purple, with the changing motions of the animal. We have seen that the vertical tubes between which the digestive cavity is enclosed, start like the cavity itself from that pole of the body where the mouth is placed, and that, as they approach the opposite pole, at a distance from the mouth of about two thirds the whole length of the body, they unite in the canal, which then extends to the other pole where the eye-speck is placed. As it is just at this point of juncture between the tubes and the canal that the two main horizontal tubes arise from which all the others branch on the same plane (Figs. 27, 28), it follows that they reach the periphery, not on a level with the pole opposite the mouth, but removed from it by about one third the height of the body. In consequence of this the eight vertical tubes arising from the horizontal ones, in order to run the entire length of the body from pole to pole, extend in opposite directions, sending a branch to each pole, though the branch running toward the mouth is of course the longer of the two. The tentacles have their roots in two sacs within the body, placed at right angles with the split of the mouth. (Figs. 27, 30.) They open at the surface on the opposite side from the mouth, though not immediately within the area at which the eye-speck is placed, but somewhat above it, and at a little distance on either side of it. The tentacles may be drawn completely within these sacs, or be extended outside, as we have seen, to a greater or less degree, and in every variety of curve or spiral.

_Bolina_. (_Bolina alata_ AG.)

The Bolina (Fig 32), like the Pleurobrachia, is slightly oval in form, with a longitudinal split at one end of the body, forming a mouth which opens into a capacious sac or digestive cavity. But it differs from the Pleurobrachia in having the oral end of the body split into two larger lobes (Fig. 31), hanging down from the mouth. These lobes may gape widely, or they may close completely over the mouth so as to hide it from view, and their different aspects under various degrees of expansion or contraction account for the discrepancies in the description of these animals. We have seen that the Pleurobrachia moves with the mouth upward; but the Bolina, on the contrary, usually carries the mouth downward, though it occasionally reverses its position, and in this attitude, with the lobes spread open, it is exceedingly graceful in form, and looks like a white flower with the crown fully expanded. These broad lobes are balanced on the other sides of the body by four smaller appendages, divided in pairs, two on each side (Fig. 32), called auricles. These so-called auricles are in fact organs of the same kind as the larger lobes, though less developed. The rows of locomotive flappers on the Bolina differ in length from each other (Fig. 31), instead of being equal, as in the Pleurobrachia. The four longest ones are opposite each other on those sides of the body where the larger lobes are developed, the four short ones being in pairs on the sides where the auricles are placed. At first sight they all seem to terminate at the margin of the body, but a closer examination shows that the circulating tubes connected with the longer row extend into the lobes, where they wind about in a variety of complicated involutions. (Fig. 32.) The movements of the Bolina are more sluggish than those of the Pleurobrachia, and the long tentacles, so graceful an ornament to the latter, are wanting in the former. With these exceptions the description given above of the Pleurobrachia will serve equally well for the Bolina. The structure is the same in all essential points, though it differs in the size and proportion of certain external features, and its play of color is less brilliant than that of the Pleurobrachia. The Bolina, with its slow, undulating motion, its broad lobes sometimes spreading widely, at other times folded over the mouth, its delicacy of tint and texture, and its rows of vibrating fringes along the surface, is nevertheless a very beautiful object, and well rewards the extreme care without which it dies at once in confinement.

_Idyia_. (_Idyia roseola_ AG.)

The lowest genus of Ctenophoræ found on our coast, the Idyia (Fig. 33), has neither the tentacles of the Pleurobrachia, nor the lobes of the Bolina. It is a simple ovate sphere, the interior of which is almost entirely occupied by an immense digestive cavity. It would seem that the reception and digestion of food is intended to be the almost exclusive function of this animal, for it has a mouth whose ample dimensions correspond with its capacious stomach. Instead of the longitudinal split serving as a mouth, in the Bolina and Pleurobrachia, one end of the body in the Idyia is completely open (Fig. 33), so that occasionally some unsuspicious victim of smaller diameter than itself may be seen to swim into this wide portal, when suddenly the door closes behind him with a quick contraction, and he finds himself a prisoner. The Idyia does not always obtain its food after this indolent fashion however, for it often attacks a Bolina or Pleurobrachia as large or even larger than itself, when it extends its mouth to the utmost, slowly overlapping the prey it is trying to swallow by frequent and repeated contractions, and even cutting off by the same process such portions as cannot be forced into the digestive cavity.

The general internal structure of the Idyia corresponds with that of the Bolina and Pleurobrachia; it has the same tubes branching horizontally from the main cavity, then ramifying as they approach the periphery till they are multiplied to eight in all, each of which gives off one of the vertical tubes connected with the eight rows of locomotive flappers. Opposite the mouth is the eye-speck, placed as in the two other genera, at the centre of a small circumscribed area, which in the Idyia is surrounded by delicate fringes, forming a rosette at this end of the body. These animals are exceedingly brilliant in color; bright pink is their prevailing hue, though pink, red, yellow, orange, green, and purple, sometimes chase each other in quick succession along their locomotive fringes. At certain seasons, when most numerous, they even give a rosy tinge to patches on the surface of the sea. Their color is brightest and deepest before the spawning season, but as this advances, and the ovaries and spermaries are emptied, they grow paler, retaining at last only a faint pink tint. They appear early in July, rapidly attain their maximum size, and are most numerous during the first half of August. Toward the end of August they spawn, and the adults are usually destroyed by the early September storms, the young disappearing at the same time, not to be seen again till the next summer. It is an interesting question, not yet solved, to know what becomes of the summer's brood in the following winter. They probably sink into deep waters during this intervening period. The Idyia, like the Pleurobrachia, moves with the mouth upward, but inclined slightly forward also, so as to give an oblique direction to the axis of the body.[4]

[Footnote 4: Until this summer only the three genera of Ctenophoræ above mentioned were supposed to exist along our coast, but during the present season I have had the good fortune to find two additional ones. One of them, the Lesueuria, resembles a Bolina with the long lobes so cut off, that they have a very stunted appearance in comparison with those of the Bolina. The other, the Mertensia, is closely allied to Pleurobrachia; it is exceedingly flattened and pear-shaped. This species was discovered long ago by Fabricius, but had escaped thus far the attention of other naturalists. (_A. Agassiz_.)]

EMBRYOLOGY OF CTENOPHORÆ.

All the Ctenophoræ are reproduced from eggs, these eggs being so transparent that one may follow with comparative ease the changes undergone by the young while still within the egg envelope. Unfortunately, however, they are so delicate that it is impossible to keep them alive for any length of time, even by supplying them constantly with fresh sea-water, and keeping them continually in motion, both of which are essential conditions to their existence. It is therefore only from eggs accidentally fished up at different stages of growth that we may hope to ascertain any facts respecting the sequence of their development. When hatched, the little Ctenophore is already quite advanced. It is small when compared with the size of the egg envelope, and long before it is set free, it swims about with great velocity within the walls of its diminutive prison (Fig. 35). The importance of studying the young stages of animals can hardly find a better illustration than among the Ctenophoræ. Before their extraordinary embryonic changes were understood, many of the younger forms had found their way into our scientific annals as distinct animals, and our nomenclature thus became burdened with long lists of names which will disappear as our knowledge advances.

The great size of their locomotive flappers in proportion to the rest of the body, is characteristic of the young Ctenophoræ. They seem like large paddles on the sides of these tiny transparent spheres, and, owing to their great power as compared with those of the adult, the young move with extraordinary rapidity. The Pleurobrachia alone retains its quickness of motion in after life, and although its long graceful streamers appear only as short stumpy tentacles in the young (Fig. 34), yet its active little body would be more easily recognized in the earlier stages of growth than that of the other Ctenophoræ. Figs. 34, 35, and 36 show the Pleurobrachia at various stages of growth; Fig. 34, with its thick stunted tentacles and short rows of flappers, is the youngest; the flappers themselves are rather long at this age, looking more like stiff hairs than like the minute fringes of the adult. In Fig. 35 the tentacles are already considerably longer and more delicate; in Fig. 36 the vertical tubes are already completed, while Figs. 27-29 present it in its adult condition.

The Idyia differs greatly in appearance at different periods of its development, and, indeed, no one would suspect, without some previous knowledge of its transformations, that the young Idyia, with its rapid gyrations, its short ambulacral tubes, like immense pouches (Fig. 37), its large pigment spots scattered over the surface (Fig. 38), was an earlier stage of the rosy-hued Idyia, which glides through the water with a scarcely perceptible motion. Figs. 37-40 represent the various stages of its growth. It will be seen how very short are the locomotive fringes (Fig. 39) in comparison with those of the full-grown ones (Fig. 33). It is only in the adult Idyia that these rows attain their full height, and the tubes, ramifying throughout the body (Fig. 40), are completed.

The Bolina, in its early condition, recalls the young Pleurobrachia. At this period it has the same rapid motion, and when somewhat more advanced, long tentacles, resembling those of the Pleurobrachia, make their appearance (Fig. 41); it is only at a later period that the tentacles become contracted, while the large lobes (Fig. 42), so characteristic of Bolina, are formed by the elongation of the oral end of the body, the auricles becoming more conspicuous at the same time (Fig. 43). A little later the lobes enlarge, the movements become more lazy; it assumes both in form and habits the character of the adult Bolina.

The series of changes through which the Ctenophoræ pass are as remarkable as any we shall have occasion to describe, though not accompanied with such absolutely different conditions of existence. The comparison of the earlier stages of life in these animals with their adult condition is important, not only with reference to their mode of development, but also because it gives us some insight into the relative standing of the different groups, since it shows us that certain features, permanent in the lower groups, are transient in the higher ones. A striking instance of this occurs in the fact mentioned above, that though the long tentacles so characteristic of the adult Pleurobrachia exist in the young Bolina, they yield in importance at a later period to the lobes which eventually become the predominant and characteristic feature of the latter.

* * * * *

DISCOPHORÆ.

The disk of the Discophoræ is by no means so delicate as that of the other Jelly-fishes. It seems indeed quite solid, and somewhat like cartilage to the touch, and yet so large a part of its bulk consists of water, that a Cyanea, weighing when alive about thirty-four pounds, being left to dry in the sun for some days, was found to have lost about 99/100 of its original weight,--only the merest film remaining on the paper upon which it had been laid. The prominence of the disk in this group of Jelly-fishes is well characterized by their German name, "Scheiben quallen," viz. disk-medusæ. We shall see hereafter that the disk, so large and seemingly solid in the Discophoræ, thins out in many of the other Jelly-fishes, and becomes exceedingly concave. This is especially the case in many of the Hydroid Medusæ, where it assumes a bell-shaped form, and is constantly spoken of as the bell. It should be remembered, however, in reading descriptions of these animals, that the so-called bell is only a modified disk, and perfectly homologous with that organ in the Discophoræ.

The Discophorous Medusæ are distinguished from all others by the peculiar nature of the reproductive organs. They are contained in pouches (Fig. 50, _o_, _o_, _o_, _o_), the contents of which are first discharged into the main cavity, and then pass out through the mouth. Pillars support the four angles of the digestive cavity, thus separating the lower from the upper floor of the disk, while the chymiferous tubes (Fig. 50) branch and run into each other near the periphery, forming a more or less complicated anastomosing network, instead of a simple circular tube, as is the case with the Hydroid Medusæ. (Fig. 74.)

_Cyanea_. (_Cyanea arctica_ PÉR. et LES.)