Zoophytes, Cœlenterata, Medusæ, Corals, Hydrozoa.
Fig. 344.—Gorgonia Nobilis.
A study of the earliest growth of the Cœlenterata has shown that their internal cavities are nothing more than regular radiate out-growths of the internal structures. The result of this development is a condition which does not occur again in the whole of the animal kingdom. There is a system of cavities all in open communication one with another, no closed blood vascular system, and no specialised respiratory apparatus. Again, all the animals that constitute this large group are radiate in structure, that is, when viewed from above they are typically star-shaped, and if cut across, every horizontal section shows a symmetrical arrangement of the several parts around a centre. There are other radiate animals, as the Echinoderms, but while in these five is the fundamental number of rays, in the Cœlenterata the rays are a multiple of four, six and upwards. The skeleton or framework of each differs, and when the Cœlenterata form calcareous structures, these are quite different from the tests of the sea-urchins; and in all cases the anterior portion of the body is crowned with one or more circles of tentacles, which remain perfectly flexible and flower-like. The most highly-developed of the free forms are the sea-anemones and the jelly-fish. These have no hard or calcareous skeleton whatever, but withal they are, in the opinion of naturalists and microscopists, the most beautiful objects among Zoophytes.
In spite of their variety of forms, the Cœlenterata seem to be as incapable of higher development as do the Echinoderms, and they have failed to make headway in fresh water, but it is not improbable that some of the simplest forms of the whole group may have given rise to higher animal forms, while the sea anemones, corals, &c., being those descendants of the primitive simple form, have retained the original type of organization almost unchanged.
Fig. 345.—Hydra viridis, adhering to a stalk of Anacharis alsinastrum.
The type of the group is the Hydra, a fresh-water polyp, commonly found attached to the leaves and stems of many aquatic plants, or floating pieces of stick. Two species are well known to microscopists, the H. viridis, or green polyp, and the H. vulgaris, somewhat darker in colour, probably dependent upon the nature of its food. The third, less common species, the H. fasca, is distinguished from both by the length of its tentacles, which, when fully extended, greatly exceed those of either of the before-mentioned. The fresh-water group measures from one-eighth to the one-third of an inch in length, and form simple stocks of one, two or more branches. They almost exactly resemble in form the polyps of the Hydractinia, which are provided with a circle of tentacles. When placed in a vessel of water and left undisturbed they often attach themselves to the side, where they may be examined with a moderate power at leisure. They are then seen to spread out their tentacles like fine threads, and seize upon any small creature that may come in their way, and by the same means convey it to a mouth capable of great extension. All Hydra possess stinging-cells, by means of which they paralyse their prey. Many Hydra attain to a large size, and shoot out long poisonous filaments; they also possess smaller kinds of smooth cells, which appear to be employed for an entirely different purpose, but for what is not positively known. Hydra usually multiply by means of buds, an out-growth from the body, and these remain attached to the mother stalk for some time, often long enough to give rise to one or two smaller buds. Single eggs are also developed in the ectoderm beneath capsules, or wart-like prominences. The adult animal can be cut to pieces, and from each piece a new individual will be developed. This method of reproduction was first tried by the naturalist Trembley in 1739, whose experiments in this direction excited the greatest interest among the naturalists of the middle of the last century. Hydra fusca in various stages of development is given in outline in [Fig. 346].
Fig. 346.
1, 2, 3. Hydra in various stages of development; 4. Group of Stentor polymorphus, many-shaped Stentor; 5. Englena; 6. Monads.
In the polyps belonging to this family the body-structure for the most part consists of a homogeneous aggregation of vesicular granules, held together by an intercellular sarcode, and capable of great extension and contraction, so that these animals can assume a variety of forms and extend their body and tentacles until the latter become almost invisible. It was the resemblance in this respect to the fabled Hydra that originated the name. Its organ of prehension is termed the hasta; this consists of a sac or opening at the terminal end of the tentacle, within which is seen a saucer-shaped vesicle, supporting a minute ovate body, which carries a sharp calcareous piece termed a sagitta or arrow. Although the fresh-water Hydra may be regarded as typical of this group of animals, marine fauna furnish a far more extensive group in the corals, jelly-fish, and sea-anemones.
A smaller group, the Ctenophora, although members of this sub-kingdom, have not yet found their true position; nevertheless they are interesting glassy, transparent creatures, either shaped like apples, melons, or Phrygian caps, or else forming bands of some considerable length; all are wonderfully transparent, with the single exception of the Beroë. These inhabit the open sea, and are only seen inshore when driven in by currents or strong winds. Their position in the water is usually more or less vertical, the mouth being turned downwards. The portion from which this group derives its name is the ribs, which are symmetrically arranged, and consist of rows of short transverse combs, each forming rows of cilia, which, as they wave to and fro, constitute a swimming or rowing plate, their activity in the water depending upon the will of the animal. They are also provided with an oral umbrella, and capturing filaments or tentacles with hair-like branches. These tentacles, attached to the sides of the animal, are capable of erection or withdrawal into pockets. Great variety is seen in these accessory organs of locomotion; for instance, the Cydippidæ ([Plate XVII].) have only arms, but these are remarkable for their length, and serve for the purpose of capturing food as well as for steering. The most interesting, if not the most beautiful of the Ctenophora, are the Beroidæ; it is this family that bear a resemblance to the Phrygian cap ([Plate XVII]., e). The mouth is wide, but it appears to have no capturing tentacles, and yet their habits are carnivorous; they will even devour their own relations. Many of the genus are phosphorescent, and in place of stinging-cells have small spherical knobs beset with sticky globules, in which their food becomes entangled, and these are apparently in constant use.
PLATE XVII.
ZOOPHYTES, ASTEROIDS, NUDIBRANCHIS, ACALEPS, ECHINOIDS, CTENOPHORA, TUNICATA, AND CRUSTACEANS.
The Stinging Series, Cnidaria, comprise sea-anemones, corals, jelly-fish among marine animals, and Hydra among the fresh-water Cœlenterata; and derive their name from a remarkably curious feature, the so-called stinging capsules. These are not only offensive, but also defensive weapons with all the animals belonging to this group; the possession of which has converted the bell-like jelly-fish into a simple Cnidarian. The principal change is in the gelatinous layer between the outer wall and the inner digesting layer of the ectoderm. But without entering further into their structure and relations, the stinging-cells and batteries claim especial attention. These cells vary considerably in size without their characteristics being essentially changed. The protoplasm of the cell is modified into a tolerably firm substance, enclosing an oval or cylindrical vesicle. Closely associated with this is a pointed process, standing up far above the level of the outer covering, known as the cnidocil. Within the vesicle is found, either spirally rolled up or in an irregular tangle, a long filament or hollow tube, a prolongation of the vesicle, but turned outside in. This tube is more than twenty times as long as the cell, is pointed at the tip, and beset with two rows of fine spirally-arranged barbed hooks. When the cnidocil is touched or irritated, this filament is violently shot out, being then turned inside-out, like the fingers of a glove. So long as the thread remains rolled up within the vesicle the barbed hooks remain in their tube, but when shot out, they change to the outside. The rolled-up filament appears to be filled with some poisonous material, which is ejected when the tube is shot out, and where the point strikes a wound is inflicted, so that unless the prey is stronger than the attacker it cannot escape. The greater the struggle, the larger the number of capsules discharged in order to kill.
Fig. 347.—The Stinging Capsules of Cnidaria.
1 and 2. Retracted filaments; 3. Partly protruded; 4. Fully protruded. Magnified × 600. (Warne.)
Polypomedusæ.—Among the higher development of the stinging group is the jelly-fish. The Siphonophora, as represented by the Portuguese man-of-war, are in their turn the highest development of swimming-bells, and exhibit many modifications and combinations of individuals. The tentacles of the Physalia, the best known, are stiff with batteries of stinging-capsules, the sting of which is more like the shock of the electric current. The Challenger soundings brought to light some remarkably interesting forms, and these have furnished much work for the microscope, as all their larval forms are extremely curious. Among the Hydromedusæ there are many different life histories. Take the jelly-fish, the eggs of which have given up forming stocks, and are hatched out at once as Medusæ. There are others, the eggs of which form stocks; others, again, in which the sexual individuals do not swim away as jelly-fish. The last were at one time described under a new name, because of one or two curious forms being taken creeping on the ground. This creeping Medusa (Clavatella prolifera) has six arms, the tips of which are provided with true suckers, and on these it walks as on stilts, while from each arm a short stalk arises, the swollen end of which is beset with stinging capsules. It has an extensile mouth-tube, and feeds upon small crustaceans found on seaweeds.
Fig. 348.—Plumularia primata. Doris tuberculata seen clinging to a fucus.
Among the forms that swim away as jelly-fish a very curious example is presented in Corymorpha mutans. These swim about for a time, and then firmly attach themselves by numerous thread-like appendages, forced into the sand, and where the young prepare for their next metamorphosis. As an example of the stocks of those representatives which do not swim away as jelly-fish, take the beautifully-feathered, plant-like creatures found erect along the seashore, the Sertularia ([Fig. 358], No. 12) and Plumularia. Plumularia primata, [Fig. 348]. Other members of these groups will be found in [Plate IV]., Nos. 95-99.
Fig. 349.—Group of female stock of Hydractinia echinata.
a, a. Nutritive individuals; b, b. Female individuals and groups of eggs. Highly magnified.—(Warne.)
In addition to the nutritive individuals, there are the egg-bearing; these do not become free-swimming individuals. One small family is neither branched nor feathered—the Hydractinia echinata, found in the North Sea and on the Norwegian coasts, where it attaches itself to the shells of gastropods, selecting those inhabited by hermit crabs. The part of the stock common to all the individuals is the skin-like portion which adheres to the surface of the shell. In some spiny processes are produced, and the nutritive canals running down the stems of the polyps are continued into the membrane belonging to the stock, as seen in [Fig. 349].
The nutritive individuals are distinguished by long tentacles, mouths, and digestive canals. The females have no mouths, and are supplied with food through the system of canals running to them from the nutritive males. These reproductive members are furnished with stinging threads instead of tentacles for the protection of their ova. The ciliated larvæ, in a very short time, swim off to found new colonies.
Fig. 350.—Medusæ, Jelly-fish.
The free-swimming jelly-fish ([Fig. 350], and Plate XVII., c and d) belong to the order Scyphomedusæ. These are characterised by their delicate colouring, and from the arrangement of their nervous system, which can only be made out by staining. Some new and curious forms were dredged from a depth of more than 6,000 feet off the coast of New Zealand, varying in size from an inch to twenty inches; many having from four to eight or ten eyes arranged along the margin.
Anthozoa.—From the free-swimming we turn to a group of permanently fixed polyp forms, the sea-anemones and corals. The development of Monoxenia commences with the egg, repeatedly dividing into many parts ([Fig. 351], C, D, and E), by a process common to the animal kingdom, termed egg-segmentation, in this particular instance proceeding from an apparently hollow sphere, A, enclosing a single layer of cells, G. Each cell sends out a long cilia, or whip-like process, F, by means of which the larva turns about and swims in the body fluid of the parent polyp. One half of the sphere now becomes enfolded into the other half, H, and forms what is termed a gastrula, I, K. The gastrula stage of Monoxenia is of the simplest kind, the larva forming a sac, with walls consisting of two layers, an outer, or ectoderm, and an inner, or endoderm. The transition from the flat dish shape, H, to the sac with a narrow mouth is at once clear, and the knowledge that all the Cœlenterates proceed from similar larvæ, and that all the complications of their various systems are developed from a simple gastrula, throws much light on their anatomy. During these transitions the endoderm, whose cells multiply, continues as an uninterrupted lining to the stomach and its appendages, while the ectoderm yields the cuticular elements.
Fig. 351.—Stages in development of Monoxenia Darwinii, × 600.—(Warne.)
A third intermediate gelatinous layer, the mesoglæa, arises between the two layers in which muscles and connective interstitial tissue appear. In the mesoglæa of one species of coral calcification takes place; this internal calcification has but a small share in the work of the great rock-making corals, their most important calcification being external. In Monoxenia, although the transition from the gastrula larva to the adult animal has not been seen, there can be no doubt as to how this is carried out, the transformations having been watched throughout in other species. The larva attaches itself with the end opposite the mouth, the cilia disappear, and after the mouth-tube has been formed by the folding in of the anterior end along the longitudinal axis of the body, and has thus become marked off from the stomach, eight hollow tentacles rise round the mouth as outgrowths of the body cavity, or as direct continuations of the stomach.
Like all other corals, Monoxenia periodically multiply by means of eggs, which are formed either in the walls of the radiating partitions or septa, or along the free edges. These are ejected through the oral opening. As a rule, the polyps are either male or female; but in stock-forming species individuals of the two sexes are often mixed. Monoxenia may be taken as the simplest type of the regularly radiate polyps; in all the different organs being repeated in regular rings round a central axis; the mouth also is circular. From this interesting account, drawn by Haeckel, of a simple polyp, it will be at once seen what kind of radiate animal it is that builds up the coral reefs. “No garden on earth can match the gardens of the sea that circle the northern part of Australia. As the tide ebbs in azure sunset, coral-reefs peer out symmetrically arranged in beds and intersected by emerald pathways coursing through corals of all hues and tints fathoms deep in the channels.”
In a growing polyp-stock the individuals usually remain in organic connection; that is to say, each first provides for itself and then shares its superfluity with others, sometimes by means of a continuous reticulated system of canals perforating the calcareous substance which often separates the members of one stock from another. The whole colony may thus be physiologically one creature with many mouths. There are others that remain single, as the inverted pyramidal-looking bodies, Fungidæ, commonly called “Sea-mushrooms,” found in great variety. The colour of the polypidom is white, of a flattened round shape, made up of thin plates or scales, imbedded in a translucent jelly-like substance, and within is concealed a polyp; the footstalk, by means of which the animal is attached to the rock, is of a calcareous nature ([Fig. 352], No. 1).
Fig. 352.—Sea-Anemones.
1. Actinia rubra, tentacles displayed and retracted; 2. Heticictis bellis; 3. H. bellis, seen from above.
Hexactinia (six-rayed polyps) are not limited to six rays, as the name given them may seem to imply; they are, in fact, very numerous in some of the largest and most gorgeous of the sea-anemones. All are distinguished by their solitary manner of life, their size, and their vivid and variedly beautiful colouring. The endoderm is firm, and when the animal withdraws its tentacles and shuts its body substance in, there is some difficulty in penetrating to the interior. It does not, however, secrete a calcareous skeleton inside or out, as do the true coral polyps. Among the Hexactinia the sea-anemone ([Fig. 352]) takes the first place.
These beautifully coloured creatures are, for the most part, found attached to the spot selected by the larvæ; a few species bore into the sand with the posterior part of the body, or build a sheath, which they inhabit. They are voracious feeders, and devour large pieces of flesh, and even mussel and oysters, sucking them in by means of their long grasping tentacles. Well-fed anemones change their skin frequently, during which process they remain closely retracted; the shed skin forms a loose girdle around the base. Actinia bellis not infrequently attach themselves to the shells of crabs and whelks, and are thus carried to pastures new.
Fig. 353.—Larvæ of Sea-Anemones, Actinia effœta, highly magnified.
On account of the ease with which anemones are kept in captivity, their mode of reproduction can be closely observed. With but few exceptions they develop from eggs, and in the course of a few weeks are hatched into ciliated infusorial larvæ, presenting most curious and exquisite representations of jugs and jars, with cover lids (as seen in [Fig. 353], Actinia effœta). These evince the handiwork of a master hand in the ceramic art. They are, however, of so translucent a nature as to permit of the internal structure being seen to consist of nerves and vessels, and which are rendered more apparent by staining. These settle down in a week or ten days, and then shed their cilia, the first tentacle appearing during the process of attachment.
In some species the young Actiniæ are seen to pass through their whole development within the body cavity of the parent. Most anemones are provided with several circles of more or less cylindrical tentacles, and there are a few specially beautiful species which, besides tentacles of the usual form, have, either within or without the ordinary circle of tentacles, lobed or leaf-like tactile and seizing organs. These belong to the family of the beautiful Crambactis of the Red Sea. Below these grasping tentacles comes a circle of thicker arms unlike the former, being spindle shaped. All the tentacles of the sea-anemones are hollow with a fine aperture at the tip, through which, on closing rapidly, it is seen to expel a jet of water.
True Corals.—It will have been noticed in the foregoing remarks that in the soft body-division of the Hexactinia there are both single individuals and colonies joined together to form stocks. The same diversity in this respect will be found among corals proper, with this difference, that the skeleton-forming polyps, by combining, build up substantial structures in the most secure and advantageous positions. Now it so happens that all the corals found about our coasts are generally small and solitary dwellers, one of the best known of which is the scarlet crisp coral, Flabellum, and is characterised by the slit-like form of the mouth. Viewed sideways it resembles a small fan fastened along the edges, and just inside a row of fully developed tentacles is seen protruding. An interesting form of budding occurs in these corals: the buds fall off, and in this budding condition the coral might pass, and indeed has been described as a different species of Flabellum. The colour of the coral is a beautifully transparent red. Remarkable as the solitary corals are, they are surpassed both in number and in form by those which form compound stocks, that is to say, in which the buds do not fall off, but go on building up coral islands and barrier reefs in the warmer seas. Some very few typical forms only are given in the group accompanying, shown in [Fig. 358].
A different kind of stock is developed in a number of forms, some producing many buds, as in the Madrepores, in which selected polyps spring up above the rest, their sides also becoming covered with small buds, each one of which is a living, feeding, coral animal surrounded by a crown of tentacles. These Madrepores play a very important part in the building up of coral reefs.
Fig. 354. Developmental stages of Larvæ, Astroides calycularis, × 40.
Another massive coral, the Astroides calycularis, has a different mode of growth, the tubes not being fused together. When seen standing out these yellowish-red polyps have been mistaken for small anemones. The larvæ of this coral leave the egg while still in the large chambered body cavity of the parent, where they swim about for a time, till they escape through the mouth. They are worm-like in form, and swim by means of cilia, which are thicker at the foremost end. The mouth first appears after leaving the parent, but as they soon become exhausted by the effort they assume a contracted form, and attach themselves, as do anemones, by pressing the thicker end of the body against a rock, the whole contracting into a thick round disc, while longitudinal furrows become visible at the upper part where the mouth sinks in. At the end of these furrows twelve tentacles appear. The accompanying illustration shows the various stages through which the larvæ pass in rapid succession ([Fig. 354]); at the same time it has already commenced to secrete its calcareous skeleton. This is not formed as a connected whole but from a number of separate centres of secretion formed between the polyp and the substance to which it has attached itself, and which become gradually fused into a perfect skeleton. A section of the polyp at this stage forms an interesting microscopical object.
The so-called eight-rayed corals consist of the one genus Tubipora, the members of which are few in number and not varied in form (Fig. 358, No. 10). In the structure, however, of skeletons they are unique among extant corals. Each individual secretes a smooth-walled tube without calcification of the vertical septa. These tubes, like the pipes of an organ, stand almost parallel, and are united to form a stock by means of transverse platforms. The formation of buds does not appear to take place in this family.
Another of the eight-rayed corals is Gorgoniidæ. These are permanently fixed to the spot on which they are found, and form a bush-like growth, giving no idea of the living coral, as it rises in graceful branching colonies, in deep water, and represents a portion of Gorgonia nobilis with polyps expanded (Figs. 344 and 358, No. 9).
Other corals present numerous other departures from the types we have been considering, but so far modified in form as that of the Sea-pen, Veretillum ([Fig. 355]), the stock part of which is surrounded by polyps continued down a portion of the cylindrical stalk. The best known of the species is Pennatula phosphorea of the Mediterranean.
Fig. 355.
1. Pennatula phosphorea; 2. Synapta chirodata; 3. Anchor-shaped spiculum and plate from the ectoderm of same.
Pennatulidæ.—This family derives its name from penna, a quill. Their spicula also resemble a penholder in appearance, shown in [Fig. 358], No. 3. The polyps are without colour, provided with eight rather long retractile tentacula, beautifully ciliated on the inner aspect with two series of short processes, and strengthened by these crystalline spicula, a row being carried up the stalk, together with a series of ciliated processes. The mouth, occupying the centre of the tentacula, is somewhat angular. The ova lie between the membranous part of the pinnæ; these are globular, of a yellowish colour, and by pressure can be made to pass through the mouth. Dr. Grant wrote:—“A more singular and beautiful spectacle could scarcely be conceived than that of a deep purple Pennatula phosphorea, with all its delicate transparent polyps expanded and emitting their usual brilliant phosphorescent light, sailing through the still and dark abyss, by the regular and synchronous pulsations of the minute fringed arms of the polyps.”
The spicula are seen to be a continuous series of cones fitting into each other.