Coelom

“ Coelomula of sagitta (gastrula with a couple of coelom-pouches. (From Kowalevsky.) bl.p primitive mouth, al primitive gut, pv coelom-folds, m permanent mouth.)
The four secondary germinal layers are so distributed in the structure of the body in all the coelomaria (or all metazoa that have a body-cavity) that the outer two, joined fast together, constitute the body-wall, and the inner two the ventral wall; the two walls are separated by the cavity of the coelom. Each of the walls is made up of a limiting layer and a middle layer. ”

“ Soc., 6, 1886, p. 27) (1) that the coelom of Peripatus gives rise to the nephridia and generative glands only, and to no other part of the body-cavity of the adult, (2) that the nephridia of the adult do not open as had been supposed into the body-cavity, (3) that the body-cavity is entirely formed of the blood-containing space, the coelom having no perivisceral portion. These results were extended by the same author (Quart. Journ. Mic. Sci., 27, 1887, pp. 486-540) to other Arthropods and to the Mollusca, and the modern theory of the coelom was finally established. ”

“ This cavity, common to all the higher animals, is known as the coelom; it is lined by cells, which cover the intestines as well as the inside of the body-wall; and upon the intestine assume the form so characteristic of the group, namely, that of large yellow cells loaded with secreted matters, and called "chloragogen-cells" by Claparède. The coelom communicates with the exterior by means of the dorsal pores, the nephridia, and the ducts of the reproductive organs. As in all animals which possess a coelom, the reproductive tissues, ova and sperm, are developed on its walls. ”

“ But however many sacs may arise from the primitive enteron, and however these sacs may ultimately divide up and arrange themselves, the important point of development common to all these animals, about which there can be no dispute, is that the coelom is a direct differentiation of a portion of the enteron. ”

“ The coelom, an organ which is found in the so-called coelomate animals, and which in the adult is usually divided up more or less completely into three parts, namely, body-cavity, renal organs, generative glands, presents in different animals both these methods of development. In certain animals it develops by the direct modification of a part of the primitive enteron, while in others it arises by the gradual shaping of a mass of tissue which consists of a compact mass of nuclei derived by nuclear proliferation from one or more of the pre-existing tissues of the body. ”

“ Several forms of cells float freely in the fluid of the coelom. In another sense also the coelom is not a closed cavity, for it communicates in several ways with the external medium. Thus, among the Oligochaeta there are often a series of dorsal pores, or a single head pore, present also among the Polychaeta (in Ammochares) . In these and other Chaetopods the coelom is also put into indirect relations with the outside world by the nephridia and by the gonad ducts. ”

“ Since the connexion between the coelom and the excretory organs is always a very close one throughout the animal kingdom, interest naturally centres upon the renal organs in Crustacea, and it has been suggested that these organs in Crustacea represent the sole remains, with the possible exception of the gonads, of the coelom. Since, at any rate, a part of the kidneys appears to be developed as a closed sac in the mesoderm, and since they possess a possible segmental value, this suggestion is plausible ”

“ Similarly an anal aperture is formed in connexion with a special invagination which meets the hinder part of the met-enteron, and is termed the proctodaeum.
The coelom is primarily and essentially the generative cavity: the reproductive cells arise from its walls, i.e. from the coelomic epithelium. True nephridia do not primarily open into the coelom, as was formerly taught, but are intra-cellular ducts in the mesoderm. Such organs are absent in Mollusca in the adult state, but a pair of nephridia usually occurs in the larva. ”

“ Regarding this as a collar-cavity, it becomes possible to agree with Masterman that the region shown in fig. 4, 1. between the tentacles and the praeoral hood, is really a collar the coelom of which develops relatively late. It will be noticed that the lophophore of Phoronis is, on this assumption, a derivative of the collar just as it is in the Pterobranchia. The epistome of the adult Phoronis cannot well be the proboscis since its cavity is continuous with the lophophoral coelom, and because the praeoral hood of Actinotrocha is entirely lost at the metamorphosis. ”

“ Since the skeleton of the corona is composed of plates immovably connected together, muscles corresponding to the ambulacral muscles of the Asteroids would be useless, and so the wall of the perihaemal canal remains thin and the side of it turned towards the general coelom develops no muscles, and that turned towards the nerve-cord no nerve-cells. Where, however, the radial nerve enters the nerve-ring, and on the ring itself, an inner layer of nerve-cells is developed from the lantern-coelom which represents the lower or oral portions of the radial perihaemal canals. ”

“ Evolution of the Echinoderms.—It is reasonable to suppose that the Coelomata—animals in which the body-cavity is divided into a gut passing from mouth to anus and a hollow (coelom) surrounding it—were derived from the simpler Coelentera, in which the primitive body-cavity (archenteron) is not so divided, and has only one aperture serving as both mouth and anus. ”

“ In the Oligochaeta there is a closer correspondence between external metamerism and the divisions of the coelom than is apparent in some Chaetopods. The external segments are usually definable by the setae; and if the setae are absent, as in the anterior segments 796 of several Geoscolicidae, the nephridiopores indicate the segments; to each segment corresponds internally a chamber of the coelom which is separated from adjacent segments by transverse septa, which are only unrecognizable in the genus Aeolosoma and in the head region of other Oligochaeta. ”

“ Relationship between protractor muscles and the peripharyngeal coelom
Because the lobes of frilled protractor muscles are entirely immersed in coelomic fluid (Fig. 2A–C) , the precise location of the protractor muscles and their interplay with the compartments of the peripharyngeal coelom are important in the context of this study. The interradial cavity is indirectly in contact with seawater through the so-called buccal sacs (Fig. 4) . ”

“ The anterior coelom communicates with the exterior by a short pore-canal, and later forms a connexion, the stone-canal, with the hydrocoel. At the apex of the prae-oral lobe there is formed a thickened patch of ectoderm, bearing stiff sensory hairs, and having at their bases nerve-fibres and ganglion cells. This larval brain corresponds to that of the Tornaria and Echinopluteus. Behind the brain there is a glandular pit, which is used for fixation, and recalls the similar organ in the Bipinnaria. ”

“ Both series of organs consist essentially of a ciliated tube leading from the coelom to the exterior. Both series of organs grow back centrifugally from the funnel. In both the cavity originally or immediately continuous with the coelom appears first in the funnel and grows backwards. In some cases, e.g. oviducts of Oligochaeta, sperm ducts of Phreoryctes, the coelomoducts occupy, like the nephridia, two segments, the funnel opening into that in front of the segment which carries the external pore. ”

“ An increased precision was given to the conception of coelom by the discovery in 1880 (Quart. Journ. Mic. Sci., 20, p. 164) that the nephridia of Elasmobranchs are a direct differentiation of a portion of it. ”

“ Analysis of Echinoderm Characters.—Regarding the Echinoderms as a whole in the light of the foregoing account, we may give the following analytic summary of the characters that distinguish them from other coelomate animals:—
They live in salt or brackish water; a primitive bilateral symmetry is still manifest in the right and left divisions of the coelom; the middle coelomic cavities are primitively transformed into two hydrocoels communicating with the exterior indirectly through a duct or ducts of the anterior coelom ”

“ Moreover, upon the intestine the coelomic cells are modified into chloragogen cells. In Acanthobdella the testes are, however, not contained in the general coelom, and the nephridia lie in the septa. It is remarkable, in view of the spaciousness of the coelom, that the funnels of the latter have not been seen. Ozobranchus possesses a coelom which is less typically chaetopodous than that of Acanthobdella, but more so than in other leeches. ”

“ In the case of the pronephros the glomeruli are as a rule fused together into a single glomus. In the mesonephros they remain separate and in this case the portion of coelom surrounding the glomerulus tends to be nipped off from the general coelom—to form a Malpighian body. The separation may be incomplete—the Malpighian coelom remaining in connexion with the general coelom by a narrow peritoneal canal. ”

“ The Starfish possesses no definite kidney for getting rid of nitrogenous waste. In most of the higher animals with a well-developed coelom it has been proved that the kidney is simply a specialised portion of the coelom, and in many cases some parts of the coelomic wall still retain their excretory functions, which apparently the whole originally possessed. ”

“ In the Cyclostomes (e.g. Petromyzon) the eggs and spermatozoa are discharged from the gonads into the coelom, whence they reach the exterior through a pair of "genital pores" leading from the hinder end of the coelom into a urinogenital sinus formed by the united extremities of the two archinephric ducts. [479] Myxine has, however, but a single median pore, opening into an integumentary cloaca, which also receives the rectal and urinary orifices. Bdellostoma has two such pores communicating with a similar cloaca. ”

“ The chief difference between the coelomation of the acrania (amphioxus) and the other vertebrates (with skulls—craniotes) is that the two coelom-folds of the primitive gut in the former are from the first hollow vesicles, filled with fluid, but in the latter are empty pouches, the layers of which (inner and outer) close with each other. In common parlance we still call a pouch or pocket by that name, whether it is full or empty. It is different in ontogeny; in some of our embryological literature ordinary logic does not count for very much. ”

“ The chief difference is that the gonad or generative portion of the coelom is single and median, opening into the pericardium by a single posterior aperture. The excretory organs or coelomoducts arise from the posterior corners of the pericardium, run forwards and then backwards to open by separate apertures lateral to the gills (fig. 5, A) . There are no accessory generative organs. ”

“ A clitellum is present. The segments of body are few (not more than thirty-four) and fixed in number. The anus is dorsal. One or two (anterior and posterior) suckers always present. Nervous system always in coelom. Coelom generally reduced to a system of tubes, sometimes communicating with vascular system; in Acanthobdella and Ozobranchus a series of metamerically arranged chambers as in Oligochaeta. Nephridia always paired, rarely (Pontobdella) forming a network communicating from segment to segment; lumen of nephridia always intracellular, funnels pervious or impervious. ”

“ The two lateral lobes contain the coelom; each separates off in front a segment which forms the head and presumably then divides again to form anteriorly the trunk, and posteriorly the tail regions. An interesting feature of the development of Chaetognaths is that, as in some insects, the cells destined to form the reproductive organs are differentiated at a very early period, being apparent even in the gastrula stage.
The great bulk of the group is pelagic, as the transparent nature of all their tissues indicates. They move by flexing their bodies. ”

“ Like so many of the latter, the Echinodermata have an anus, that is, a second opening to the alimentary canal through which indigestible material is rejected; like them also, they have a body-cavity or coelom surrounding the alimentary canal—from the lining of which the genital cells are developed. On the other hand, there is no definite circulatory system, nor any specialised excretory organ, and the nervous system exhibits no concentration which could be called a brain, and is, moreover, in close connexion with the skin. In all these points the Echinodermata resemble the Coelenterata. ”

“ To this category belong certain sacs and pouches in many, perhaps most, genera of the Oligochaeta family, Eudrilidae, and possibly the gonad ducts in the Hirudinea. As an example of the former it has been shown (Beddard) that a large median sac in Lybiodrilus is at first freely open to the coelom, that it later becomes shut off from the same, that it then acquires an external orifice, and, finally, that it encloses the ovary or ovaries, between which and the exterior a passage is thus effected. ”

“ Inasmuch as the first rudiment of the coelom nearly always makes its appearance at an early stage, when the ectoderm and endoderm are almost the only tissues present, and as it then bulks relatively very large and frequently contains within itself the potential centres of growth of other organs, e.g. ”