A further fact must be considered in deciding this question, which is the discovery of ramifying coelomic tubes, approaching close to, but not entering, the epidermis in the Polychaete Arenicola.
A general nerve-plexus probably exists over considerable parts of the skin, and there are special nervous concentrations in the region of the epistome and along a double crescent (N) which follows the parietal attachment of the coelomic septum.
All living Echinoderms have a lacunar, haemal system of diverse origin; this, the ambulacral system, and the coelomic cavities, contain a fluid holding albumen in solution and carrying numerous amoebocytes, which are developed in special lymph-glands and are capable of wandering through all tissues.
Although within the atrial cavity, it is separated from the latter by a narrow coelomic space, bounded towards the atrium by coelomic and atrial epithelium.
Among the Amphineura we find one pair of coelomic ducts in the Aplacophora, two pairs in the Chitons.
Among the Oligochaeta the dorsal vessel in Dinodrilus and Megascolides is enclosed in a separate coelomic chamber which may or may not communicate with the main coelomic cavity.
As each genital gland enlarges it remains attached to the rest of the intermediate cell mass by a constricted fold of the coelomic membrane, known as the mesorchium in the male, and the mesovarium in the female.
Blochmann regards as coelomic, but it must be remembered that his interpretation rests largely on histological grounds, and at present embryological confirmation is wanting.
Even if it be admitted that the postseptal space may be the metasomatic cavity, the praeseptal space can hardly be regarded as coelomic in nature, since it is in continuity with the vascular system; while Masterman's conclusion that the cavity of the praeoral hood (the supposed proboscis-cavity) is separated from that of the supposed collar has received no confirmation.
Even the ectoderm can rarely be recognized as an obvious epithelium except in regions where budding is taking place, while muscular layers are always absent and a coelomic epithelium can seldom be observed.
From this the blood passes into two lateral vessels which pierce the coelomic septum (s.), the right vessel proceeding on the anterior side of the oesophagus, as shown in fig.
I, Coelom of the first somite It also shows us that the neurowhich carries the anten meres, no less than the embryonic nae and is in front of the coelomic cavities, point to the existmouth.
In 1883-1886 Bateson showed by his embryological researches that the Enteropneusta exhibit chordate (vertebrate) affinities in respect of the coelomic, skeletal and nervous systems as well as in regard to the respiratory system, and, further, that the gill-slits are formed upon a plan similar to that of the gillslits of Amphioxus, being subdivided by tongue-bars which depend from the dorsal borders of the slits.
In animals which exhibit typical segmentation or metamerism, such as segmented worms (Chaetopoda), each segment or metamere possesses its own coelomic cavity, a pair of coelomic ducts, and a pair of nephridia.
In any case the cavity of the prostomium is single, and not formed, as is the cavity of the segments of the body, by paired coelomic chambers.
In favour of regarding the vascular system as totally absent, is the fact that the median coelomic channels contain no dorsal and ventral vessel.
In Heliodrilus the blind extremity of the spermatheca is enclosed in a coelomic sac which is in connexion with the sacs envolving the ovaries and oviducts.
In the male there are a right and a left protrusible penis in every genital segment, into which opens the nephridium and a sperm-sac. The wide funnels of the nephridia of this region are possibly of coelomic origin.
In the Phylactolaemata the outermost layer of the bodywall is a flexible, uncalcified cuticle or "ectocyst," beneath which follow in succession the ectoderm, the muscular layers and the coelomic epithelium.
In these features, and in the fact that the gonads are local proliferations of the coelomic epithelium, which have undergone no further changes in the simpler forms, the coelom of this group shows in a particularly clear fashion the general characters of the coelom in the higher Metazoa.
Into the space between the walls of the coelom and the outer body-wall, originally filled with jelly, definite cells now wandered, chiefly derived from the coelomic walls.
It does not seem to be coelomic.
It has, moreover, been shown (see especially Goodrich, 5) that shortly before its metamorphosis, Actinotrocha develops a coelomic space which lies immediately in front of the oblique septum, and gives rise later to the cavity of the lophophore and tentacles.
It is possible therefore to regard the latter condition as secondary, and to conclude that the separate genital ducts have been derived from the original single pair of coelomic ducts, as in Lamellibranchs.
It seems likely that the coelomic pore-canals were originally excretory organs, but in the existing Enteropneusta the pore-canals (especially the collar canals) have, as we have seen, acquired new functions or become vestigial, and the function of excretion is now mainly accomplished by a structure peculiar to the Enteropneusta called the glomerulus, a vascular complex placed on either side of the anterior portion of the stomochord, projecting into the proboscis-coelom.
Montgomery has also described certain spaces which may be coelomic lying between the alimentary canal and the inner longitudinal layer of muscles in the Heteronemertini.
More usually, and indeed in nearly every other case among the Oligochaeta and Hirudinea, the coelomic aperture of the nephridium consists of several cells, ciliated like the nephridium itself for a greater or less extent, forming a funnel.
Moreover, upon the intestine the coelomic cells are modified into chloragogen cells.
No doubt can be entertained that the greater part of the inner layer corresponds to the mesoderm of more ordinary embryos, for the coelomic pouches, the germ-cells, the musculature and the vascular system all arise from it.
On the whole, then, the most probable conclusion is that the original ancestral form of the Mollusca was unsegmented, possessed one pair of true nephridia, and one pair of coelomic ducts whose function was to conduct the generative products to the exterior.
Previously to this, Lankester's pupil Gulland had shown (1885) that in the embryo the coxal gland is a comparatively simple tube, which opens to the exterior in this position and by its other extremity into a coelomic space.
Quite independent of these are certain large dorsally situate funnel-like folds of the coelomic epithelium, ciliated, but of which no duct has been discovered leading to the exterior.
Rarely the nephridium does not communicate with the coelom; in such cases the nephridium ends in a single cell, like the "flame cell" of a Platyhelminth worm, in which there is a lumen blocked at the coelomic end by a tuft of fine cilia projecting into the lumen.
Recent research, however, points to these cords of the rete testis et ovarii as being derived from the coelomic epithelium instead of from the mesonephros.
Segmentation is complete, a gastrula is formed, the blastopore closes, the archenteron gives off two coelomic sacs which, as far as is known, are unaffected by the super ficial segmentation of the body that divides the larva into three segments.
Since the proboscis is a purely larval organ in this genus, it may be supposed that the coelomic space which properly belongs to it fails to develop, but that the praeoral hood itself is none the less the morphological representative of the proboscis.
The first prosthomere has its appendages represented by the compound eyes and a protocerebrum, the second has the antennae for its appendages and a deutocerebral neuromere, the third has suffered suppression of its appendages (which corresponded to the second pair of antennae of Crustacea), but has a tritocerebrum and coelomic chamber.
The genital products were derived from the lining of the coelomic cavities, but it would not be safe to say that any particular region was as yet specialized for generation.
The gonad ducts of Lumbricus, &c., must perform one function of nephridia; they must convey to the exterior some of the coelomic fluid with its disintegrated products of waste.
The gut has no coelomic space surrounding it.
The large sacs which have been termed vagina are suggestive of the large coelomic spermathecae in Eudrilids, a comparison which needs, however, embryological data, not at present forthcoming, for its justification.
The ligamentum denticulatum is thus lined on one side by the epiblastic atrial epithelium, and on the other by mesoblastic coelomic epithelium.
The nephridia, in fact, on this view, are ectodermic ingrowths, the coelomoducts coelomic outgrowths.
The network is, however, of the duct of the nephridium, possibly ectodermic in origin, and does not affect the glandular tubes which remain undivided and with one coelomic funnel each.
The postseptal division is a coelomic space, partially subdivided by a ventral mesentery.
The structure of the Mollusca in the greater number of cases agrees with the hypothesis that the primitive form was unsegmented, and therefore had but one pair of coelomic ducts and one pair of nephridia.
The system of blood-vessels is entirely separate from the coelomic cavities.
The three prosthomeres or prae-oral somites of Crustacea due to the sinking back of the mouth one somite farther than in Arachnida are not clearly indicated by coelomic cavities in the embryo, but their existence is clearly established by the development and position of the appendages and by the neuromeres.
There are thus two pairs of coelomic ducts.
There is further a great tendency for the endothelial cells to form muscles, and this is especially pronounced in the small arm-sinus, where a conspicuous muscle is built up. The mantle-sinuses which form the chief spaces in the mantle are diverticula of the main coelomic cavity.
There is no blood system, and the coelomic corpuscles contain FIG.
There is no possibility that sperm and ova can escape by these tubes not in company with coelomic fluid.
There is usually one pair of coelomic ducts leading from the pericardium to the exterior, and these are the excretory organs or kidneys, formerly known as the organs of Bojanus.
There should also be only a single coelom, or a pair of lateral coelomic cavities.
These are suspended to the muscular bodywall by a double membrane, called the ligamentum denticulatum, which forms at once the roof of the atrial chamber and the floor of a persistent portion of the original body-cavity or coelom (the dorsal coelomic canal on each side of the pharynx).
These are the atrial coelomic funnels or brown funnels, so called on account of the characteristic pigmentation of their walls.
These break off and, lying in the coelomic fluid, break up into spermatozoa.
These cavities communicate with the exterior through the gonad ducts, which have nothing to do with them, but whose coelomic funnels are taken up by them in the course of their growth.
These ducts, however, as well as the coelomic epithelium, may assume excretory functions.
These often have the form of prisms of calcite surrounded by a cuti cular meshwork; the whole is nourished and kept alive by processes, which in Crania are branched; these perforate the shell and permit the access of the coelomic fluid throughout its substance.
These regions of the coelom end at the ends of the body and communicate with each other by means of a branched system of coelomic sinuses, which are in places very fine tubes.
These sinuses are completely shut off from all other cavities, they do not open into the main coelomic space nor into the small arm-sinus, nor does the right sinus communicate with the left.
They are essentially spherical, pear-shaped or oval sacs opening on to the exterior but closed at the coelomic end.
They are said to be developed from the coelomic epithelium which covers the efferent vessel or its caeca.
They invariably disappear before the adult stage is reached, but their presence in the larva is evidence that the ancestral mollusc possessed a pair of true nephridia quite distinct from the coelomic excretory organs, which are so characteristic of existing forms in the adult condition.
This segment is indicated embryologically by its paired coelomic cavities.
This segregation of the germ cells and their independence of the intestinal sac is an indication that the origin of these cells is not coelomic nor enteric, and until we possess further information as to the evolution of the complex genitalia of the higher Turbellaria we cannot hope to understand the presence of such highly modified structures in animals of an otherwise low grade or organization.
Thus in the Chaetopoda the perivisceral cavity is coelomic; in this respect the group contrasts with the Arthropoda and Molluscs, where the perivisceral cavity is, mainly at least, part of the vascular or haemal system, and agrees with the Vertebrata.
Which of the coelomic cavities this last is connected with is uncertain, for there is considerable doubt as to the origin of the genital glands in the embryonic development of recent echinoderms. It seems clear, however, that there was but a single duct and a single bunch of reproductive cells, as in the holothurians, though perhaps bifurcate, as in some of those animals.