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THE CLASS AS A WHOLE

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Originally appearing in Volume V05, Page 792 of the 1911 Encyclopedia Britannica.
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THE CLASS AS A WHOLE.—The Chaetopoda are with but few exceptions (Myzostomida in part, Sternaspis) elongated worms, flattened or, more usually, cylindrical, and bilaterally symmetrical. The body consists of a number of exactly similar or closely similar segments, which are never fused and metamorphosed, as in the Arthropoda, to form specialized regions of the body. It is, however, always possible to recognize a head, which consists at least of the peristomial segment with a forward projection of the same, the prostomium. A thorax also is sometimes to be distingui,hed from an abdomen. Where locomotive appendages (the parapodia of the Polychaeta) exist, they are never jointed, as always in the Arthropoda; nor are they modified anteriorly to form jaws, as in that group. The prostomium overhangs the mouth, and is often of considerable size and, as a rule, quite distinct from the segment following, being E, Eye. M, Mouth. d.c, Dorsal cirrus. per, Peristomium, probably equal to two segments, per.c, Peristomial cirri. separated by an external groove, and containing, at least temporarily, the brain, which always arises there. Its cavity also is at first independent of the coelom though later invaded by the latter. 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. It has, however, been alleged that this cavity is formed by a pair of mesoblastic somites (N. Kleinenberg), in which case there is more reason for favouring the view that would assign an equality between the prostomium and the (in that case) other segments of the body. The peculiar prostomium of Tomopteris is described below. The body wall of the Chaetopoda consists of a " dermo-muscular " tube which is separated from the gut by the coelom and its peritoneal walls, except in most leeches. A single layer of epidermic cells, some of which are glandular, forms the outer layer. Rarely are these ciliated, and then only in limited tracts. They secrete a cuticle which never approaches in thickness the often calcified cuticle of Arthropods. Below this is a circular, and below that again a longitudinal, layer of muscle fibres. These muscles are not striated, as they are in the Arthropoda. Setae.—These chitinous, rod-like, rarely squat and then hook-like structures are found in the majority of the Chaetopoda, being absent only in certain Archiannelida, most leeches, and a very few Oligochaeta. They exist in the Brachiopoda (which are probably not unrelated to the Chaetopoda), but otherwise are absolutely distinctive of the Chaetopods. The setae are invariably formed each within an epidermic cell, and they are sheathed in involutions of the epidermis. Their shape and size varies greatly and is often of use in classification. The setae are organs of locomotion, though their large size and occasionally jagged edges in some of the Polychaeta suggest an aggressive function. They are disposed in two groups on either side, corresponding in the Polychaeta to the parapodia; the two bundles are commonly reduced among the earthworms to two pairs of setae or even to a single seta. On the other hand, incertain Polychaeta the bundles of setae are so extensive that they nearly form a complete circle surrounding the body; and in the Oligochaet genus Perichaeta (=Pheretima), and some allies, there is actually a complete circle of setae in each segment broken only by minute gaps, one dorsal, the other ventral. Coelom.—The Chaetopoda are characterized by a spacious coelom, which is divided into a series of chambers in accordance with the general metamerism of the body. This is the typical arrangement, which is exhibited in the majority of the Polychaeta and Oligochaeta; in these the successive chambers of the coelom are separated by the intersegmental septa, sheets of muscle fibres extending from the body wall to the gut and thus forming partitions across the body. The successive cavities are not, however, completely closed from each other; there is some communication between adjoining segments, and the septa are sometimes deficient here and there. 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. The coelom is lined throughout by cells, which upon the intestine become large and loaded with excretory granules, and are known as chloragogen cells. 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. 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: It has been indeed largely upon the conditions characterizing the Chaetopoda that the conception of the coelom in the Coelomocoela has been based. Among the simpler Chaetopoda the coelom retains the character of a series of paired chambers, showing the above relations to the exterior and to the gonads. There are, however, further complications in some forms. Especially are these to be seen in the more modified Oligochaeta and in the much more modified Hirudinea. In the Polychaeta, which are to be regarded as structurally simpler forms than the two groups just referred to, there is but little sub-division of the cqelom of the segments, indeed a tendency in the reverse direction, owing to the suppression of septa. 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. To this pericardial coelom is frequently added a gonocoel enclosing the gonads and the funnels of their ducts. This condition is more fully dealt with below in the description of the Oligochaeta. The division and, indeed, partial suppression of the coelom culminates in the leeches, which in this, as in some other respects, are the most modified of Annelids. Nervous System.—In all Chaetopods this system consists of cerebral ganglia connected by a circumoesophageal commissure with a ventral ganglionated cord. The plan of the central nervous system is therefore that of the Arthropoda. Among the Archiannelida, in Aeolosoma and some Polychaetes, the whole central nervous system remains imbedded in the epidermis. In others, it lies in the coelom, often surrounded by a special and occasionally rather thick sheath. The cerebral ganglia constitute an archicerebrum for the most part, there being no evidence that, as in the Arthropoda, a movement forward of post-oral ganglia has taken place. In the leeches, however, there seems to be the commencement of the formation of a syncerebrum. In the latter, the segmentally arranged ganglia are more sharply marked off from the connectives than in other Chaetopods, where nerve cells exist along the whole ventral chain, though more numerous in segmentally disposed swellings. Vascular System.—In addition to the coelom, another system of fluid-holding spaces lies between the body wall and the gut in the Chaetopoda. This is the vascular or haemal system (formerly and unnecessarily termed pseudhaemal). With a few exceptions among the Polychaeta the vascular system is always present among the Chaetopoda, and always consists of a system of vessels with definite walls, which rarely communicate with the coelom. It is in fact typically a closed system. The larger trunks open into each other either directly by cross branches, or a capillary system is formed. There are no lacunar blood spaces with ill-defined or absent walls except for a sinus surrounding the intestine, which is at least frequently present. The principal trunks consist of a dorsal vessel lying above the gut, and a ventral vessel below the gut but above the nervous cord. These two vessels in the Oligochaeta are united in the anterior region of the body by a smaller or greater number of branches which surround the oesophagus and are, some of them at least, contractile and in that case wider than the rest. The dorsal vessel also communicates with the ventral vessel indirectly by the intestinal sinus, which gives off branches to both the longitudinal trunks, and by tegementary vessels and capillaries which supply the skin and the nephridia. In the smaller and simpler forms the capillary networks are much reduced, but the dorsal and ventral vessels are usually present. The former, however, is frequently A f s /s/ rc. B ^\ B, dorsal view of the same. pt, Prostomial palp. pp, Parapodium. pr, Prostomium. pr.t, Prostomial tentacle. t.s, Trunk segment. v.c, Ventral cirrus. developed only in the anterior region of the body where it emerges from the peri-intestinal blood sinus. On the other hand, additional longitudinal trunks are sometimes developed, the chief one of which is a supra-intestinal vessel lying below the dorsal vessel and closely adherent to the walls of the oesophagus in which region it appears. The capillaries sometimes (in many leeches and Oligochaeta) extend into the epidermis itself. Usually they do not extend outwards of the muscular layers of the body wall. The main trunks of the vascular system often possess valves at the origin of branches which regulate the direction of the blood flow. Among many Oligochaeta the dorsal blood-vessel is partly or entirely a double tube, which is a retention of a character shown by F. Vezhdovsky to exist in the embryo of certain forms. The blood in the Chaetopoda consists of a plasma in which float a few corpuscles. The plasma is coloured red by haemoglobin: it is sometimes (in Sabella and a few other Polychaeta) green, which tint is due to another respiratory pigment. The plasma may be pink (Magelona) or yellow (Aphrodite) in which cases the colour is owing to another pigment. In Aeolosoma it is usually colourless. The vascular system is in the majority of Chaetopods a closed system. It has been asserted (and denied) that the cellular rod which is known as the " Heart-body " (Herzkorper), and is to be found in the dorsal vessel of many Oligochaeta and Polychaeta, is formed of cells which are continuous with the chloragogen cells, thus implying the existence of apertures of communication with the coelom. The statement has been often made and denied, but it now seems to have been placed on a firm basis (E. S. Goodrich), that among the Hirudinea the coelom, which is largely broken up into narrow tubes, may be confluent with the tubes of the vascular system. This state of affairs has no antecedent improbability about it, since in the Vertebrata the coelom is unquestionably confluent with the haemal system through the lymphatic vessels. Finally, there are certain Polychaeta, e.g. the Capitellidae, in which the vascular system has vanished altogether, leaving a coelom containing haemoglobin-impregnated corpuscles. It has been suggested (E. Ray Lankester) that this condition has been arrived at through some such intermediate stage as that offered by Polychaet Magelona. In this worm the ventral blood-vessel is so swollen as to occupy nearly the whole of the available coelom. Carry the process but a little farther and the coelom disappears and its place is taken by a blood space or haemocoel. It has been held that the condition shown in certain leeches tend to prove that the coelom and haemocoel are primitively one series of spaces which have been gradually differentiated. The facts of development, however, prove their distinctness, though those same facts do not speak clearly as to the true nature of the blood system. One view of the origin of the latter (largely based upon observations upon the development of Polygordius) sees in the blood system a persistent blastocoel. F. Vezhdovsky has lately seen reasons for regarding the blood system as originating entirely from the hypoblast by the secretion of fluid, the blood, from particular intestinal cells and the consequent formation of spaces through pressure, which become lined with these cells. Nephridia and Coelomoducts.—The name " Nephridium " was originally given by Sir E. Ray Lankester to the members of a series of tubes, proved in some cases to be excretory in nature, which exist typically to the number of a single pair in most of the segments of the Chaetopod body, and open each by a ciliated orifice into the coelom on the one hand, and by a pore on to the exterior of the body on the other. In its earlier conception, this view embraced as homologous organs (so far as the present group is concerned) not only the nephridia of Oligochaeta and Hirudinea, which are obviously closely similar, but the wide tubes with an intercellular lumen and large funnels of certain Polychaeta, and (though with less assurance) the gonad ducts in Oligochaeta and Hirudinea. The function of nitrogenous excretion was not therefore a necessary part of the view—though it may be pointed out that there are grounds for believing that the gonad ducts are to some extent also organs of excretion (see below). Later, the investigations of E. Meyer and organs are present to the number of a single pair per somite, and are E. S. Goodrich, endorsed by Lankester, led to the opinion that under commonly present in the majority of the segments of the body, the general morphological conception of " nephridium " were failing often among the Oligochaeta in a varying number of the included two distinct sets of organs, viz. nephridia and coelomo- anterior segments. They are considerably reduced in number in ducts. The former (represented by, e.g. the " segmental organs " certain Polychaeta. Essentially,a nephridium is a tube,generally very of Lumbricus) have been asserted to be " ultimately, though not long and much folded upon itself, composed of a string of cells placed always, actually traceable to the ectoderm "; the latter (repre- end to end in which the continuous lumen is excavated. Such cells sented by, e.g. the oviduct of Lumbricus) are parts of the coelomic wall itself, which have grown out to the exterior. The nephridia, in fact, on this view, are ectodermic ingrowths, the coelomoducts coelomic outgrowths. The cavity of the former has nothing to do with coelom. The cavity of the latter is coelom. The embryological facts upon which this view has been based, however, have been differently interpreted. According to C. O. Whitman the entire nephridial system (in the leech Clepsine) is formed by the differentiation of a continuous epiblastic band on each side. The exact opposite is maintained by R. S. Bergh (for Lumbricus and Criodrilus), whose figures show a derivation of the entire nephridium from mesoblast, and an absence of any connexion between successive nephridia by any continuous band, epiblastic or mesoblastic. A midway position is taken up by Wilson, who asserts the mesoblastic formation of the funnel, but also asserts the presence of a continuous band of epiblast from which certainly the terminal vesicle of the nephridium, and doubtfully the glandular part of the tube is derived. Vezhdovsky's figures of Rhynchelmis agree with those of Bergh in showing the backward growth of the nephridium from the funnel cell. There are thus substantial reasons for believing that the nephridium grows backwards from a funnel as does the coelomoduct. It is therefore by no means certain that so profound a difference embryologically can be asserted to exist between the excretory nephridia and the ducts leading from the coelom to the exterior, which are usually associated with the extrusion of the genital products among the Chaetopoda. There are, however, anatomical and histological differences to be seen at any rate at the extremes between the undoubted nephridia of Goodrich, Meyer and Lankester, and the coelomoducts of the same authors. i. Nephridia.—Excretory organs which are undisputed nephridia are practically universal among the Oligochaeta, Hirudinea and Archiannelida, and occur in many Polychaeta. Their total absence has been asserted definitely only in Paranais littoralis. Usually these rr e es C A, Diagram of the nephridium the nephridium of Nephthys of Nereis diversicolor. scolopendroides. B, Diagram of the nephridium of c.s, Cqt surface. Alciope, into which opens the cst, Coelomostome. large genital funnel (coelo- f, Flagellum. mostome). g.f, Genital funnel. C, Small portion of the nephri- n, Neck of solenocyte. dium of Glycera siphono- n.c, Nephridial canal. stoma, showing the canal cut n.p, Nephridiopore. through, and the solenocytes nst, Nephridiostome. on the outer surface. nu, Nucleus of solenocyte. D, Optical section of a branch of s, Solenocytes. t, Tube. are termed " drain pipe " cells. Frequently the lumen is branched and may form a complicated anastomosing network in these cells. Externally, the nephridium opens by a straight part of the tube, which is often very wide, and here the intracellular lumen becomes intercellular. 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. This is so with Aeolosoma (Vezhdovsky). The condition is interesting as a persistence of the conditions obtaining in the provisional nephridia of e.g. Rhynchelmis, which afterwards become by an enlargement and opening up of the funnel the permanent nephridia of the adult worm. In some Polychaets (e.g. Glycera, see fig. 2) there are many of these flame cells to a single nephridium which are specialized in form, and have been termed " solenocytes " (Goodrich). They are repeated in Polygordius, and are exactly 792 to be compared with similarly-placed cells in the nephridia of Amphioxus. 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. The funnel varies greatly in size and number of its component cells. There are so many differences of detail that no line can be drawn between the one-celled funnel of Aeolosoma and the extraordinarily large and folded funnel of the posterior nephridia in the Oligochaete Thamnodrilus. In the last-mentioned worm the funnels of the anterior nephridia are small and but few celled; it is only the nephridia in and behind the 17th segment of the body which are particularly large and with a sinuous margin, which recall the funnels of the gonad ducts (i.e. coelomoducts). Among the Polychaeta the nephridium of Nereis (see fig. z) is like that of the Oligochaeta and Hirudinea in that the coiled glandular tube has an intracellular duct which is ciliated in the same way in parts. The Polychaeta, however, present us with another form of nephridium seen, for example, in Arenicola, where a large funnel leads into a short and wide excretory tube whose lumen is inter-cellular. In the young stages of this worm which have been investigated by W. B. Benham, the tube, though smaller, and with a but little pronounced funnel, has still an intercellular duct. That these organs in Polychaeta serve for the removal of the generative products to the exterior is proved not only by the correspondence in number to them of the gonads, but by actual observation of the generative products in transit. This form of nephridia leads to the shorter but essentially similar organs in the Polychaete Sternaspis, and to those of the Echiuroidea (q.v.) and of the Gephyrea (q.v.). Though the paired arrangement of the nephridia is the prevalent one in the Chaetopoda, there are many examples, among the Oligochaeta, of species and genera in which there are several, even many, nephridia in each segment of the body, which may or may not be connected among themselves, but have in any case separate orifices on to the exterior. 2. Coelomoducts.—In this category are included (by Goodrich and Lankester) the gonad ducts of the Oligochaeta, certain funnels without any aperture to the exterior that have been detected in Nereis, &c., funnels with wide and short ducts attached to nephridia in other Polychaeta, gonad ducts in the Capitellidae, the gonad ducts of the leeches. In all these cases we have a duct which has a usually wide, always intercellular, lumen, generally, if not always, ciliated, which opens directly into the coelom on the one hand and on to the exterior of the body on fhe other. These characters are plain in all the cases cited, excepting only the leeches which will be considered separately. There is not a great deal of difference between most of these structures and true nephridia. It is not clear, for example, to which category it is necessary to refer the excretory_ organs of Arenicola, or Polynoe. 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 origin-ally 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. It is by no means certain that a hard and fast line can be drawn between intra- and intercellular lumina. Finally, in function there are some points of likeness. 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. There is no possibility that sperm and ova can escape by these tubes not in company with coelomic fluid. In the case of many Oligochaeta where there is no vascular network surrounding the nephridium, this function must be the chief one of those glands, the more elaborate process of excretion taking place in the case of nephridia surrounded by a rich plexus of blood capillaries. A consideration of the mode of development and appearance of the coelomoducts that have thus far been enumerated (with the possible exception of those of the leeches) seems to show that there is a distinct though varying relation between them and the nephridia. It has been shown that in Tubifex, and some other aquatic Oligochaeta, the genital segments are at first provided with nephridia, and that these disappear on the appearance of the generative ducts, which are coelomeducts. In Lumbricus the connexion is a little closer; the funnel of the nephridium, in the segments in which the funnels of the gonad ducts are to be developed, persists and is continuous with the gonad duct funnels on their first appearance. In the development of the Acanthodrilid earthworm Octochaetus (F. E. Beddard) the funnels of the pronephridia disappear except in the genital segments, where they seem to be actually converted into the genital funnels. At the least there is no doubt that the genital funnels are developed precisely where the nephridial funnels formerly existed. If the genital funnels are not wholly or partly formed out of the nephridial funnels they have replaced them. In the genital segments of Eudrilus the nephridia are present, but the funnels have not been found though they are obvious in other segments. Here also the genital funnels have either replaced or been formed out of nephridial funnels. In Haplotaxis helerogyne(W. B. Benham) the sperm ducts are hardly to be distinguished from nephridia; they are sinuous tubes with an intra-cellular duct. But the funnel is large and thus differs from the funnels of the nephridia in adjoining segments. Here again the nephridial funnel seems to have been converted into or certainly replaced by a secondarily developed funnel. This example is similar to cases among the Po1vchaeta where a true nephridium is provided with a large funnel, coelomostome, according to the nomenclature of Lankester. The whole organ, having, as is thought but not known, this double origin, is termed a nephromixium. The various facts, however, seem to be susceptible of another interpretation. It may be pointed out that the several examples described recall a phenomenon which is not uncommon and is well known to anatomists. That is the replacement of an organ by, sometimes coupled with its partial conversion into, a similar or slightly different organ performing the same or an analogous function. Thus the postcaval vein of the higher vertebrata is partly a new structure altogether, and is partly formed out of the pre-existing posterior cardinals. The more complete replacements, such as the nephridia of the genital segment of Tubifex by a subsequently formed genital duct, may be compared with the succession of the nesonephros to the pronephros in vertebrates, and of the metanephros to the mesonephros in'the higher vertebrates. It might be well to term these structures, mostly serving as gonad ducts, which have an undoubted resemblance to nephridia, and for the most part an undoubted connexion with nephridia, " Nephrodinia," to distinguish them from another category of " ducts " which are communications between the coelom and the exterior, and which have no relation whatever to nephridia or to the organs just discussed. For these latter, the term coelomoducts might well be reserved. 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. To this category will belong the oviducts in Teleostean fishes and probably the gonad ducts in several groups of invertebrates.
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