The Nephridium of Lumbricus and Its Blood- Supply; with Remarks on the Nephridia in Other Chsetopoda
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THE NEPHRIDITJM OF LUMBRIOUS. 293 The Nephridium of Lumbricus and its Blood- supply; with Remarks on the Nephridia in other Chsetopoda. By W, Blaxland Benham, D.Sc. (Loud.), Anatomical Department, Oxford. With Plates XXIII—XXV. THE nephridium of the common earthworm has recently been the subject of a contribution by Dr. Groehlich (20), in which he remarks on various statements made by Gegenbaur in his classical paper on the subject, published in 1853 (19), in which he corrected the then prevailing view that the seg- mentally arranged tubes were respiratory, and suggested their excretory function. As several of Goehlich's statements are at variance with those of Gegenbaur I was led to look into the details of the organ, and I then found that on those points the more recent writer was in error, and that Gegenbaur's description is much more nearly correct than Goehlich's. In fact, I have not very much to add to the description of the earlier writer beyond matters of histological detail, and more especially the blood-supply, which Professor Lankester suggested that I should work out. The work was carried out in the Zoological Laboratory of University College during the past eighteen months. 294 W. BLAXLAND BENHAM. CONTENTS OF PAPER. 1. Nomenclature of the parts of the nephridium, and the course of the various regions. 2. Histology of the various regions, and suggestions as to their function. 3. Comparison with the nephridium in other genera. 4. The nephrostome of Perichseta malamaniensis, n. sp., and other genera. 5. Circulation in the nephridium. 6. The nephridium of Arenicola. 1. NOMENCLATURE OF THE PARTS AND COURSE OF THE VARIOUS REGIONS. The long winding tube which constitutes the excretory organ of Lumbricus is for the most part embedded in a coating of vesicular cells, which are in their turn surrounded by the pavement-cells of the coelomic epithelium. The tube is, in its adult condition, as in its development, divisible into two portions: (I) a praseptal portion, consisting of the in- ternal funnel or nephrostome and a short ciliated tube; and (II) a much more extensive post-septal portion. The latter is readily distinguishable into four regions: (1) the very long but narrow tube in continuity with the praeseptal tube; (2) the short brownish ciliated middle tube; (3) the wide large tube; and (4) the muscular tube or duct, which opens to the exterior. The first three of these tubes (1, 2, 3) are twined about in rather a complicated but quite constant manner, and are bound together by the coat of vesicular cells in such a way as to form two great " loops," visible fairly distinctly with the naked eye when an opened worm is covered by spirit. Another "loop" is formed by the muscular duct (4). This last I will call the "first loop" (E, figs. 1, 2, 3); the "second loop" (F) contains the greater part of the narrow tube (1), and of the large tube (3); finally, the " third loop" (a) consists of a part of the narrow tube, a part of the large tube, and the whole of the middle tube (2). The course taken by these various regions, from the funnel to the external aperture, and the relations of the tubes to the THE NEPHBIDIUM OP LUMBBIOUS. 295 loops, will be readily seen by a glance at fig. 2, which will convey more information than any verbal description. I will, therefore, pass on to the histology of the various parts. I will premise that, with the exception of the funnel itself, or rather a part of the funnel, together with probably the mus- cular duct, the canal of the nephridium is " intra-cellular," i. e. the nephridium consists of a large number of perforated or "drain-pipe" cells, placed end to end (see PI. XXIV, figs. 30, 31, 32). The size of the cell and the relative proportion occupied by the lumen constitute the main morphological differences observable in the various regions; but the character of the protoplasm is of very great physiological importance, although we do not fully know the exact function of each region. 2. THE STRUCTURE OF THE VARIOUS REGIONS. I. The Prseseptal Portion.—The nephrostome or funnel is carried at the end of the short portion of the " narrow tube " which passes through the anterior septum bounding a given somite. The character of this portion of the tube will be described below. This prseseptal portion of the tube is sur- rounded by a mass of vesicular cells, the outlines of which are readily seen in the living condition, and frequently in stained preparations. These cells resemble those coating the post- septal portion, and appear to be similar to those described and figured by Kukenthal round the vessels and nephridia of Tubifex (24), and not improbably they have the same meaning. The superficial cells are usually flattened, and form a coelomic epithelium continuous with that covering the septum. The funnel (PI. XXIII, figs. 4—7, and 32) has recently been figured by Goehlich (20) in some detail, but he has alto- gether misunderstood the appearance represented. As is well known, the funnel consists of a number of very long, narrow, somewhat wedge-shaped cells—which I shall call "marginal cells"—arranged around a central point. The nucleus in each is placed near the outer end of the cell, which is rounded TOL. XXXII, PART III.—NEW SER. U 296 W. BLAXLAND BENHAM. FIG. 1.—A nephridium treated with nitrate of silver, to show the pavement coelomic epithelium covering the vesicular tissue, and forming in parts a suspensory fenestrated mem- brane, which carries blood-vessels. E. First loop. F F. Second loop. G G. Third loop. N. Nerve-cord. S. Septum. SN, Subneural blood-vessel. V. Subintestinal blood-vessel, a. Somatic branch from subintestinal blood-vessel, d. Ventral part of commissural vessel, h, h, h. Suspensory membranes, carrying and covering the nephridial blood-vessels, i. The group of coelomic epithelial cells around the prseseptal region of the narrow tube. FIG. 1. 298 W. BLAXLAND BBNHAM. off peripherally, and ciliated all over one surface. Goehlich describes and figures one or two rows of specially long cilia radiating from the centre. These I have never seen, although I have had under observation hundreds of nephridial funnels in demonstrating to my students, and have examined carefully dozens for this very purpose. The appearance figured by Goehlich is due to a crumpling of the funnel—an action to which it is liable on being covered by a glass slip,—so that there will be caused radial folds, the cilia of which will then be seen side- ways instead of from above, will therefore be more distinct, and will appear longer. There are normally no such longer cilia or such radiating lines. Another phenomenon, which, indeed, renders the interpreta- tion of the central portion of the funnel difficult, is the col- lection of a mass of ccelomic corpuscles in the funnel. This gives rise to the appearance represented in Goehlich's figure, and which was considered by him to form part of the wall of the funnel; but a careful examination, both of fresh specimens and stained preparations, has convinced me that these cells do not belong to the funnel, but are cells of the coelomic fluid, in some cases dead or dying, which will probably be carried to the exterior, as Kukenthal has suggested. In the fresh state this group of cells or " debris " appears darker and less transparent than the cells of the funnel. Stained with borax carmine the nuclei have an appearance distinct from those of the nephridial cells, being brighter, more granular, and much smaller. In fact, many have the appearance of cells which have died, and have begun to disintegrate. This mass of cells is shown in figs. 5 and 6. In one case (that represented by fig. 6), in which the funnel was examined twenty-four hours after the worm had been killed, not only were the cilia still active, but some of the cells forming the debris were still amoeboid, one or two short pointed pseudo- podia being present (fig. 6, a). Another case (fig. 5), where the cells of the debris are relatively few, shows the shape of the real- mouth of the funnel, which is marked out by the aggregation of the cells into a crescentic mass. It was this THE NEPHRIDIUM OP IiTJMBRlCTJS. 299 specimen that first led me to a better understanding of the real nature of the funnel, for no existing drawings or description give any clue to the real mode of communication between the ccelom and the nephridial tube. I will now proceed to describe the true structure of the nephrostome. If we follow the " narrow tube " forwards from the septum to the funnel, we see in optical section the finely granular wall on each side, with the nuclei of the component cells alternately on this side and on that. Arrived at the centre of the funnel, or thereabouts, the two walls suddenly diverge, each bending outwards, and then sharply backwards nearly parallel to its former course (fig. 4). The true " drain-pipe " cells cease at this point of divergence. The backwardly directed, or " centrifugal cells," as they may be termed, are merely grooved1 (fig. 32), but otherwise resemble the drain- pipe cells in structure, being granular and ciliated. These grooved or " gutter " cells, in reality, lie in a different plane from the narrow tube, a higher plane when the funnel is viewed from in front; and they frequently hide the wall of the tube, as is the case in fig.