All recent observers, including Braun[428] for Reptilia, and Egli[429] for Mammalia, have stated that the Müllerian duct develops as a groove in the peritoneal epithelium, which is continued backward as a primitively solid rod in the space between the Wolffian duct and peritoneal epithelium.
In our preliminary account we stated[430], in accordance with the general view, that the Müllerian duct was formed as a groove, or elongated involution of the peritoneal epithelium adjoining the Wolffian duct. We have now reason to believe that this is not the case. In the earliest condition of the Müllerian duct which we have been able to observe, it consists of three successive open involutions of the peritoneal epithelium, connected together by more or less well-defined ridge-like thickenings of the epithelium. We believe, on grounds hereafter to be stated, that the whole of this formation is equivalent to the head-kidney of the Ichthyopsida. The head-kidney, as we shall continue to call it, takes its origin from the layer of thickened epithelium situated near the dorsal angle of the body-cavity, close to the Wolffian duct, which has been known since the publication of Waldeyer's important researches as the germinal epithelium. The anterior of the three open involutions or grooves is situated some little distance behind the front end of the Wolffian duct. It is simply a shallow groove in the thickest part of the germinal epithelium, and forms a corresponding projection into the adjacent stroma. In front the projection is separated by a considerable interval from the Wolffian duct; but near its hindermost part it almost comes into contact with the Wolffian duct. The groove extends in all for about five of our sections, and then terminates by its walls becoming gradually continued into a slight ridge-like thickening of the germinal epithelium. The groove arises as a simple depression in a linear area of thickened germinal epithelium. The linear area is, however, continued very considerably further forward than the groove, and sometimes exhibits a slight central depression, which might be regarded as a forward continuation of the groove. The passage from the groove to the ridge may best be conceived by supposing the groove to be suddenly filled up, so as to form a solid ridge pointing inwards towards the Wolffian duct.
The ridge succeeding the first groove is continued for about six sections, and is considerably more prominent at its posterior extremity than in front. It is replaced by groove number two, which appears as if formed by the reverse process to that by which the ridge arose, viz., by a hollowing out of the ridge on the side towards the body-cavity. The wall of the second groove is, after a few sections, continued into a second ridge or thickening of the germinal epithelium, which, however, is so faintly marked as to be hardly visible in its middle part. In its turn this ridge is replaced by the third and last groove. This vanishes after one or two sections, and behind the point of its disappearance we have failed to find any further traces of the head-kidney. The whole formation extends through about twenty-four of our sections and one and a half segments (muscle-plates).
We have represented (Plate 27, Series A, Nos. 1-10) a fairly complete series of sections through part of the head-kidney of an embryo slightly older than that last described, containing the second and third grooves and accessory parts. The connection between the grooves and the ridges is very well illustrated in Nos. 3, 4, and 5 of this series. In No. 3 we have a prominent ridge, in the interior of which there appears in No. 4 a groove, which becomes gradually wider in Nos. 5 and 6. Both the grooves and ridges are better marked in this than in the younger stage; but the chief difference between the two stages consists in the third groove no longer forming the hindermost limit of the head-kidney. Instead of this, the last groove (No. 7) terminates by the upper part of its walls becoming constricted off as a separate rod, which appears at first to contain a lumen continuous with the open groove. This rod (Nos. 7, 8, 9, 10) situated between the germinal epithelium and Wolffian duct is continued backward for some sections. It finally terminates by a pointed extremity, composed of not more than two cells abreast (Nos. 8-10).
Our third stage, sections of which are represented in series B (Plate 27), is considerably advanced beyond that last described. The most important change which has been effected concerns the ridges connecting the successive grooves. A lumen has appeared in each of these, which seems to open at both ends into the adjacent grooves. At the same time the cells, which previously constituted the ridge, have become (except where they are continuous with the walls of the grooves) partially constricted off from the germinal epithelium. The ridges, in fact, now form ducts situated in the stroma of the ovarian ridge, in the space between the Wolffian duct and the germinal epithelium. The duct continuous with the last groove is somewhat longer than before. In a general way, the head-kidney may now be described as a duct opening into the body-cavity by three groove-like apertures, and continuous behind with the rudiment of the true Müllerian duct. Although the general constitution of the head-kidney at this stage is fairly simple, there are a few features in our sections which we do not fully understand, and a few points about the organ which deserve a rather fuller description than we have given in this general sketch.
The anterior groove (Nos. 1-3, series B, Pl. 27) is at first somewhat separated from the Wolffian duct, but approaches close to it in No. 3. In Nos. 2 and 3 there appears a rod-like body on the outer side of the walls of the groove. In No. 2 this body is disconnected with the walls of the groove, and even appears as if formed by a second invagination of the germinal epithelium. In No. 3 this body becomes partially continuous with the walls of the groove, and finally in No. 4 it becomes completely continuous with the walls of the groove, and its lumen communicates freely with the groove[431].
The last trace of this body is seen on the upper wall of the groove in No. 5. We believe that the body (r1) represents the ridge between the first and second grooves of the earlier stage; so that in passing from No. 3 to No. 5 we pass from the first to the second groove. The meaning of the features of the body r1 in No. 2 we do not fully understand, but cannot regard them as purely accidental, since we have met with more or less similar features in other series of sections. The second groove becomes gradually narrower, and finally is continued into the second ridge (No. 8). The ridge contains a lumen, and is only connected with the germinal epithelium by a narrow wall of cells. A narrow passage from the body-cavity leads into that wall for a short distance in No. 8, but it is probably merely the hinder end of the groove of No. 7. The third groove appears in No. 11, and opens into the lumen of the second ridge (r2) in No. 12. In No. 13 the groove is closed, and there is present in its place a duct (r3) connected with the germinal epithelium by a wall of cells. This duct is the further development of the third ridge of the last stage; its lumen opens into the body-cavity through the third and last groove (gr3). In the next section this duct (r3) is entirely separated from the germinal epithelium, and it may be traced backwards through several sections until it terminates by a solid point, very much as in the last stage.
In the figures of this series (B) there may be noticed on the outer side of the Müllerian duct a fold of the germinal epithelium (x) forming a second groove. It is especially conspicuous in the first six sections of the series. This fold sometimes becomes much deeper, and then forms a groove, the upper end of which is close to the grooves of the head-kidney. It is very often much deeper than these are, and without careful study might easily be mistaken for one of these grooves. Fig. C, taken from a series slightly younger than B, shews this groove (x) in its most exaggerated form.
The stage we have just described is that of the fullest development of the head-kidney. In it, as in all the previous stages, there appear to be only three main openings into the body-cavity; but we have met in some of our sections with indications of the possible presence of one or two extra rudimentary grooves.
In an embryo not very much older than the one last described the atrophy of the head-kidney is nearly completed, and there is present but a single groove opening into the body-cavity.