Fig. 15.—Egg of Bilharzia, with the shell ad­her­ing to the es­caped cili­ated em­bry­o. Orig­inal.     Fig. 16.—Cili­ated em­bry­o of Bil­harzia in the act of dy­ing from the es­cape of the sar­co­dic con­tents. Orig­inal.

The behaviour of the embryo under the action of reagents of various kinds is remarkable. Thus, when on the 5th of Sept., 1870, I placed some ova in brackish water, of the strength of two parts of fresh water to one of pure sea-water, their contents were readily developed, though the escaping embryos did not swim vigorously. When again I placed some other eggs in pure sea-water, their contained embryos became instantly transfixed, the vibratile cilia of the head being rigid and motionless. At first I naturally concluded that the embryos were killed outright; but, to my great surprise, the shock passed away in about half an hour, when they revived and were soon afterwards hatched. One of the larvæ thus set free carried off several of the loose intra-chorional globules which had, during the period of transfixion, become firmly adherent to the ends of the caudal cilia. Here I may remark upon a decided difference observable between the cilia of the head and body respectively. The former are at all times vibratile, active, and conspicuous, whilst the latter are more delicate, capable of comparatively little motion, and partaking more of the character of fine setæ. In length their general measurement varies from 1/2500″ to 1/2000″. The action of pure sea-water on the free animalcules, previously immersed in fresh or brackish water, was equally striking. All, without exception, immediately became paralysed and almost motionless; nevertheless, on again adding fresh water, several entirely recovered. It is worthy of notice that in these cases the cephalic cilia furnished the first indications of returning viability. I was particularly struck with the behaviour of one embryo, which, under the stimulus of the sudden shock, retracted its cone-shaped head almost entirely within the general cavity of the body (fig. [14], lower specimen). In their moribund condition, whatever shape the embryo retained, the sarcodic contents gradually faded away; the outline of the creature, however, becoming more marked (fig. [16]). Usually the body of the animalcule became elongated whilst expiring in sea-water. Under other circumstances the embryo frequently bursts; the sarcodic contents escaping in the form of amœba-like bodies and the cilia retaining their powers of movement long after all traces of the sarcode have disappeared.

Fig. 17.—Cili­a­ted em­bryo of Fa­sci­o­la he­pa­tica, show­ing the so-​called eye-spot. After Leuc­kart.

The larvæ of Bilharzia closely resemble those of Fasciola hepatica, which latter may be appropriately noticed in this place. The ciliated embryo of the common liver fluke has the form of a long cone inverted; the anterior end or head being flatly convex. In the centre is a short proboscis-like papilla destitute of cilia (fig. 17). The general covering of cilia rests on a well-defined granular epidermis; this latter being succeeded by a dense peripheral layer of large nucleated cells, each of them measuring about 1/2500″ in diameter. The epidermis measures 1/6250″ in thickness. In the central mass of parenchyma no internal organs are recognisable, but Leuckart observed indications of a canal which he thought might open at the tail, though the opening itself was not actually visible.

As long as the ciliated covering remains intact the embryo, like other animalcules, displays great activity, whirling round and round on its own axis, and also describing gyrations and circles of different degrees of range in the water, the latter movements being accomplished by bending the body upon itself to a greater or lesser curvature. The embryos of Bilharzia and other infusoria exhibit the same behaviour, and, as Leuckart observes, when these embryos knock against any obstruction, they pause after the blow, as if to consider the nature of the substance they have touched. As in the case of fluke embryos generally, the ciliated covering eventually falls off and the embryo reassumes a more or less oval figure, at the same time changing its swimming mode of progression for the less dignified method of creeping. In the free ciliated condition the embryo of the common liver-fluke measures, according to Leuckart, 1/190″ in length, the anterior broad end being 1/500″. The cilia have a longitudinal measurement of 1/1388″.

Fig. 18.—‌Cili­ated em­bryo of Dis­toma lan­ce­o­la­tum. After Leuckart.

According to the observations of Dr Willemoes-Suhm, the cilia of the embryos of the Distoma megastoma are limited to the anterior pole of the body. This is also the arrangement, as Leuckart first pointed out, in Distoma lanceolatum (fig. 18). On the other hand, Pagenstecher has shown that the embryos of Distoma cygnoides and Amphistoma (Diplodiscus) subclavatum are ciliated all over, an observation which, as regards the latter species, has been confirmed by Wagener and others. Dr Pagenstecher’s original statement to the effect that “intrachorional germs of trematodes offer no distinctive characters,” must, therefore, in the present state of our knowledge, be accepted as a general conclusion admitting of many exceptions. In the early stages of development the embryo of Distoma lanceolatum occupies the centre of the egg, and according to Leuckart has its conical head invariably directed towards the upper pole of the shell, or, in other words, to that end of the egg which is furnished with a lid-like operculum. Leuckart describes the embryo itself as “finely granular and armed at the tip with a dagger-like spine, which, with the simultaneous displacement of the adjacent granular mass, can be pushed forward and drawn back again.” Besides this so-called cephalic granular mass, there are within the embryonic body two other granular masses widely separated from each other, but occupying the posterior half of the embryo. These Leuckart supposes to be the rudiments of a future brood, to be developed at the time when the free embryo shall have lost its ciliated swimming apparatus, shall have bored its way by means of the cephalic spine into the tissues of a mollusk, and shall have become metamorphosed into a sac-like larva (Nurse, Sporocyst, or Redia, as the case may be). Whatever be the full significance of these internal developments, we have at least satisfactory evidence that the complete and free embryo is a globe-shaped animalcule, having the anterior third or cephalic end of the body covered with cilia, and armed with a central boring spine. In consequence of this limitation of the ciliated covering, its swimming movements are less vivacious than those of the embryo of Fasciola hepatica; it will, therefore, probably take up its residence in a less active host than that chosen by the embryo of Fasciola, selecting one of those mollusks which either move slowly or are prone to keep at the bottom of the water. The mature eggs have a length of 1/625 to 1/555 of an inch, and a breadth of 1/833″. The long diameter of the free embryo varies from 1/990″ to 1/833″, the transverse diameter being 1/1562″. Whilst the embryos were still in the egg Leuckart could see no ciliary motion. With most observers, both the ciliary apparatus and the boring spine appear at this stage to have altogether escaped observation.