on their origin and partly on their fate, had great influence on the science of comparative anatomy during the last thirty years of the 19th century, for the homology of the layers being admitted, they afforded a kind of final court of appeal in determining questions of doubtful homologies between adult organs. Great importance was therefore attached to them by embryologists, and both their mode of development and the part which they play in forming the adult organs were examined with the greatest care. It is very unusual for all the layers to be established at the same time. As a general rule the ectoderm and endoderm, which may be called the primary layers, come first, and later the mesoderm is developed from one or other of them. There are two main methods in which the first two are differentiated—invagination and delamination. The former is generally found in small eggs, in which the embryo at the close of cleavage assumes the form of a sphere, having a fluid or gelatinous material in its centre, and bounded externally by a thin layer of protoplasm, in which all the nuclei are contained. Such a sphere is called a blastosphere, and may be regarded as a spherical mass of protoplasm, of which the central portion is so much vacuolated that it seems to consist entirely of fluid. The central part of the blastosphere is called the segmentation cavity or blastocoel. The blastosphere soon gives rise, by the invagination of one part of its wall upon the other, and a consequent obliteration of the segmentation cavity, to a double-walled cup with a wide opening, which, however, soon becomes narrowed to a small pore. This cup-stage is called the gastrula stage; the outer wall of the gastrula is the ectoderm, and its inner the endoderm; while its cavity is the enteron, and the opening to the exterior the blastopore. Origin of the primary layers by delamination occurs universally in eggs with large yolks (Cephalopoda and many Vertebrata), and occasionally in others. In it cleavage gives rise to a solid mass, which divides by delamination into two layers, the ectoderm and endoderm. The main difference between the two methods of development lies in the fact that in the first of them the endoderm at its first origin shows the relations which it possesses in the adult, namely, of forming the epithelial wall of the enteric space, whereas in the second method the endoderm is at first a solid mass, in which the enteric space makes its appearance later by excavation. In the delaminate method the enteric space is at first without a blastopore, and sometimes it never acquires this opening, but a blastopore is frequently formed, and the two-layered gastrula stage is reached, though by a very different route from that taken in the formation of the invaginate gastrula. According to the layer-theory, these two layers are homologous throughout the series of Metazoa; their limits can always be accurately defined, they give rise to the same organs in all cases, and the adult organs (excluding the mesodermal organs) can be traced back to one or other of them with absolute precision. Thus the ectoderm gives rise to the epidermis, to the nervous system, and to the lining of the stomodaeum and proctodaeum, if such parts of the alimentary canal are present. The endoderm, on the other hand, gives rise to the lining of the enteron, and of the glands which open into it.
So far as these two layers are concerned, and excluding the mesoderm, it would appear that the layer-theory does apply in a very remarkable manner to the whole of the Metazoa. But even here, when the actual facts are closely scanned, there are found to be difficulties, which appear to indicate that the theory may not perhaps be such an infallible guide as it seems at first sight. Leaving out of consideration the case of the Mammalia, in which the differentiation of the segmented ovum is not into ectoderm and endoderm, and the case of the sponges, the most important of these difficulties concern the stomodaeum and proctodaeum. The best case to examine is that of Peripatus capensis, in which the blastopore is at first a long slit, and gives rise to both the mouth and the anus of the adult. Here there is always found at the lips of the blastopore, and extending for a short distance inwards as enteric lining, a certain amount of tissue, which by its characters must be regarded as ectoderm. Now, in the closure of the blastopore between the mouth and anus, this tissue, which at the mouth and anus develops into the lining of the stomodaeum and proctodaeum, is left inside, and actually gives rise to the median ventral epithelium of the alimentary canal. Hence the development of Peripatus capensis suggests the conclusion, if we strictly apply the layer-theory, that a considerable portion of the true mesenteron is lined by ectoderm, and is not homologous with the corresponding portion of the mesenteron of other animals—a conclusion which will on all hands be admitted to be absurd. The difficulties in the application of the layer-theory become vastly greater when the Mesoderm. origin and fate of the mesoderm is considered. The mesoderm is, if we may judge from the number of organs which are derived from it, much the most important of the three layers. It generally arises later than the others, and in its very origin presents difficulties to the theory, which are much increased when we consider its history. It is generally, though not always, developed from the endoderm, either as hollow outgrowths containing prolongations of the enteric cavity, which become the coelom, or as solid proliferations. But in some groups the mesoderm is actually laid down in cleavage, and is present at the end of that process. In others it is entirely derived from the ectoderm (Peripatus capensis). In yet others it is partly derived from endoderm and partly from ectoderm (primitive streak of amniotic Vertebrates). Finally, in whatever manner the first rudiments are developed, it frequently receives considerable reinforcements from one of the primary layers. For instance, the structure known as the nerve crest of the vertebrate embryo is not, as was formerly supposed, exclusively concerned with the formation of the spinal nerves and ganglia, but contributes largely to the mesoderm of the axial region of the body. This is particularly clearly seen in the case of the anterior part of the head of Elasmobranch and probably of other vertebrate embryos, where all the mesoderm present is derived from the anterior part of the neural crest (Quart. Journ. Mic. Science, xxxvii. p. 92).
The layer-theory, then, will not bear critical examination. It is clear, both from their origin and history, that the layers or masses of cells called ectoderm, endoderm and mesoderm have not the same value in different animals; indeed, it is misleading to speak of three layers. At the most we can only speak of two, for the mesoderm is formed after the others, has a composite origin, and has no more claim to be considered an embryonic layer than has the rudiment of the central nervous system, which in some animals, indeed, appears as soon as the mesoderm. Arguments as to homology, based on derivation or non-derivation from the same embryonic layer, have therefore in themselves but little value.
It has frequently been asserted that the reproductive cells are marked off at a very early stage of the development (Sagitta, certain Crustacea, Scorpio). Recently it has been asserted that in Ascaris (T. Boveri, Kuppfer’s Festschrift, 1899, p. 383) the reproductive cells are set apart after the first cleavage, and that they can be traced by certain peculiarities of their nuclei into the adult reproductive glands.
It has been already stated that the mesoderm is a composite tissue. This fact is frequently conspicuous at its first establishment. In many Coelomata it is present under two forms from the beginning. One of these is epithelial in character, while the other has the form of a network of protoplasm, Mesenchyme. with nuclei at the nodes. The former is called simply epithelial mesoderm, the latter mesenchyme. Sometimes the epithelial mesoderm is the first formed, and what little mesenchyme there is is developed from it (Amphioxus, Balanoglossus, &c.) Sometimes the mesenchyme is the first to arise, the epithelial mesoderm developing from it (most, if not all, Vertebrates). Finally, it sometimes happens that these two kinds of tissue arise separately from one or other of the primary layers (Echinodermata). As already hinted, in Balanoglossus and Amphioxus the whole of the mesoderm of the body is at first in an epithelial condition, being developed as an outgrowth of the gut-wall. In Peripatus capensis also, and possibly in other Arthropods, it has at first an intermediate form, being derived from a primitive streak and not from the gut-wall, but it rapidly assumes an epithelial structure, from which all the mesodermal tissues are developed. In Annelids the bulk of the mesoderm has at first a modified epithelial form similar to that of Arthropods, but it is formed, not from a primitive streak, but from some peculiar cells produced in cleavage, called pole-cells. In Annelids with trochosphere larvae a certain amount of mesenchyme is formed at an earlier
