Another case of practical gynecomastism has been described by Alexander von Humboldt. In a South American forest he found a solitary settler whose wife had died in child-birth. The man had laid the new-born child on his own breast in despair; and the continuous stimulus of the child's sucking movements had revived the activity of the mammary glands. It is possible that nervous suggestion had some share in it. Similar cases have been often observed in recent years, even among other male mammals (such as sheep and goats).

The great scientific interest of these facts is in their bearing on the question of heredity. The stem-history of the mammarium rests partly on its embryology (Chapter 2.24.) and partly on the facts of comparative anatomy and physiology. As in the lower and higher mammals (the monotremes, and most of the marsupials) the whole lactiferous apparatus is only found in the female; and as there are traces of it in the male only in a few younger marsupials, there can be no doubt that these important organs were originally found only in the female mammal, and that they were acquired by these through a special adaptation to habits of life.

Later, these female organs were communicated to both sexes by heredity; and they have been maintained in all persons of either sex, although they are not physiologically active in the males. This normal permanence of the female lactiferous organs in BOTH sexes of the higher mammals and man is independent of any selection, and is a fine instance of the much-disputed "inheritance of acquired characters."

CHAPTER 1.12. EMBRYONIC SHIELD AND GERMINATIVE AREA.

The three higher classes of vertebrates which we call the amniotes—the mammals, birds, and reptiles—are notably distinguished by a number of peculiarities of their development from the five lower classes of the stem—the animals without an amnion (the anamnia). All the amniotes have a distinctive embryonic membrane known as the amnion (or "water-membrane"), and a special embryonic appendage—the allantois. They have, further, a large yelk-sac, which is filled with food-yelk in the reptiles and birds, and with a corresponding clear fluid in the mammals. In consequence of these later-acquired structures, the original features of the development of the amniotes are so much altered that it is very difficult to reduce them to the palingenetic embryonic processes of the lower amnion-less vertebrates. The gastraea theory shows us how to do this, by representing the embryology of the lowest vertebrate, the skull-less amphioxus, as the original form, and deducing from it, through a series of gradual modifications, the gastrulation and coelomation of the craniota.

It was somewhat fatal to the true conception of the chief embryonic processes of the vertebrate that all the older embryologists, from Malpighi (1687) and Wolff (1750) to Baer (1828) and Remak (1850), always started from the investigation of the hen's egg, and transferred to man and the other vertebrates the impressions they gathered from this. This classical object of embryological research is, as we have seen, a source of dangerous errors. The large round food-yelk of the bird's egg causes, in the first place, a flat discoid expansion of the small gastrula, and then so distinctive a development of this thin round embryonic disk that the controversy as to its significance occupies a large part of embryological literature.

(FIGURE 1.105. Severance of the discoid mammal embryo from the yelk-sac, in transverse section (diagrammatic). A The germinal disk (h, hf) lies flat on one side of the branchial-gut vesicle (kb). B In the middle of the germinal disk we find the medullary groove (mr), and underneath it the chorda (ch). C The gut-fibre-layer (df) has been enclosed by the gut-gland-layer (dd). D The skin-fibre-layer (hf) and gut-fibre-layer (df) divide at the periphery; the gut (d) begins to separate from the yelk-sac or umbilical vesicle (nb). E The medullary tube (mr) is closed; the body-cavity (c) begins to form. F The provertebrae (w) begin to grow round the medullary tube (mr) and the chorda (ch): the gut (d) is cut off from the umbilical vesicle (nb). H The vertebrae (w) have grown round the medullary tube (mr) and chorda; the body-cavity is closed, and the umbilical vesicle has disappeared. The amnion and serous membrane are omitted. The letters have the same meaning throughout: h horn-plate, mr medullary tube, hf skin-fibre-layer, w provertebrae, ch chorda, c body-cavity or coeloma, df gut-fibre-layer, dd gut-gland-layer, d gut-cavity, nb umbilical vesicle.)

One of the most unfortunate errors that this led to was the idea of an original antithesis of germ and yelk. The latter was regarded as a foreign body, extrinsic to the real germ, whereas it is properly a part of it, an embryonic organ of nutrition. Many authors said there was no trace of the embryo until a later stage, and outside the yelk; sometimes the two-layered embryonic disk itself, at other times only the central portion of it (as distinguished from the germinative area, which we will describe presently), was taken to be the first outline of the embryo. In the light of the gastraea theory it is hardly necessary to dwell on the defects of this earlier view and the erroneous conclusions drawn from it. In reality, the first segmentation-cell, and even the stem-cell itself and all that issues therefrom, belong to the embryo. As the large original yelk-mass in the undivided egg of the bird only represents an inclosure in the greatly enlarged ovum, so the later contents of its embryonic yelk-sac (whether yet segmented or not) are only a part of the entoderm which forms the primitive gut. This is clearly shown by the ova of the amphibia and cyclostoma, which explain the transition from the yelk-less ova of the amphioxus to the large yelk-filled ova of the reptiles and birds.

It is precisely in the study of these difficult features that we see the incalculable value of phylogenetic considerations in explaining complex ontogenetic facts, and the need of separating cenogenetic phenomena from palingenetic. This is particularly clear as regards the comparative embryology of the vertebrates, because here the phylogenetic unity of the stem has been already established by the well-known facts of paleontology and comparative anatomy. If this unity of the stem, on the basis of the amphioxus, were always borne in mind, we should not have these errors constantly recurring.

In many cases the cenogenetic relation of the embryo to the food-yelk has until now given rise to a quite wrong idea of the first and most important embryonic processes in the higher vertebrates, and has occasioned a number of false theories in connection with them. Until thirty years ago the embryology of the higher vertebrates always started from the position that the first structure of the embryo is a flat, leaf-shaped disk; it was for this reason that the cell-layers that compose this germinal disk (also called germinative area) are called "germinal layers." This flat germinal disk, which is round at first and then oval, and which is often described as the tread or cicatricula in the laid hen's egg, is found at a certain part of the surface of the large globular food-yelk. I am convinced that it is nothing else than the discoid, flattened gastrula of the birds. At the beginning of germination the flat embryonic disk curves outwards, and separates on the inner side from the underlying large yelk-ball. In this way the flat layers are converted into tubes, their edges folding and joining together (Figure 1.105). As the embryo grows at the expense of the food-yelk, the latter becomes smaller and smaller; it is completely surrounded by the germinal layers. Later still, the remainder of the food-yelk only forms a small round sac, the yelk-sac or umbilical vesicle (Figure 1.105 nb). This is enclosed by the visceral layer, is connected by a thin stalk, the yelk-duct, with the central part of the gut-tube, and is finally, in most of the vertebrates, entirely absorbed by this (H). The point at which this takes place, and where the gut finally closes, is the visceral navel. In the mammals, in which the remainder of the yelk-sac remains without and atrophies, the yelk-duct at length penetrates the outer ventral wall. At birth the umbilical cord proceeds from here, and the point of closure remains throughout life in the skin as the navel.

As the older embryology of the higher vertebrates was mainly based on the chick, and regarded the antithesis of embryo (or formative-yelk) and food-yelk (or yelk-sac) as original, it had also to look upon the flat leaf-shaped structure of the germinal disk as the primitive embryonic form, and emphasise the fact that hollow grooves were formed of these flat layers by folding, and closed tubes by the joining together of their edges.

This idea, which dominated the whole treatment of the embryology of the higher vertebrates until thirty years ago, was totally false. The gastraea theory, which has its chief application here, teaches us that it is the very reverse of the truth. The cup-shaped gastrula, in the body-wall of which the two primary germinal layers appear from the first as closed tubes, is the original embryonic form of all the vertebrates, and all the multicellular invertebrates; and the flat germinal disk with its superficially expanded germinal layers is a later, secondary form, due to the cenogenetic formation of the large food-yelk and the gradual spread of the germ-layers over its surface. Hence the actual folding of the germinal layers and their conversion into tubes is not an original and primary, but a much later and tertiary, evolutionary process. In the phylogeny of the vertebrate embryonic process we may distinguish the following three stages:—

A. First Stage: Primary (palingenetic) embryonic process.

The germinal layers form from the first closed tubes, the one-layered blastula being converted into the two-layered gastrula by invagination. No food-yelk. (Amphioxus.)

B. Second Stage: Secondary (cenogenetic) embryonic process.

The germinal layers spread out leaf-wise, food-yelk gathering in the ventral entoderm, and a large yelk-sac being formed from the middle of the gut-tube. (Amphibia.)

C. Third Stage: Tertiary (cenogenetic) embryonic process.

The germinal layers form a flat germinal disk, the borders of which join together and form closed tubes, separating from the central yelk-sac. (Amniotes.)

As this theory, a logical conclusion from the gastraea theory, has been fully substantiated by the comparative study of gastrulation in the last few decades, we must exactly reverse the hitherto prevalent mode of treatment. The yelk-sac is not to be treated, as was done formerly, as if it were originally antithetic to the embryo, but as an essential part of it, a part of its visceral tube. The primitive gut of the gastrula has, on this view, been divided into two parts in the higher animals as a result of the cenogenetic formation of the food-yelk—the permanent gut (metagaster), or permanent alimentary canal, and the yelk-sac (lecithoma), or umbilical vesicle. This is very clearly shown by the comparative ontogeny of the fishes and amphibia. In these cases the whole yelk undergoes cleavage at first, and forms a yelk-gland, composed of yelk-cells, in the ventral wall of the primitive gut. But it afterwards becomes so large that a part of the yelk does not divide, and is used up in the yelk-sac that is cut off outside.

(FIGURE 1.106. The visceral embryonic vesicle (blastocystis or gastrocystis) of a rabbit (the "blastula" or vesicula blastodermica of other writers), a outer envelope (ovolemma), b skin-layer or ectoderm, forming the entire wall of the yelk-vesicle, c groups of dark cells, representing the visceral layer or entoderm.

FIGURE 1.107. The same in section. Letters as above. d cavity of the vesicle. (From Bischoff.))

When we make a comparative study of the embryology of the amphioxus, the frog, the chick, and the rabbit, there cannot, in my opinion, be any further doubt as to the truth of this position, which I have held for thirty years. Hence in the light of the gastraea theory we must regard the features of the amphioxus as the only and real primitive structure among all the vertebrates, departing very little from the palingenetic embryonic form. In the cyclostoma and the frog these features are, on the whole, not much altered cenogenetically, but they are very much so in the chick, and most of all in the rabbit. In the bell-gastrula of the amphioxus and in the hooded gastrula of the lamprey and the frog the germinal layers are found to be closed tubes or vesicles from the first. On the other hand, the chick-embryo (in the new laid, but not yet hatched, egg) is a flat circular disk, and it was not easy to recognise this as a real gastrula. Rauber and Goette have, however, achieved this. As the discoid gastrula grows round the large globular yelk, and the permanent gut then separates from the outlying yelk-sac, we find all the processes which we have shown (diagrammatically) in Figure 1.108—processes that were hitherto regarded as principal acts, whereas they are merely secondary.

The oldest, oviparous mammals, the monotremes, behave in the same way as the reptiles and birds. But the corresponding embryonic processes in the viviparous mammals, the marsupials and placentals, are very elaborate and distinctive. They were formerly quite misinterpreted; it was not until the publication of the studies of Edward van Beneden (1875) and the later research of Selenka, Kuppfer, Rabl, and others, that light was thrown on them, and we were in a position to bring them into line with the principles of the gastraea theory and trace them to the embryonic forms of the lower vertebrates. Although there is no independent food-yelk, apart from the formative yelk, in the mammal ovum, and although its segmentation is total on that account, nevertheless a large yelk-sac is formed in their embryos, and the "embryo proper" spreads leaf-wise over its surface, as in the reptiles and birds, which have a large food-yelk and partial segmentation. In the mammals, as well as in the latter, the flat, leaf-shaped germinal disk separates from the yelk-sac, and its edges join together and form tubes.

How can we explain this curious anomaly? Only as a result of very characteristic and peculiar cenogenetic modifications of the embryonic process, the real causes of which must be sought in the change in the rearing of the young on the part of the viviparous mammals. These are clearly connected with the fact that the ancestors of the viviparous mammals were oviparous amniotes like the present monotremes, and only gradually became viviparous. This can no longer be questioned now that it has been shown (1884) that the monotremes, the lowest and oldest of the mammals, still lay eggs, and that these develop like the ova of the reptiles and birds. Their nearest descendants, the marsupials, formed the habit of retaining the eggs, and developing them in the oviduct; the latter was thus converted into a womb (uterus). A nutritive fluid that was secreted from its wall, and passed through the wall of the blastula, now served to feed the embryo, and took the place of the food-yelk. In this way the original food-yelk of the monotremes gradually atrophied, and at last disappeared so completely that the partial ovum-segmentation of their descendants, the rest of the mammals, once more became total. From the discogastrula of the former was evolved the distinctive epigastrula of the latter.

It is only by this phylogenetic explanation that we can understand the formation and development of the peculiar, and hitherto totally misunderstood, blastula of the mammal. The vesicular condition of the mammal embryo was discovered 200 years ago (1677) by Regner de Graaf. He found in the uterus of a rabbit four days after impregnation small, round, loose, transparent vesicles, with a double envelope. However, Graaf's discovery passed without recognition. It was not until 1827 that these vesicles were rediscovered by Baer, and then more closely studied in 1842 by Bischoff in the rabbit (Figures 1.106 and 1.107). They are found in the womb of the rabbit, the dog, and other small mammals, a few days after copulation. The mature ova of the mammal, when they have left the ovary, are fertilised either here or in the oviduct immediately afterwards by the invading sperm-cells.* (* In man and the other mammals the fertilisation of the ova probably takes place, as a rule, in the oviduct; here the ova, which issue from the female ovary in the shape of the Graafian follicle, and enter the inner aperture of the oviduct, encounter the mobile sperm-cells of the male seed, which pass into the uterus at copulation, and from this into the external aperture of the oviduct. Impregnation rarely takes place in the ovary or in the womb.) (As to the womb and oviduct see Chapter 2.29.) The cleavage and formation of the gastrula take place in the oviduct. Either here in the oviduct or after the mammal gastrula has passed into the uterus it is converted into the globular vesicle which is shown externally in Figure 1.106, and in section in Figure 1.107. The thick, outer, structureless envelope that encloses it is the original ovolemma or zona pellucida, modified, and clothed with a layer of albumin that has been deposited on the outside. From this stage the envelope is called the external membrane, the primary chorion or prochorion (a). The real wall of the vesicle enclosed by it consists of a simple layer of ectodermic cells (b), which are flattened by mutual pressure, and generally hexagonal; a light nucleus shines through their fine-grained protoplasm (Figure 1.108). At one part (c) inside this hollow ball we find a circular disc, formed of darker, softer, and rounder cells, the dark-grained entodermic cells (Figure 1.109).

(FIGURE 1.108. Four entodermic cells from the embryonic vesicle of the rabbit.

FIGURE 1.109. Two entodermic cells from the embryonic vesicle of the rabbit.)

The characteristic embryonic form that the developing mammal now exhibits has up to the present usually been called the "blastula" (Bischoff), "sac-shaped embryo" (Baer), "vesicular embryo" (vesicula blastodermica, or, briefly, blastosphaera). The wall of the hollow vesicle, which consists of a single layer of cells, was called the "blastoderm," and was supposed to be equivalent to the cell-layer of the same name that forms the wall of the real blastula of the amphioxus and many of the invertebrates (such as Monoxenia, Figure 1.29 F, G). Formerly this real blastula was generally believed to be equivalent to the embryonic vesicle of the mammal. However, this is by no means the case. What is called the "blastula" of the mammal and the real blastula of the amphioxus and many of the invertebrates are totally different embryonic structures. The latter (blastula) is palingenetic, and precedes the formation of the gastrula. The former (blastodermic vesicle) is cenogenetic, and follows gastrulation. The globular wall of the blastula is a real blastoderm, and consists of homogeneous (blastodermic) cells; it is not yet differentiated into the two primary germinal layers. But the globular wall of the mammal vesicle is the differentiated ectoderm, and at one point in it we find a circular disk of quite different cells—the entoderm. The round cavity, filled with fluid, inside the real blastula is the segmentation-cavity. But the similar cavity within the mammal vesicle is the yelk-sac cavity, which is connected with the incipient gut-cavity. This primitive gut-cavity passes directly into the segmentation-cavity in the mammals, in consequence of the peculiar cenogenetic changes in their gastrulation, which we have considered previously (Chapter 1.9). For these reasons it is very necessary to recognise the secondary embryonic vesicle in the mammal (gastrocystis or blastocystis) as a characteristic structure peculiar to this class, and distinguish it carefully from the primary blastula of the amphioxus and the invertebrates.

(FIGURE 1.110. Ovum of a rabbit from the uterus, one sixth of an inch in diameter. The embryonic vesicle (b) has withdrawn a little from the smooth ovolemma (a). In the middle of the ovolemma we see the round germinal disk (blastodiscus, c), at the edge of which (at d) the inner layer of the embryonic vesicle is already beginning to expand. (Figures 1.110 to 1.114 from Bischoff.)

FIGURE 1.111. The same ovum, seen in profile. Letters as in Figure 1.110.

FIGURE 1.112. Ovum of a rabbit from the uterus, one-fourth of an inch in diameter. The blastoderm is already for the most part two-layered (b). The ovolemma, or outer envelope, is tufted (a).

FIGURE 1.113. The same ovum, seen in profile. Letters as in Figure 1.112.

FIGURE 1.114. Ovum of a rabbit from the uterus, one-third of an inch in diameter. The embryonic vesicle is now nearly everywhere two-layered (k) only remaining one-layered below (at d).

FIGURE 1.115. Round germinative area of the rabbit, divided into the central light area (area pellucida) and the peripheral dark area (area opaca). The light area seems darker on account of the dark ground appearing through it.)

The small, circular, whitish, and opaque spot which the gastric disk (Figure 1.106) forms at a certain part of the surface of the clear and transparent embryonic vesicle has long been known to science, and compared to the germinal disk of the birds and reptiles. Sometimes it has been called the germinal disk, sometimes the germinal spot, and usually the germinative area. From the area the further development of the embryo proceeds. However, the larger part of the embryonic vesicle of the mammal is not directly used for building up the later body, but for the construction of the temporary umbilical vesicle. The embryo separates from this in proportion as it grows at its expense; the two are only connected by the yelk-duct (the stalk of the yelk-sac), and this maintains the direct communication between the cavity of the umbilical vesicle and the forming visceral cavity (Figure 1.105).

The germinative area or gastric disk of the animal consists at first (like the germinal disk of birds and reptiles) merely of the two primary germinal layers, the ectoderm and entoderm. But soon there appears in the middle of the circular disk between the two a third stratum of cells, the rudiment of the middle layer or fibrous layer (mesoderm). This middle germinal layer consists from the first, as we have seen in Chapter 1.10, of two separate epithelial plates, the two layers of the coelom-pouches (parietal and visceral). However, in all the amniotes (on account of the large formation of yelk) these thin middle plates are so firmly pressed together that they seem to represent a single layer. It is thus peculiar to the amniotes that the middle of the germinative area is composed of four germinal layers, the two limiting (or primary) layers and the middle layers between them (Figures 1.96 and 1.97). These four secondary germinal layers can be clearly distinguished as soon as what is called the sickle-groove (or "embryonic sickle") is seen at the hind border of the germinative area. At the borders, however, the germinative area of the mammal only consists of two layers. The rest of the wall of the embryonic vesicle consists at first (but only for a short time in most of the mammals) of a single layer, the outer germinal layer.

(FIGURE 1.116. Oval area, with the opaque whitish border of the dark area without.)

From this stage, however, the whole wall of the embryonic vesicle becomes two-layered. The middle of the germinative area is much thickened by the growth of the cells of the middle layers, and the inner layer expands at the same time, and increases at the border of the disk all round. Lying close on the outer layer throughout, it grows over its inner surface at all points, covers first the upper and then the lower hemisphere, and at last closes in the middle of the inner layer (Figures 1.110 to 1.114). The wall of the embryonic vesicle now consists throughout of two layers of cells, the ectoderm without and the entoderm within. It is only in the centre of the circular area, which becomes thicker and thicker through the growth of the middle layers, that it is made up of all four layers. At the same time, small structureless tufts or warts are deposited on the surface of the outer ovolemma or prochorion, which has been raised above the embryonic vesicle (Figures 1.112 to 1.114 a).

(FIGURE 1.117. Oval germinal disk of the rabbit, magnified about ten times. As the delicate, half-transparent disk lies on a black ground, the pellucid area looks like a dark ring, and the opaque area (lying outside it) like a white ring. The oval shield in the centre also looks whitish, and in its axis we see the dark medullary groove. (From Bischoff.))

We may now disregard both the outer ovolemma and the greater part of the vesicle, and concentrate our attention on the germinative area and the four-layered embryonic disk. It is here alone that we find the important changes which lead to the differentiation of the first organs. It is immaterial whether we examine the germinative area of the mammal (the rabbit, for instance) or the germinal disk of a bird or a reptile (such as a lizard or tortoise). The embryonic processes we are now going to consider are essentially the same in all members of the three higher classes of vertebrates which we call the amniotes. Man is found to agree in this respect with the rabbit, dog, ox, etc.; and in all these animals the germinative area undergoes essentially the same changes as in the birds and reptiles. They are most frequently and accurately studied in the chick, because we can have incubated hens' eggs in any quantity at any stage of development. Moreover, the round germinal disk of the chick passes immediately after the beginning of incubation (within a few hours) from the two-layered to the four-layered stage, the two-layered mesoderm developing from the median primitive groove between the ectoderm and entoderm (Figures 1.82 to 1.95).

The first change in the round germinal disk of the chick is that the cells at its edges multiply more briskly, and form darker nuclei in their protoplasm. This gives rise to a dark ring, more or less sharply set off from the lighter centre of the germinal disk (Figure 1.115). From this point the latter takes the name of the "light area" (area pellucida), and the darker ring is called the "dark area" (area opaca). (In a strong light, as in Figures 1.115 to 1.117, the light area seems dark, because the dark ground is seen through it; and the dark area seems whiter). The circular shape of the area now changes into elliptic, and then immediately into oval (Figures 1.116 and 1.117). One end seems to be broader and blunter, the other narrower and more pointed; the former corresponds to the anterior and the latter to the posterior section of the subsequent body. At the same time, we can already trace the characteristic bilateral form of the body, the antithesis of right and left, before and behind. This will be made clearer by the "primitive streak," which appears at the posterior end.

(FIGURE 1.118. Pear-shaped germinal shield of the rabbit (eight days old), magnified twenty times. rf medullary groove. pr primitive groove (primitive mouth). (From Kolliker.)

FIGURE 1.119. Median longitudinal section of the gastrula of four vertebrates. (From Rabl.) A discogastrula of a shark (Pristiurus). B amphigastrula of a sturgeon (Accipenser). C amphigastrula of an amphibium (Triton). D epigastrula of an amniote (diagram). a ventral, b dorsal lip of the primitive mouth.)

At an early stage an opaque spot is seen in the middle of the clear germinative area, and this also passes from a circular to an oval shape. At first this shield-shaped marking is very delicate and barely perceptible; but it soon becomes clearer, and now stands out as an oval shield, surrounded by two rings or areas (Figure 1.117). The inner and brighter ring is the remainder of the pellucid area, and the dark outer ring the remainder of the opaque area; the opaque shield-like spot itself is the first rudiment of the dorsal part of the embryo. We give it briefly the name of embryonic shield or dorsal shield. In most works this embryonic shield is described as "the first rudiment or trace of the embryo," or "primitive embryo." But this is wrong, though it rests on the authority of Baer and Bischoff. As a matter of fact, we already have the embryo in the stem-cell, the gastrula, and all the subsequent stages. The embryonic shield is simply the first rudiment of the dorsal part, which is the earliest to develop. As the older names of "embryonic rudiment" and "germinative area" are used in many different senses—and this has led to a fatal confusion in embryonic literature—we must explain very clearly the real significance of these important embryonic parts of the amniote. It will be useful to do so in a series of formal principles:—

1. The so-called "first trace of the embryo" in the amniotes, or the embryonic shield, in the centre of the pellucid area, consists merely of an early differentiation and formation of the middle dorsal parts.

2. Hence the best name for it is "the dorsal shield," as I proposed long ago.

3. The germinative area, in which the first embryonic blood-vessels appear at an early stage, is not opposed as an external area to the "embryo proper," but is a part of it.

4. In the same way, the yelk-sac or the umbilical vesicle is not a foreign external appendage of the embryo, but an outlying part of its primitive gut.

5. The dorsal shield gradually separates from the germinative area and the yelk-sac, its edges growing downwards and folding together to form ventral plates.

6. The yelk-sac and vessels of the germinative area, which soon spread over its whole surface, are, therefore, real embryonic organs, or temporary parts of the embryo, and have a transitory importance in connection with the nutrition of the growing later body; the latter may be called the "permanent body" in contrast to them.

The relation of these cenogenetic features of the amniotes to the palingenetic structures of the older non-amniotic vertebrates may be expressed in the following theses: The original gastrula, which completely passes into the embryonic body in the acrania, cyclostoma, and amphibia, is early divided into two parts in the amniotes—the embryonic shield, which represents the dorsal outline of the permanent body; and the temporary embryonic organs of the germinative area and its blood-vessels, which soon grow over the whole of the yelk-sac. The differences which we find in the various classes of the vertebrate stem in these important particulars can only be fully understood when we bear in mind their phylogenetic relations on the one hand, and, on the other, the cenogenetic modifications of structure that have been brought about by changes in the rearing of the young and the variation in the mass of the food-yelk.

We have already described in Chapter 1.9 the changes which this increase and decrease of the nutritive yelk causes in the form of the gastrula, and especially in the situation and shape of the primitive mouth. The primitive mouth or prostoma is originally a simple round aperture at the lower pole of the long axis; its dorsal lip is above and ventral lip below. In the amphioxus this primitive mouth is a little eccentric, or shifted to the dorsal side (Figure 1.39). The aperture increases with the growth of the food-yelk in the cyclostoma and ganoids; in the sturgeon it lies almost on the equator of the round ovum, the ventral lip (a) in front and the dorsal lip (b) behind (Figure 1.119 b). In the wide-mouthed, circular discoid gastrula of the selachii or primitive fishes, which spreads quite flat on the large food-yelk, the anterior semi-circle of the border of the disk is the ventral, and the posterior semicircle the dorsal lip (Figure 1.119 A). The amphiblastic amphibia are directly connected with their earlier fish-ancestors, the dipneusts and ganoids, and further the oldest selachii (Cestracion); they have retained their total unequal segmentation, and their small primitive mouth (Figure 1.119 C, ab), blocked up by the yelk-stopper, lies at the limit of the dorsal and ventral surface of the embryo (at the lower pole of its equatorial axis), and there again has an upper dorsal and a lower ventral lip (a, b). The formation of a large food-yelk followed again in the stem-forms of the amniotes, the protamniotes or proreptilia, descended from the amphibia (Figure 1.119 D). But here the accumulation of the food-yelk took place only in the ventral wall of the primitive-gut, so that the narrow primitive mouth lying behind was forced upwards, and came to lie on the back of the discoid "epigastrula" in the shape of the "primitive groove"; thus (in contrast to the case of the selachii, Figure 1.119 A) the dorsal lip (b) had to be in front, and the ventral lip (a) behind (Figure 1.119 D). This feature was transmitted to all the amniotes, whether they retained the large food-yelk (reptiles, birds, and monotremes), or lost it by atrophy (the viviparous mammals).

This phylogenetic explanation of gastrulation and coelomation, and the comparative study of them in the various vertebrates, throw a clear and full light on many ontogenetic phenomena, as to which the most obscure and confused opinions were prevalent thirty years ago. In this we see especially the high scientific value of the biogenetic law and the careful separation of palingenetic from cenogenetic processes. To the opponents of this law the real explanation of these remarkable phenomena is impossible. Here, and in every other part of embryology, the true key to the solution lies in phylogeny.

CHAPTER 1.13. DORSAL BODY AND VENTRAL BODY.

The earliest stages of the human embryo are, for the reasons already given, either quite unknown or only imperfectly known to us. But as the subsequent embryonic forms in man behave and develop just as they do in all the other mammals, there cannot be the slightest doubt that the preceding stages also are similar. We have been able to see in the coelomula of the human embryo (Figure 1.97), by transverse sections through its primitive mouth, that its two coelom-pouches are developed in just the same way as in the rabbit (Figure 1.96); moreover, the peculiar course of the gastrulation is just the same.

(FIGURE 1.120. Embryonic vesicle of a seven-days-old rabbit with oval embryonic shield (ag). A seen from above, B from the side. (From Kolliker.) ag dorsal shield or embryonic spot. In B the upper half of the vesicle is made up of the two primary germinal layers, the lower (up to ge) only from the outer layer.)

The germinative area forms in the human embryo in the same way as in the other mammals, and in the middle part of this we have the embryonic shield, the purport of which we considered in Chapter 1.12. The next changes in the embryonic disk, or the "embryonic spot," take place in corresponding fashion. These are the changes we are now going to consider more closely.

The chief part of the oval embryonic shield is at first the narrow hinder end; it is in the middle line of this that the primitive streak appears (Figure 1.121 ps). The narrow longitudinal groove in it—the so-called "primitive groove"—is, as we have seen, the primitive mouth of the gastrula. In the gastrula-embryos of the mammals, which are much modified cenogenetically, this cleft-shaped prostoma is lengthened so much that it soon traverses the whole of the hinder half of the dorsal shield; as we find in a rabbit embryo of six to eight days (Figure 1.122 pr). The two swollen parallel borders that limit this median furrow are the side lips of the primitive mouth, right and left. In this way the bilateral-symmetrical type of the vertebrate becomes pronounced. The subsequent head of the amniote is developed from the broader and rounder fore-half of the dorsal shield.

In this fore-half of the dorsal shield a median furrow quickly makes its appearance (Figure 1.123 rf). This is the broader dorsal furrow or medullary groove, the first beginning of the central nervous system. The two parallel dorsal or medullary swellings that enclose it grow together over it afterwards, and form the medullary tube. As is seen in transverse sections, it is formed only of the outer germinal layer (Figures 1.95 and 1.136). The lips of the primitive mouth, however, lie, as we know, at the important point where the outer layer bends over the inner, and from which the two coelom pouches grow between the primary germinal layers.

(FIGURE 1.121. Oval embryonic shield of the rabbit (A of six days eighteen hours, B of eight days). (From Kolliker.) ps primitive streak, pr primitive groove, arg area germinalis, sw sickle-shaped germinal growth.

FIGURE 1.122. Dorsal shield (ag) and germinative area of a rabbit-embryo of eight days. (From Kolliker.) pr primitive groove, rf dorsal furrow.

FIGURE 1.123. Embryonic shield of a rabbit of eight days. (From Van Beneden.) pr primitive groove, cn canalis neurentericus, nk nodus neurentericus (or "Hensen's ganglion"), kf head-process (chorda).

FIGURE 1.124. Longitudinal section of the coelomula of amphioxus (from the left). i entoderm, d primitive gut, cn medullary duct, n nerve tube, m mesoderm, s first primitive segment, c coelom-pouches. (From Hatschek.))

Thus the median primitive furrow (pr) in the hind-half and the median medullary furrow (rf) in the fore-half of the oval shield are totally different structures, although the latter seems to a superficial observer to be merely the forward continuation of the former. Hence they were formerly always confused. This error was the more pardonable as immediately afterwards the two grooves do actually pass into each other in a very remarkable way. The point of transition is the remarkable neurenteric canal (Figure 1.124 cn). But the direct connection which is thus established does not last long; the two are soon definitely separated by a partition.

The enigmatic neurenteric canal is a very old embryonic organ, and of great phylogenetic interest, because it arises in the same way in all the chordonia (both tunicates and vertebrates). In every case it touches or embraces like an arch the posterior end of the chorda, which has been developed here in front out of the middle line of the primitive gut (between the two coelom-folds of the sickle groove) ("head-process," Figure 1.123 kf). These very ancient and strictly hereditary structures, which have no physiological significance to-day, deserve (as "rudimentary organs") our closest attention. The tenacity with which the useless neurenteric canal has been transmitted down to man through the whole series of vertebrates is of equal interest for the theory of descent in general, and the phylogeny of the chordonia in particular.

The connection which the neurenteric canal (Figure 1.123 cn) establishes between the dorsal nerve-tube (n) and the ventral gut-tube (d) is seen very plainly in the amphioxus in a longitudinal section of the coelomula, as soon as the primitive mouth is completely closed at its hinder end. The medullary tube has still at this stage an opening at the forward end, the neuroporus (Figure 1.83 np). This opening also is afterwards closed. There are then two completely closed canals over each other—the medullary tube above and the gastric tube below, the two being separated by the chorda. The same features as in the acrania are exhibited by the related tunicates, the ascidiae.

Again, we find the neurenteric canal in just the same form and situation in the amphibia. A longitudinal section of a young tadpole (Figure 1.125) shows how we may penetrate from the still open primitive mouth (x) either into the wide primitive gut-cavity (al) or the narrow overlying nerve-tube. A little later, when the primitive mouth is closed, the narrow neurenteric canal (Figure 1.126 ne) represents the arched connection between the dorsal medullary canal (mc) and the ventral gastric canal.

(FIGURE 1.125. Longitudinal section of the chordula of a frog. (From Balfour.) nc nerve-tube, x canalis neurentericus, al alimentary canal, yk yelk-cells, m mesoderm.

FIGURE 1.126. Longitudinal section of a frog-embryo. (From Goette.) m mouth, l liver, an anus, ne canalis neurentericus, mc medullary-tube, pn pineal body (epiphysis), ch chorda.

FIGURES 1.127 AND 1.128. Dorsal shield of the chick. (From Balfour.) The medullary furrow (me), which is not yet visible in Figure 1.130, encloses with its hinder end the fore end of the primitive groove (pr) in Figure 1.131.)

In the amniotes this original curved form of the neurenteric canal cannot be found at first, because here the primitive mouth travels completely over to the dorsal surface of the gastrula, and is converted into the longitudinal furrow we call the primitive groove. Hence the primitive groove (Figure 1.128 pr), examined from above, appears to be the straight continuation of the fore-lying and younger medullary furrow (me). The divergent hind legs of the latter embrace the anterior end of the former. Afterwards we have the complete closing of the primitive mouth, the dorsal swellings joining to form the medullary tube and growing over it. The neurenteric canal then leads directly, in the shape of a narrow arch-shaped tube (Figure 1.129 ne), from the medullary tube (sp) to the gastric tube (pag). Directly in front of it is the latter end of the chorda (cli).

While these important processes are taking place in the axial part of the dorsal shield, its external form also is changing. The oval form (Figure 1.117) becomes like the sole of a shoe or sandal, lyre-shaped or finger-biscuit shaped (Figure 1.130). The middle third does not grow in width as quickly as the posterior, and still less than the anterior third; thus the shape of the permanent body becomes somewhat narrow at the waist. At the same time, the oval form of the germinative area returns to a circular shape, and the inner pellucid area separates more clearly from the opaque outer area (Figure 1.131 a). The completion of the circle in the area marks the limit of the formation of blood-vessels in the mesoderm.

(FIGURE 1.129. Longitudinal section of the hinder end of a chick. (From Balfour.) sp medullary tube, connected with the terminal gut (pag) by the neurenteric canal (ne), ch chorda, pr neurenteric (or Hensen's) ganglion, al allantois, ep ectoderm, hy entoderm, so parietal layer, sp visceral layer, an anus-pit, am amnion.)

The characteristic sandal-shape of the dorsal shield, which is determined by the narrowness of the middle part, and which is compared to a violin, lyre, or shoe-sole, persists for a long time in all the amniotes. All mammals, birds, and reptiles have substantially the same construction at this stage, and even for a longer or shorter period after the division of the primitive segments into the coelom-folds has begun (Figure 1.132). The human embryonic shield assumes the sandal-form in the second week of development; towards the end of the week our sole-shaped embryo has a length of about one-twelfth of an inch (Figure 1.133).

The complete bilateral symmetry of the vertebrate body is very early indicated in the oval form of the embryonic shield (Figure 1.117) by the median primitive streak; in the sandal-form it is even more pronounced (Figures 1.131 to 1.135). In the lateral parts of the embryonic shield a darker central and a lighter peripheral zone become more obvious; the former is called the stem-zone (Figure 1.134 stz), and the latter the parietal zone (pz); from the first we get the dorsal and from the second the ventral half of the body-wall. The stem-zone of the amniote embryo would be called more appropriately the dorsal zone or dorsal shield; from it develops the whole of the dorsal half of the later body (or permanent body)—that is to say, the dorsal body (episoma). Again, it would be better to call the "parietal zone" the ventral zone or ventral shield; from it develop the ventral "lateral plates," which afterwards separate from the embryonic vesicle and form the ventral body (hyposoma)—that is to say, the ventral half of the permanent body, together with the body-cavity and the gastric canal that it encloses.

(FIGURE 1.130. Germinal area or germinal disk of the rabbit, with sole-shaped embryonic shield, magnified about ten times. The clear circular field (d) is the opaque area. The pellucid area (c) is lyre-shaped, like the embryonic shield itself (b). In its axis is seen the dorsal furrow or medullary furrow (a). (From Bischoff.))

The sole-shaped germinal shields of all the amniotes are still, at the stage of construction which Figure 1.134 illustrates in the rabbit and Figure 1.135 in the opossum, so like each other that we can either not distinguish them at all or only by means of quite subordinate peculiarities in the size of the various parts. Moreover, the human sandal-shaped embryo cannot at this stage be distinguished from those of other mammals, and it particularly resembles that of the rabbit. On the other hand, the outer form of these flat sandal-shaped embryos is very different from the corresponding form of the lower animals, especially the acrania (amphioxus). Nevertheless, the body is just the same in the essential features of its structure as that we find in the chordula of the latter (Figures 1.83 to 1.86), and in the embryonic forms which immediately develop from it. The striking external difference is here again due to the fact that in the palingenetic embryos of the amphioxus (Figures 1.83 and 1.84) and the amphibia (Figures 1.85 and 1.86) the gut-wall and body-wall form closed tubes from the first, whereas in the cenogenetic embryos of the amniotes they are forced to expand leaf-wise on the surface owing to the great extension of the food-yelk.

(FIGURE 1.131. Embryo of the opossum, sixty hours old, one-sixth of an inch in diameter. (From Selenka) b the globular embryonic vesicle, a the round germinative area, b limit of the ventral plates, r dorsal shield, v its fore part, u the first primitive segment, ch chorda, chr its fore-end, pr primitive groove (or mouth).

FIGURE 1.132. Sandal-shaped embryonic shield of a rabbit of eight days, with the fore part of the germinative area (ao opaque, ap pellucid area). (From Kolliker.) rf dorsal furrow, in the middle of the medullary plate, h, pr primitive groove (mouth), stz dorsal (stem) zone, pz ventral (parietal) zone. In the narrow middle part the first three primitive segments may be seen.)

It is all the more notable that the early separation of dorsal and ventral halves takes place in the same rigidly hereditary fashion in all the vertebrates. In both the acrania and the craniota the dorsal body is about this period separated from the ventral body. In the middle part of the body this division has already taken place by the construction of the chorda between the dorsal nerve-tube and the ventral canal. But in the outer or lateral part of the body it is only brought about by the division of the coelom-pouches into two sections—a dorsal episomite (dorsal segment or provertebra) and a ventral hyposomite (or ventral segment) by a frontal constriction. In the amphioxus each of the former makes a muscular pouch, and each of the latter a sex-pouch or gonad.

These important processes of differentiation in the mesoderm, which we will consider more closely in the next chapter, proceed step by step with interesting changes in the ectoderm, while the entoderm changes little at first. We can study these processes best in transverse sections, made vertically to the surface through the sole-shaped embryonic shield. Such a transverse section of a chick embryo, at the end of the first day of incubation, shows the gut-gland layer as a very simple epithelium, which is spread like a leaf over the outer surface of the food-yelk (Figure 1.92). The chorda (ch) has separated from the dorsal middle line of the entoderm; to the right and left of it are the two halves of the mesoderm, or the two coelom-folds. A narrow cleft in the latter indicates the body-cavity (uwh); this separates the two plates of the coelom-pouches, the lower (visceral) and upper (parietal). The broad dorsal furrow (Rf) formed by the medullary plate (m) is still wide open, but is divided from the lateral horn-plate (h) by the parallel medullary swellings, which eventually close.

(FIGURE 1.133. Human embryo at the sandal-stage, one-twelfth of an inch long, from the end of the second week, magnified twenty-five times. (From Count Spee.)

FIGURE 1.134. Sandal-shaped embryonic shield of a rabbit of nine days. (From Kolliker.) (Back view from above.) stz stem-zone or dorsal shield (with eight pairs of primitive segments), pz parietal or ventral zone, ap pellucid area, af amnion-fold, h heart, ph pericardial cavity, vo omphalo-mesenteric vein, ab eye-vesicles, vh fore brain, mh middle brain, hh hind brain, uw primitive segments (or vertebrae).)

During these processes important changes are taking place in the outer germinal layer (the "skin-sense layer"). The continued rise and growth of the dorsal swellings causes their higher parts to bend together at their free borders, approach nearer and nearer (Figure 1.136 w), and finally unite. Thus in the end we get from the open dorsal furrow, the upper cleft of which becomes narrower and narrower, a closed cylindrical tube (Figure 1.137 mr). This tube is of the utmost importance; it is the beginning of the central nervous system, the brain and spinal marrow, the medullary tube. This embryonic fact was formerly looked upon as very mysterious. We shall see presently that in the light of the theory of descent it is a thoroughly natural process. The phylogenetic explanation of it is that the central nervous system is the organ by means of which all intercourse with the outer world, all psychic action and sense-perception, are accomplished; hence it was bound to develop originally from the outer and upper surface of the body, or from the outer skin. The medullary tube afterwards separates completely from the outer germinal layer, and is surrounded by the middle parts of the provertebrae and forced inwards (Figure 1.146). The remaining portion of the skin-sense layer (Figure 1.93 h) is now called the horn-plate or horn-layer, because from it is developed the whole of the outer skin or epidermis, with all its horny appendages (nails, hair, etc.).

(FIGURE 1.135. Sandal-shaped embryonic shield of an opossum (Didelphys), three days old. (From Selenka.) (Back view from above.) stz stem-zone or dorsal shield (with eight pairs of primitive segments), pz parietal or ventral zone, ap pellucid area, ao opaque area, hh halves of the heart, v fore-end, h hind-end. In the median line we see the chorda (ch) through the transparent medullary tube (m). u primitive segment, pr primitive streak (or primitive mouth).)

A totally different organ, the prorenal (primitive kidney) duct (ung), is found to be developed at an early stage from the ectoderm. This is originally a quite simple, tube-shaped, lengthy duct, or straight canal, which runs from front to rear at each side of the provertebrae (on the outer side, Figure 1.93 ung). It originates, it seems, out of the horn-plate at the side of the medullary tube, in the gap that we find between the provertebral and the lateral plates. The prorenal duct is visible in this gap even at the time of the severance of the medullary tube from the horn-plate. Other observers think that the first trace of it does not come from the skin-sense layer, but the skin-fibre layer.

The inner germinal layer, or the gut-fibre layer (Figure 1.93 dd), remains unchanged during these processes. A little later, however, it shows a quite flat, groove-like depression in the middle line of the embryonic shield, directly under the chorda. This depression is called the gastric groove or furrow. This at once indicates the future lot of this germinal layer. As this ventral groove gradually deepens, and its lower edges bend towards each other, it is formed into a closed tube, the alimentary canal, in the same way as the medullary groove grows into the medullary tube. The gut-fibre layer (Figure 1.137 f), which lies on the gut-gland layer (d), naturally follows it in its folding. Moreover, the incipient gut-wall consists from the first of two layers, internally the gut-gland layer and externally the gut-fibre layer.

The formation of the alimentary canal resembles that of the medullary tube to this extent—in both cases a straight groove or furrow arises first of all in the middle line of a flat layer. The edges of this furrow then bend towards each other, and join to form a tube (Figure 1.137). But the two processes are really very different. The medullary tube closes in its whole length, and forms a cylindrical tube, whereas the alimentary canal remains open in the middle, and its cavity continues for a long time in connection with the cavity of the embryonic vesicle. The open connection between the two cavities is only closed at a very late stage, by the construction of the navel. The closing of the medullary tube is effected from both sides, the edges of the groove joining together from right and left. But the closing of the alimentary canal is not only effected from right and left, but also from front and rear, the edges of the ventral groove growing together from every side towards the navel. Throughout the three higher classes of vertebrates the whole of this process of the construction of the gut is closely connected with the formation of the navel, or with the separation of the embryo from the yelk-sac or umbilical vesicle.

In order to get a clear idea of this, we must understand carefully the relation of the embryonic shield to the germinative area and the embryonic vesicle. This is done best by a comparison of the five stages which are shown in longitudinal section in Figures 1.138 to 1.142. The embryonic shield (c), which at first projects very slightly over the surface of the germinative area, soon begins to rise higher above it, and to separate from the embryonic vesicle. At this point the embryonic shield, looked at from the dorsal surface, shows still the original simple sandal-shape (Figures 1.133 to 1.135). We do not yet see any trace of articulation into head, neck, trunk, etc., or limbs. But the embryonic shield has increased greatly in thickness, especially in the anterior part. It now has the appearance of a thick, oval swelling, strongly curved over the surface of the germinative area. It begins to sever completely from the embryonic vesicle, with which it is connected at the ventral surface. As this severance proceeds, the back bends more and more; in proportion as the embryo grows the embryonic vesicle decreases, and at last it merely hangs as a small vesicle from the belly of the embryo (Figure 1.142 ds). In consequence of the growth-movements which cause this severance, a groove-shaped depression is formed at the surface of the vesicle, the limiting furrow, which surrounds the vesicle in the shape of a pit, and a circular mound or dam (Figure 1.139 ks) is formed at the outside of this pit by the elevation of the contiguous parts of the germinal vesicle.

(FIGURE 1.136. Transverse section of the embryonic disk of a chick at the end of the first day of incubation, magnified about twenty times. The edges of the medullary plate (m), the medullary swellings (w), which separate the medullary from the horn-plate (h), are bending towards each other. At each side of the chorda (ch) the primitive segment plates (u) have separated from the lateral plates (sp). A gut-gland layer. (From Remak.))

In order to understand clearly this important process, we may compare the embryo to a fortress with its surrounding rampart and trench. The ditch consists of the outer part of the germinative area, and comes to an end at the point where the area passes into the vesicle. The important fold of the middle germinal layer that brings about the formation of the body-cavity spreads beyond the borders of the embryo over the whole germinative area. At first this middle layer reaches as far as the germinative area; the whole of the rest of the embryonic vesicle consists in the beginning only of the two original limiting layers, the outer and inner germinal layers. Hence, as far as the germinative area extends the germinal layer splits into the two plates we have already recognised in it, the outer skin-fibre layer and the inner gut-fibre layer. These two plates diverge considerably, a clear fluid gathering between them (Figure 1.140 am). The inner plate, the gut-fibre layer, remains on the inner layer of the embryonic vesicle (on the gut-gland layer). The outer plate, the skin-fibre layer, lies close on the outer layer of the germinative area, or the skin-sense layer, and separates together with this from the embryonic vesicle. From these two united outer plates is formed a continuous membrane. This is the circular mound that rises higher and higher round the whole embryo, and at last joins above it (Figures 1.139 to 1.142 am). To return to our illustration of the fortress, we must imagine the circular rampart to be extraordinarily high and towering far above the fortress. Its edges bend over like the combs of an overhanging wall of rock that would enclose the fortress; they form a deep hollow, and at last join together above. In the end the fortress lies entirely within the hollow that has been formed by the growth of the edges of this large rampart.

(FIGURE 1.137. Three diagrammatic transverse sections of the embryonic disk of the higher vertebrate, to show the origin of the tubular organs from the bending germinal layers. In Figure A the medullary tube (n) and the alimentary canal (a) are still open grooves. In Figure B the medullary tube (n) and the dorsal wall are closed, but the alimentary canal (a) and the ventral wall are open; the prorenal ducts (u) are cut off from the horn-plate (h) and internally connected with segmental prorenal canals. In Figure C both the medullary tube and the dorsal wall above and the alimentary canal and ventral wall below are closed. All the open grooves have become closed tubes; the primitive kidneys are directed inwards. The letters have the same meaning in all three figures: h skin-sense layer, n medullary tube, u prorenal ducts, x axial rod, s primitive-vertebra, r dorsal wall, b ventral wall, c body-cavity or coeloma, f gut-fibre layer, t primitive artery (aorta), v primitive vein (subintestinal vein), d gut-fibre layer, a alimentary canal.)

As the two outer layers of the germinative area thus rise in a fold about the embryo, and join above it, they come at last to form a spacious sac-like membrane about it. This envelope takes the name of the germinative membrane, or water-membrane, or amnion (Figure 1.142 am). The embryo floats in a watery fluid, which fills the space between the embryo and the amnion, and is called the amniotic fluid (Figures 1.141 and 1.142 ah). We will deal with this remarkable formation and with the allantois later on (Chapter 1.15). In front of the allantois the yelk-sac or umbilical vesicle (ds), the remainder of the original embryonic vesicle, starts from the open belly of the embryo (Figure 1.138 kh). In more advanced embryos, in which the gastric wall and the ventral wall are nearly closed, it hangs out of the navel-opening in the shape of a small vesicle with a stalk (Figures 1.141 and 1.142 ds). The more the embryo grows, the smaller becomes the vitelline (yelk) sac. At first the embryo looks like a small appendage of the large embryonic vesicle. Afterwards it is the yelk-sac, or the remainder of the embryonic vesicle, that seems a small pouch-like appendage of the embryo (Figure 1.142 ds). It ceases to have any significance in the end. The very wide opening, through which the gastric cavity at first communicates with the umbilical vesicle, becomes narrower and narrower, and at last disappears altogether. The navel, the small pit-like depression that we find in the developed man in the middle of the abdominal wall, is the spot at which the remainder of the embryonic vesicle (the umbilical vesicle) originally entered into the ventral cavity, and joined on to the growing gut.

(FIGURES 1.138 TO 1.142. Five diagrammatic longitudinal sections of the maturing mammal embryo and its envelopes. In Figures 1.138 to 1.141 the longitudinal section passes through the sagittal or middle plane of the body, dividing the right and left halves; in Figure 1.142 the embryo is seen from the left side. In Figure 1.138 the tufted it prochorion (dd apostrophe) encloses the germinal vesicle, the wall of which consists of the two primary layers. Between the outer (a) and inner (i) layer the middle layer (m) has been developed in the region of the germinative area. In Figure 1.139 the embryo (e) begins to separate from the embryonic vesicle (ds), while the wall of the amnion-fold rises about it (in front as head-sheath, ks, behind as tail-sheath, ss). In Figure 1.140 the edges of the amniotic fold (am) rise together over the back of the embryo, and form the amniotic cavity (ah); as the embryo separates more completely from the embryonic vesicle (ds) the alimentary canal (dd) is formed, from the hinder end of which the allantois grows (al). In Figure 1.141 the allantois is larger; the yelk-sac (ds) smaller. In Figure 1.142 the embryo shows the gill-clefts and the outline of the two legs; the chorion has formed branching villi (tufts.) In all four figures e = embryo, a outer germinal layer, m middle germinal layer, i inner germinal layer, am amnion (ks head-sheath, ss tail-sheath), ah amniotic cavity, as amniotic sheath of the umbilical cord, kh embryonic vesicle, ds yelk-sac (umbilical vesicle), dg vitelline duct, df gut-fibre layer, dd gut-gland layer, al allantois, vl = hh place of heart, d vitelline membrane (ovolemma or prochorion), d apostrophe tufts or villi of same, sh serous membrane (serolemma), sz tufts of same, ch chorion, chz tufts or villi, st terminal vein, r pericoelom or serocoelom (the space, filled with fluid, between the amnion and chorion). (From Kolliker.))

The origin of the navel coincides with the complete closing of the external ventral wall. In the amniotes the ventral wall originates in the same way as the dorsal wall. Both are formed substantially from the skin-fibre layer, and externally covered with the horn-plate, the border section of the skin-sense layer. Both come into existence by the conversion of the four flat germinal layers of the embryonic shield into a double tube by folding from opposite directions; above, at the back, we have the vertebral canal which encloses the medullary tube, and below, at the belly, the wall of the body-cavity which contains the alimentary canal (Figure 1.137).

We will consider the formation of the dorsal wall first, and that of the ventral wall afterwards (Figures 1.143 to 1.147). In the middle of the dorsal surface of the embryo there is originally, as we already know, the medullary (mr) tube directly underneath the horn-plate (h), from the middle part of which it has been developed. Later, however, the provertebral plates (uw) grow over from the right and left between these originally connected parts (Figures 1.145 and 1.146). The upper and inner edges of the two provertebral plates push between the horn-plate and medullary tube, force them away from each other, and finally join between them in a seam that corresponds to the middle line of the back. The coalescence of these two dorsal plates and the closing in the middle of the dorsal wall take place in the same way as the medullary tube, which is henceforth enclosed by the vertebral tube. Thus is formed the dorsal wall, and the medullary tube takes up a position inside the body. In the same way the provertebral mass grows afterwards round the chorda, and forms the vertebral column. Below this the inner and outer edge of the provertebral plate splits on each side into two horizontal plates, of which the upper pushes between the chorda and medullary tube, and the lower between the chorda and gastric tube. As the plates meet from both sides above and below the chorda, they completely enclose it, and so form the tubular, outer chord-sheath, the sheath from which the vertebral column is formed (perichorda, Figure 1.137 C, s; Figures 1.145 uwh, 1.146).

(FIGURES 1.143 TO 1.146. Transverse sections of embryos (of chicks). Figure 1.143 of the second, Figure 1.144 of the third, Figure 1.145 of the fourth, and Figure 1.146 of the fifth day of incubation. Figures 1.143 to 1.145 from Kolliker, magnified about 100 times; Figure 1.146 from Remak, magnified about twenty times. h horn-plate, mr medullary tube, ung prorenal duct, un prorenal vesicles, hp skin-fibre layer, m = mu = mp muscle-plate, uw provertebral plate (wh cutaneous rudiment of the body of the vertebra, wb of the arch of the vertebra, wq the rib or transverse continuation), uwh provertebral cavity, ch axial rod or chorda, sh chorda-sheath, bh ventral wall, g hind and v fore root of the spinal nerves, a = af = am amniotic fold, p body-cavity or coeloma, df gut-fibre layer, ao primitive aortas, sa secondary aorta, vc cardinal veins, d = dd gut-gland layer, dr gastric groove. In Figure 1.143 the larger part of the right half, in Figure 1.144 the larger part of the left half, of the section is omitted. Of the yelk-sac or remainder of the embryonic vesicle only a small piece of the wall is indicated below.)

We find in the construction of the ventral wall precisely the same processes as in the formation of the dorsal wall (Figure 1.137 B, Figure 1.144 hp, Figure 1.146 bh). It is formed on the flat embryonic shield of the amniotes from the upper plates of the parietal zone. The right and left parietal plates bend downwards towards each other, and grow round the gut in the same way as the gut itself closes. The outer part of the lateral plates forms the ventral wall or the lower wall of the body, the two lateral plates bending considerably on the inner side of the amniotic fold, and growing towards each other from right and left. While the alimentary canal is closing, the body-wall also closes on all sides. Hence the ventral wall, which encloses the whole ventral cavity below, consists of two parts, two lateral plates that bend towards each other. These approach each other all along, and at last meet at the navel. We ought, therefore, really to distinguish two navels, an inner and an outer one. The internal or intestinal navel is the definitive point of the closing of the gut wall, which puts an end to the open communication between the ventral cavity and the cavity of the yelk-sac (Figure 1.105). The external navel in the skin is the definitive point of the closing of the ventral wall; this is visible in the developed body as a small depression.

(FIGURE 1.147. Median longitudinal section of the embryo of a chick (fifth day of incubation), seen from the right side (head to the right, tail to the left). Dorsal body dark, with convex outline. d gut, o mouth, a anus, l lungs, h liver, g mesentery, v auricle of the heart, k ventricle of the heart, b arch of the arteries, t aorta, c yelk-sac, m vitelline (yelk) duct, u allantois, r pedicle (stalk) of the allantois, n amnion, w amniotic cavity, s serous membrane. (From Baer.))

With the formation of the internal navel and the closing of the alimentary canal is connected the formation of two cavities, which we call the capital and the pelvic sections of the visceral cavity. As the embryonic shield lies flat on the wall of the embryonic vesicle at first, and only gradually separates from it, its fore and hind ends are independent in the beginning; on the other hand, the middle part of the ventral surface is connected with the yelk-sac by means of the vitelline or umbilical duct (Figure 1.147 m). This leads to a notable curving of the dorsal surface; the head-end bends downwards towards the breast and the tail-end towards the belly. We see this very clearly in the excellent old diagrammatic illustration given by Baer (Figure 1.147), a median longitudinal section of the embryo of the chick, in which the dorsal body or episoma is deeply shaded. The embryo seems to be trying to roll up, like a hedgehog protecting itself from its pursuers. This pronounced curve of the back is due to the more rapid growth of the convex dorsal surface, and is directly connected with the severance of the embryo from the yelk-sac. The further bending of the embryo leads to the formation of the "head-cavity" of the gut (Figure 1.148 above D) and a similar one at the tail, known as its "pelvic cavity."

As a result of these processes the embryo attains a shape that may be compared to a wooden shoe, or, better still, to an overturned canoe. Imagine a canoe or boat with both ends rounded and a small covering before and behind; if this canoe is turned upside down, so that the curved keel is uppermost, we have a fair picture of the canoe-shaped embryo (Figure 1.147). The upturned convex keel corresponds to the middle line of the back; the small chamber underneath the fore-deck represents the capital cavity, and the small chamber under the rear-deck the pelvic chamber of the gut (cf. Figure 1.140).

The embryo now, as it were, presses into the outer surface of the embryonic vesicle with its free ends, while it moves away from it with its middle part. As a result of this change the yelk-sac becomes henceforth only a pouch-like outer appendage at the middle of the ventral wall. The ventral appendage, growing smaller and smaller, is afterwards called the umbilical (navel) vesicle. The cavity of the yelk-sac or umbilical vesicle communicates with the corresponding visceral cavity by a wide opening, which gradually contracts into a narrow and long canal, the vitelline (yelk) duct (ductus vitellinus, Figure 1.147 m). Hence, if we were to imagine ourselves in the cavity of the yelk-sac, we could get from it through the yelk-duct into the middle and still wide open part of the alimentary canal. If we were to go forward from there into the head-part of the embryo, we should reach the capital cavity of the gut, the fore-end of which is closed up.

The reader will ask: "Where are the mouth and the anus?" These are not at first present in the embryo. The whole of the primitive gut-cavity is completely closed, and is merely connected in the middle by the vitelline duct with the equally closed cavity of the embryonic vesicle (Figure 1.140). The two later apertures of the alimentary canal—the anus and the mouth—are secondary constructions, formed from the outer skin. In the horn-plate, at the spot where the mouth is found subsequently, a pit-like depression is formed, and this grows deeper and deeper, pushing towards the blind fore-end of the capital cavity; this is the mouth-pit. In the same way, at the spot in the outer skin where the anus is afterwards situated a pit-shaped depression appears, grows deeper and deeper, and approaches the blind hind-end of the pelvic cavity; this is the anus-pit. In the end these pits touch with their deepest and innermost points the two blind ends of the primitive alimentary canal, so that they are now only separated from them by thin membranous partitions. This membrane finally disappears, and henceforth the alimentary canal opens in front at the mouth and in the rear by the anus (Figures 1.141 and 1.147). Hence at first, if we penetrate into these pits from without, we find a partition cutting them off from the cavity of the alimentary canal, which gradually disappears. The formation of mouth and anus is secondary in all the vertebrates.

(FIGURE 1.148. Longitudinal section of the fore half of a chick-embryo at the end of the first day of incubation (seen from the left side). k head-plates, ch chorda. Above it is the blind fore-end of the ventral tube (m); below it the capital cavity of the gut. d gut-gland layer, df gut-fibre layer, h horn plate, hh cavity of the heart, hk heart-capsule, ks head-sheath, kk head-capsule. (From Remak.))

During the important processes which lead to the formation of the navel, and of the intestinal wall and ventral wall, we find a number of other interesting changes taking place in the embryonic shield of the amniotes. These relate chiefly to the prorenal ducts and the first blood-vessels. The prorenal (primitive kidney) ducts, which at first lie quite flat under the horn-plate or epiderm (Figure 1.93 ung), soon back towards each other in consequence of special growth movements (Figures 1.143 to 1.145 ung). They depart more and more from their point of origin, and approach the gut-gland layer. In the end they lie deep in the interior, on either side of the mesentery, underneath the chorda, (Figure 1.145 ung). At the same time, the two primitive aortas change their position (cf. Figures 1.138 to 1.145 ao); they travel inwards underneath the chorda, and there coalesce at last to form a single secondary aorta, which is found under the rudimentary vertebral column (Figure 1.145 ao). The cardinal veins, the first venous blood-vessels, also back towards each other, and eventually unite immediately above the rudimentary kidneys (Figures 1.145 vc, 152 cav). In the same spot, at the inner side of the fore-kidneys, we soon see the first trace of the sexual organs. The most important part of this apparatus (apart from all its appendages) is the ovary in the female and the testicle in the male. Both develop from a small part of the cell-lining of the body-cavity, at the spot where the skin-fibre layer and gut-fibre layer touch. The connection of this embryonic gland with the prorenal ducts, which lie close to it and assume most important relations to it, is only secondary.

(FIGURE 1.149. Longitudinal section of a human embryo of the fourth week, one-fifth of an inch long, magnified fifteen times. Showing: bend of skull, yelk-sac, umbilical cord, terminal gut, rudimentary kidneys, mesoderm, head-gut (with gill-clefts), primitive lungs, liver, stomach, pancreas, mesentery, primitive kidneys, allantoic duct, rectum. (From Kollmann.)

FIGURE 1.150. Transverse section of a human embryo of fourteen days. mr medullary tube, ch chorda. vu umbilical vein, mt myotome, mp middle plate, ug prorenal duct, lh body-cavity, e ectoderm, bh ventral skin, hf skin-fibre layer, df gut-fibre layer. (From Kollmann.)

FIGURE 1.151. Transverse section of a shark-embryo (or young selachius). mr medullary tube, ch chorda, a aorta, d gut, vp principal (or subintestinal) vein, mt myotome, mm muscular mass of the provertebra, mp middle plate, ug prorenal duct, lh body-cavity, e ectoderm of the rudimentary extremities, mz mesenchymic cells, z point where the myotome and nephrotome separate. (From H.E. Ziegler.)

FIGURE 1.152. Transverse section of a duck-embryo with twenty-four primitive segments. (From Balfour.) From a dorsal lateral joint of the medullary tube (spc) the spinal ganglia (spg) grow out between it and the horn-plate. ch chorda, ao double aorta, hy gut-gland layer, sp gut-fibre layer, with blood-vessels in section, ms muscle plate, in the dorsal wall of the myocoel (episomite). Below the cardinal vein (cav) is the prorenal duct (wd) and a segmental prorenal canal (st). The skin-fibre layer of the body-wall (so) is continued in the amniotic fold (am). Between the four secondary germinal layers and the structures formed from them there is formed embryonic connective matter with stellate cells and vascular structures (Hertwig's "mesenchym").)

CHAPTER 1.14. THE ARTICULATION OF THE BODY.*

(* The term articulation is used in this chapter to denote both "segmentation" and "articulation" in the ordinary sense.—Translator.)

The vertebrate stem, to which our race belongs as one of the latest and most advanced outcomes of the natural development of life, is rightly placed at the head of the animal kingdom. This privilege must be accorded to it, not only because man does in point of fact soar far above all other animals, and has been lifted to the position of "lord of creation"; but also because the vertebrate organism far surpasses all the other animal-stems in size, in complexity of structure, and in the advanced character of its functions. From the point of view of both anatomy and physiology, the vertebrate stem outstrips all the other, or invertebrate, animals.

There is only one among the twelve stems of the animal kingdom that can in many respects be compared with the vertebrates, and reaches an equal, if not a greater, importance in many points. This is the stem of the articulates, composed of three classes: 1, the annelids (earth-worms, leeches, and cognate forms); 2, the crustacea (crabs, etc.); 3, the tracheata (spiders, insects, etc.). The stem of the articulates is superior not only to the vertebrates, but to all other animal-stems, in variety of forms, number of species, elaborateness of individuals, and general importance in the economy of nature.

When we have thus declared the vertebrates and the articulates to be the most important and most advanced of the twelve stems of the animal kingdom, the question arises whether this special position is accorded to them on the ground of a peculiarity of organisation that is common to the two. The answer is that this is really the case; it is their segmental or transverse articulation, which we may briefly call metamerism. In all the vertebrates and articulates the developed individual consists of a series of successive members (segments or metamera = "parts"); in the embryo these are called primitive segments or somites. In each of these segments we have a certain group of organs reproduced in the same arrangement, so that we may regard each segment as an individual unity, or a special "individual" subordinated to the entire personality.

The similarity of their segmentation, and the consequent physiological advance in the two stems of the vertebrates and articulates, has led to the assumption of a direct affinity between them, and an attempt to derive the former directly from the latter. The annelids were supposed to be the direct ancestors, not only of the crustacea and tracheata, but also of the vertebrates. We shall see later (Chapter 2.20) that this annelid theory of the vertebrates is entirely wrong, and ignores the most important differences in the organisation of the two stems. The internal articulation of the vertebrates is just as profoundly different from the external metamerism of the articulates as are their skeletal structure, nervous system, vascular system, and so on. The articulation has been developed in a totally different way in the two stems. The unarticulated chordula (Figures 1.83 to 1.86), which we have recognised as one of the chief palingenetic embryonic forms of the vertebrate group, and from which we have inferred the existence of a corresponding ancestral form for all the vertebrates and tunicates, is quite unthinkable as the stem-form of the articulates.

All articulated animals came originally from unarticulated ones. This phylogenetic principle is as firmly established as the ontogenetic fact that every articulated animal-form develops from an unarticulated embryo. But the organisation of the embryo is totally different in the two stems. The chordula-embryo of all the vertebrates is characterised by the dorsal medullary tube, the neurenteric canal, which passes at the primitive mouth into the alimentary canal, and the axial chorda between the two. None of the articulates, either annelids or arthropods (crustacea and tracheata), show any trace of this type of organisation. Moreover, the development of the chief systems of organs proceeds in the opposite way in the two stems. Hence the segmentation must have arisen independently in each. This is not at all surprising; we find analogous cases in the stalk-articulation of the higher plants and in several groups of other animal stems.

The characteristic internal articulation of the vertebrates and its importance in the organisation of the stem are best seen in the study of the skeleton. Its chief and central part, the cartilaginous or bony vertebral column, affords an obvious instance of vertebrate metamerism; it consists of a series of cartilaginous or bony pieces, which have long been known as vertebrae (or spondyli). Each vertebra is directly connected with a special section of the muscular system, the nervous system, the vascular system, etc. Thus most of the "animal organs" take part in this vertebration. But we saw, when we were considering our own vertebrate character (in Chapter 1.11), that the same internal articulation is also found in the lowest primitive vertebrates, the acrania, although here the whole skeleton consists merely of the simple chorda, and is not at all articulated. Hence the articulation does not proceed primarily from the skeleton, but from the muscular system, and is clearly determined by the more advanced swimming-movements of the primitive chordonia-ancestors.

(FIGURES 1.153 TO 1.155. Sole-shaped embryonic disk of the chick, in three successive stages of development, looked at from the dorsal surface, magnified about twenty times, somewhat diagrammatic. Figure 1.153 with six pairs of somites. Brain a simple vesicle (hb). Medullary furrow still wide open from x; greatly widened at z. mp medullary plates, sp lateral plates, y limit of gullet-cavity (sh) and fore-gut (vd). Figure 1.154 with ten pairs of somites. Brain divided into three vesicles: v fore-brain, m middle-brain, h hind-brain, c heart, dv vitelline-veins. Medullary furrow still wide open behind (z). mp medullary plates. Figure 1.155 with sixteen pairs of somites. Brain divided into five vesicles: v fore-brain, z intermediate-brain, m middle-brain, h hind-brain, n after-brain, a optic vesicles, g auditory vesicles, c heart, dv vitelline veins, mp medullary plate, uw primitive vertebra.)

It is, therefore, wrong to describe the first rudimentary segments in the vertebrate embryo as primitive vertebrae or provertebrae; the fact that they have been so called for some time has led to much error and misunderstanding. Hence we shall give the name of "somites" or primitive segments to these so-called "primitive vertebrae." If the latter name is retained at all, it should only be used of the sclerotom—i.e., the small part of the somites from which the later vertebra does actually develop.

Articulation begins in all vertebrates at a very early embryonic stage, and this indicates the considerable phylogenetic age of the process. When the chordula (Figures 1.83 to 1.86) has completed its characteristic composition, often even a little earlier, we find in the amniotes, in the middle of the sole-shaped embryonic shield, several pairs of dark square spots, symmetrically distributed on both sides of the chorda (Figures 1.131 to 1.135). Transverse sections (Figure 1.93 uw) show that they belong to the stem-zone (episoma) of the mesoderm, and are separated from the parietal zone (hyposoma) by the lateral folds; in section they are still quadrangular, almost square, so that they look something like dice. These pairs of "cubes" of the mesoderm are the first traces of the primitive segments or somites, the so-called "protovertebrae." (Figures 1.153 to 1.155 uw).

(FIGURE 1.156. Embryo of the amphioxus, sixteen hours old, seen from the back. (From Hatschek.) d primitive gut, u primitive mouth, p polar cells of the mesoderm, c coelom-pouches, m their first segment, n medullary tube, i entoderm, e ectoderm, s first segment-fold.

FIGURE 1.157. Embryo of the amphioxus, twenty hours old, with five somites. (Right view; for left view see Figure 1.124.) (From Hatschek.) V fore end, H hind end. ak, mk, ik outer, middle, and inner germinal layers; dh alimentary canal, n neural tube, cn canalis neurentericus, ush coelom-pouches (or primitive-segment cavities), us1 first (and foremost) primitive segment.)

Among the mammals the embryos of the marsupials have three pairs of somites (Figure 1.131) after sixty hours, and eight pairs after seventy-two hours (Figure 1.135). They develop more slowly in the embryo of the rabbit; this has three somites on the eighth day (Figure 1.132), and eight somites a day later (Figure 1.134). In the incubated hen's egg the first somites make their appearance thirty hours after incubation begins (Figure 1.153). At the end of the second day the number has risen to sixteen or eighteen (Figure 1.155). The articulation of the stem-zone, to which the somites owe their origin, thus proceeds briskly from front to rear, new transverse constrictions of the "protovertebral plates" forming continuously and successively. The first segment, which is almost half-way down in the embryonic shield of the amniote, is the foremost of all; from this first somite is formed the first cervical vertebra with its muscles and skeletal parts. It follows from this, firstly, that the multiplication of the primitive segments proceeds backwards from the front, with a constant lengthening of the hinder end of the body; and, secondly, that at the beginning of segmentation nearly the whole of the anterior half of the sole-shaped embryonic shield of the amniote belongs to the later head, while the whole of the rest of the body is formed from its hinder half. We are reminded that in the amphioxus (and in our hypothetic primitive vertebrate, Figures 1.98 to 1.102) nearly the whole of the fore half corresponds to the head, and the hind half to the trunk.