The Evolution of Man

by Ernst Haeckel

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 XII.
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 gastræa 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 cœlomation 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.

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 gastræa 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.

Fig. 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 provertebræ (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 vertebræ (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 provertebræ, ch chorda, c body-cavity or cœloma, df gut-fibre-layer, dd gut-gland-layer, d gut-cavity, nb umbilical vesicle.

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 (Fig. 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 (Fig. 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.