THE EVOLUTION OF MAN
Volume I

CHAPTER XV
FŒTAL MEMBRANES AND CIRCULATION

Among the many interesting phenomena that we have encountered in the course of human embryology, there is an especial importance in the fact that the development of the human body follows from the beginning just the same lines as that of the other viviparous mammals. As a fact, all the embryonic peculiarities that distinguish the mammals from other animals are found also in man; even the ovum with its distinctive membrane (zona pellucida, Fig. 14) shows the same typical

Fig. 179—Human embryos from the second to the fifteenth week, seen from the left, the curved back turned towards the right. (Mostly from Ecker.) II of fourteen days. III of three weeks. IV of four weeks. V of five weeks. VI of six weeks. VII of seven weeks. VIII of eight weeks. XII of twelve weeks. XV of fifteen weeks.

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structure in all mammals (apart from the older oviparous monotremes). It has long since been deduced from the structure of the developed man that his natural place in the animal kingdom is among the mammals. Linné (1735) placed him in this class with the apes, in one and the same order (primates), in his Systema Naturæ. This position is fully confirmed by comparative embryology. We see that man entirely resembles the higher mammals, and most of all the apes, in embryonic development as well as in anatomic structure. And if we seek to understand this ontogenetic agreement in the light of the biogenetic law, we find that it proves clearly and necessarily the descent of man from a series of other mammals, and proximately from the primates. The common origin of man and the other mammals from a single ancient stem-form can no longer be questioned; nor can the immediate blood-relationship of man and the ape.

The essential agreement in the whole bodily form and inner structure is still visible in the embryo of man and the other mammals at the late stage of development at which the mammal-body can be recognised as such. But at a somewhat earlier stage, in which the limbs, gill-arches, sense-organs, etc., are already outlined, we cannot yet recognise the mammal embryos as such, or distinguish them from those of birds and reptiles. When we consider still earlier stages of development, we are unable to discover any essential difference in bodily structure between the embryos of these higher vertebrates and those of the lower, the amphibia and fishes. If, in fine, we go back to the construction of the body out of the four germinal layers, we are astonished to perceive that these four layers are the same in all vertebrates, and everywhere take a similar part in the building-up of the fundamental organs of the body. If we inquire as to the origin of these four secondary layers, we learn that they always arise in the same way from the two primary layers; and the latter have the same significance in all the metazoa (i.e., all animals except the unicellulars). Finally, we see that the cells which make up the primary germinal layers owe their origin in every case to the repeated cleavage of a single simple cell, the stem-cell or fertilised ovum.

Fig. 180—Very young human embryo of the fourth week, one-fourth of an inch long (taken from the womb of a suicide eight hours after death). (From Rabl.) n nasal pits, a eye, u lower jaw, z arch of hyoid bone, k3 and k4 third and fourth gill-arch, h heart; s primitive segments, vg fore-limb (arm), hg hind-limb (leg), between the two the ventral pedicle.

It is impossible to lay too much stress on this remarkable agreement in the chief embryonic features in man and the other animals. We shall make use of it later on for our monophyletic theory of descent—the hypothesis of a common descent of man and all the metazoa from the gastræa. The first rudiments of the principal parts of the body, especially the oldest organ, the alimentary canal, are the same everywhere; they have always the same extremely simple form. All the peculiarities that distinguish the various groups of animals from each other only appear gradually in the course of embryonic development; and the closer the relation of the various groups, the later they are found. We may formulate this phenomenon in a definite law, which may in a sense be regarded as an appendix to our biogenetic law. This is the law of the ontogenetic connection of related animal forms. It runs: The closer the

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relation of two fully-developed animals in respect of their whole bodily structure, and the nearer they are connected in the classification of the animal kingdom, the longer do their embryonic forms retain their identity, and the longer is it impossible (or only possible on the ground of subordinate features) to distinguish between their embryos. This law applies to all animals whose embryonic development is, in the main, an hereditary summary of their ancestral history, or in which the original form of development has been faithfully preserved by heredity. When, on the other hand, it has been altered by cenogenesis, or disturbance of development, we find a limitation of the law, which increases in proportion to the introduction of new features by adaptation (cf. Chapter I, pp. 4–6). Thus the apparent exceptions to the law can always be traced to cenogenesis.

Fig. 181—Human embryo of the middle of the fifth week, one-third of an inch long. (From Rabl.) Letters as in Fig. 180, except sk curve of skull, ok upper jaw, hb neck-indentation.

When we apply to man this law of the ontogenetic connection of related forms, and run rapidly over the earliest stages of human development with an eye to it, we notice first of all the structural identity of the ovum in man and the other mammals at the very beginning (Figs. 1, 14). The human ovum possesses all the distinctive features of the ovum of the viviparous mammals, especially the characteristic formation of its membrane (zona pellucida), which clearly distinguishes it from the ovum of all other animals. When the human fœtus has attained the age of fourteen days, it forms a round vesicle (or “embryonic vesicle”) about a quarter of an inch in diameter. A thicker part of its border forms a simple sole-shaped embryonic shield one-twelfth of an inch long (Fig. 133). On its dorsal side we find in the middle line the straight medullary furrow, bordered by the two parallel dorsal or medullary swellings. Behind, it passes by the neurenteric canal into the primitive gut or primitive groove. From this the folding of the two cœlom-pouches proceeds in the same way as in the other mammals (cf. Fig. 96, 97). In the middle of the sole-shaped embryonic shield the first primitive segments immediately begin to make their appearance. At this age the human embryo cannot be distinguished from that of other mammals, such as the hare or dog.

A week later (or after the twenty-first day) the human embryo has doubled its length; it is now about one-fifth of an inch long, and, when seen from the side, shows the characteristic bend of the back, the swelling of the head-end, the first outline of the three higher sense-organs, and the rudiments of the gill-clefts, which pierce the sides of the neck (Fig. 179, III). The allantois has grown out of the gut behind. The embryo is already entirely enclosed in the amnion, and is only connected in the middle of the belly by the vitelline duct with the embryonic vesicle, which changes into the yelk-sac. There are no extremities or limbs at this stage, no trace of arms or legs. The head-end has been strongly differentiated from the tail-end; and the first outlines of the cerebral vesicles in front, and the heart below, under the fore-arm, are already more or less clearly seen. There is as yet no real face. Moreover, we seek in vain at this stage a special character that may distinguish the human embryo from that of other mammals.

A week later (after the fourth week, on the twenty-eighth to thirtieth day of development) the human embryo has

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Fig. 182—Median longitudinal section of the tail of a human embryo, two-thirds of an inch long. (From Ross Granville Harrison.) Med medullary tube, Ca.fil caudal filament, ch chorda, ao caudal artery, V.c.i caudal vein, an anus, S.ug sinus urogenitalis.

reached a length of about one-third of an inch (Fig 179 IV). We can now clearly distinguish the head with its various parts; inside it the five primitive cerebral vesicles (fore-brain, middle-brain, intermediate-brain, hind-brain, and after-brain); under the head the gill-arches, which divide the gill-clefts; at the sides of the head the rudiments of the eyes, a couple of pits in the outer skin, with a pair of corresponding simple vesicles growing out of the lateral wall of the fore-brain (Figs. 180, 181 a). Far behind the eyes, over the last gill-arches, we see a vesicular rudiment of the auscultory organ. The rudimentary limbs are now clearly outlined—four simple buds of the shape of round plates, a pair of fore (vg) and a pair of hind legs (hg), the former a little larger than the latter. The large head bends over the trunk, almost at a right angle. The latter is still connected in the middle of its ventral side with the embryonic vesicle; but the embryo has still further severed itself from it, so that it already hangs out as the yelk-sac. The hind part of the body is also very much curved, so that the pointed tail-end is directed towards the head. The head and face-part are sunk entirely on the still open breast. The bend soon increases so much that the tail almost touches the forehead (Fig. 179 V.; Fig. 181). We may then distinguish three or four special curves on the round dorsal surface—namely, a skull-curve in the region of the second cerebral vesicle, a neck-curve at the beginning of the spinal cord, and a tail-curve at the fore-end. This pronounced curve is only shared by man and

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the higher classes of vertebrates (the amniotes); it is much slighter, or not found at all, in the lower vertebrates. At this age (four weeks) man has a considerable tail, twice as long as his legs. A vertical longitudinal section through the middle plane of this tail (Fig. 182) shows that the hinder end of the spinal marrow extends to the point of the tail, as also does the underlying chorda (ch), the terminal continuation of the vertebral column. Of the latter, the rudiments of the seven coccygeal (or lowest) vertebræ are visible—thirty-two indicates the third and thirty-six the seventh of these. Under the vertebral column we see the hindmost ends of the two large blood-vessels of the tail, the principal artery (aorta caudalis or arteria sacralis media, Ao), and the principal vein (vena caudalis or sacralis media). Underneath is the opening of the anus (an) and the urogenital sinus (S.ug). From this anatomic structure of the human tail it is perfectly clear that it is the rudiment of an ape-tail, the last hereditary relic of a long hairy tail, which has been handed down from our tertiary primate ancestors to the present day.

Fig. 183—Human embryo, four weeks old, opened on the ventral side. Ventral and dorsal walls are cut away, so as to show the contents of the pectoral and abdominal cavities. All the appendages are also removed (amnion, allantois, yelk-sac), and the middle part of the gut. n eye, 3 nose, 4 upper jaw, 5 lower jaw, 6 second, 6'' third gill-arch, ov heart (o right, o' left auricle; v right, v' left ventricle), b origin of the aorta, f liver (u umbilical vein), e gut (with vitelline artery, cut off at a'), j' vitelline vein, m primitive kidneys, t rudimentary sexual glands, r terminal gut (cut off at the mesentery z), n umbilical artery, u umbilical vein, 9 fore-leg, 9' hind-leg. (From Coste.) | Fig. 184—Human embryo, five weeks old, opened from the ventral side (as in Fig. 183). Breast and belly-wall and liver are removed. 3 outer nasal process, 4 upper jaw, 5 lower jaw, z tongue, v right, v' left ventricle of heart, o' left auricle, b origin of aorta, b', b'', b''' first, second, and third aorta-arches, c, c', c'' vena cava, ae lungs (y pulmonary artery), e stomach, m primitive kidneys (j left vitelline vein, s cystic vein, a right vitelline artery, n umbilical artery, u umbilical vein), x vitelline duct, i rectum, 8 tail, 9 fore-leg, 9' hind-leg. (From Coste.)

It sometimes happens that we find even external relics of this tail growing. According to the illustrated works of

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Surgeon-General Bernhard Ornstein, of Greece, these tailed men are not uncommon; it is not impossible that they gave rise to the ancient fables of the satyrs. A great number of such cases are given by Max Bartels in his essay on “Tailed Men” (1884, in the Archiv für Anthropologie, Band XV), and critically examined. These atavistic human tails are often mobile; sometimes they contain only muscles and fat, sometimes also rudiments of caudal vertebræ. They have a length of eight to ten inches and more. Granville Harrison has very carefully studied one of these cases of “pigtail,” which he removed by operation from a six months old child in 1901. The tail moved briskly when the child cried or was excited, and was drawn up when at rest.

Fig. 185—The head of Miss Julia Pastrana. (From a photograph by Hintze.) | Fig. 186—Human ovum of twelve to thirteen days (?). (From Allen Thomson.) 1. Not opened. 2. Opened and magnified. Within the outer chorion the tiny curved fœtus lies on the large embryonic vesicle, to the left above.

Fig. 187—Human ovum of ten days. (From Allen Thomson.) Opened; the small fœtus in the right half, above. Fig. 188—Human fœtus of ten days, taken from the preceding ovum, magnified, a yelk-sac, b neck (the medullary groove already closed), c head (with open medullary groove), d hind part (with open medullary groove), e a shred of the amnion. | Fig. 189—Human ovum of twenty to twenty-two days. (From Allen Thomson.) Opened. The chorion forms a spacious vesicle, to the inner wall of which the small fœtus (to the right above) is attached by a short umbilical cord. Fig. 190—Human fœtus of twenty to twenty-two days, taken from the preceding ovum, magnified. a amnion, b yelk-sac, c lower-jaw process of the first gill-arch, d upper-jaw process of same, e second gill-arch (two smaller ones behind). Three gill-clefts are clearly seen. f rudimentary fore-leg, g auditory vesicle, h eye, i heart.

In the opinion of some travellers and anthropologists, the atavistic tail-formation is hereditary in certain isolated tribes (especially in south-eastern Asia and the archipelago), so that we might speak of a special race or “species” of tailed men

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(Homo caudatus). Bartels has “no doubt that these tailed men will be discovered in the advance of our geographical and ethnographical knowledge of the lands in question” (Archiv für Anthropologie, Band XV, p. 129).

Fig. 191—Human embryo of sixteen to eighteen days. (From Coste.) Magnified. The embryo is surrounded by the amnion, (a), and lies free with this in the opened embryonic vesicle. The belly is drawn up by the large yelk-sac (d), and fastened to the inner wall of the embryonic membrane by the short and thick pedicle (b). Hence the normal convex curve of the back (Fig. 190) is here changed into an abnormal concave surface. h heart, m parietal mesoderm. The spots on the outer wall of the serolemma are the roots of the branching chorion-villi, which are free at the border.

When we open a human embryo of one month (Fig. 183), we find the alimentary canal formed in the body-cavity, and for the most part cut off from the embryonic vesicle. There are both mouth and anus apertures. But the mouth-cavity is not yet separated from the nasal cavity, and the face not yet shaped. The heart shows all its four sections; it is very large, and almost fills the whole of the pectoral cavity (Fig. 183 ov). Behind it are the very small rudimentary lungs. The primitive kidneys (m) are very large; they fill the greater part of the abdominal cavity, and extend from the liver (f) to the pelvic gut. Thus at the end of the first month all the chief organs are already outlined. But there are at this stage no features by which the human embryo materially differs from that of the dog, the hare, the ox, or the horse—in a word, of any other higher mammal. All these embryos have the same, or at least a very similar, form; they can at the most be

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distinguished from the human embryo by the total size of the body or some other insignificant difference in size. Thus, for instance, in man the head is larger in proportion to the trunk than in the ox. The tail is rather longer in the dog than in man. These are all negligible differences. On the other hand, the whole internal organisation and the form and arrangement of the various organs are essentially the same in the human embryo of four weeks as in the embryos of the other mammals at corresponding stages.

Fig. 192—Human embryo of the fourth week, one-third of an inch long, lying in the dissected chorion.

Fig. 193—Human embryo of the fourth week, with its membranes, like Fig. 192, but a little older. The yelk-sac is rather smaller, the amnion and chorion larger.

It is otherwise in the second month of human development. Fig. 179 represents a human embryo of six weeks (VI), one of seven weeks (VII), and one of eight weeks (VIII), at natural size. The differences which mark off the human embryo from that of the dog and the lower mammals now begin to be more pronounced. We can see important differences at the sixth, and still more at the eighth week, especially in the formation of the head. The size of the various sections of the brain is greater in man, and the tail is shorter.

Other differences between man and the lower mammals are found in the relative size of the internal organs. But even at this stage the human embryo differs very little from that of the nearest related mammals—the apes, especially the anthropomorphic apes.

The features by means of which we distinguish between them are not clear until later on. Even at a much more advanced stage of development, when we can distinguish the human fœtus from that of the ungulates at a glance, it still closely resembles that of the higher apes. At last we get the distinctive features, and

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we can distinguish the human embryo confidently at the first glance from that of all other mammals during the last four months of fœtal life—from the sixth to the ninth month of pregnancy. Then we begin to find also the differences between the various races of men, especially in regard to the formation of the skull and the face. (Cf. Chapter XXIII.)

| Fig. 194—Human embryo with its membranes, six weeks old. The outer envelope of the whole ovum is the chorion, thickly covered with its branching villi, a product of the serous membrane. The embryo is enclosed in the delicate amnion-sac. The yelk-sac is reduced to a small pear-shaped umbilical vesicle; its thin pedicle, the long vitelline duct, is enclosed in the umbilical cord. In the latter, behind the vitelline duct, is the much shorter pedicle of the allantois, the inner lamina of which (the gut-gland layer) forms a large vesicle in most of the mammals, while the outer lamina is attached to the inner wall of the outer embryonic coat, and forms the placenta there. (Half diagrammatic.)">

The striking resemblance that persists so long between the embryo of man and of the higher apes disappears much earlier in the lower apes. It naturally remains longest in the large anthropomorphic apes (gorilla, chimpanzee, orang, and gibbon). The physiognomic similarity of these animals, which we find so great in their earlier years, lessens with the increase of age. On the other hand, it remains throughout life in the remarkable long-nosed ape of Borneo (Nasalis larvatus). Its finely-shaped nose would be regarded with envy by many a man who has too little of that organ. If we compare the face of the long-nosed ape with that of abnormally ape-like human beings (such as the famous Miss Julia Pastrana, Fig. 185), it will be admitted to represent a higher stage of development. There are still people among us who look especially to the face for the “image of God in man.” The long-nosed ape would have more claim to this than some of the stumpy-nosed human individuals one meets.

This progressive divergence of the human from the animal form, which is based on the law of the ontogenetic connection between related forms, is found in the structure of the internal organs as well as in external form. It is also expressed in the construction of the envelopes and appendages that we find surrounding the fœtus externally, and that we will now consider more closely. Two of these appendages—the amnion and the allantois—are only found in the three higher classes of vertebrates, while the third, the yelk-sac, is found in most of the vertebrates. This is a circumstance of great importance, and it gives us valuable data for constructing man’s genealogical tree.

As regards the external membrane that encloses the ovum in the mammal womb,

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we find it just the same in man as in the higher mammals. The ovum is, the reader will remember, first surrounded by the transparent structureless ovolemma or zona pellucida (Figs. 1, 14). But very soon, even in the first week of development, this is replaced by the permanent chorion. This is formed from the external layer of the amnion, the serolemma, or “serous membrane,” the formation of which we shall consider presently; it surrounds the fœtus and its appendages as a broad, completely closed sac; the space between the two, filled with clear watery fluid, is the serocœlom, or interamniotic cavity (“extra-embryonic body-cavity”). But the smooth surface of the sac is quickly covered with numbers of tiny tufts, which are really hollow outgrowths like the fingers of a glove (Figs. 186, 191, 198 chz). They ramify and push into the corresponding depressions that are formed by the tubular glands of the mucous membrane of the maternal womb. Thus, the ovum secures its permanent seat (Fig. 186–194).

Fig. 195—Diagram of the embryonic organs of the mammal (fœtal membranes and appendages). (From Turner.) E, M, H outer, middle, and inner germ layer of the embryonic shield, which is figured in median longitudinal section, seen from the left. am amnion. AC amniotic cavity, UV yelk-sac or umbilical vesicle, ALC allantois, al pericœlom or serocœlom (inter-amniotic cavity), sz serolemma (or serous membrane), pc prochorion (with villi).)

In human ova of eight to twelve days this external membrane, the chorion, is already covered with small tufts or villi, and forms a ball or spheroid of one-fourth to one-third of an inch in diameter (Figs. 186–188). As a large quantity of fluid gathers inside it, the chorion expands more and more, so that the embryo only occupies a small part of the space within the vesicle. The villi of the chorion grow larger and more numerous. They branch out more and more. At first the villi cover the whole surface, but they afterwards disappear from the greater part of it; they then develop with proportionately greater vigour at a spot where the placenta is formed from the allantois.

When we open the chorion of a human embryo of three weeks, we find on the ventral side of the fœtus a large round sac, filled with fluid. This is the yelk-sac, or “umbilical vesicle,” the origin of which we have considered previously. The larger the embryo becomes the smaller we find the yelk-sac. In the end we find the remainder of it in the shape of a small pear-shaped vesicle, fastened to a long thin stalk (or pedicle), and hanging from the open belly of the fœtus (Fig. 194). This pedicle is the vitelline duct, and is separated from the body at the closing of the navel.

Behind the yelk-sac a second appendage,

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of much greater importance, is formed at an early stage at the belly of the mammal embryo. This is the allantois or “primitive urinary sac,” an important embryonic organ, only found in the three higher classes of vertebrates. In all the amniotes the allantois quickly appears at the hinder end of the alimentary canal, growing out of the cavity of the pelvic gut (Fig. 147 r, u, Fig. 195 ALC).

The further development of the allantois varies considerably in the three sub-classes of the mammals. The two lower sub-classes, monotremes and marsupials, retain the simpler structure of their ancestors, the reptiles. The wall of the allantois and the enveloping serolemma remains smooth and without villi, as in the birds. But in the third sub-class of the mammals the serolemma forms, by invagination at its outer surface, a number of hollow tufts or villi, from which it takes the name of the chorion or mallochorion. The gut-fibre layer of the allantois, richly supplied with branches of the umbilical vessel, presses into these tufts of the primary chorion, and forms the “secondary chorion.” Its embryonic blood-vessels are closely correlated to the contiguous maternal blood-vessels of the environing womb, and thus is formed the important nutritive apparatus of the embryo which we call the placenta.

Fig. 196—Diagrammatic frontal section of the pregnant human womb. (From Longet.) The embryo hangs by the umbilical cord, which encloses the pedicle of the allantois (al). nb umbilical vessel, am amnion, ch chorion, ds decidua serotina, dv decidua vera, dr decidua reflexa, z villi of the placenta, c cervix uteri, u uterus.)

The pedicle of the allantois, which connects the embryo with the placenta and conducts the strong umbilical vessels from the former to the latter, is covered by the amnion, and, with this amniotic sheath and the pedicle of the yelk-sac, forms what is called the umbilical cord (Fig. 196 al). As the large and blood-filled vascular network of the fœtal allantois attaches itself closely to the mucous lining of the maternal womb, and the partition between the blood-vessels of mother and child becomes much thinner, we get that remarkable nutritive apparatus of the fœtal body which is characteristic of the placentalia (or choriata). We shall return afterwards to the closer consideration of this (cf. Chapter XXIII).

In the various orders of mammals the placenta undergoes many modifications, and these are in part of great evolutionary importance and useful in classification. There is only one of these that need be specially mentioned—the important fact, established by Selenka in 1890, that the distinctive human placentation is confined to the anthropoids. In this most advanced group of the mammals the allantois is very small, soon loses its cavity, and then, in common with the amnion, undergoes certain peculiar changes. The umbilical cord develops in this case from what is called the “ventral pedicle.” Until very recently this was regarded as a structure peculiar to man. We now know from Selenka that the much-discussed ventral pedicle is merely the pedicle of the allantois, combined with the pedicle of the amnion and the rudimentary pedicle of the yelk-sac. It has just the same structure in the orang and gibbon (Fig. 197) and very probably in the chimpanzee and gorilla, as in man; it is, therefore, not a disproof, but a striking fresh proof, of the blood-relationship of man and the anthropoid apes.

We find only in the anthropoid apes—the gibbon and orang of Asia and the chimpanzee and gorilla of Africa—the peculiar and elaborate formation of the placenta that characterises man (Fig. 198).

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In this case there is at an early stage an intimate blending of the chorion of the embryo and the part of the mucous lining of the womb to which it attaches. The villi of the chorion with the blood-vessels they contain grow so completely into the tissue of the uterus, which is rich in blood, that it becomes impossible to separate them, and they form together a sort of cake. This comes away as the “afterbirth” at parturition; at the same time, the part of the mucous lining of the womb that has united inseparably with the chorion is torn away; hence it is called the decidua (“falling-away membrane”), and also the “sieve-membrane,” because it is perforated like a sieve. We find a decidua of this kind in most of the higher placentals; but it is only in man and the anthropoid apes that it divides into three parts—the outer, inner, and placental decidua. The external or true decidua (Fig. 196 du, Fig. 199 g) is the part of the mucous lining of the womb that clothes the inner surface of the uterine cavity wherever it is not connected with the placenta. The placental or spongy decidua (placentalis or serotina, Fig. 196 ds, Fig. 199 d) is really the placenta itself, or the maternal part of it (placenta uterina)—namely, that part of the mucous lining of the womb which unites intimately with the chorion-villi of the fœtal placenta. The internal or false decidua (interna or reflexa, Fig. 196 dr, Fig. 199 f) is that part of the mucous lining of the womb which encloses the remaining surface of the ovum, the smooth chorion (chorion læve), in the shape of a special thin membrane. The origin of these three different deciduous membranes, in regard to which quite erroneous views (still retained in their names) formerly prevailed, is now quite clear, The external decidua vera is the specially modified and subsequently detachable superficial stratum of the original mucous lining of the womb. The placental decidua serotina is that part of the preceding which is completely transformed by the ingrowth of the chorion-villi, and is used for constructing the placenta. The inner decidua reflexa is formed by the rise of a circular fold of the mucous lining (at the border of the decidua vera and serotina), which grows over the fœtus (like the anmnion) to the end.

Fig. 197—Male embryo of the Siamang-gibbon (Hylobates siamanga) of Sumatra; to the left the dissected uterus, of which only the dorsal half is given. The embryo has been taken out, and the limbs folded together; it is still connected by the umbilical cord with the centre of the circular placenta which is attached to the inside of the womb. This embryo takes the head-position in the womb, and this is normal in man also.

The peculiar anatomic features that characterise the human fœtal membranes are found in just the same way in the higher

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apes. Until recently it was thought that the human embryo was distinguished by its peculiar construction of a solid allantois and a special ventral pedicle, and that the umbilical cord developed from this in a different way than in the other mammals. The opponents of the unwelcome “ape-theory” laid great stress on this, and thought they had at last discovered an important indication that separated man from all the other placentals. But the remarkable discoveries published by the distinguished zoologist Selenka in 1890 proved that man shares these peculiarities of placentation with the anthropoid apes, though they are not found in the other apes. Thus the very feature which was advanced by our critics as a disproof became a most important piece of evidence in favour of our pithecoid origin.)

Fig. 198—Frontal section of the pregnant human womb. (From Turner.) The embryo (a month old) hangs in the middle of the amniotic cavity by the ventral pedicle or umbilical cord, which connects it with the placenta (above).

Of the three vesicular appendages of the amniote embryo which we have now described the amnion has no blood-vessels at any moment of its existence. But the other two vesicles, the yelk-sac and the allantois, are equipped with large blood-vessels, and these effect the nourishment of the embryonic body. We may take the opportunity to make a few general observations on the first circulation in the embryo and its central organ, the heart. The first blood-vessels, the heart, and the first blood itself, are formed from the gut-fibre layer. Hence it was called by earlier embryologists the “vascular layer.” In a sense the term is quite correct. But it must not be understood as if all the blood-vessels in the body came from this layer, or as if the whole of this layer were taken up only with the formation of blood-vessels. Neither of these suppositions is true. Blood-vessels may be formed independently in other parts, especially in the various products of the skin-fibre layer.

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Fig. 199—Human fœtus, twelve weeks old, with its membranes. The umbilical cord goes from its navel to the placenta. b amnion, c chorion, d placenta, d apostrophe, relics of villi on smooth chorion, f internal or reflex decidua, g external or true decidua. (From B. Schultze.)

Fig. 200—Mature human fœtus (at the end of pregnancy, in its natural position, taken out of the uterine cavity). On the inner surface of the latter (to the left) is the placenta, which is connected by the umbilical cord with the child’s navel. (From Bernhard Schultze.)

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The first blood-vessels of the mammal embryo have been considered by us previously, and we shall study the development of the heart in the second volume.

In every vertebrate it lies at first in the ventral wall of the fore-gut, or in the ventral (or cardiac) mesentery, by which it is connected for a time with the wall of the body. But it soon severs itself from the place of its origin, and lies freely in a cavity—the cardiac cavity. For a short time it is still connected with the former by the thin plate of the mesocardium. Afterwards it lies quite free in the cardiac cavity, and is only directly connected with the gut-wall by the vessels which issue from it.

Fig. 201—Vitelline vessels in the germinative area of a chick-embryo, at the close of the third day of incubation. (From Balfour.) The detached germinative area is seen from the ventral side: the arteries are dark, the veins light. H heart, AA aorta-arches, Ao aorta, R.of.A right omphalo-mesenteric artery, S.T. sinus terminalis, L.Of and R.Of right and left omphalo-mesenteric veins, S.V. sinus venosus, D.C. ductus Cuvieri, S.Ca.V. and V.Ca. fore and hind cardinal veins.

The fore-end of the spindle-shaped tube, which soon bends into an S-shape (Figure 1.202), divides into a right and left branch. These tubes are bent upwards arch-wise, and represent the first arches of the aorta. They rise in the wall of the fore-gut, which they enclose in a sense, and then unite above, in the upper wall of the fore gut-cavity, to form a large single artery, that runs backward immediately under the chorda, and is called the aorta (Fig. 201 Ao). The first pair of aorta-arches rise on the inner wall of the first pair of gill-arches, and so lie between the first gill-arch (k) and the fore-gut (d), just as we find them throughout life in the fishes. The single aorta, which results from the conjunction of these two first vascular arches, divides again immediately into two parallel branches, which run backwards on either side of the chorda. These are the primitive aortas which we have already mentioned; they are also called the posterior vertebral arteries. These two arteries now give off at each side, behind, at right angles, four or five branches, and these pass from the embryonic body to the germinative area, they

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are called omphalo-mesenteric or vitelline arteries. They represent the first beginning of a fœtal circulation. Thus, the first blood-vessels pass over the embryonic body and reach as far as the edge of the germinative area. At first they are confined to the dark or “vascular” area. But they afterwards extend over the whole surface of the embryonic vesicle. In the end, the whole of the yelk-sac is covered with a vascular net-work. These vessels have to gather food from the contents of the yelk-sac and convey it to the embryonic body. This is done by the veins, which pass first from the germinative area, and afterwards from the yelk-sac, to the farther end of the heart. They are called vitelline, or, frequently, omphalo-mesenteric, veins.

These vessels naturally atrophy with the degeneration of the umbilical vesicle, and the vitelline circulation is replaced by a second, that of the allantois. Large blood-vessels are developed in the wall of the urinary sac or the allantois, as before, from the gut-fibre layer. These vessels grow larger and larger, and are very closely connected with the vessels that develop in the body of the embryo itself. Thus, the secondary, allantoic circulation gradually takes the place of the original vitelline circulation. When the allantois has attached itself to the inner wall of the chorion and been converted into the placenta, its blood-vessels alone effect the nourishment of the embryo. They are called umbilical vessels, and are originally double—a pair of umbilical arteries and a pair of umbilical veins. The two umbilical veins (Fig. 183 u), which convey blood from the placenta to the heart, open it first into the united vitelline veins. The latter then disappear, and the right umbilical vein goes with them, so that henceforth a single large vein, the left umbilical vein, conducts all the blood from the placenta to the heart of the embryo. The two arteries of the allantois, or the umbilical arteries (Figs. 183 n, 184 n), are merely the ultimate terminations of the primitive aortas, which are strongly developed afterwards. This umbilical circulation is retained until the nine months of embryonic life are over, and the human embryo enters into the world as the independent individual. The umbilical cord (Fig. 196 al), in which these large blood-vessels pass from the embryo to the placenta, comes away, together with the latter, in the after-birth, and with the use of the lungs begins an entirely new form of circulation, which is confined to the body of the infant.

Fig. 202—Boat-shaped embryo of the dog, from the ventral side, magnified. In front under the forehead we can see the first pair of gill-arches; underneath is the S-shaped heart, at the sides of which are the auditory vesicles. The heart divides behind into the two vitelline veins, which expand in the germinative area (which is torn off all round). On the floor of the open belly lie, between the protovertebræ, the primitive aortas, from which five pairs of vitelline arteries are given off. (From Bischoff.)

There is a great phylogenetic significance in the perfect agreement which we find between man and the anthropoid apes in these important features of embryonic circulation, and the special construction of the placenta and the umbilical cord. We must infer from it a close blood-relationship of man and the anthropomorphic apes—a common descent of them from one and the same extinct group of lower apes. Huxley’s “pithecometra-principle” applies to these ontogenetic features as much as to any other morphological relations: “The differences in construction of any part of the body are less between man and the anthropoid apes than between the latter and the lower apes.”

This important Huxleian law, the chief consequence of which is “the descent of man from the ape,” has lately been confirmed in an interesting and unexpected way from the side of the experimental

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physiology of the blood. The experiments of Hans Friedenthal at Berlin have shown that human blood, mixed with the blood of lower apes, has a poisonous effect on the latter; the serum of the one destroys the blood-cells of the other. But this does not happen when human blood is mixed with that of the anthropoid ape. As we know from many other experiments that the mixture of two different kinds of blood is only possible without injury in the case of two closely related animals of the same family, we have another proof of the close blood-relationship, in the literal sense of the word, of man and the anthropoid ape.

Fig. 203—Lar or white-handed gibbon (Hylobates lar or albimanus), from the Indian mainland (From Brehm.)

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Fig. 204—Young orang (Satyrus orang), asleep.

The existing anthropoid apes are only a small remnant of a large family of eastern apes (or Catarrhinæ), from which man was evolved about the end of the Tertiary period. They fall into two geographical groups—the Asiatic and the African anthropoids. In each group we can distinguish two genera. The oldest of these four genera is the gibbon Hylobates, Fig. 203); there are from eight to twelve species of it in the East Indies. I made observations of four of them during my voyage in the East Indies (1901), and had a specimen of the ash-grey gibbon (Hylobates leuciscus) living for several months in the garden of my house in Java. I have described the interesting habits of this ape (regarded by the Malays as the wild descendant of men who had lost their way) in my Malayischen

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Reisebriefen (chap. xi). Psychologically, he showed a good deal of resemblance to the children of my Malay hosts, with whom he played and formed a very close friendship.

Fig. 205—Wild orang (Dyssatyrus auritius). (From R. Fick and Leutemann.).

The second, larger and stronger, genus of Asiatic anthropoid ape is the orang (Satyrus); he is now found only in the islands of Borneo and Sumatra. Selenka, who has published a very thorough Study of the Development and Cranial Structure of the Anthropoid Apes (1899), distinguishes ten races of the orang, which may, however, also be regarded as “local varieties or species.” They fall into two sub-genera or genera: one group, Dyssatyrus (orang-bentang, Fig. 205), is distinguished for the strength of its limbs, and the formation of very peculiar and salient cheek-pads in the elderly male; these are wanting in the other group, the ordinary orang-outang (Eusatyrus).

Several species have lately been distinguished in the two genera of the black African anthropoid apes (chimpanzee and gorilla). In the genus Anthropithecus (or Anthropopithecus, formerly Troglodytes), the bald-headed chimpanzee, A. calvus (Fig. 206), and the gorilla-like A. mafuca differ very strikingly from the ordinary Anthropithecus niger (Fig. 207), not only in the size and proportion of many parts of the body, but also in the peculiar shape of the head, especially the ears and lips, and in the hair and colour. The controversy that still continues as to whether these different forms of

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Fig. 206—The bald-headed chimpanzee (Anthropithecus calvus). Female. This fresh species, described by Frank Beddard in 1897 as Troglodytes calvus, differs considerably from the ordinary A. niger Fig. 207) in the structure of the head, the colouring, and the absence of hair in parts.

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chimpanzee and orang are “merely local varieties” or “true species” is an idle one; as in all such disputes of classifiers there is an utter absence of clear ideas as to what a species really is.

Of the largest and most famous of all the anthropoid apes, the gorilla, Paschen has lately discovered a giant-form in the interior of the Cameroons, which seems to differ from the ordinary species (Gorilla gina Fig. 208), not only by its unusual size and strength, but also by a special formation of the skull. This giant gorilla (Gorilla gigas, Fig. 209) is six feet eight inches long; the span of its great arms is about nine feet; its powerful chest is twice as broad as that of a strong man.

Fig. 207—Female chimpanzee (Anthropithecus niger). (From Brehm.)

The whole structure of this huge anthropoid ape is not merely very similar to that of man, but it is substantially the same. “The same 200 bones, arranged in the same way, form our internal skeleton; the same 300 muscles effect our movements; the same hair covers our skin; the same groups of ganglionic cells compose the ingenious mechanism of our brain; the same four-chambered heart is the central pump of our circulation.” The really existing differences in the shape and size of the various parts are explained by differences in their growth, due to adaptation to different habits of life and unequal use of the various organs. This of itself proves morphologically the descent of man from the ape. We will return to the point in Chapter XXIII. But I wanted to point already to this important solution of “the question of questions,” because that agreement

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in the formation of the embryonic membranes and in fœtal circulation which I have described affords a particularly weighty proof of it. It is the more instructive as even cenogenetic structures may in certain circumstances have a high phylogenetic value. In conjunction with the other facts, it affords a striking confirmation of our biogenetic law.

Fig. 208—Female gorilla. (From Brehm.).

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Fig. 209—Male giant-gorilla (Gorilla gigas), from Yaunde, in the interior of the Cameroons. Killed by H. Paschen, stuffed by Umlauff.

Title and Contents
Glossary
Chapter XIV
Vol. II Title
Figs. 1–209
Figs. 210–408