The Evolution of Man Volume I Part 13

Closely related morphologically and physiologically to the alimentary ca.n.a.l is the vascular system of the vertebrate, the chief sections of which develop from the fibrous gut-layer. It consists of two different but directly connected parts, the system of blood-vessels and that of lymph-vessels. In the pa.s.sages of the one we find red blood, and in the other colourless lymph. To the lymphatic system belong, first of all, the lymphatic ca.n.a.ls proper or absorbent veins, which are distributed among all the organs, and absorb the used-up juices from the tissues, and conduct them into the venous blood; but besides these there are the chyle-vessels, which absorb the white chyle, the milky fluid prepared by the alimentary ca.n.a.l from the food, and conduct this also to the blood.

The blood-vessel system of the vertebrate has a very elaborate construction, but seems to have had a very simple form in the primitive vertebrate, as we find it to-day permanently in the annelids (for instance, earth-worms) and the amphioxus. We accordingly distinguish first of all as essential, original parts of it two large single blood-ca.n.a.ls, which lie in the fibrous wall of the gut, and run along the alimentary ca.n.a.l in the median plane of the body, one above and the other underneath the ca.n.a.l. These princ.i.p.al ca.n.a.ls give out numerous branches to all parts of the body, and pa.s.s into each other by arches before and behind; we will call them the primitive artery and the primitive vein. The first corresponds to the dorsal vessel, the second to the ventral vessel, of the worms. The primitive or princ.i.p.al artery, usually called the aorta (Figure 1.98 a), lies above the gut in the middle line of its dorsal side, and conducts oxidised or arterial blood from the gills to the body. The primitive or princ.i.p.al vein (Figure 1.100 v) lies below the gut, in the middle line of its ventral side, and is therefore also called the vena subintestinalis; it conducts carbonised or venous blood back from the body to the gills. At the branchial section of the gut in front the two ca.n.a.ls are connected by a number of branches, which rise in arches between the gill-clefts. These "branchial vascular arches" (kg) run along the gill-arches, and have a direct share in the work of respiration. The anterior continuation of the princ.i.p.al vein which runs on the ventral wall of the gill-gut, and gives off these vascular arches upwards, is the branchial artery (ka). At the border of the two sections of the ventral vessel it enlarges into a contractile spindle-shaped tube (Figures 1.98 and 1.100 h). This is the first outline of the heart, which afterwards becomes a four-chambered pump in the higher vertebrates and man. There is no heart in the amphioxus, probably owing to degeneration. In prospondylus the ventral gill-heart probably had the simple form in which we still find it in the ascidia and the embryos of the craniota (Figures 1.98 and 1.100 h).

The kidneys, which act as organs of excretion or urinary organs in all vertebrates, have a very different and elaborate construction in the various sections of this stem; we will consider them further in Chapter 2.29. Here I need only mention that in our hypothetical primitive vertebrate they probably had the same form as in the actual amphioxus--the primitive kidneys (protonephra). These are originally made up of a double row of little ca.n.a.ls, which directly convey the used-up juices or the urine out of the body-cavity (Figure 1.102 n).

The inner aperture of these p.r.o.nephridial ca.n.a.ls opens with a ciliated funnel into the body-cavity; the external aperture opens in lateral grooves of the epidermis, a couple of longitudinal grooves in the lateral surface of the outer skin (Figure 1.102 b). The p.r.o.nephridial duct is formed by the closing of this groove to the right and left at the sides. In all the craniota it develops at an early stage in the h.o.r.n.y plate; in the amphioxus it seems to be converted into a wide cavity, the atrium, or peribranchial s.p.a.ce.

Next to the kidneys we have the s.e.xual organs of the vertebrate. In most of the members of this stem the two are united in a single urogenital system; it is only in a few groups that the urinary and s.e.xual organs are separated (in the amphioxus, the cyclostoma, and some sections of the fish-cla.s.s). In man and all the higher vertebrates the s.e.xual apparatus is made up of various parts, which we will consider in Chapter 2.29. But in the two lowest cla.s.ses of our stem, the acrania and cyclostoma, they consist merely of simple s.e.xual glands or gonads, the ovaries of the female s.e.x and the t.e.s.t.i.c.l.es (spermaria) of the male; the former provide the ova, the latter the sperm. In the craniota we always find only one pair of gonads; in the amphioxus several pairs, arranged in succession. They must have had the same form in our hypothetical prospondylus (Figures 1.98 and 1.100 s). These segmental pairs of gonads are the original ventral halves of the coelom-pouches.

The organs which we have now enumerated in this general survey, and of which we have noted the characteristic disposition, are those parts of the organism that are found in all vertebrates without exception in the same relation to each other, however much they may be modified. We have chiefly had in view the transverse section of the body (Figures 1.101 and 1.102), because in this we see most clearly the distinctive arrangement of them. But to complete our picture we must also consider the segmentation or metamera-formation of them, which has yet been hardly noticed, and which is seen best in the longitudinal section. In man and all the more advanced vertebrates the body is made up of a series or chain of similar members, which succeed each other in the long axis of the body--the segments or metamera of the organism. In man these h.o.m.ogeneous parts number thirty-three in the trunk, but they run to several hundred in many of the vertebrates (such as serpents or eels). As this internal articulation or metamerism is mainly found in the vertebral column and the surrounding muscles, the sections or metamera were formerly called pro-vertebrae. As a fact, the articulation is by no means chiefly determined and caused by the skeleton, but by the muscular system and the segmental arrangement of the kidneys and gonads. However, the composition from these pro-vertebrae or internal metamera is usually, and rightly, put forward as a prominent character of the vertebrate, and the manifold division or differentiation of them is of great importance in the various groups of the vertebrates. But as far as our present task--the derivation of the simple body of the primitive vertebrate from the chordula--is concerned, the articulate parts or metamera are of secondary interest, and we need not go into them just now.

(FIGURE 1.103 A, B, C, D. Instances of redundant mammary glands and nipples (hypermastism). A a pair of small redundant b.r.e.a.s.t.s (with two nipples on the left) above the large normal ones; from a 45-year-old Berlin woman, who had had children 17 times (twins twice). (From Hansemann.) B the highest number: ten nipples (all giving milk), three pairs above, one pair below, the large normal b.r.e.a.s.t.s; from a 22-year-old servant at Warschau. (From Neugebaur.) C three pairs of nipples: two pairs on the normal glands and one pair above; from a 19-year-old j.a.panese girl. D four pairs of nipples: one pair above the normal and two pairs of small accessory nipples underneath; from a 22-year-old Bavarian soldier. (From Wiedersheim.))

The characteristic composition of the vertebrate body develops from the embryonic structure in the same way in man as in all the other vertebrates. As all competent experts now admit the monophyletic origin of the vertebrates on the strength of this significant agreement, and this "common descent of all the vertebrates from one original stem-form" is admitted as an historical fact, we have found the answer to "the question of questions." We may, moreover, point out that this answer is just as certain and precise in the case of the origin of man from the mammals. This advanced vertebrate cla.s.s is also monophyletic, or has evolved from one common stem-group of lower vertebrates (reptiles, and, earlier still, amphibia). This follows from the fact that the mammals are clearly distinguished from the other cla.s.ses of the stem, not merely in one striking particular, but in a whole group of distinctive characters.

It is only in the mammals that we find the skin covered with hair, the breast-cavity separated from the abdominal cavity by a complete diaphragm, and the larynx provided with an epiglottis. The mammals alone have three small auscultory bones in the tympanic cavity--a feature that is connected with the characteristic modification of their maxillary joint. Their red blood-cells have no nucleus, whereas this is retained in all other vertebrates. Finally, it is only in the mammals that we find the remarkable function of the breast structure which has given its name to the whole cla.s.s--the feeding of the young by the mother's milk. The mammary glands which serve this purpose are interesting in so many ways that we may devote a few lines to them here.

As is well known, the lower mammals, especially those which beget a number of young at a time, have several mammary glands at the breast.

Hedgehogs and sows have five pairs, mice four or five pairs, dogs and squirrels four pairs, cats and bears three pairs, most of the ruminants and many of the rodents two pairs, each provided with a teat or nipple (mastos). In the various genera of the half-apes (lemurs) the number varies a good deal. On the other hand, the bats and apes, which only beget one young at a time as a rule, have only one pair of mammary glands, and these are found at the breast, as in man.

These variations in the number or structure of the mammary apparatus (mammarium) have become doubly interesting in the light of recent research in comparative anatomy. It has been shown that in man and the apes we often find redundant mammary glands (hyper-mastism) and corresponding teats (hyper-thelism) in both s.e.xes. Figure 1.103 shows four cases of this kind--A, B, and C of three women, and D of a man.

They prove that all the above-mentioned numbers may be found occasionally in man. Figure 1.103 A shows the breast of a Berlin woman who had had children seventeen times, and who has a pair of small accessory b.r.e.a.s.t.s (with two nipples on the left one) above the two normal b.r.e.a.s.t.s; this is a common occurrence, and the small soft pad above the breast is not infrequently represented in ancient statues of Venus. In Figure 1.103 C we have the same phenomenon in a j.a.panese girl of nineteen, who has two nipples on each breast besides (three pairs altogether). Figure 1.103 D is a man of twenty-two with four pairs of nipples (as in the dog), a small pair above and two small pairs beneath the large normal teats. The maximum number of five pairs (as in the sow and hedgehog) was found in a Polish servant of twenty-two who had had several children; milk was given by each nipple; there were three pairs of redundant nipples above and one pair underneath the normal and very large b.r.e.a.s.t.s (Figure 1.103 B).

A number of recent investigations (especially among recruits) have shown that these things are not uncommon in the male as well as the female s.e.x. They can only be explained by evolution, which attributes them to atavism and latent heredity. The earlier ancestors of all the primates (including man) were lower placentals, which had, like the hedgehog (one of the oldest forms of the living placentals), several mammary glands (five or more pairs) in the abdominal skin. In the apes and man only a couple of them are normally developed, but from time to time we get a development of the atrophied structures. Special notice should be taken of the arrangement of these accessory mammae; they form, as is clearly seen in Figure 1.103 B and D, two long rows, which diverge forward (towards the arm-pit), and converge behind in the middle line (towards the loins). The milk-glands of the polymastic lower placentals are arranged in similar lines.

The phylogenetic explanation of polymastism, as given in comparative anatomy, has lately found considerable support in ontogeny. Hans Strahl, E. Schmitt, and others, have found that there are always in the human embryo at the sixth week (when it is three-fifths of an inch long) the microscopic traces of five pairs of mammary glands, and that they are arranged at regular distances in two lateral and divergent lines, which correspond to the mammary lines. Only one pair of them--the central pair--are normally developed, the others atrophying.

Hence there is for a time in the human embryo a normal hyperthelism, and this can only be explained by the descent of man from lower primates (lemurs) with several pairs.

But the milk-gland of the mammal has a great morphological interest from another point of view. This organ for feeding the young in man and the higher mammals is, as is known, found in both s.e.xes. However, it is usually active only in the female s.e.x, and yields the valuable "mother's milk"; in the male s.e.x it is small and inactive, a real rudimentary organ of no physiological interest. Nevertheless, in certain cases we find the breast as fully developed in man as in woman, and it may give milk for feeding the young.

(FIGURE 1.104. A Greek gynecomast.)

We have a striking instance of this gynecomastism (large milk-giving b.r.e.a.s.t.s in a male) in Figure 1.104. I owe the photograph (taken from life) to the kindness of Dr. Ornstein, of Athens, a German physician, who has rendered service by a number of anthropological observations, (for instance, in several cases of tailed men). The gynecomast in question is a Greek recruit in his twentieth year, who has both normally developed male organs and very p.r.o.nounced female b.r.e.a.s.t.s. It is noteworthy that the other features of his structure are in accord with the softer forms of the female s.e.x. It reminds us of the marble statues of hermaphrodites which the ancient Greek and Roman sculptors often produced. But the man would only be a real hermaphrodite if he had ovaries internally besides the (externally visible) t.e.s.t.i.c.l.es.

I observed a very similar case during my stay in Ceylon (at Belligemma) in 1881. A young Cinghalese in his twenty-fifth year was brought to me as a curious hermaphrodite, half-man and half-woman. His large b.r.e.a.s.t.s gave plenty of milk; he was employed as "male nurse" to suckle a new-born infant whose mother had died at birth. The outline of his body was softer and more feminine than in the Greek shown in Figure 1.104. As the Cinghalese are small of stature and of graceful build, and as the men often resemble the women in clothing (upper part of the body naked, female dress on the lower part) and the dressing of the hair (with a comb), I first took the beardless youth to be a woman. The illusion was greater, as in this remarkable case gynecomastism was a.s.sociated with cryptorchism--that is to say, the t.e.s.t.i.c.l.es had kept to their original place in the visceral cavity, and had not travelled in the normal way down into the s.c.r.o.t.u.m. (Cf.

Chapter 2.29.) Hence the latter was very small, soft, and empty.

Moreover, one could feel nothing of the t.e.s.t.i.c.l.es in the inguinal ca.n.a.l. On the other hand, the male organ was very small, but normally developed. It was clear that this apparent hermaphrodite also was a real male.

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 s.e.xes by heredity; and they have been maintained in all persons of either s.e.x, although they are not physiologically active in the males. This normal permanence of the female lactiferous organs in BOTH s.e.xes 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 cla.s.ses 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 cla.s.ses 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 cla.s.sical 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 ant.i.thesis 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-ma.s.s 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 ant.i.thesis 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 inv.a.g.i.n.ation. 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 ant.i.thetic 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 ca.n.a.l, 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 princ.i.p.al 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 pa.s.sed 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 pa.s.sed 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 pa.s.s 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 pa.s.sed 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 alb.u.min 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.)