(FIGURE 1.13. Ova of various animals, executing amoeboid movements, highly magnified. All the ova are naked cells of varying shape. In the dark fine-grained protoplasm (yelk) is a large vesicular nucleus (the germinal vesicle), and in this is seen a nuclear body (the germinal spot), in which again we often see a germinal point. Figures A1 to A4 represent the ovum of a sponge (Leuculmis echinus) in four successive movements. B1 to B8 are the ovum of a parasitic crab (Chondracanthus cornutus), in eight successive movements. (From Edward von Beneden.) C1 to C5 show the ovum of the cat in various stages of movement (from Pfluger); Figure P the ovum of a trout; E the ovum of a chicken; F a human ovum.)
The fertilised bird-ovum (Figure 1.15) is notably different. It is true that in its earliest stage (Figure 1.13 E) this ovum also is very like that of the mammal (Figure 1.13 F). But afterwards, while still within the oviduct, it takes up a quantity of nourishment and works this into the familiar large yellow yelk. When we examine a very young ovum in the hen's oviduct, we find it to be a simple, small, naked, amoeboid cell, just like the young ova of other animals (Figure 1.13). But it then grows to the size we are familiar with in the round yelk of the egg. The nucleus of the ovum, or the germinal vesicle, is thus pressed right to the surface of the globular ovum, and is embedded there in a small quantity of transparent matter, the so-called white yelk. This forms a round white spot, which is known as the "tread" (cicatricula) (Figure 1.15 b). From the tread a thin column of the white yelk penetrates through the yellow yelk to the centre of the globular cell, where it swells into a small, central globule (wrongly called the yelk-cavity, or latebra, Figure 1.15 d apostrophe). The yellow yelk-matter which surrounds this white yelk has the appearance in the egg (when boiled hard) of concentric layers (c). The yellow yelk is also enclosed in a delicate structureless membrane (the membrana vitellina, a).
As the large yellow ovum of the bird attains a diameter of several inches in the bigger birds, and encloses round yelk-particles, there was formerly a reluctance to consider it as a simple cell. This was a mistake. Every animal that has only one cell-nucleus, every amoeba, every gregarina, every infusorium, is unicellular, and remains unicellular whatever variety of matter it feeds on. So the ovum remains a simple cell, however much yellow yelk it afterwards accumulates within its protoplasm. It is, of course, different, with the bird's egg when it has been fertilised. The ovum then consists of as many cells as there are nuclei in the tread. Hence, in the fertilised egg which we eat daily, the yellow yelk is already a multicellular body. Its tread is composed of several cells, and is now commonly called the germinal disc. We shall return to this discogastrula in Chapter 1.9.
(FIGURE 1.14. The human ovum, taken from the female ovary, magnified 500 times. The whole ovum is a simple round cell. The chief part of the globular mass is formed by the nuclear yelk (deutoplasm), which is evenly distributed in the active protoplasm, and consists of numbers of fine yelk-granules. In the upper part of the yelk is the transparent round germinal vesicle, which corresponds to the nucleus. This encloses a darker granule, the germinal spot, which shows a nucleolus. The globular yelk is surrounded by the thick transparent germinal membrane (ovolemma, or zona pellucida). This is traversed by numbers of lines as fine as hairs, which are directed radially towards the centre of the ovum. These are called the pore-canals; it is through these that the moving spermatozoa penetrate into the yelk at impregnation.
FIGURE 1.15. A fertilised ovum from the oviduct of a hen. the yellow yelk (c) consists of several concentric layers (d), and is enclosed in a thin yelk-membrane (a). The nucleus or germinal vesicle is seen above in the cicatrix or "tread" (b). From that point the white yelk penetrates to the central yelk-cavity (d apostrophe). The two kinds of yelk do not differ very much.
FIGURE 1.16. A creeping amoeba (highly magnified). The whole organism is a simple naked cell, and moves about by means of the changing arms which it thrusts out of and withdraws into its protoplasmic body. Inside it is the roundish nucleus with its nucleolus.)
When the mature bird-ovum has left the ovary and been fertilised in the oviduct, it covers itself with various membranes which are secreted from the wall of the oviduct. First, the large clear albuminous layer is deposited around the yellow yelk; afterwards, the hard external shell, with a fine inner skin. All these gradually forming envelopes and processes are of no importance in the formation of the embryo; they serve merely for the protection of the original simple ovum. We sometimes find extraordinarily large eggs with strong envelopes in the case of other animals, such as fishes of the shark type. Here, also, the ovum is originally of the same character as it is in the mammal; it is a perfectly simple and naked cell. But, as in the case of the bird, a considerable quantity of nutritive yelk is accumulated inside the original yelk as food for the developing embryo; and various coverings are formed round the egg. The ovum of many other animals has the same internal and external features. They have, however, only a physiological, not a morphological, importance; they have no direct influence on the formation of the foetus. They are partly consumed as food by the embryo, and partly serve as protective envelopes. Hence we may leave them out of consideration altogether here, and restrict ourselves to material points—TO THE SUBSTANTIAL IDENTITY OF THE ORIGINAL OVUM IN MAN AND THE REST OF THE ANIMALS (Figure 1.13).
Now, let us for the first time make use of our biogenetic law; and directly apply this fundamental law of evolution to the human ovum. We reach a very simple, but very important, conclusion. FROM THE FACT THAT THE HUMAN OVUM AND THAT OF ALL OTHER ANIMALS CONSISTS OF A SINGLE CELL, IT FOLLOWS IMMEDIATELY, ACCORDING TO THE BIOGENETIC LAW, THAT ALL THE ANIMALS, INCLUDING MAN, DESCEND FROM A UNICELLULAR ORGANISM. If our biogenetic law is true, if the embryonic development is a summary or condensed recapitulation of the stem-history—and there can be no doubt about it—we are bound to conclude, from the fact that all the ova are at first simple cells, that all the multicellular organisms originally sprang from a unicellular being. And as the original ovum in man and all the other animals has the same simple and indefinite appearance, we may assume with some probability that this unicellular stem-form was the common ancestor of the whole animal world, including man. However, this last hypothesis does not seem to me as inevitable and as absolutely certain as our first conclusion.
This inference from the unicellular embryonic form to the unicellular ancestor is so simple, but so important, that we cannot sufficiently emphasise it. We must, therefore, turn next to the question whether there are to-day any unicellular organisms, from the features of which we may draw some approximate conclusion as to the unicellular ancestors of the multicellular organisms. The answer is: Most certainly there are. There are assuredly still unicellular organisms which are, in their whole nature, really nothing more than permanent ova. There are independent unicellular organisms of the simplest character which develop no further, but reproduce themselves as such, without any further growth. We know to-day of a great number of these little beings, such as the gregarinae, flagellata, acineta, infusoria, etc. However, there is one of them that has an especial interest for us, because it at once suggests itself when we raise our question, and it must be regarded as the unicellular being that approaches nearest to the real ancestral form. This organism is the amoeba.
For a long time now we have comprised under the general name of amoebae a number of microscopic unicellular organisms, which are very widely distributed, especially in fresh-water, but also in the ocean; in fact, they have lately been discovered in damp soil. There are also parasitic amoebae which live inside other animals. When we place one of these amoebae in a drop of water under the microscope and examine it with a high power, it generally appears as a roundish particle of a very irregular and varying shape (Figures 1.16 and 1.17). In its soft, slimy, semi-fluid substance, which consists of protoplasm, we see only the solid globular particle it contains, the nucleus. This unicellular body moves about continually, creeping in every direction on the glass on which we are examining it. The movement is effected by the shapeless body thrusting out finger-like processes at various parts of its surface; and these are slowly but continually changing, and drawing the rest of the body after them. After a time, perhaps, the action changes. The amoeba suddenly stands still, withdraws its projections, and assumes a globular shape. In a little while, however, the round body begins to expand again, thrusts out arms in another direction, and moves on once more. These changeable processes are called "false feet," or pseudopodia, because they act physiologically as feet, yet are not special organs in the anatomic sense. They disappear as quickly as they come, and are nothing more than temporary projections of the semi-fluid and structureless body.
(FIGURE 1.17. Division of a unicellular amoeba (Amoeba polypodia) in six stages. (From F.E. Schultze.) the dark spot is the nucleus, the lighter spot a contractile vacuole in the protoplasm. The latter reforms in one of the daughter-cells.)
FIGURE 1.18. Ovum of a sponge (Olynthus). The ovum creeps about in a body of the sponge by thrusting out ever-changing processes. It is indistinguishable from the common amoeba.)
If you touch one of these creeping amoebae with a needle, or put a drop of acid in the water, the whole body at once contracts in consequence of this mechanical or physical stimulus. As a rule, the body then resumes its globular shape. In certain circumstances—for instance, if the impurity of the water lasts some time—the amoeba begins to develop a covering. It exudes a membrane or capsule, which immediately hardens, and assumes the appearance of a round cell with a protective membrane. The amoeba either takes its food directly by imbibition of matter floating in the water, or by pressing into its protoplasmic body solid particles with which it comes in contact. The latter process may be observed at any moment by forcing it to eat. If finely ground colouring matter, such as carmine or indigo, is put into the water, you can see the body of the amoeba pressing these coloured particles into itself, the substance of the cell closing round them. The amoeba can take in food in this way at any point on its surface, without having any special organs for intussusception and digestion, or a real mouth or gut.
The amoeba grows by thus taking in food and dissolving the particles eaten in its protoplasm. When it reaches a certain size by this continual feeding, it begins to reproduce. This is done by the simple process of cleavage (Figure 1.17). First, the nucleus divides into two parts. Then the protoplasm is separated between the two new nuclei, and the whole cell splits into two daughter-cells, the protoplasm gathering about each of the nuclei. The thin bridge of protoplasm which at first connects the daughter-cells soon breaks. Here we have the simple form of direct cleavage of the nuclei. Without mitosis, or formation of threads, the homogeneous nucleus divides into two halves. These move away from each other, and become centres of attraction for the enveloping matter, the protoplasm. The same direct cleavage of the nuclei is also witnessed in the reproduction of many other protists, while other unicellular organisms show the indirect division of the cell.
Hence, although the amoeba is nothing but a simple cell, it is evidently able to accomplish all the functions of the multicellular organism. It moves, feels, nourishes itself, and reproduces. Some kinds of these amoebae can be seen with the naked eye, but most of them are microscopically small. It is for the following reasons that we regard the amoebae as the unicellular organisms which have special phylogenetic (or evolutionary) relations to the ovum. In many of the lower animals the ovum retains its original naked form until fertilisation, develops no membranes, and is then often indistinguishable from the ordinary amoeba. Like the amoebae, these naked ova may thrust out processes, and move about as travelling cells. In the sponges these mobile ova move about freely in the maternal body like independent amoebae (Figure 1.17). They had been observed by earlier scientists, but described as foreign bodies—namely, parasitic amoebae, living parasitically on the body of the sponge. Later, however, it was discovered that they were not parasites, but the ova of the sponge. We also find this remarkable phenomenon among other animals, such as the graceful, bell-shaped zoophytes, which we call polyps and medusae. Their ova remain naked cells, which thrust out amoeboid projections, nourish themselves, and move about. When they have been fertilised, the multicellular organism is formed from them by repeated segmentation.
It is, therefore, no audacious hypothesis, but a perfectly sound conclusion, to regard the amoeba as the particular unicellular organism which offers us an approximate illustration of the ancient common unicellular ancestor of all the metazoa, or multicellular animals. The simple naked amoeba has a less definite and more original character than any other cell. Moreover, there is the fact that recent research has discovered such amoeba-like cells everywhere in the mature body of the multicellular animals. They are found, for instance, in the human blood, side by side with the red corpuscles, as colourless blood-cells; and it is the same with all the vertebrates. They are also found in many of the invertebrates—for instance, in the blood of the snail. I showed, in 1859, that these colourless blood-cells can, like the independent amoebae, take up solid particles, or "eat" (whence they are called phagocytes = "eating-cells," Figure 1.19). Lately, it has been discovered that many different cells may, if they have room enough, execute the same movements, creeping about and eating. They behave just like amoebae (Figure 1.12). It has also been shown that these "travelling-cells," or planocytes, play an important part in man's physiology and pathology (as means of transport for food, infectious matter, bacteria, etc.).
The power of the naked cell to execute these characteristic amoeba-like movements comes from the contractility (or automatic mobility) of its protoplasm. This seems to be a universal property of young cells. When they are not enclosed by a firm membrane, or confined in a "cellular prison," they can always accomplish these amoeboid movements. This is true of the naked ova as well as of any other naked cells, of the "travelling-cells," of various kinds in connective tissue, lymph-cells, mucus-cells, etc.
We have now, by our study of the ovum and the comparison of it with the amoeba, provided a perfectly sound and most valuable foundation for both the embryology and the evolution of man. We have learned that the human ovum is a simple cell, that this ovum is not materially different from that of other mammals, and that we may infer from it the existence of a primitive unicellular ancestral form, with a substantial resemblance to the amoeba.
The statement that the earliest progenitors of the human race were simple cells of this kind, and led an independent unicellular life like the amoeba, has not only been ridiculed as the dream of a natural philosopher, but also been violently censured in theological journals as "shameful and immoral." But, as I observed in my essay On the Origin and Ancestral Tree of the Human Race in 1870, this offended piety must equally protest against the "shameful and immoral" fact that each human individual is developed from a simple ovum, and that this human ovum is indistinguishable from those of the other mammals, and in its earliest stage is like a naked amoeba. We can show this to be a fact any day with the microscope, and it is little use to close one's eyes to "immoral" facts of this kind. It is as indisputable as the momentous conclusions we draw from it and as the vertebrate character of man (see Chapter 1.11).
(FIGURE 1.19. Blood-cells that eat, or phagocytes, from a naked sea-snail (Thetis), greatly magnified. I was the first to observe in the blood-cells of this snail the important fact that "the blood-cells of the invertebrates are unprotected pieces of plasm, and take in food, by means of their peculiar movements, like the amoebae." I had (in Naples, on May 10th, 1859) injected into the blood-vessels of one of these snails an infusion of water and ground indigo, and was greatly astonished to find the blood-cells themselves more or less filled with the particles of indigo after a few hours. After repeated injections I succeeded in "observing the very entrance of the coloured particles in the blood-cells, which took place just in the same way as with the amoeba." I have given further particulars about this in my Monograph on the Radiolaria.)
We now see very clearly how extremely important the cell theory has been for our whole conception of organic nature. "Man's place in nature" is settled beyond question by it. Apart from the cell theory, man is an insoluble enigma to us. Hence philosophers, and especially physiologists, should be thoroughly conversant with it. The soul of man can only be really understood in the light of the cell-soul, and we have the simplest form of this in the amoeba. Only those who are acquainted with the simple psychic functions of the unicellular organisms and their gradual evolution in the series of lower animals can understand how the elaborate mind of the higher vertebrates, and especially of man, was gradually evolved from them. The academic psychologists who lack this zoological equipment are unable to do so.
This naturalistic and realistic conception is a stumbling-block to our modern idealistic metaphysicians and their theological colleagues. Fenced about with their transcendental and dualistic prejudices, they attack not only the monistic system we establish on our scientific knowledge, but even the plainest facts which go to form its foundation. An instructive instance of this was seen a few years ago, in the academic discourse delivered by a distinguished theologian, Willibald Beyschlag, at Halle, January 12th, 1900, on the occasion of the centenary festival. The theologian protested violently against the "materialistic dustmen of the scientific world who offer our people the diploma of a descent from the ape, and would prove to them that the genius of a Shakespeare or a Goethe is merely a distillation from a drop of primitive mucus." Another well-known theologian protested against "the horrible idea that the greatest of men, Luther and Christ, were descended from a mere globule of protoplasm." Nevertheless, not a single informed and impartial scientist doubts the fact that these greatest men were, like all other men—and all other vertebrates—developed from an impregnated ovum, and that this simple nucleated globule of protoplasm has the same chemical constitution in all the mammals.
CHAPTER 1.7. CONCEPTION.
The recognition of the fact that every man begins his individual existence as a simple cell is the solid foundation of all research into the genesis of man. From this fact we are forced, in virtue of our biogenetic law, to draw the weighty phylogenetic conclusion that the earliest ancestors of the human race were also unicellular organisms; and among these protozoa we may single out the vague form of the amoeba as particularly important (cf. Chapter 1.6). That these unicellular ancestral forms did once exist follows directly from the phenomena which we perceive every day in the fertilised ovum. The development of the multicellular organism from the ovum, and the formation of the germinal layers and the tissues, follow the same laws in man and all the higher animals. It will, therefore, be our next task to consider more closely the impregnated ovum and the process of conception which produces it.
The process of impregnation or sexual conception is one of those phenomena that people love to conceal behind the mystic veil of supernatural power. We shall soon see, however, that it is a purely mechanical process, and can be reduced to familiar physiological functions. Moreover, this process of conception is of the same type, and is effected by the same organs, in man as in all the other mammals. The pairing of the male and female has in both cases for its main purpose the introduction of the ripe matter of the male seed or sperm into the female body, in the sexual canals of which it encounters the ovum. Conception then ensues by the blending of the two.
We must observe, first, that this important process is by no means so widely distributed in the animal and plant world as is commonly supposed. There is a very large number of lower organisms which propagate unsexually, or by monogamy; these are especially the sexless monera (chromacea, bacteria, etc.) but also many other protists, such as the amoebae, foraminifera, radiolaria, myxomycetae, etc. In these the multiplication of individuals takes place by unsexual reproduction, which takes the form of cleavage, budding, or spore-formation. The copulation of two coalescing cells, which in these cases often precedes the reproduction, cannot be regarded as a sexual act unless the two copulating plastids differ in size or structure. On the other hand, sexual reproduction is the general rule with all the higher organisms, both animal and plant; very rarely do we find asexual reproduction among them. There are, in particular, no cases of parthenogenesis (virginal conception) among the vertebrates.
Sexual reproduction offers an infinite variety of interesting forms in the different classes of animals and plants, especially as regards the mode of conception, and the conveyance of the spermatozoon to the ovum. These features are of great importance not only as regards conception itself, but for the development of the organic form, and especially for the differentiation of the sexes. There is a particularly curious correlation of plants and animals in this respect. The splendid studies of Charles Darwin and Hermann Muller on the fertilisation of flowers by insects have given us very interesting particulars of this.* (* See Darwin's work, On the Various Contrivances by which Orchids are Fertilised (1862).) This reciprocal service has given rise to a most intricate sexual apparatus. Equally elaborate structures have been developed in man and the higher animals, serving partly for the isolation of the sexual products on each side, partly for bringing them together in conception. But, however interesting these phenomena are in themselves, we cannot go into them here, as they have only a minor importance—if any at all—in the real process of conception. We must, however, try to get a very clear idea of this process and the meaning of sexual reproduction.
In every act of conception we have, as I said, to consider two different kinds of cells—a female and a male cell. The female cell of the animal organism is always called the ovum (or ovulum, egg, or egg-cell); the male cells are known as the sperm or seed-cells, or the spermatozoa (also spermium and zoospermium). The ripe ovum is, on the whole, one of the largest cells we know. It attains colossal dimensions when it absorbs great quantities of nutritive yelk, as is the case with birds and reptiles and many of the fishes. In the great majority of the animals the ripe ovum is rich in yelk and much larger than the other cells. On the other hand, the next cell which we have to consider in the process of conception, the male sperm-cell or spermatozoon, is one of the smallest cells in the animal body. Conception usually consists in the bringing into contact with the ovum of a slimy fluid secreted by the male, and this may take place either inside or out of the female body. This fluid is called sperm, or the male seed. Sperm, like saliva or blood, is not a simple fluid, but a thick agglomeration of innumerable cells, swimming about in a comparatively small quantity of fluid. It is not the fluid, but the independent male cells that swim in it, that cause conception.
(FIGURE 1.20. Spermia or spermatozoa of various mammals. The pear-shaped flattened nucleus is seen from the front in I and sideways in II. k is the nucleus, m its middle part (protoplasm), s the mobile, serpent-like tail (or whip); M four human spermatozoa, A spermatozoa from the ape; K from the rabbit; H from the mouse; C from the dog; S from the pig.
FIGURE 1.21. Spermatozoa or spermidia of various animals. (From Lang). a of a fish, b of a turbellaria worm (with two side-lashes), c to e of a nematode worm (amoeboid spermatozoa), f from a craw fish (star-shaped), g from the salamander (with undulating membrane), h of an annelid (a and h are the usual shape).
FIGURE 1.22. A single human spermatozoon magnified 2000 times; a shows it from the broader and b from the narrower side. k head (with nucleus), m middle-stem, h long-stem, and e tail. (From Retzius.))
The spermatozoa of the great majority of animals have two characteristic features. Firstly, they are extraordinarily small, being usually the smallest cells in the body; and, secondly, they have, as a rule, a peculiarly lively motion, which is known as spermatozoic motion. The shape of the cell has a good deal to do with this motion. In most of the animals, and also in many of the lower plants (but not the higher) each of these spermatozoa has a very small, naked cell-body, enclosing an elongated nucleus, and a long thread hanging from it (Figure 1.20). It was long before we could recognise that these structures are simple cells. They were formerly held to be special organisms, and were called "seed animals" (spermato-zoa, or spermato-zoidia); they are now scientifically known as spermia or spermidia, or as spermatosomata (seed-bodies) or spermatofila (seed threads). It took a good deal of comparative research to convince us that each of these spermatozoa is really a simple cell. They have the same shape as in many other vertebrates and most of the invertebrates. However, in many of the lower animals they have quite a different shape. Thus, for instance, in the craw fish they are large round cells, without any movement, equipped with stiff outgrowths like bristles (Figure 1.21 f). They have also a peculiar form in some of the worms, such as the thread-worms (filaria); in this case they are sometimes amoeboid and like very small ova (Figure 1.21 c to e). But in most of the lower animals (such as the sponges and polyps) they have the same pine-cone shape as in man and the other animals (Figure 1.21 a, h).
When the Dutch naturalist Leeuwenhoek discovered these thread-like lively particles in 1677 in the male sperm, it was generally believed that they were special, independent, tiny animalcules, like the infusoria, and that the whole mature organism existed already, with all its parts, but very small and packed together, in each spermatozoon (see Chapter 1.2). We now know that the mobile spermatozoa are nothing but simple and real cells, of the kind that we call "ciliated" (equipped with lashes, or cilia). In the previous illustrations we have distinguished in the spermatozoon a head, trunk, and tail. The "head" (Figure 1.20 k) is merely the oval nucleus of the cell; the body or middle-part (m) is an accumulation of cell-matter; and the tail (s) is a thread-like prolongation of the same.
Moreover, we now know that these spermatozoa are not at all a peculiar form of cell; precisely similar cells are found in various other parts of the body. If they have many short threads projecting, they are called ciliated; if only one long, whip-shaped process (or, more rarely, two or four), caudate (tailed) cells.
Very careful recent examination of the spermia, under a very high microscopic power (Figure 1.22 a, b), has detected some further details in the finer structure of the ciliated cell, and these are common to man and the anthropoid ape. The head (k) encloses the elliptic nucleus in a thin envelope of cytoplasm; it is a little flattened on one side, and thus looks rather pear-shaped from the front (b). In the central piece (m) we can distinguish a short neck and a longer connective piece (with central body). The tail consists of a long main section (h) and a short, very fine tail (e).
The process of fertilisation by sexual conception consists, therefore, essentially in the coalescence and fusing together of two different cells. The lively spermatozoon travels towards the ovum by its serpentine movements, and bores its way into the female cell (Figure 1.23). The nuclei of both sexual cells, attracted by a certain "affinity," approach each other and melt into one.
The fertilised cell is quite another thing from the unfertilised cell. For if we must regard the spermia as real cells no less than the ova, and the process of conception as a coalescence of the two, we must consider the resultant cell as a quite new and independent organism. It bears in the cell and nuclear matter of the penetrating spermatozoon a part of the father's body, and in the protoplasm and caryoplasm of the ovum a part of the mother's body. This is clear from the fact that the child inherits many features from both parents. It inherits from the father by means of the spermatozoon, and from the mother by means of the ovum. The actual blending of the two cells produces a third cell, which is the germ of the child, or the new organism conceived. One may also say of this sexual coalescence that the STEM-CELL IS A SIMPLE HERMAPHRODITE; it unites both sexual substances in itself.
(FIGURE 1.23. The fertilisation of the ovum by the spermatozoon (of a mammal). One of the many thread-like, lively spermidia pierces through a fine pore-canal into the nuclear yelk. The nucleus of the ovum is invisible.
FIGURE 1.24. An impregnated echinoderm ovum, with small homogeneous nucleus (e k). (From Hertwig.))
I think it necessary to emphasise the fundamental importance of this simple, but often unappreciated, feature in order to have a correct and clear idea of conception. With that end, I have given a special name to the new cell from which the child develops, and which is generally loosely called "the fertilised ovum," or "the first segmentation sphere." I call it "the stem-cell" (cytula). The name "stem-cell" seems to me the simplest and most suitable, because all the other cells of the body are derived from it, and because it is, in the strictest sense, the stem-father and stem-mother of all the countless generations of cells of which the multicellular organism is to be composed. That complicated molecular movement of the protoplasm which we call "life" is, naturally, something quite different in this stem-cell from what we find in the two parent-cells, from the coalescence of which it has issued. THE LIFE OF THE STEM-CELL OR CYTULA IS THE PRODUCT OR RESULTANT OF THE PATERNAL LIFE-MOVEMENT THAT IS CONVEYED IN THE SPERMATOZOON AND THE MATERNAL LIFE-MOVEMENT THAT IS CONTRIBUTED BY THE OVUM.
The admirable work done by recent observers has shown that the individual development, in man and the other animals, commences with the formation of a simple "stem-cell" of this character, and that this then passes, by repeated segmentation (or cleavage), into a cluster of cells, known as "the segmentation sphere" or "segmentation cells." The process is most clearly observed in the ova of the echinoderms (star-fishes, sea-urchins, etc.). The investigations of Oscar and Richard Hertwig were chiefly directed to these. The main results may be summed up as follows:—
Conception is preceded by certain preliminary changes, which are very necessary—in fact, usually indispensable—for its occurrence. They are comprised under the general heading of "Changes prior to impregnation." In these the original nucleus of the ovum, the germinal vesicle, is lost. Part of it is extruded, and part dissolved in the cell contents; only a very small part of it is left to form the basis of a fresh nucleus, the pronucleus femininus. It is the latter alone that combines in conception with the invading nucleus of the fertilising spermatozoon (the pronucleus masculinus).
The impregnation of the ovum commences with a decay of the germinal vesicle, or the original nucleus of the ovum (Figure 1.8). We have seen that this is in most unripe ova a large, transparent, round vesicle. This germinal vesicle contains a viscous fluid (the caryolymph). The firm nuclear frame (caryobasis) is formed of the enveloping membrane and a mesh-work of nuclear threads running across the interior, which is filled with the nuclear sap. In a knot of the network is contained the dark, stiff, opaque nuclear corpuscle or nucleolus. When the impregnation of the ovum sets in, the greater part of the germinal vesicle is dissolved in the cell; the nuclear membrane and mesh-work disappear; the nuclear sap is distributed in the protoplasm; a small portion of the nuclear base is extruded; another small portion is left, and is converted into the secondary nucleus, or the female pro-nucleus (Figure 1.24 e k).
The small portion of the nuclear base which is extruded from the impregnated ovum is known as the "directive bodies" or "polar cells"; there are many disputes as to their origin and significance, but we are as yet imperfectly acquainted with them. As a rule, they are two small round granules, of the same size and appearance as the remaining pro-nucleus. They are detached cell-buds; their separation from the large mother-cell takes place in the same way as in ordinary "indirect cell-division." Hence, the polar cells are probably to be conceived as "abortive ova," or "rudimentary ova," which proceed from a simple original ovum by cleavage in the same way that several sperm-cells arise from one "sperm-mother-cell," in reproduction from sperm. The male sperm-cells in the testicles must undergo similar changes in view of the coming impregnation as the ova in the female ovary. In this maturing of the sperm each of the original seed-cells divides by double segmentation into four daughter-cells, each furnished with a fourth of the original nuclear matter (the hereditary chromatin); and each of these four descendant cells becomes a spermatozoon, ready for impregnation. Thus is prevented the doubling of the chromatin in the coalescence of the two nuclei at conception. As the two polar cells are extruded and lost, and have no further part in the fertilisation of the ovum, we need not discuss them any further. But we must give more attention to the female pro-nucleus which alone remains after the extrusion of the polar cells and the dissolving of the germinal vesicle (Figure 1.23 e k). This tiny round corpuscle of chromatin now acts as a centre of attraction for the invading spermatozoon in the large ripe ovum, and coalesces with its "head," the male pro-nucleus. The product of this blending, which is the most important part of the act of impregnation, is the stem-nucleus, or the first segmentation nucleus (archicaryon)—that is to say, the nucleus of the new-born embryonic stem-cell or "first segmentation cell." This stem-cell is the starting point of the subsequent embryonic processes.
Hertwig has shown that the tiny transparent ova of the echinoderms are the most convenient for following the details of this important process of impregnation. We can, in this case, easily and successfully accomplish artificial impregnation, and follow the formation of the stem-cell step by step within the space of ten minutes. If we put ripe ova of the star-fish or sea-urchin in a watch glass with sea-water and add a drop of ripe sperm-fluid, we find each ovum impregnated within five minutes. Thousands of the fine, mobile ciliated cells, which we have described as "sperm-threads" (Figure 1.20), make their way to the ova, owing to a sort of chemical sensitive action which may be called "smell." But only one of these innumerable spermatozoa is chosen—namely, the one that first reaches the ovum by the serpentine motions of its tail, and touches the ovum with its head. At the spot where the point of its head touches the surface of the ovum the protoplasm of the latter is raised in the form of a small wart, the "impregnation rise" (Figure 1.25 A). The spermatozoon then bores its way into this with its head, the tail outside wriggling about all the time (Figure 1.25 B, C). Presently the tail also disappears within the ovum. At the same time the ovum secretes a thin external yelk-membrane (Figure 1.25 C), starting from the point of impregnation; and this prevents any more spermatozoa from entering.
Inside the impregnated ovum we now see a rapid series of most important changes. The pear-shaped head of the sperm-cell, or the "head of the spermatozoon," grows larger and rounder, and is converted into the male pro-nucleus (Figure 1.26 s k). This has an attractive influence on the fine granules or particles which are distributed in the protoplasm of the ovum; they arrange themselves in lines in the figure of a star. But the attraction or the "affinity" between the two nuclei is even stronger. They move towards each other inside the yelk with increasing speed, the male (Figure 1.27 s k) going more quickly than the female nucleus (e k). The tiny male nucleus takes with it the radiating mantle which spreads like a star about it. At last the two sexual nuclei touch (usually in the centre of the globular ovum), lie close together, are flattened at the points of contact, and coalesce into a common mass. The small central particle of nuclein which is formed from this combination of the nuclei is the stem-nucleus, or the first segmentation nucleus; the new-formed cell, the product of the impregnation, is our stem-cell, or "first segmentation sphere" (Figure 1.2).
(FIGURE 1.25. Impregnation of the ovum of a star-fish. (From Hertwig.) Only a small part of the surface of the ovum is shown. One of the numerous spermatozoa approaches the "impregnation rise" (A), touches it (B), and then penetrates into the protoplasm of the ovum (C).
FIGURES 1.26 AND 1.27. Impregnation of the ovum of the sea-urchin. (From Hertwig.) In Figure 1.26 the little sperm-nucleus (sk) moves towards the larger nucleus of the ovum (ek). In Figure 1.27 they nearly touch, and are surrounded by the radiating mantle of protoplasm.)
Hence the one essential point in the process of sexual reproduction or impregnation is the formation of a new cell, the stem-cell, by the combination of two originally different cells, the female ovum and the male spermatozoon. This process is of the highest importance, and merits our closest attention; all that happens in the later development of this first cell and in the life of the organism that comes of it is determined from the first by the chemical and morphological composition of the stem-cell, its nucleus and its body. We must, therefore, make a very careful study of the rise and structure of the stem-cell.
The first question that arises is as to the two different active elements, the nucleus and the protoplasm, in the actual coalescence. It is obvious that the nucleus plays the more important part in this. Hence Hertwig puts his theory of conception in the principle: "Conception consists in the copulation of two cell-nuclei, which come from a male and a female cell." And as the phenomenon of heredity is inseparably connected with the reproductive process, we may further conclude that these two copulating nuclei "convey the characteristics which are transmitted from parents to offspring." In this sense I had in 1866 (in the ninth chapter of the General Morphology) ascribed to the reproductive nucleus the function of generation and heredity, and to the nutritive protoplasm the duties of nutrition and adaptation. As, moreover, there is a complete coalescence of the mutually attracted nuclear substances in conception, and the new nucleus formed (the stem-nucleus) is the real starting-point for the development of the fresh organism, the further conclusion may be drawn that the male nucleus conveys to the child the qualities of the father, and the female nucleus the features of the mother. We must not forget, however, that the protoplasmic bodies of the copulating cells also fuse together in the act of impregnation; the cell-body of the invading spermatozoon (the trunk and tail of the male ciliated cell) is dissolved in the yelk of the female ovum. This coalescence is not so important as that of the nuclei, but it must not be overlooked; and, though this process is not so well known to us, we see clearly at least the formation of the star-like figure (the radial arrangement of the particles in the plasma) in it (Figures 1.26 to 1.27).
The older theories of impregnation generally went astray in regarding the large ovum as the sole base of the new organism, and only ascribed to the spermatozoon the work of stimulating and originating its development. The stimulus which it gave to the ovum was sometimes thought to be purely chemical, at other times rather physical (on the principle of transferred movement), or again a mystic and transcendental process. This error was partly due to the imperfect knowledge at that time of the facts of impregnation, and partly to the striking difference in the sizes of the two sexual cells. Most of the earlier observers thought that the spermatozoon did not penetrate into the ovum. And even when this had been demonstrated, the spermatozoon was believed to disappear in the ovum without leaving a trace. However, the splendid research made in the last three decades with the finer technical methods of our time has completely exposed the error of this. It has been shown that the tiny sperm-cell is NOT SUBORDINATED TO, BUT COORDINATED WITH, the large ovum. The nuclei of the two cells, as the vehicles of the hereditary features of the parents, are of equal physiological importance. In some cases we have succeeded in proving that the mass of the active nuclear substance which combines in the copulation of the two sexual nuclei is originally the same for both.
These morphological facts are in perfect harmony with the familiar physiological truth that the child inherits from both parents, and that on the average they are equally distributed. I say "on the average," because it is well known that a child may have a greater likeness to the father or to the mother; that goes without saying, as far as the primary sexual characters (the sexual glands) are concerned. But it is also possible that the determination of the latter—the weighty determination whether the child is to be a boy or a girl—depends on a slight qualitative or quantitative difference in the nuclein or the coloured nuclear matter which comes from both parents in the act of conception.
The striking differences of the respective sexual cells in size and shape, which occasioned the erroneous views of earlier scientists, are easily explained on the principle of division of labour. The inert, motionless ovum grows in size according to the quantity of provision it stores up in the form of nutritive yelk for the development of the germ. The active swimming sperm-cell is reduced in size in proportion to its need to seek the ovum and bore its way into its yelk. These differences are very conspicuous in the higher animals, but they are much less in the lower animals. In those protists (unicellular plants and animals) which have the first rudiments of sexual reproduction the two copulating cells are at first quite equal. In these cases the act of impregnation is nothing more than a sudden GROWTH, in which the originally simple cell doubles its volume, and is thus prepared for reproduction (cell-division). Afterwards slight differences are seen in the size of the copulating cells; though the smaller ones still have the same shape as the larger ones. It is only when the difference in size is very pronounced that a notable difference in shape is found: the sprightly sperm-cell changes more in shape and the ovum in size.
Quite in harmony with this new conception of the EQUIVALENCE OF THE TWO GONADS, or the equal physiological importance of the male and female sex-cells and their equal share in the process of heredity, is the important fact established by Hertwig (1875), that in normal impregnation only one single spermatozoon copulates with one ovum; the membrane which is raised on the surface of the yelk immediately after one sperm-cell has penetrated (Figure 1.25 C) prevents any others from entering. All the rivals of the fortunate penetrator are excluded, and die without. But if the ovum passes into a morbid state, if it is made stiff by a lowering of its temperature or stupefied with narcotics (chloroform, morphia, nicotine, etc.), two or more spermatozoa may penetrate into its yelk-body. We then witness polyspermism. The more Hertwig chloroformed the ovum, the more spermatozoa were able to bore their way into its unconscious body.
(FIGURE 1.28. Stem-cell of a rabbit, magnified 200 times. In the centre of the granular protoplasm of the fertilised ovum (d) is seen the little, bright stem-nucleus, z is the ovolemma, with a mucous membrane (h). s are dead spermatozoa.)
These remarkable facts of impregnation are also of the greatest interest in psychology, especially as regards the theory of the cell-soul, which I consider to be its chief foundation. The phenomena we have described can only be understood and explained by ascribing a certain lower degree of psychic activity to the sexual principles. They FEEL each other's proximity, and are drawn together by a SENSITIVE impulse (probably related to smell); they MOVE towards each other, and do not rest until they fuse together. Physiologists may say that it is only a question of a peculiar physico-chemical phenomenon, and not a psychic action; but the two cannot be separated. Even the psychic functions, in the strict sense of the word, are only complex physical processes, or "psycho-physical" phenomena, which are determined in all cases exclusively by the chemical composition of their material substratum.
The monistic view of the matter becomes clear enough when we remember the radical importance of impregnation as regards heredity. It is well known that not only the most delicate bodily structures, but also the subtlest traits of mind, are transmitted from the parents to the children. In this the chromatic matter of the male nucleus is just as important a vehicle as the large caryoplasmic substance of the female nucleus; the one transmits the mental features of the father, and the other those of the mother. The blending of the two parental nuclei determines the individual psychic character of the child.
But there is another important psychological question—the most important of all—that has been definitely answered by the recent discoveries in connection with conception. This is the question of the immortality of the soul. No fact throws more light on it and refutes it more convincingly than the elementary process of conception that we have described. For this copulation of the two sexual nuclei (Figures 1.26 and 1.27) indicates the precise moment at which the individual begins to exist. All the bodily and mental features of the new-born child are the sum-total of the hereditary qualities which it has received in reproduction from parents and ancestors. All that man acquires afterwards in life by the exercise of his organs, the influence of his environment, and education—in a word, by adaptation—cannot obliterate that general outline of his being which he inherited from his parents. But this hereditary disposition, the essence of every human soul, is not "eternal," but "temporal"; it comes into being only at the moment when the sperm-nucleus of the father and the nucleus of the maternal ovum meet and fuse together. It is clearly irrational to assume an "eternal life without end" for an individual phenomenon, the commencement of which we can indicate to a moment by direct visual observation.
The great importance of the process of impregnation in answering such questions is quite clear. It is true that conception has never been studied microscopically in all its details in the human case—notwithstanding its occurrence at every moment—for reasons that are obvious enough. However, the two cells which need consideration, the female ovum and the male spermatozoon, proceed in the case of man in just the same way as in all the other mammals; the human foetus or embryo which results from copulation has the same form as with the other animals. Hence, no scientist who is acquainted with the facts doubts that the processes of impregnation are just the same in man as in the other animals.
The stem-cell which is produced, and with which every man begins his career, cannot be distinguished in appearance from those of other mammals, such as the rabbit (Figure 1.28). In the case of man, also, this stem-cell differs materially from the original ovum, both in regard to form (morphologically), in regard to material composition (chemically), and in regard to vital properties (physiologically). It comes partly from the father and partly from the mother. Hence it is not surprising that the child who is developed from it inherits from both parents. The vital movements of each of these cells form a sum of mechanical processes which in the last analysis are due to movements of the smallest vital parts, or the molecules, of the living substance. If we agree to call this active substance plasson, and its molecules plastidules, we may say that the individual physiological character of each of these cells is due to its molecular plastidule-movement. HENCE, THE PLASTIDULE-MOVEMENT OF THE CYTULA IS THE RESULTANT OF THE COMBINED PLASTIDULE-MOVEMENTS OF THE FEMALE OVUM AND THE MALE SPERM-CELL.* (* The plasson of the stem-cell or cytula may, from the anatomical point of view, be regarded as homogeneous and structureless, like that of the monera. This is not inconsistent with our hypothetical ascription to the plastidules (or molecules of the plasson) of a complex molecular structure. The complexity of this is the greater in proportion to the complexity of the organism that is developed from it and the length of the chain of its ancestry, or to the multitude of antecedent processes of heredity and adaptation.)
CHAPTER 1.8. THE GASTRAEA THEORY.
There is a substantial agreement throughout the animal world in the first changes which follow the impregnation of the ovum and the formation of the stem-cell; they begin in all cases with the segmentation of the ovum and the formation of the germinal layers. The only exception is found in the protozoa, the very lowest and simplest forms of animal life; these remain unicellular throughout life. To this group belong the amoebae, gregarinae, rhizopods, infusoria, etc. As their whole organism consists of a single cell, they can never form germinal layers, or definite strata of cells. But all the other animals—all the tissue-forming animals, or metazoa, as we call them, in contradistinction to the protozoa—construct real germinal layers by the repeated cleavage of the impregnated ovum. This we find in the lower cnidaria and worms, as well as in the more highly-developed molluscs, echinoderms, articulates, and vertebrates.
In all these metazoa, or multicellular animals, the chief embryonic processes are substantially alike, although they often seem to a superficial observer to differ considerably. The stem-cell that proceeds from the impregnated ovum always passes by repeated cleavage into a number of simple cells. These cells are all direct descendants of the stem-cell, and are, for reasons we shall see presently, called segmentation-cells. The repeated cleavage of the stem-cell, which gives rise to these segmentation-spheres, has long been known as "segmentation." Sooner or later the segmentation-cells join together to form a round (at first, globular) embryonic sphere (blastula); they then form into two very different groups, and arrange themselves in two separate strata—the two primary germinal layers. These enclose a digestive cavity, the primitive gut, with an opening, the primitive mouth. We give the name of the gastrula to the important embryonic form that has these primitive organs, and the name of gastrulation to the formation of it. This ontogenetic process has a very great significance, and is the real starting-point of the construction of the multicellular animal body.
The fundamental embryonic processes of the cleavage of the ovum and the formation of the germinal layers have been very thoroughly studied in the last thirty years, and their real significance has been appreciated. They present a striking variety in the different groups, and it was no light task to prove their essential identity in the whole animal world. But since I formulated the gastraea theory in 1872, and afterwards (1875) reduced all the various forms of segmentation and gastrulation to one fundamental type, their identity may be said to have been established. We have thus mastered the law of unity which governs the first embryonic processes in all the animals.
Man is like all the other higher animals, especially the apes, in regard to these earliest and most important processes. As the human embryo does not essentially differ, even at a much later stage of development—when we already perceive the cerebral vesicles, the eyes, ears, gill-arches, etc.—from the similar forms of the other higher mammals, we may confidently assume that they agree in the earliest embryonic processes, segmentation and the formation of germinal layers. This has not yet, it is true, been established by observation. We have never yet had occasion to dissect a woman immediately after impregnation and examine the stem-cell or the segmentation-cells in her oviduct. However, as the earliest human embryos we have examined, and the later and more developed forms, agree with those of the rabbit, dog, and other higher mammals, no reasonable man will doubt but that the segmentation and formation of layers are the same in both cases.
But the special form of segmentation and layer formation which we find in the mammal is by no means the original, simple, palingenetic form. It has been much modified and cenogenetically altered by a very complex adaptation to embryonic conditions. We cannot, therefore, understand it altogether in itself. In order to do this, we have to make a COMPARATIVE study of segmentation and layer-formation in the animal world; and we have especially to seek the original, PALINGENETIC form from which the modified CENOGENETIC (see Chapter 1.1) form has gradually been developed.
This original unaltered form of segmentation and layer-formation is found to-day in only one case in the vertebrate-stem to which man belongs—the lowest and oldest member of the stem, the wonderful lancelet or amphioxus (cf. Chapters 2.16 and 2.17). But we find a precisely similar palingenetic form of embryonic development in the case of many of the invertebrate animals, as, for instance, the remarkable ascidia, the pond-snail (Limnaeus), and arrow-worm (Sagitta), and many of the echinoderms and cnidaria, such as the common star-fish and sea-urchin, many of the medusae and corals, and the simpler sponges (Olynthus). We may take as an illustration the palingenetic segmentation and germinal layer-formation in an eight-fold insular coral, which I discovered in the Red Sea, and described as Monoxenia Darwinii.
(FIGURE 1.29. Gastrulation of a coral (Monoxenia Darwinii). A, B, stem-cell (cytula) or impregnated ovum. In Figure A (immediately after impregnation) the nucleus is invisible. In Figure B (a little later) it is quite clear. C two segmentation-cells. D four segmentation-cells. E mulberry-formation (morula). F blastosphere (blastula). G blastula (transverse section). H depula, or hollowed blastula (transverse section). I gastrula (longitudinal section). K gastrula, or cup-sphere, external appearance.)
The impregnated ovum of this coral (Figure 1.29 A, B) first splits into two equal cells (C). First, the nucleus of the stem-cell and its central body divide into two halves. These recede from and repel each other, and act as centres of attraction on the surrounding protoplasm; in consequence of this, the protoplasm is constricted by a circular furrow, and, in turn, divides into two halves. Each of the two segmentation-cells thus produced splits in the same way into two equal cells. The four segmentation-cells (grand-daughters of the stem-cell) lie in one plane. Now, however, each of them subdivides into two equal halves, the cleavage of the nucleus again preceding that of the surrounding protoplasm. The eight cells which thus arise break into sixteen, these into thirty-two, and then (each being constantly halved) into sixty-four, 128, and so on.* (* The number of segmentation-cells thus produced increases geometrically in the original gastrulation, or the purest palingenetic form of cleavage. However, in different animals the number reaches a different height, so that the morula, and also the blastula, may consist sometimes of thirty-two, sometimes of sixty-four, and sometimes of 128, or more, cells.) The final result of this repeated cleavage is the formation of a globular cluster of similar segmentation-cells, which we call the mulberry-formation or morula. The cells are thickly pressed together like the parts of a mulberry or blackberry, and this gives a lumpy appearance to the surface of the sphere (Figure E).* (* The segmentation-cells which make up the morula after the close of the palingenetic cleavage seem usually to be quite similar, and to present no differences as to size, form, and composition. That, however, does not prevent them from differentiating into animal and vegetative cells, even during the cleavage.)
When the cleavage is thus ended, the mulberry-like mass changes into a hollow globular sphere. Watery fluid or jelly gathers inside the globule; the segmentation-cells are loosened, and all rise to the surface. There they are flattened by mutual pressure, and assume the shape of truncated pyramids, and arrange themselves side by side in one regular layer (Figures F, G). This layer of cells is called the germinal membrane (or blastoderm); the homogeneous cells which compose its simple structure are called blastodermic cells; and the whole hollow sphere, the walls of which are made of the preceding, is called the blastula or blastosphere.* (* The blastula of the lower animals must not be confused with the very different blastula of the mammal, which is properly called the gastrocystis or blastocystis. This cenogenetic gastrocystis and the palingenetic blastula are sometimes very wrongly comprised under the common name of blastula or vesicula blastodermica.)
In the case of our coral, and of many other lower forms of animal life, the young embryo begins at once to move independently and swim about in the water. A fine, long, thread-like process, a sort of whip or lash, grows out of each blastodermic cell, and this independently executes vibratory movements, slow at first, but quicker after a time (Figure F). In this way each blastodermic cell becomes a ciliated cell. The combined force of all these vibrating lashes causes the whole blastula to move about in a rotatory fashion. In many other animals, especially those in which the embryo develops within enclosed membranes, the ciliated cells are only formed at a later stage, or even not formed at all. The blastosphere may grow and expand by the blastodermic cells (at the surface of the sphere) dividing and increasing, and more fluid is secreted in the internal cavity. There are still to-day some organisms that remain throughout life at the structural stage of the blastula—hollow vesicles that swim about by a ciliary movement in the water, the wall of which is composed of a single layer of cells, such as the volvox, the magosphaera, synura, etc. We shall speak further of the great phylogenetic significance of this fact in Chapter 2.19.
A very important and remarkable process now follows—namely, the curving or invagination of the blastula (Figure H). The vesicle with a single layer of cells for wall is converted into a cup with a wall of two layers of cells (cf. Figures G, H, I). A certain spot at the surface of the sphere is flattened, and then bent inward. This depression sinks deeper and deeper, growing at the cost of the internal cavity. The latter decreases as the hollow deepens. At last the internal cavity disappears altogether, the inner side of the blastoderm (that which lines the depression) coming to lie close on the outer side. At the same time, the cells of the two sections assume different sizes and shapes; the inner cells are more round and the outer more oval (Figure I). In this way the embryo takes the form of a cup or jar-shaped body, with a wall made up of two layers of cells, the inner cavity of which opens to the outside at one end (the spot where the depression was originally formed). We call this very important and interesting embryonic form the "cup-embryo" or "cup-larva" (gastrula, Figure 1.29, I longitudinal section, K external view). I have in my Natural History of Creation given the name of depula to the remarkable intermediate form which appears at the passage of the blastula into the gastrula. In this intermediate stage there are two cavities in the embryo—the original cavity (blastocoel) which is disappearing, and the primitive gut-cavity (progaster) which is forming.
I regard the gastrula as the most important and significant embryonic form in the animal world. In all real animals (that is, excluding the unicellular protists) the segmentation of the ovum produces either a pure, primitive, palingenetic gastrula (Figure 1.29 I, K) or an equally instructive cenogenetic form, which has been developed in time from the first, and can be directly reduced to it. It is certainly a fact of the greatest interest and instructiveness that animals of the most different stems—vertebrates and tunicates, molluscs and articulates, echinoderms and annelids, cnidaria and sponges—proceed from one and the same embryonic form. In illustration I give a few pure gastrula forms from various groups of animals (Figures 1.30 to 1.35, explanation given below each).
(FIGURES 1.30 TO 1.35. In each figure d is the primitive-gut cavity, o primitive mouth, s segmentation-cavity, i entoderm (gut-layer), e ectoderm (skin layer).
FIGURE 1.30. (A) Gastrula of a very simple primitive-gut animal or gastraead (gastrophysema). (Haeckel.)
FIGURE 1.31. (B) Gastrula of a worm (Sagitta). (From Kowalevsky.)
FIGURE 1.32. (C) Gastrula of an echinoderm (star-fish, Uraster), not completely folded in (depula). (From Alexander Agassiz.)
FIGURE 1.33. (D) Gastrula of an arthropod (primitive crab, Nauplius) (as 32).
FIGURE 1.34. (E) Gastrula of a mollusc (pond-snail, Linnaeus). (From
Karl Rabl.)
FIGURE 1.35. (F) Gastrula of a vertebrate (lancelet, Amphioxus). (From
Kowalevsky.) (Front view.))
In view of this extraordinary significance of the gastrula, we must make a very careful study of its original structure. As a rule, the typical gastrula is very small, being invisible to the naked eye, or at the most only visible as a fine point under very favourable conditions, and measuring generally 1/500 to 1/250 of an inch (less frequently 1/50 inch, or even more) in diameter. In shape it is usually like a roundish drinking-cup. Sometimes it is rather oval, at other times more ellipsoid or spindle-shaped; in some cases it is half round, or even almost round, and in others lengthened out, or almost cylindrical.
I give the name of primitive gut (progaster) and primitive mouth (prostoma) to the internal cavity of the gastrula-body and its opening; because this cavity is the first rudiment of the digestive cavity of the organism, and the opening originally served to take food into it. Naturally, the primitive gut and mouth change very considerably afterwards in the various classes of animals. In most of the cnidaria and many of the annelids (worm-like animals) they remain unchanged throughout life. But in most of the higher animals, and so in the vertebrates, only the larger central part of the later alimentary canal develops from the primitive gut; the later mouth is a fresh development, the primitive mouth disappearing or changing into the anus. We must therefore distinguish carefully between the primitive gut and mouth of the gastrula and the later alimentary canal and mouth of the fully developed vertebrate.* (* My distinction (1872) between the primitive gut and mouth and the later permanent stomach (metagaster) and mouth (metastoma) has been much criticised; but it is as much justified as the distinction between the primitive kidneys and the permanent kidneys. Professor E. Ray-Lankester suggested three years afterwards (1875) the name archenteron for the primitive gut, and blastoporus for the primitive mouth.)
(FIGURE 1.36. Gastrula of a lower sponge (olynthus). A external view, B longitudinal section through the axis, g primitive-gut cavity, a primitive mouth-aperture, i inner cell-layer (entoderm, endoblast, gut-layer), e external cell-layer (outer germinal layer, ectoderm, ectoblast, or skin-layer).
The two layers of cells which line the gut-cavity and compose its wall are of extreme importance. These two layers, which are the sole builders of the whole organism, are no other than the two primary germinal layers, or the primitive germ-layers. I have spoken in the introductory section (Chapter 1.3.) of their radical importance. The outer stratum is the skin-layer, or ectoderm (Figures 1.30 to 1.35 e); the inner stratum is the gut-layer, or entoderm (i). The former is often also called the ectoblast, or epiblast, and the latter the endoblast, or hypoblast. FROM THESE TWO PRIMARY GERMINAL LAYERS ALONE IS DEVELOPED THE ENTIRE ORGANISM OF ALL THE METAZOA OR MULTICELLULAR ANIMALS. The skin-layer forms the external skin, the gut-layer forms the internal skin or lining of the body. Between these two germinal layers are afterwards developed the middle germinal layer (mesoderma) and the body-cavity (coeloma) filled with blood or lymph.
The two primary germinal layers were first distinguished by Pander in 1817 in the incubated chick. Twenty years later (1849) Huxley pointed out that in many of the lower zoophytes, especially the medusae, the whole body consists throughout life of these two primary germinal layers. Soon afterwards (1853) Allman introduced the names which have come into general use; he called the outer layer the ectoderm ("outer-skin"), and the inner the entoderm ("inner-skin"). But in 1867 it was shown, particularly by Kowalevsky, from comparative observation, that even in invertebrates, also, of the most different classes—annelids, molluscs, echinoderms, and articulates—the body is developed out of the same two primary layers. Finally, I discovered them (1872) in the lowest tissue-forming animals, the sponges, and proved in my gastraea theory that these two layers must be regarded as identical throughout the animal world, from the sponges and corals to the insects and vertebrates, including man. This fundamental "homology [identity] of the primary germinal layers and the primitive gut" has been confirmed during the last thirty years by the careful research of many able observers, and is now pretty generally admitted for the whole of the metazoa.
As a rule, the cells which compose the two primary germinal layers show appreciable differences even in the gastrula stage. Generally (if not always) the cells of the skin-layer or ectoderm (Figures 1.36 c and 1.37 e) are the smaller, more numerous, and clearer; while the cells of the gut-layer, or entoderm (i), are larger, less numerous, and darker. The protoplasm of the ectodermic (outer) cells is clearer and firmer than the thicker and softer cell-matter of the entodermic (inner) cells; the latter are, as a rule, much richer in yelk-granules (albumen and fatty particles) than the former. Also the cells of the gut-layer have, as a rule, a stronger affinity for colouring matter, and take on a tinge in a solution of carmine, aniline, etc., more quickly and appreciably than the cells of the skin-layer. The nuclei of the entoderm-cells are usually roundish, while those of the ectoderm-cells are oval.
When the doubling-process is complete, very striking histological differences between the cells of the two layers are found (Figure 1.37). The tiny, light ectoderm-cells (e) are sharply distinguished from the larger and darker entoderm-cells (i). Frequently this differentiation of the cell-forms sets in at a very early stage, during the segmentation-process, and is already very appreciable in the blastula.
We have, up to the present, only considered that form of segmentation and gastrulation which, for many and weighty reasons, we may regard as the original, primordial, or palingenetic form. We might call it "equal" or homogeneous segmentation, because the divided cells retain a resemblance to each other at first (and often until the formation of the blastoderm). We give the name of the "bell-gastrula," or archigastrula, to the gastrula that succeeds it. In just the same form as in the coral we considered (Monoxenia, Figure 1.29), we find it in the lowest zoophyta (the gastrophysema, Figure 1.30), and the simplest sponges (olynthus, Figure 1.36); also in many of the medusae and hydrapolyps, lower types of worms of various classes (brachiopod, arrow-worm, Figure 1.31), tunicates (ascidia), many of the echinoderms (Figure 1.32), lower articulates (Figure 1.33), and molluscs (Figure 1.34), and, finally, in a slightly modified form, in the lowest vertebrate (the amphioxus, Figure 1.35).
(FIGURE 1.37. Cells from the two primary germinal layers of the mammal (from both layers of the blastoderm). i larger and darker cells of the inner stratum, the vegetal layer or entoderm. e smaller and clearer cells from the outer stratum, the animal layer or ectoderm.
FIGURE 1.38. Gastrulation of the amphioxus, from Hatschek (vertical section through the axis of the ovum). A, B, C three stages in the formation of the blastula; D, E curving of the blastula; F complete gastrula. h segmentation-cavity. g primitive gut-cavity.))
The gastrulation of the amphioxus is especially interesting because this lowest and oldest of all the vertebrates is of the highest significance in connection with the evolution of the vertebrate stem, and therefore with that of man (compare Chapters 2.16 and 2.17). Just as the comparative anatomist traces the most elaborate features in the structures of the various classes of vertebrates to divergent development from this simple primitive vertebrate, so comparative embryology traces the various secondary forms of vertebrate gastrulation to the simple, primary formation of the germinal layers in the amphioxus. Although this formation, as distinguished from the cenogenetic modifications of the vertebrate, may on the whole be regarded as palingenetic, it is nevertheless different in some features from the quite primitive gastrulation such as we have, for instance, in the Monoxenia (Figure 1.29) and the Sagitta. Hatschek rightly observes that the segmentation of the ovum in the amphioxus is not strictly equal, but almost equal, and approaches the unequal. The difference in size between the two groups of cells continues to be very noticeable in the further course of the segmentation; the smaller animal cells of the upper hemisphere divide more quickly than the larger vegetal cells of the lower (Figure 1.38 A, B). Hence the blastoderm, which forms the single-layer wall of the globular blastula at the end of the cleavage-process, does not consist of homogeneous cells of equal size, as in the Sagitta and the Monoxenia; the cells of the upper half of the blastoderm (the mother-cells of the ectoderm) are more numerous and smaller, and the cells of the lower half (the mother-cells of the entoderm) less numerous and larger. Moreover, the segmentation-cavity of the blastula (Figure 1.38 C, h) is not quite globular, but forms a flattened spheroid with unequal poles of its vertical axis. While the blastula is being folded into a cup at the vegetal pole of its axis, the difference in the size of the blastodermic cells increases (Figure 1.38 D, E); it is most conspicuous when the invagination is complete and the segmentation-cavity has disappeared (Figure 1.38 F). The larger vegetal cells of the entoderm are richer in granules, and so darker than the smaller and lighter animal cells of the ectoderm.
But the unequal gastrulation of the amphioxus diverges from the typical equal cleavage of the Sagitta, the Monoxenia (Figure 1.29), and the Olynthus (Figure 1.36), in another important particular. The pure archigastrula of the latter forms is uni-axial, and it is round in its whole length in transverse section. The vegetal pole of the vertical axis is just in the centre of the primitive mouth. This is not the case in the gastrula of the amphioxus. During the folding of the blastula the ideal axis is already bent on one side, the growth of the blastoderm (or the increase of its cells) being brisker on one side than on the other; the side that grows more quickly, and so is more curved (Figure 1.39 v), will be the anterior or belly-side, the opposite, flatter side will form the back (d). The primitive mouth, which at first, in the typical archigastrula, lay at the vegetal pole of the main axis, is forced away to the dorsal side; and whereas its two lips lay at first in a plane at right angles to the chief axis, they are now so far thrust aside that their plane cuts the axis at a sharp angle. The dorsal lip is therefore the upper and more forward, the ventral lip the lower and hinder. In the latter, at the ventral passage of the entoderm into the ectoderm, there lie side by side a pair of very large cells, one to the right and one to the left (Figure 1.39 p): these are the important polar cells of the primitive mouth, or "the primitive cells of the mesoderm." In consequence of these considerable variations arising in the course of the gastrulation, the primitive uni-axial form of the archigastrula in the amphioxus has already become tri-axial, and thus the two-sidedness, or bilateral symmetry, of the vertebrate body has already been determined. This has been transmitted from the amphioxus to all the other modified gastrula-forms of the vertebrate stem.
Apart from this bilateral structure, the gastrula of the amphioxus resembles the typical archigastrula of the lower animals (Figures 1.30 to 1.36) in developing the two primary germinal layers from a single layer of cells. This is clearly the oldest and original form of the metazoic embryo. Although the animals I have mentioned belong to the most diverse classes, they nevertheless agree with each other, and many more animal forms, in having retained to the present day, by a conservative heredity, this palingenetic form of gastrulation which they have from their earliest common ancestors. But this is not the case with the great majority of the animals. With these the original embryonic process has been gradually more or less altered in the course of millions of years by adaptation to new conditions of development. Both the segmentation of the ovum and the subsequent gastrulation have in this way been considerably changed. In fact, these variations have become so great in the course of time that the segmentation was not rightly understood in most animals, and the gastrula was unrecognised. It was not until I had made an extensive comparative study, lasting a considerable time (in the years 1866 to 1875), in animals of the most diverse classes, that I succeeded in showing the same common typical process in these apparently very different forms of gastrulation, and tracing them all to one original form. I regard all those that diverge from the primary palingenetic gastrulation as secondary, modified, and cenogenetic. The more or less divergent form of gastrula that is produced may be called a secondary, modified gastrula, or a metagastrula. The reader will find a scheme of these different kinds of segmentation and gastrulation at the close of this chapter.