The Evolution of Man, V.1.
by Ernst Haeckel
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Here, I think, I have given an analytic solution of the problem of the origin of species, and so met the demand of certain opponents of evolution for an actual instance of descent from a stem-form. Those who are not satisfied with the synthetic proofs of the theory of evolution which are provided by comparative anatomy, embryology, paleontology, dysteleology, chorology, and classification, may try to refute the analytic proof given in my treatise on the sponge, the outcome of five years of assiduous study. I repeat: It is now impossible to oppose evolution on the ground that we have no convincing example of the descent of all the species of a group from a common ancestor. The monograph on the sponges furnishes such a proof, and, in my opinion, an indisputable proof. Any man of science who will follow the protracted steps of my inquiry and test my assertions will find that in the case of the sponges we can follow the actual evolution of species in a concrete case. And if this is so, if we can show the origin of all the species from a common form in one single class, we have the solution of the problem of man's origin, because we are in a position to prove clearly his descent from the lower animals.

At the same time, we can now reply to the often-repeated assertion, even heard from scientists of our own day, that the descent of man from the lower animals, and proximately from the apes, still needs to be "proved with certainty." These "certain proofs" have been available for a long time; one has only to open one's eyes to see them. It is a mistake to seek them in the discovery of intermediate forms between man and the ape, or the conversion of an ape into a human being by skilful education. The proofs lie in the great mass of empirical material we have already collected. They are furnished in the strongest form by the data of comparative anatomy and embryology, completed by paleontology. It is not a question now of detecting new proofs of the evolution of man, but of examining and understanding the proofs we already have.

I was almost alone thirty-six years ago when I made the first attempt, in my General Morphology, to put organic science on a mechanical foundation through Darwin's theory of descent. The association of ontogeny and phylogeny and the proof of the intimate causal connection between these two sections of the science of evolution, which I expounded in my work, met with the most spirited opposition on nearly all sides. The next ten years were a terrible "struggle for life" for the new theory. But for the last twenty-five years the tables have been turned. The phylogenetic method has met with so general a reception, and found so prolific a use in every branch of biology, that it seems superfluous to treat any further here of its validity and results. The proof of it lies in the whole morphological literature of the last three decades. But no other science has been so profoundly modified in its leading thoughts by this adoption, and been forced to yield such far-reaching consequences, as that science which I am now seeking to establish—monistic anthropogeny.

This statement may seem to be rather audacious, since the very next branch of biology, anthropology in the stricter sense, makes very little use of these results of anthropogeny, and sometimes expressly opposes them.* (*This does not apply to English anthropologists, who are almost all evolutionists.) This applies especially to the attitude which has characterised the German Anthropological Society (the Deutsche Gesellschaft fur Anthropologie) for some thirty years. Its powerful president, the famous pathologist, Rudolph Virchow, is chiefly responsible for this. Until his death (September 5th, 1902) he never ceased to reject the theory of descent as unproven, and to ridicule its chief consequence—the descent of man from a series of mammal ancestors—as a fantastic dream. I need only recall his well-known expression at the Anthropological Congress at Vienna in 1894, that "it would be just as well to say man came from the sheep or the elephant as from the ape."

Virchow's assistant, the secretary of the German Anthropological Society, Professor Johannes Ranke of Munich, has also indefatigably opposed transformism: he has succeeded in writing a work in two volumes (Der Mensch), in which all the facts relating to his organisation are explained in a sense hostile to evolution. This work has had a wide circulation, owing to its admirable illustrations and its able treatment of the most interesting facts of anatomy and physiology—exclusive of the sexual organs! But, as it has done a great deal to spread erroneous views among the general public, I have included a criticism of it in my History of Creation, as well as met Virchow's attacks on anthropogeny.

Neither Virchow, nor Ranke, nor any other "exact" anthropologist, has attempted to give any other natural explanation of the origin of man. They have either set completely aside this "question of questions" as a transcendental problem, or they have appealed to religion for its solution. We have to show that this rejection of the rational explanation is totally without justification. The fund of knowledge which has accumulated in the progress of biology in the nineteenth century is quite adequate to furnish a rational explanation, and to establish the theory of the evolution of man on the solid facts of his embryology.


In order to understand clearly the course of human embryology, we must select the more important of its wonderful and manifold processes for fuller explanation, and then proceed from these to the innumerable features of less importance. The most important feature in this sense, and the best starting-point for ontogenetic study, is the fact that man is developed from an ovum, and that this ovum is a simple cell. The human ovum does not materially differ in form and composition from that of the other mammals, whereas there is a distinct difference between the fertilised ovum of the mammal and that of any other animal.

(FIGURE 1.1. The human ovum, magnified 100 times. The globular mass of yelk (b) is enclosed by a transparent membrane (the ovolemma or zona pellucida [a]), and contains a noncentral nucleus (the germinal vesicle, c). Cf. Figure 1.14.)

This fact is so important that few should be unaware of its extreme significance; yet it was quite unknown in the first quarter of the nineteenth century. As we have seen, the human and mammal ovum was not discovered until 1827, when Carl Ernst von Baer detected it. Up to that time the larger vesicles, in which the real and much smaller ovum is contained, had been wrongly regarded as ova. The important circumstance that this mammal ovum is a simple cell, like the ovum of other animals, could not, of course, be recognised until the cell theory was established. This was not done, by Schleiden for the plant and Schwann for the animal, until 1838. As we have seen, this cell theory is of the greatest service in explaining the human frame and its embryonic development. Hence we must say a few words about the actual condition of the theory and the significance of the views it has suggested.

In order properly to appreciate the cellular theory, the most important element in our science, it is necessary to understand in the first place that the cell is a UNIFIED ORGANISM, a self-contained living being. When we anatomically dissect the fully-formed animal or plant into its various organs, and then examine the finer structure of these organs with the microscope, we are surprised to find that all these different parts are ultimately made up of the same structural element or unit. This common unit of structure is the cell. It does not matter whether we thus dissect a leaf, flower, or fruit, or a bone, muscle, gland, or bit of skin, etc.; we find in every case the same ultimate constituent, which has been called the cell since Schleiden's discovery. There are many opinions as to its real nature, but the essential point in our view of the cell is to look upon it as a self-contained or independent living unit. It is, in the words of Brucke, "an elementary organism." We may define it most precisely as the ultimate organic unit, and, as the cells are the sole active principles in every vital function, we may call them the "plastids," or "formative elements." This unity is found in both the anatomic structure and the physiological function. In the case of the protists, the entire organism usually consists of a single independent cell throughout life. But in the tissue-forming animals and plants, which are the great majority, the organism begins its career as a simple cell, and then grows into a cell-community, or, more correctly, an organised cell-state. Our own body is not really the simple unity that it is generally supposed to be. On the contrary, it is a very elaborate social system of countless microscopic organisms, a colony or commonwealth, made up of innumerable independent units, or very different tissue-cells.

In reality, the term "cell," which existed long before the cell theory was formulated, is not happily chosen. Schleiden, who first brought it into scientific use in the sense of the cell theory, gave this name to the elementary organisms because, when you find them in the dissected plant, they generally have the appearance of chambers, like the cells in a bee-hive, with firm walls and a fluid or pulpy content. But some cells, especially young ones, are entirely without the enveloping membrane, or stiff wall. Hence we now generally describe the cell as a living, viscous particle of protoplasm, enclosing a firmer nucleus in its albuminoid body. There may be an enclosing membrane, as there actually is in the case of most of the plants; but it may be wholly lacking, as is the case with most of the animals. There is no membrane at all in the first stage. The young cells are usually round, but they vary much in shape later on. Illustrations of this will be found in the cells of the various parts of the body shown in Figures 1.3 to 1.7.

Hence the essential point in the modern idea of the cell is that it is made up of two different active constituents—an inner and an outer part. The smaller and inner part is the nucleus (or caryon or cytoblastus, Figure 1.1 c and Figure 1.2 k). The outer and larger part, which encloses the other, is the body of the cell (celleus, cytos, or cytosoma). The soft living substance of which the two are composed has a peculiar chemical composition, and belongs to the group of the albuminoid plasma-substances ("formative matter"), or protoplasm. The essential and indispensable element of the nucleus is called nuclein (or caryoplasm); that of the cell body is called plastin (or cytoplasm). In the most rudimentary cases both substances seem to be quite simple and homogeneous, without any visible structure. But, as a rule, when we examine them under a high power of the microscope, we find a certain structure in the protoplasm. The chief and most common form of this is the fibrous or net-like "thready structure" (Frommann) and the frothy "honeycomb structure" (Butschli).

(FIGURE 1.2. Stem-cell of one of the echinoderms (cytula, or "first segmentation-cell" = fertilised ovum), after Hertwig. k is the nucleus or caryon.)

The shape or outer form of the cell is infinitely varied, in accordance with its endless power of adapting itself to the most diverse activities or environments. In its simplest form the cell is globular (Figure 1.2). This normal round form is especially found in cells of the simplest construction, and those that are developed in a free fluid without any external pressure. In such cases the nucleus also is not infrequently round, and located in the centre of the cell-body (Figure 1.2 k). In other cases, the cells have no definite shape; they are constantly changing their form owing to their automatic movements. This is the case with the amoebae (Figures 1.15 and 1.16) and the amoeboid travelling cells (Figure 1.11), and also with very young ova (Figure 1.13). However, as a rule, the cell assumes a definite form in the course of its career. In the tissues of the multicellular organism, in which a number of similar cells are bound together in virtue of certain laws of heredity, the shape is determined partly by the form of their connection and partly by their special functions. Thus, for instance, we find in the mucous lining of our tongue very thin and delicate flat cells of roundish shape (Figure 1.3). In the outer skin we find similar, but harder, covering cells, joined together by saw-like edges (Figure 1.4). In the liver and other glands there are thicker and softer cells, linked together in rows (Figure 1.5).

The last-named tissues (Figures 1.3 to 1.5) belong to the simplest and most primitive type, the group of the "covering-tissues," or epithelia. In these "primary tissues" (to which the germinal layers belong) simple cells of the same kind are arranged in layers. The arrangement and shape are more complicated in the "secondary tissues," which are gradually developed out of the primary, as in the tissues of the muscles, nerves, bones, etc. In the bones, for instance, which belong to the group of supporting or connecting organs, the cells (Figure 1.6) are star-shaped, and are joined together by numbers of net-like interlacing processes; so, also, in the tissues of the teeth (Figure 1.7), and in other forms of supporting-tissue, in which a soft or hard substance (intercellular matter, or base) is inserted between the cells.

(FIGURE 1.3. Three epithelial cells from the mucous lining of the tongue.

FIGURE 1.4. Five spiny or grooved cells, with edges joined, from the outer skin (epidermis): one of them (b) is isolated.

FIGURE 1.5. Ten liver-cells: one of them (b) has two nuclei.)

The cells also differ very much in size. The great majority of them are invisible to the naked eye, and can be seen only through the microscope (being as a rule between 1/2500 and 1/250 inch in diameter). There are many of the smaller plastids—such as the famous bacteria—which only come into view with a very high magnifying power. On the other hand, many cells attain a considerable size, and run occasionally to several inches in diameter, as do certain kinds of rhizopods among the unicellular protists (such as the radiolaria and thalamophora). Among the tissue-cells of the animal body many of the muscular fibres and nerve fibres are more than four inches, and sometimes more than a yard, in length. Among the largest cells are the yelk-filled ova; as, for instance, the yellow "yolk" in the hen's egg, which we shall describe later (Figure 1.15).

Cells also vary considerably in structure. In this connection we must first distinguish between the active and passive components of the cell. It is only the former, or active parts of the cell, that really live, and effect that marvellous world of phenomena to which we give the name of "organic life." The first of these is the inner nucleus (caryoplasm), and the second the body of the cell (cytoplasm). The passive portions come third; these are subsequently formed from the others, and I have given them the name of "plasma-products." They are partly external (cell-membranes and intercellular matter) and partly internal (cell-sap and cell-contents).

The nucleus (or caryon), which is usually of a simple roundish form, is quite structureless at first (especially in very young cells), and composed of homogeneous nuclear matter or caryoplasm (Figure 1.2 k). But, as a rule, it forms a sort of vesicle later on, in which we can distinguish a more solid nuclear base (caryobasis) and a softer or fluid nuclear sap (caryolymph). In a mesh of the nuclear network (or it may be on the inner side of the nuclear envelope) there is, as a rule, a dark, very opaque, solid body, called the nucleolus. Many of the nuclei contain several of these nucleoli (as, for instance, the germinal vesicle of the ova of fishes and amphibia). Recently a very small, but particularly important, part of the nucleus has been distinguished as the central body (centrosoma)—a tiny particle that is originally found in the nucleus itself, but is usually outside it, in the cytoplasm; as a rule, fine threads stream out from it in the cytoplasm. From the position of the central body with regard to the other parts it seems probable that it has a high physiological importance as a centre of movement; but it is lacking in many cells.

The cell-body also consists originally, and in its simplest form, of a homogeneous viscid plasmic matter. But, as a rule, only the smaller part of it is formed of the living active cell-substance (protoplasm); the greater part consists of dead, passive plasma-products (metaplasm). It is useful to distinguish between the inner and outer of these. External plasma-products (which are thrust out from the protoplasm as solid "structural matter") are the cell-membranes and the intercellular matter. The internal plasma-products are either the fluid cell-sap or hard structures. As a rule, in mature and differentiated cells these various parts are so arranged that the protoplasm (like the caryoplasm in the round nucleus) forms a sort of skeleton or framework. The spaces of this network are filled partly with the fluid cell-sap and partly by hard structural products.

(FIGURE 1.6. Nine star-shaped bone-cells, with interlaced branches.

FIGURE 1.7. Eleven star-shaped cells from the enamel of a tooth, joined together by their branchlets.)

The simple round ovum, which we take as the starting-point of our study (Figures 1.1 and 1.2), has in many cases the vague, indifferent features of the typical primitive cell. As a contrast to it, and as an instance of a very highly differentiated plastid, we may consider for a moment a large nerve-cell, or ganglionic cell, from the brain. The ovum stands potentially for the entire organism—in other words, it has the faculty of building up out of itself the whole multicellular body. It is the common parent of all the countless generations of cells which form the different tissues of the body; it unites all their powers in itself, though only potentially or in germ. In complete contrast to this, the neural cell in the brain (Figure 1.9) develops along one rigid line. It cannot, like the ovum, beget endless generations of cells, of which some will become skin-cells, others muscle-cells, and others again bone-cells. But, on the other hand, the nerve-cell has become fitted to discharge the highest functions of life; it has the powers of sensation, will, and thought. It is a real soul-cell, or an elementary organ of the psychic activity. It has, therefore, a most elaborate and delicate structure. Numbers of extremely fine threads, like the electric wires at a large telegraphic centre, cross and recross in the delicate protoplasm of the nerve cell, and pass out in the branching processes which proceed from it and put it in communication with other nerve-cells or nerve-fibres (a, b). We can only partly follow their intricate paths in the fine matter of the body of the cell.

Here we have a most elaborate apparatus, the delicate structure of which we are just beginning to appreciate through our most powerful microscopes, but whose significance is rather a matter of conjecture than knowledge. Its intricate structure corresponds to the very complicated functions of the mind. Nevertheless, this elementary organ of psychic activity—of which there are thousands in our brain—is nothing but a single cell. Our whole mental life is only the joint result of the combined activity of all these nerve-cells, or soul-cells. In the centre of each cell there is a large transparent nucleus, containing a small and dark nuclear body. Here, as elsewhere, it is the nucleus that determines the individuality of the cell; it proves that the whole structure, in spite of its intricate composition, amounts to only a single cell.

(FIGURE 1.8. Unfertilised ovum of an echinoderm (from Hertwig). The vesicular nucleus (or "germinal vesicle") is globular, half the size of the round ovum, and encloses a nuclear framework, in the central knot of which there is a dark nucleolus (the "germinal spot").

FIGURE 1.9. A large branching nerve-cell, or "soul-cell," from the brain of an electric fish (Torpedo), magnified 600 times. In the middle of the cell is the large transparent round nucleus, one nucleolus, and, within the latter again, a nucleolinus. The protoplasm of the cell is split into innumerable fine threads (or fibrils), which are embedded in intercellular matter, and are prolonged into the branching processes of the cell (b). One branch (a) passes into a nerve-fibre. (From Max Schultze.))

In contrast with this very elaborate and very strictly differentiated psychic cell (Figure 1.9), we have our ovum (Figures 1.1 and 1.2), which has hardly any structure at all. But even in the case of the ovum we must infer from its properties that its protoplasmic body has a very complicated chemical composition and a fine molecular structure which escapes our observation. This presumed molecular structure of the plasm is now generally admitted; but it has never been seen, and, indeed, lies far beyond the range of microscopic vision. It must not be confused—as is often done—with the structure of the plasm (the fibrous network, groups of granules, honey-comb, etc.) which does come within the range of the microscope.

But when we speak of the cells as the elementary organisms, or structural units, or "ultimate individualities," we must bear in mind a certain restriction of the phrases. I mean, that the cells are not, as is often supposed, the very lowest stage of organic individuality. There are yet more elementary organisms to which I must refer occasionally. These are what we call the "cytodes" (cytos = cell), certain living, independent beings, consisting only of a particle of plasson—an albuminoid substance, which is not yet differentiated into caryoplasm and cytoplasm, but combines the properties of both. Those remarkable beings called the monera—especially the chromacea and bacteria—are specimens of these simple cytodes. (Compare Chapter 2.19.) To be quite accurate, then, we must say: the elementary organism, or the ultimate individual, is found in two different stages. The first and lower stage is the cytode, which consists merely of a particle of plasson, or quite simple plasm. The second and higher stage is the cell, which is already divided or differentiated into nuclear matter and cellular matter. We comprise both kinds—the cytodes and the cells—under the name of plastids ("formative particles"), because they are the real builders of the organism. However, these cytodes are not found, as a rule, in the higher animals and plants; here we have only real cells with a nucleus. Hence, in these tissue-forming organisms (both plant and animal) the organic unit always consists of two chemically and anatomically different parts—the outer cell-body and the inner nucleus.

In order to convince oneself that this cell is really an independent organism, we have only to observe the development and vital phenomena of one of them. We see then that it performs all the essential functions of life—both vegetal and animal—which we find in the entire organism. Each of these tiny beings grows and nourishes itself independently. It takes its food from the surrounding fluid; sometimes, even, the naked cells take in solid particles at certain points of their surface—in other words, "eat" them—without needing any special mouth and stomach for the purpose (cf. Figure 1.19).

Further, each cell is able to reproduce itself. This multiplication, in most cases, takes the form of a simple cleavage, sometimes direct, sometimes indirect; the simple direct (or "amitotic") division is less common, and is found, for instance, in the blood cells (Figure 1.10). In these the nucleus first divides into two equal parts by constriction. The indirect (or "mitotic") cleavage is much more frequent; in this the caryoplasm of the nucleus and the cytoplasm of the cell-body act upon each other in a peculiar way, with a partial dissolution (caryolysis), the formation of knots and loops (mitosis), and a movement of the halved plasma-particles towards two mutually repulsive poles of attraction (caryokinesis, Figure 1.11.)

(FIGURE 1.10. Blood-cells, multiplying by direct division, from the blood of the embryo of a stag. Originally, each blood-cell has a nucleus and is round (a). When it is going to multiply, the nucleus divides into two (b, c, d). Then the protoplasmic body is constricted between the two nuclei, and these move away from each other (e). Finally, the constriction is complete, and the cell splits into two daughter-cells (f). (From Frey.))

FIGURE 1.11. Indirect or mitotic cell-division (with caryolysis and caryokinesis) from the skin of the larva of a salamander. (From Rabl.). A. Mother-cell (Knot, spirema), with Nuclear threads (chromosomata) (coloured nuclear matter, chromatin), Cytosoma, Nuclear membrane, Protoplasm of the cell-body and Nuclear sap. B. Mother-star, the loops beginning to split lengthways (nuclear membrane gone), with Star-like appearance in cytoplasm, Centrosoma (sphere of attraction), Nuclear spindle (achromin, colourless matter) and Nuclear loops (chromatin, coloured matter). C. The two daughter-stars, produced by the breaking of the loops of the mother-star (moving away), with Upper daughter-crown, Connecting threads of the two crowns (achromin), Lower daughter-crown and Double-star (amphiaster). D. The two daughter-cells, produced by the complete division of the two nuclear halves (cytosomata still connected at the equator) (Double-knot, Dispirema), with Upper daughter-nucleus, Equatorial constriction of the cell-body and Lower daughter-nucleus.)

The intricate physiological processes which accompany this "mitosis" have been very closely studied of late years. The inquiry has led to the detection of certain laws of evolution which are of extreme importance in connection with heredity. As a rule, two very different parts of the nucleus play an important part in these changes. They are: the chromatin, or coloured nuclear substance, which has a peculiar property of tingeing itself deeply with certain colouring matters (carmine, haematoxylin, etc.), and the achromin (or linin, or achromatin), a colourless nuclear substance that lacks this property. The latter generally forms in the dividing cell a sort of spindle, at the poles of which there is a very small particle, also colourless, called the "central body" (centrosoma). This acts as the centre or focus in a "sphere of attraction" for the granules of protoplasm in the surrounding cell-body, and assumes a star-like appearance (the cell-star, or monaster). The two central bodies, standing opposed to each other at the poles of the nuclear spindle, form "the double-star" (or amphiaster, Figure 1.11, BC). The chromatin often forms a long, irregularly-wound thread—"the coil" (spirema, Figure A). At the commencement of the cleavage it gathers at the equator of the cell, between the stellar poles, and forms a crown of U-shaped loops (generally four or eight, or some other definite number). The loops split lengthwise into two halves (B), and these back away from each other towards the poles of the spindle (C). Here each group forms a crown once more, and this, with the corresponding half of the divided spindle, forms a fresh nucleus (D). Then the protoplasm of the cell-body begins to contract in the middle, and gather about the new daughter-nuclei, and at last the two daughter-cells become independent beings.

Between this common mitosis, or indirect cell-division—which is the normal cleavage-process in most cells of the higher animals and plants—and the simple direct division (Figure 1.10) we find every grade of segmentation; in some circumstances even one kind of division may be converted into another.

The plastid is also endowed with the functions of movement and sensation. The single cell can move and creep about, when it has space for free movement and is not prevented by a hard envelope; it then thrusts out at its surface processes like fingers, and quickly withdraws them again, and thus changes its shape (Figure 1.12). Finally, the young cell is sensitive, or more or less responsive to stimuli; it makes certain movements on the application of chemical and mechanical irritation. Hence we can ascribe to the individual cell all the chief functions which we comprehend under the general heading of "life"—sensation, movement, nutrition, and reproduction. All these properties of the multicellular and highly developed animal are also found in the single animal-cell, at least in its younger stages. There is no longer any doubt about this, and so we may regard it as a solid and important base of our physiological conception of the elementary organism.

Without going any further here into these very interesting phenomena of the life of the cell, we will pass on to consider the application of the cell theory to the ovum. Here comparative research yields the important result that EVERY OVUM IS AT FIRST A SIMPLE CELL. I say this is very important, because our whole science of embryology now resolves itself into the problem: "How does the multicellular organism arise from the unicellular?" Every organic individual is at first a simple cell, and as such an elementary organism, or a unit of individuality. This cell produces a cluster of cells by segmentation, and from these develops the multicellular organism, or individual of higher rank.

When we examine a little closer the original features of the ovum, we notice the extremely significant fact that in its first stage the ovum is just the same simple and indefinite structure in the case of man and all the animals (Figure 1.13). We are unable to detect any material difference between them, either in outer shape or internal constitution. Later, though the ova remain unicellular, they differ in size and shape, enclose various kinds of yelk-particles, have different envelopes, and so on. But when we examine them at their birth, in the ovary of the female animal, we find them to be always of the same form in the first stages of their life. In the beginning each ovum is a very simple, roundish, naked, mobile cell, without a membrane; it consists merely of a particle of cytoplasm enclosing a nucleus (Figure 1.13). Special names have been given to these parts of the ovum; the cell-body is called the yelk (vitellus), and the cell-nucleus the germinal vesicle. As a rule, the nucleus of the ovum is soft, and looks like a small pimple or vesicle. Inside it, as in many other cells, there is a nuclear skeleton or frame and a third, hard nuclear body (the nucleolus). In the ovum this is called the germinal spot. Finally, we find in many ova (but not in all) a still further point within the germinal spot, a "nucleolin," which goes by the name of the germinal point. The latter parts (germinal spot and germinal point) have, apparently, a minor importance, in comparison with the other two (the yelk and germinal vesicle). In the yelk we must distinguish the active formative yelk (or protoplasm = first plasm) from the passive nutritive yelk (or deutoplasm = second plasm).

(FIGURE 1.12. Mobile cells from the inflamed eye of a frog (from the watery fluid of the eye, the humor aqueus). The naked cells creep freely about, by (like the amoeba or rhizopods) protruding fine processes from the uncovered protoplasmic body. These bodies vary continually in number, shape, and size. The nucleus of these amoeboid lymph-cells ("travelling cells," or planocytes) is invisible, because concealed by the numbers of fine granules which are scattered in the protoplasm. (From Frey.))

In many of the lower animals (such as sponges, polyps, and medusae) the naked ova retain their original simple appearance until impregnation. But in most animals they at once begin to change; the change consists partly in the formation of connections with the yelk, which serve to nourish the ovum, and partly of external membranes for their protection (the ovolemma, or prochorion). A membrane of this sort is formed in all the mammals in the course of the embryonic process. The little globule is surrounded by a thick capsule of glass-like transparency, the zona pellucida, or ovolemma pellucidum (Figure 1.14). When we examine it closely under the microscope, we see very fine radial streaks in it, piercing the zona, which are really very narrow canals. The human ovum, whether fertilised or not, cannot be distinguished from that of most of the other mammals. It is nearly the same everywhere in form, size, and composition. When it is fully formed, it has a diameter of (on an average) about 1/120 of an inch. When the mammal ovum has been carefully isolated, and held against the light on a glass-plate, it may be seen as a fine point even with the naked eye. The ova of most of the higher mammals are about the same size. The diameter of the ovum is almost always between 1/250 to 1/125 inch. It has always the same globular shape; the same characteristic membrane; the same transparent germinal vesicle with its dark germinal spot. Even when we use the most powerful microscope with its highest power, we can detect no material difference between the ova of man, the ape, the dog, and so on. I do not mean to say that there are no differences between the ova of these different mammals. On the contrary, we are bound to assume that there are such, at least as regards chemical composition. Even the ova of different men must differ from each other; otherwise we should not have a different individual from each ovum. It is true that our crude and imperfect apparatus cannot detect these subtle individual differences, which are probably in the molecular structure. However, such a striking resemblance of their ova in form, so great as to seem to be a complete similarity, is a strong proof of the common parentage of man and the other mammals. From the common germ-form we infer a common stem-form. On the other hand, there are striking peculiarities by which we can easily distinguish the fertilised ovum of the mammal from the fertilised ovum of the birds, amphibia, fishes, and other vertebrates (see the close of Chapter 2.29).

(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.


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.)


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.

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