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.

Fig. 1—The human ovum. 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. Fig. 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.

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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 Figs. 3–7.

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, Fig. 1c and Fig. 2k). 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” (Bütschli).

Fig. 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 (Fig. 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 (Fig. 2k). 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 amœbæ (Fig. 15 and 16) and the amœboid travelling cells (Fig. 11), and also with very young ova (Fig. 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 (Fig. 3). In the outer skin we find similar, but harder, covering cells, joined together by saw-like edges (Fig. 4). In the liver and other glands there are thicker and softer cells, linked together in rows (Fig. 5).

The last-named tissues (Figs. 3–5) 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,

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the cells (Fig. 6) are star-shaped, and are joined together by numbers of net-like interlacing processes; so, also, in the tissues of the teeth (Fig. 7), and in other forms of supporting-tissue, in which a soft or hard substance (intercellular matter, or base) is inserted between the cells.

Fig. 3—Three epithelial cells from the mucous lining of the tongue. Fig. 4—Five spiny or grooved cells, with edges joined, from the outer skin (epidermis): one of them (b) is isolated. Fig. 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 (Fig. 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 (Fig. 2k). 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,

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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.

Fig. 6—Nine star-shaped bone-cells, with interlaced branches.

The simple round ovum, which we take as the starting-point of our study (Figs. 1 and 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 (Fig. 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.

Fig. 7—Eleven star-shaped cells from the enamel of a tooth, joined together by their branchlets.

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

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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.

Fig. 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”).

In contrast with this very elaborate and very strictly differentiated psychic cell (Fig. 9), we have our ovum (Figs. 1 and 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 XIX.) 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. Fig. 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 (Fig. 10). In these the nucleus first divides into two equal parts by constriction. The indirect (or “mitotic”)

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Fig. 9—A large branching nerve-cell, or “soul-cell”, from the brain of an electric fish (Torpedo). 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.)

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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, Fig. 11.)

Fig. 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.)

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, hæmatoxylin, 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, Fig. 11, B C). The chromatin often forms a long, irregularly-wound thread—“the coil” (spirema, Fig. 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 (Fig. 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 (Fig. 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

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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.

A. Mother-cell (Knot, spirema) 1. Nuclear threads (chromosomata) (coloured nuclear matter, chromatin) 2. Nuclear membrane 3. Nuclear sap 4. Cytosoma 5. Protoplasm of the cell-body B. Mother-star, the loops beginning to split lengthways (nuclear membrane gone) 1. Star-like appearance in cytoplasm 2. Centrosoma (sphere of attraction) 3. Nuclear spindle (achromin, colourless matter) 4. Nuclear loops (chromatin, coloured matter) C. The two daughter-stars, produced by the breaking of the loops of the mother-star (moving away) 1. Upper daughter-crown 2. Connecting threads of the two crowns (achromin) 3. Lower daughter-crown 4. 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) 1. Upper daughter-nucleus 2. Equatorial constriction of the cell-body 3. Lower daughter-nucleus.

Fig. 11—Indirect or mitotic cell-division (with caryolysis and caryokinesis) from the skin of the larva of a salamander. (From Rabl.).

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 (Fig. 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 (Fig. 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

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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).

Fig. 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 amœba or rhizopods) protruding fine processes from the uncovered protoplasmic body. These bodies vary continually in number, shape, and size. The nucleus of these amœboid 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 medusæ) 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 (Fig. 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 Chap. XXIX).

The fertilised bird-ovum (Fig. 15) is notably different. It is true that in its earliest stage (Fig. 13 E) this ovum also is very like that of the mammal (Fig. 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

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Fig. 13—Ova of various animals, executing amœboid movements, 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. Figs. A1–A4 represent the ovum of a sponge (Leuculmis echinus) in four successive movements. B1–B8 are the ovum of a parasitic crab (Chondracanthus cornutus), in eight successive movements. (From Edward von Beneden.) C1–C5 show the ovum of the cat in various stages of movement (from Pflüger); Fig. D the ovum of a trout; E the ovum of a chicken; F a human ovum.

find it to be a simple, small, naked, amœboid cell, just like the young ova of other animals (Fig. 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) (Fig. 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, Fig. 15 d'). 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

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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 amœba, 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 Chap. IX.

Fig. 14—The human ovum, taken from the female ovary, magnified. 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.

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 fœtus. 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 (Fig. 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

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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.

Fig. 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'). The two kinds of yelk do not differ very much.

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 gregarinæ, 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 Amœba.

Fig. 16—A creeping amœba (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.

For a long time now we have comprised under the general name of amœbæ 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 amœbæ which live inside other animals. When we place one of these amœbæ 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 (Figs. 16 and 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 amœba 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

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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.

Fig. 17—Division of a unicellular amœba (Amœba 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.)

If you touch one of these creeping amœbæ 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 amœba 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 amœba 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 amœba pressing these coloured particles into itself, the substance of the cell closing round them. The amœba 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 amœba 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 (Fig. 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 amœba 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 amœbæ can be seen with the naked eye, but most of them are microscopically small. It is for the following reasons that we regard the amœbæ as the unicellular organisms which have

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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 amœba. Like the amœbæ, 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 amœbæ (Fig. 17). They had been observed by earlier scientists, but described as foreign bodies—namely, parasitic amœbæ, 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 medusæ. Their ova remain naked cells, which thrust out amœboid 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 amœba 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 amœba has a less definite and more original character than any other cell. Moreover, there is the fact that recent research has discovered such amœba-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 amœbæ, take up solid particles, or “eat” (whence they are called phagocytes = “eating-cells,” Fig. 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 amœbæ (Fig. 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 amœba-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 amœboid 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.

Fig. 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 amœba.)

We have now, by our study of the ovum and the comparison of it with the amœba, 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 amœba.

The statement that the earliest progenitors of the human race were simple cells of this kind, and led an independent unicellular life like the amœba, 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 amœba.

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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 Chap. XI).

Fig. 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 amœbæ.” 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 amœba.” 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 amœba. 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.

Title and Contents
Glossary
Chapter V
Chapter VII
Figs. 1–209
Figs. 210–408