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- The Evolution of Man, V.1. - 20/54 -


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 Evolution of Man, V.1. - 20/54

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