FIG. 41.—The chromosomes in the male and female pronucleii have resolved into a network. The male centrosome begins to show signs of activity.
FIG. 42.—The centrosome has divided, and the two pronucleii have been brought together. The network in each nucleus has again resolved itself into two chromosomes which are now brought together near the centre of the egg but do not fuse; mcr, represents the chromosomes from the male nucleus; fcr, the chromosomes from the female nucleus.

In the cases where the process has been most carefully studied, the further changes are as follows: The head of the spermatozoon, after entrance into the egg, lies dormant until the egg has thrown off its polar cells, and thus gotten rid of part of its chromosomes. Close to it lies its centrosomes (Fig. 35, ce), and there is thus formed what is known as the male pronucleus (Fig. 35-40, mn). The remains of the egg nucleus, after having discharged the polar cells, form the female nucleus (Fig. 40, fn). The chromatin material, in both the male and female pronucleus, soon breaks up into a network in which it is no longer possible to see that each contains two chromosomes (Fig. 41). Now the centrosome, which is beside the male pronucleus, shows signs of activity. It becomes surrounded by prominent rays to form an aster (Fig. 41, ce), and then it begins to move toward the female pronucleus, apparently dragging the male pronucleus after it. In this way the centrosome approaches the female pronucleus, and thus finally the two nucleii are brought into close proximity. Meantime the chromatin material in each has once more broken up into short threads or chromosomes, and once more we find that each of the nucleii contains two of these bodies (Fig. 42). In the subsequent figures the chromosomes of the male nucleus are lightly shaded, while those of the female are black in order to distinguish them. As these two nucleii finally come together their membranes disappear, and the chromatic material comes to lie freely in the egg, the male and female chromosomes, side by side, but distinct forming the segmentation nucleus. The egg plainly now contains once more the number of chromosomes normal for the cells of the animal, but half of them have been derived from each parent. It is very suggestive to find further that the chromosomes in this fertilized egg do not fuse with each other, but remain quite distinct, so that it can be seen that the new nucleus contains chromosomes derived from each parent (Fig. 42). Nor does there appear to be, in the future history of this egg, any actual fusion of the chromatic material, the male and female chromosomes perhaps always remaining distinct.

FIG. 43.—An equatorial plate is formed and each chromosome has split into two halves by longitudinal division.
FIG. 44.—The halves of the chromosomes have separated to form two nucleii, each with male and female chromosomes. The egg has divided into two cells.

While this mixture of chromosomes has been taking place the centrosome has divided into two parts, each of which becomes surrounded by an aster and travels to opposite ends of the nucleus (Fig. 42). There now follows a division of the nucleus exactly similar to that which occurs in the normal cell division already described in Figs. 28-34. Each of the chromosomes splits lengthwise (Fig. 43), and one half of each then travels toward each centrosome to form a new nucleus (Fig. 44). Since each of the four chromosomes thus splits, it follows that each of the two daughter nucleii will, of course, contain four chromosomes; two of which have been derived from the male and two from the female parent. From now the divisions of the egg follow rapidly by the normal process of cell division until from this one egg cell there are eventually derived hundreds of thousands of cells which are gradually moulded into the adult. All of these cells will, of course, contain four chromosomes; and, what is more important, half of the chromosomes will have been derived directly from the male and half from the female parent. Even into adult life, therefore, the cells of the animal probably contain chromatin derived by direct descent from each of its parents.

The Significance of Fertilization.—From this process of fertilization a number of conclusions, highly important for our purpose, can be drawn. In the first place, it is evident that the chromosomes form the part of the cell which contain the hereditary traits handed down from parent to child. This follows from the fact that the chromosomes are the only part of the cell which, in the fertilized egg, is derived from both parents. Now the offspring can certainly inherit from each parent, and hence the hereditary traits must be associated with some part of the cell which is derived from both. But the egg substance is derived from the mother alone; the centrosome, at least in some cases and perhaps in all, is derived only from the father, while the chromosomes are derived from both parents. Hence it follows that the hereditary traits must be particularly associated with the chromosomes.

With this understanding we can, at least, in part understand the purpose of fertilization. As we shall see later, it is very necessary in the building of the living machine for each individual to inherit characters from more than one individual. This is necessary to produce the numerous variations which contribute to the construction of the machine. For this purpose there has been developed the process of sexual union of reproductive cells, which introduces into the offspring chromatic material from two parents. But if the two reproductive cells should unite at once the number of chromosomes would be doubled in each generation, and hence be constantly increasing. To prevent this the polar cells are cast out, which reduces the amount of chromatic material. The union of the two pronucleii is plainly to produce a nucleus which shall contain chromosomes, and hence hereditary traits from each parent and the subsequent splitting of these chromosomes and the separation of the two halves into daughter nucleii insures that all the nucleii, and hence all cells of the adult, shall possess hereditary traits derived from both parents. Thus it comes that, even in the adult, every body cell is made up of chromosomes from each parent, and may hence inherit characters from each.

The cell of an animal thus consists of three somewhat distinct but active parts—the cell substance, the chromosomes, and the centrosome. Of these the cell substance appears to be handed down from the mother; the centrosome comes, at least in some cases, from the father, and the chromosomes from both parents. It is not yet certain, however, whether the centrosome is a constant part of the cell. In some cells it cannot yet be found, and there are some reasons for believing that it may be formed out of other parts of the cell. The nucleus is always a direct descendant from the nucleus of pre-existing cells, so that there is an absolute continuity of descent between the nucleii of the cells of an individual and those of its antecedents back for numberless generations. It is not certain that there is any such continuity of descent in the case of the centrosomes; for, while in the process of fertilization the centrosome is handed down from parent to child, there are some reasons for believing that it may disappear in subsequent cells, and later be redeveloped out of other parts. The only part of the cell in which complete continuity from parent to child is demonstrated, is the nucleus and particularly the chromosomes. All of these facts simply emphasize the importance of the chromosomes, and tell us that these bodies must be regarded as containing the most important features of the cell which constitute its individuality.