FIG. 36—The egg centrosomes have changed their position. The male cell with its centrosome remains inactive until the stage represented in FIG. 42.
FIG. 37—Beginning of the first division for removing superfluous chromosomes.
This is done by a process shown in Fig. 35. The centrosome divides as in ordinary cell division (Fig. 35), and after rotating on its axis it approaches the surface of the egg (Figs. 36 and 37). The egg now divides (Fig. 38), but the division is of a peculiar kind. Although the chromosomes divide equally the egg itself divides into two very unequal parts, one part still appearing as the egg and the other as a minute protuberance called the polar cell (pc' in Fig. 38). The chromosomes do not split as they do in the cell division already described, but each of these two cells, the egg and the polar body, receives four chromosomes (Fig. 38). The result is that the egg has now the normal number of chromosomes for the ordinary cells of the animal in question. But this is still too many, for the egg is soon to unite with the male cell; and this male cell, as we shall see, is to bring in its own quota of chromosomes. Hence the egg must get rid of still more of its chromatin material. Consequently, the first division is followed by a second (Fig. 39), in which there is again produced a large and a small cell. This division, like the first, occurs without any splitting of the chromosomes, one half of the remaining chromosomes being ejected in this new cell, the second polar cell (pc") leaving the larger cell, the egg, with just one half the number of chromosomes normal for the cells of the animal in question. Meantime the first pole cell has also divided, so that we have now, as shown in Fig. 40, four cells, three small and one large, but each containing one half the normal number of chromosomes. In the example figured, four is the normal number for the cells of the animal. The egg at the beginning of the process contained eight, but has now been reduced to two. In the further history of the egg the smaller cells, called polar cells, take no part, since they soon disappear and have nothing to do with the animal which is to result from the further division of the egg. This process of the formation of the polar cells is thus simply a device for getting rid of some of the chromatin material in the egg cell, so that it may unite with a second cell without doubling the normal number of chromosomes.
FIG.38—First division complete and first polar cell formed, pc'.
FIG.39.—Formation of the second polar cell, pc".
FIG.40.—Completion of the process of extrusion of the chromatic material; fn shows the two chromosomes retained in the egg forming the female pronucleus. The centrosome has disappeared.
Previously to this process the other sexual cell, the spermatozoon, or male reproductive cell, has been undergoing a somewhat similar process. This is also a true cell (Fig. 34, mc), although it is of a decidedly smaller size than the egg and of a very different shape. It contains cell substance, a nucleus with chromosomes, and a centrosome, the number of chromosomes, as shown later, being however only half that normal for the ordinary cells of the animals. The study of the development of the spermatozoon shows that it has come from cells which contained the normal number of four, but that this number has been reduced to one half by a process which is equivalent to that which we have just noticed in the egg. Thus it comes about that each of the sexual elements, the egg and the spermatozoon, now contains one half the normal number of chromosomes.
Now by some mechanical means these two reproductive cells are brought in contact with each other, shown in Fig. 34, and as soon as they are brought into each other's vicinity the male cell buries its head in the body of the egg. The tail by which it has been moving is cast off, and the head containing the chromosomes and the centrosome enters the egg, forming what is called the male pronucleus (Fig. 35-38, mn). This entrance of the male cell occurs either before the formation of the polar cells of the egg or afterward. If, however, it takes place before, the male pronucleus simply remains dormant in the egg while the polar cells are being protruded, and not until after that process is concluded does it begin again to show signs of activity which result in the cell union.
The further steps in this process appear to be controlled by the centrosome, although it is not quite certain whence this centrosome is derived. Originally, as we have seen, the egg contained a centrosome, and the male cell has also brought a second into the egg (Fig. 35, ce). In some cases, and this is true for the worm we are describing, it is certain that the egg centrosome disappears while that of the spermatozoon is retained alone to direct the further activities (Fig. 41). Possibly this may be the case in all eggs, but it is not sure. It is a matter of some little interest to have this settled, for if it should prove true, then it would evidently follow that the machinery for cell division, in the case of sexual reproduction, is derived from the father, although the bulk of the cell comes from the mother, while the chromosomes come from both parents.