chloride in sea-water,[1] as also the possibility in many cases that even in normal fertilization the cleavage centrosomes may arise de novo,[2] make it no longer possible to regard the centrosome as a permanent cell-structure.
Significance of Mitosis.—Whatever may be the nature of the chemico-physical changes occurring during cell-division, of which the achromatic spindle and astral rays are the visible expression, it is certain that the whole of this complicated process has for its function, not the division of the chromatin, for that has already occurred on the spireme thread or even earlier, but the distribution of the divided chromatin granules to the two daughter nuclei. It is indeed usually assumed that the mitotic mechanism is not merely for the distribution, but for the equal distribution, of the sister granules to the two daughter nuclei. The conspicuous part the chromatin is seen to play in the whole mechanism of heredity—in maturation, fertilization and development—indicating as it does that the chromatin is the chief, if not the only, bearer of the specific qualities of the organism, sufficiently clearly emphasizes the importance of the equal distribution of this substance between the daughter cells at successive cell-divisions. There are, however, serious objections to the interpretation of mitosis as an adaptation to ensure this equal distribution of the chromatin. Not only does the occurrence of amitosis show that the mitotic mechanism is not essential for either nuclear or cytoplasmic division, but direct division may occur[3] in the life-history of the germ cells, the very point at which it should not occur had mitosis the significance usually attached to it. On the other hand, the most elaborate mitosis occurs in cell-tissues (e.g. skin of salamander larva) which can take no possible share in the reproduction of the species. Moreover, we have no reason for supposing that the division of the chromatin in amitosis is not as meristic, and its subsequent distribution as equal, as is so visibly the case in mitosis.[4] It is necessary, therefore, to seek for some other explanation of the elaborate mechanism of mitosis than that which assumes it necessary for the equal distribution of the divided chromatin granules. The present writer believes the true explanation to be found in that great economic law of nature, “division of labour.” The same economy which, working under the control of natural selection, has produced the complexly differentiated tissues of the higher metazoa, which has led to the sexual differentiation between the conjugating gametes and thus to the sexual differentiation of the parents, has resulted in the production of mitosis. Only here the economy finds expression in division of labour, not in space, but in time. The work of the self-propagating chromatin granules is so ordered that periods of undisturbed metabolic activity alternate with periods of reproductive activity. The brief space of time occupied by the latter process has necessitated a more elaborate specialization of the forces—whatever their nature—controlling cell-division; a specialization which has resulted, just as a similar specialization in so many other cases has resulted, in a visible differentiation of the cell-protoplasm. This explanation is in harmony with the occurrence of typical mitosis in active tissue cells on the one hand, and of amitosis in the relatively quiescent primary germ cells on the other.
Individuality of the Chromosomes.—The most striking feature in the behaviour of the chromatin in mitosis is its resolution, at each division, into a—for any particular species—constant number of chromosomes. This constant recurrence of the specific number of chromosomes at every cell-division is capable of explanation in two radically different ways. One explanation assumes for the organism a specific peculiarity determining the segmentation of the spireme thread into a definite number of segments (Delage, 1899 and 1901).[5] The other regards chromosomes as independent units of the cell, retaining their identity between successive cell-divisions. The latter “Individualitäts Hypothese” was originally put forward by Theodor Boveri in 1887 as a result of C. Rabl’s observation (1885) that in epidermal cells of the salamander larva the chromosomes reappear in the mitosis of the daughter cells with the same arrangement as they possessed in the prophase of the mother cell—the angles of the U-shaped chromosomes being all directed towards one pole (Rabl’s “Poleseite”) of the nucleus. In the formation of the “resting” nucleus, the chromatin, becoming metabolically active, flows out on to the linin reticulum, all trace of the chromosomes being for the time lost. In Ascaris, Boveri (1888) obtained similar but still more striking results. The thickened ends of the four elongated chromosomes cause projections on the nuclear surface throughout the resting period, and the ends of the reappearing chromosomes always coincided with these protuberances; cf. also Sutton (1902) on locust spermatagonia. Moreover, the arrangement of the chromosomes must follow one of three well-marked groupings, and this is determined for each individual in the cleavage spindle of the egg and maintained throughout later development (fig. 8).
From Boveri’s Ergebnisse ü. d. Konstitution der chromatischen Substanz des Zellkerns, by permission of Gustav Fischer.
Fig. 8.—Preparation for Mitosis. a, Nucleus of “12 blastomere” of Ascaris megalocephala bivalens in resting condition; b and c, nuclei from sister 12 blastomeres in preparation for mitosis.
In the same worm (var. univalens) Boveri (1888 and 1899) found that occasional abnormalities in maturation resulted in the suppression of the first polar body and the inclusion of its chromosomes in the second maturation spindle; the egg-nucleus at the time of fertilization thus having two chromosomes instead of one, while the spermatozoon nucleus has only one. Three chromosomes instead of two reappear in subsequent divisions. Boveri’s “Individualitäts Hypothese” received striking support from the work of Herla (1893), L. R. Zoja (1895) and O. zur Strassen (1898). Herla and Zoja showed that if the egg of Ascaris megalocephala (var. bivalens), which possesses two chromosomes, be fertilized with the spermatozoon of var. univalens, in which the germ cell has only one chromosome and that smaller than either of the two in the other variety, three chromosomes reappear, two large and one small, in the cleavage divisions of the resulting hybrid embryo. Zur Strassen’s observations on the giant embryos of Ascaris also support Boveri’s theory. These embryos arise by the fusion of eggs, either before or after fertilization. The number of chromosomes in the subsequent cleavage-figures is proportional to the number of nuclei that have fused together. Similar results are given by Boveri’s (1893–1895) and T. H. Morgan’s (1895) experiments on the fertilization of enucleated sea-urchin egg-fragments; all the nuclei of the resulting embryo having only half the number of chromosomes characteristic of the species (e.g. in Echinus 9 instead of 18). All the above facts point to the conclusion that, as Boveri expressed it in his Grundgesetz der Zahlenkonstanz (1888), “the number of chromosomes arising from a resting nucleus is solely dependent on the number which originally entered into its composition.”[6]
- ↑ The force of this evidence is admitted by Boveri himself. Meves, however, maintains the possibility that the numerous centrosomes appearing in the egg arise by the rapid fragmentation of a centrosome already present.
- ↑ Cf. especially the behaviour of the centrosomes in the fertilization of the egg of Pleurophyllidia (MacFarland, 1897) and that of Cerebratulus (Coe, 1901). Not only may the sperm centrosomes totally disappear before reaching the egg-nucleus, but in the latter type the definitive centrosomes appear while the last traces of the sperm asters are still visible.
- ↑ e.g. Meves; Spermatagonia of Salamandra.
- ↑ Cf. especially the artificial production of amitosis in Spirogyra; W. Pfeffer, 1899.
- ↑ Cf. Boveri, 1904, p. 13. (For Boveri’s criticism of Delage’s views, cf. Boveri, 1901 and 1902.)
- ↑ It should, however, be noted that the assumption that a particular group of characters remains always associated in a particular chromosome is one that is very difficult to reconcile with the mode of inheritance of Mendelian pairs of characters in the case of organisms with a relatively small chromosome number.
