| Fig. 7.—Centrosomes. |
| From Prof. E. B. Wilson’s The Cell in Development and Inheritance, by permission of the author and of The Macmillan Co., New York. |
| a, Leucocyte from a Salamander, showing permanent aster and
centrosome. |
| From A. Gurwitsch, Morphologie u. Biologie der Zelle, by permission of Gustav Fischer. |
| b, Sperm-mother cell of Salamandra maculata, showing Hermann’s “central spindle.” |
The Achromatic Figure.—The mode of origin of the achromatic figure varies greatly. In some cases a distinct and continuous spindle, the “central spindle” of F. Hermann, is visible from the very first separation of the daughter centrosomes (e.g. salamander spermatogenic cell)[20] (fig. 7, b). In other cases the rays only invade the nuclear area and become continuous in the equatorial plane after the centrosomes have assumed their definitive positions at the two poles of the nucleus, and may even appear to indent the disappearing nuclear membrane as they invade the nuclear area.[21] In the salamander testis cell (fig. 7, b), and in many other cases, the whole of the achromatic figure is obviously of cytoplasmic origin. In many cases, however, it equally obviously arises within the nucleus,[22] while in yet other cases[23] the spindle fibres are of mixed origin. The question, therefore, of the cytoplasmic or nuclear origin of the achromatic figure, at one time regarded as of considerable importance, is wholly immaterial. Various elaborate theories have been propounded to explain the mechanism of the mitotic figure. H. Fol (1873) regarded the centrosomes as centres of attractive forces, and compared the mitotic figure to the lines of force in the magnetic field, a comparison made by numerous subsequent workers. E. Klein’s hypotheses of two opposing systems of contractile fibrillae, elaborated by van Beneden (1883, 1887) and accepted by Boveri (1888), was still further extended by R. Heidenhain in relation to the leucocytes of the salamander, in which there is a permanent centrosome and astral rays to which the contractile movements of the cell appear to be due[24] (fig. 7, a). Hermann on the other hand confined the contractility to the astral and mantle fibres; while L. Druner regarded the spindle as exerting a pushing force, for not only do the interzonal spindle fibres elongate during the anaphase, but they were often at this period contorted, while on the other hand astral rays may be entirely absent (e.g. Infusoria), and in some cases the spindle pole may be caused to project at the surface of the cell. The futility of these attempted mechanical explanations of mitosis is sufficiently clearly shown, not only by the contradictory nature of the explanations themselves, but by the fact that, in amitosis, nuclear and cytoplasmic division occur without any fibrillar mechanism whatever.
Centrosome.[25]—This minute body was first detected at the spindle poles by Flemming in 1875, and independently by P. J. van Beneden in 1876. The important part played by the centrosome in fertilization,[26] first described by van Beneden and Theodor Boveri in their papers of 1887-1888, together with the behaviour of this structure in mitosis, led these authors to regard the centrosome not only as the dynamic centre of the cell but as a permanent cell-organ, which, like the nucleus, passed by division from one cell-generation to the next. This conclusion appeared to receive considerable support from the recognition of the centrosome in various kinds of resting cells,[27] and especially from the relation this structure frequently shows to the locomotor apparatus of the cell (e.g. its position in the centre of the radiating fibrillae in the contractile lymph and pigment cells, and its relation to the vibratile flagellum in spermatozoa and some protozoa, e.g. Trypanosoma).[28] In almost all cases the centrosome of the resting cell, when this can be detected, lies in the cytoplasm, and is often already divided in preparation for the next mitotic division (e.g. spermatogenic cells of the salamander; Meves). In some cases, however, it resides in, or arises from, the nucleus (Brauer; spermatogenesis of Ascaris, var. univalens). This indifferent nuclear or cytoplasmic position for the centrosome is paralleled by the attraction sphere or homologue of the centrosome in many Protozoa. Thus in many forms, e.g. Euglena (Keuten), it lies within the nucleus, while in other forms, e.g. Noctiluca (Ishikawa, 1894, 1898; Calkins, 1898) and Paramoeba (F. Schaudinn, 1896), it lies in the cytoplasm, while in Tetramitus it coexists with a “distributed” nucleus. In the Heliozoa conditions are exceptionally interesting; not only is the centrosome—here resembling in appearance that of the higher forms—permanently visible and extranuclear, lying at the centre of the radiations characteristic of these forms, but there is the strongest possible evidence for its formation de novo. For Schaudinn has shown in Acanthocystis that, in the formation of the swarm spores, the nucleus divides amitotically, the centrosome remaining visible and unchanged at the centre of the radiating processes. Yet a centrosome appears later in the nucleus of the swarm spores and migrates into the cytoplasm. The experiments of T. H. Morgan and E. B. Wilson, in which numerous centrosomes and asters (“cytasters”) are caused to appear in unfertilized sea-urchin eggs by a brief immersion in a 13% solution of magnesium chloride in sea-water,[29] as also the possibility in many cases that even in normal fertilization the cleavage centrosomes may arise de novo,[30] 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[31] 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.[32] 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).[33] 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 “½ blastomere” of Ascaris megalocephala bivalens in resting condition; b and c, nuclei from sister ½ 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.”[34]
Boveri’s Law of Proportional Nuclear Growth.—The chromatin in the nucleus is exactly halved at every cell-division. As the bulk of the chromatin remains constant from one cell-generation to another, it must double its bulk between successive divisions. That this proportional growth of the chromatin is dependent solely on the chromatin mass, and not on that of the cell, is very clearly indicated by cases where the normal chromatin mass has been artificially increased or reduced,[35] the chromatin in either case doubling its bulk between successive cell-divisions, and neither the mass of the chromatin nor the number of the chromosomes undergoing any readjustment. By double or partial fertilization, different regions in the same embryo may show nuclei of different sizes (Boveri). We must therefore distinguish in the cell between “young” and “adult” chromatin. In other words the chromatin must be regarded as being composed of individual units, each with a definite constant structure and maximum growth (Boveri, 1904). This conclusion is strongly suggested, not only by the evidence in favour of the individuality of the chromosomes considered above, but also by the independent reproductive activity of the chromatin granules in the prophase of mitosis.
| From Boveri’s Ergebnisse ü. d. Konstitution der chromatischen Substanz des Zellkerns, by permission of Gustav Fischer. |
| Fig. 9.—Preparation for Mitosis. a, Spermatogonium of Brachystola magna with resting nucleus; b, Same with prophase for mitosis. (After Sutton.) |