This memoir, written in 1904, clearly asserts the homopolarity of the centrosomes, and shows that the nuclear division is the result of a bipolar action, two poles of the same sign exerting their influence on opposite sides of the nucleus. It also emphasizes the important fact that diffusion,
and as far as we know diffusion alone, is able to produce a spindle between homologous poles.
A glance at the photograph is enough to show that the spindle is formed between poles of the same sign. The lines of diffusion radiate from one centre and converge towards the other centre in curves, giving the double convergence characteristic of a spindle. The central drop merely supplies the necessary material, and should have a concentration but slightly less than that of the plasma, so as not to set up its own lines of diffusion. The photograph shows clearly that the rays of the spindle traverse the equator without any break. It has been objected that these lines form not so much a spindle as two hemi-spindles, but it is clear that these two hemi-spindles are continuous and form a single sheaf of rays uniting the two poles of concentration. This is a phenomenon entirely unknown in the magnetic or electric fields, where two poles of the same sign, one on either side of a pole of the contrary sign, give two separate spindles. In a magnetic field it is impossible to make the lines emanating from one pole converge, except to a pole of opposite sign. Hence if we admit the homopolarity of the centrosomes, we must also admit that diffusion is the vera causa of karyokinesis, since, as I showed at the Grenoble Congress in 1904, diffusion and diffusion alone is capable of producing a spindle between two poles of the same sign.
Nuclear Division.—In order to reproduce artificially the phenomena attending the division of the nucleus, we may proceed as follows. We cover a perfectly horizontal glass plate with a semi-saturated solution of potassium nitrate to represent the cytoplasm of the cell. The nucleus in the centre is reproduced by a drop of the same solution coloured by a trace of Indian ink, the solid particles of which will represent the chromatin granules of the nucleus. The addition of the Indian ink will have slightly lowered the concentration of the central drop, and this is in accordance with nature, since the osmotic pressure of the nucleus is somewhat less than that of the plasma. We next place on either side of the drop which represents the
nucleus a coloured drop of solution more concentrated than the cytoplasm solution. The particles of Indian ink in the central drop arrange themselves in a long coloured ribbon, apparently rolled up in a coil, the edges of the ribbon having a beaded appearance. After a short time the ribbon loses its beaded appearance and becomes smooth, with a double outline, as is shown in A, Fig. 32. This coil or skein of ribbon subsequently divides, forming a nuclear spindle, while the chromatin substance collects together in the equatorial plane as in B, Fig. 32.
A more advanced stage of the nuclear division is shown at C, Fig. 32, where the chromatin bands of artificial chromosomes are grouped in two conical sheafs converging towards the two centrosomes. For some considerable time these conical bundles remain united by fine filaments, the last vestiges of the nuclear spindle. The final stage is that of two artificial cells in juxtaposition, whose nuclei are formed by the original centrosomes augmented by the chromatin bands or chromosomes (Fig. 32, D).
The resemblance of these successive phenomena to those of natural karyokinesis is of the closest. The experiment shows that diffusion is quite sufficient to produce organic karyokinesis, and that the only physical force required is that of osmotic pressure. If in the cytoplasm of a cell there are two points of molecular concentration greater than that of the general mass, the nucleus must necessarily divide with all the phenomena which accompany karyokinesis. In nature these two centres of positive concentration are introduced into the protoplasm of the cell by fecundation—that is, by the entrance of the centrosomes of the sperm cell. In certain abnormal cases the concentration may be produced in the cell itself by the formation of two centres of catabolism or molecular disintegration, since, as we have seen, molecular disintegration raises the osmotic pressure. This phenomenon, namely the production of karyokinesis from centres of catabolism, may account for the abnormal karyokinesis of cancer cells and the like. The subject is one which would well repay further investigation.
