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Darwin and Modern Science [75]

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are doubled and this produces a band-like flattening and leads to the longitudinal splitting by which each chromosome is divided into two exactly equal halves. The nuclear membrane then disappears and fibrillar cell-plasma or cytoplasm invades the nuclear area. In animal cells these fibrillae in the cytoplasm centre on definite bodies (Their existence and their multiplication by fission were demonstrated by E. van Beneden and Th. Boveri in 1887.), which it is customary to speak of as Centrosomes. Radiating lines in the adjacent cell-plasma suggest that these bodies constitute centres of force. The cells of the higher plants do not possess such individualised centres; they have probably disappeared in the course of phylogenetic development: in spite of this, however, in the nuclear division-figures the fibrillae of the cell-plasma are seen to radiate from two opposite poles. In both animal and plant cells a fibrillar bipolar spindle is formed, the fibrillae of which grasp the longitudinally divided chromosomes from two opposite sides and arrange them on the equatorial plane of the spindle as the so- called nuclear or equatorial plate. Each half-chromosome is connected with one of the spindle poles only and is then drawn towards that pole. (These important facts, suspected by W. Flemming in 1882, were demonstrated by E. Heuser, L. Guignard, E. van Beneden, M. Nussbaum, and C. Rabl.)

The formation of the daughter-nuclei is then effected. The changes which the daughter-chromosomes undergo in the process of producing the daughter- nuclei repeat in the reverse order the changes which they went through in the course of their progressive differentiation from the mother-nucleus. The division of the cell-body is completed midway between the two daughter- nuclei. In animal cells, which possess no chemically differentiated membrane, separation is effected by simple constriction, while in the case of plant cells provided with a definite wall, the process begins with the formation of a cytoplasmic separating layer.

The phenomena observed in the course of the division of the nucleus show beyond doubt that an exact halving of its substance is of the greatest importance. (First shown by W. Roux in 1883.) Compared with the method of division of the nucleus, that of the cytoplasm appears to be very simple. This led to the conception that the cell-nucleus must be the chief if not the sole carrier of hereditary characters in the organism. It is for this reason that the detailed investigation of fertilisation phenomena immediately followed researches into the nucleus. The fundamental discovery of the union of two nuclei in the sexual act was then made (By O. Hertwig in 1875.) and this afforded a new support for the correct conception of the nuclear functions. The minute study of the behaviour of the other constituents of sexual cells during fertilisation led to the result, that the nucleus alone is concerned with handing on hereditary characters (This was done by O. Hertwig and the author of this essay simultaneously in 1884.) from one generation to another. Especially important, from the point of view of this conclusion, is the study of fertilisation in Angiosperms (Flowering plants); in these plants the male sexual cells lose their cell-body in the pollen-tube and the nucleus only-- the sperm-nucleus--reaches the egg. The cytoplasm of the male sexual cell is therefore not necessary to ensure a transference of hereditary characters from parents to offspring. I lay stress on the case of the Angiosperms because researches recently repeated with the help of the latest methods failed to obtain different results. As regards the descendants of angiospermous plants, the same laws of heredity hold good as for other sexually differentiated organisms; we may, therefore, extend to the latter what the Angiosperms so clearly teach us.

The next advance in the hitherto rapid progress in our knowledge of nuclear division was delayed, because it was not at once recognised that there are two absolutely different methods of nuclear
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