Years ago Ischikawa observed that, in the conjugation of _Noctiluca_,
the nuclei of the two animals become closely apposed, but that they do
not fuse, although they behave like a single nucleus in the division
which follows. In this case _paternal and maternal nuclear substance
remain separate_ (Fig. 83, vol. i. p. 317). The same phenomenon has
since been repeatedly observed in many-celled animals, first by Häcker,
then by Rückert in the Copepods, and afterwards by Conklin in the eggs
of a Gastropod (_Crepidula_). But all these observations referred only
to the earlier stages of ovum-segmentation up to twenty-nine cells,
and it could not be affirmed that the distinctiveness of the paternal
and maternal chromosomes lasted till farther on into the ontogeny.
Professor Häcker now informs me, however, that he has been able to
trace this separateness in a Copepod (_Canthocamptus_) not only from
the beginning of segmentation on to the primitive genital cell, but
also through the divisions of this up to the mother-egg-cell[6].
Thus we may now assume that the paternal and maternal hereditary
bodies remain distinct, not only for a time, but throughout the whole
development, a fact which confirms our assumption of the independence
of the nuclear rods, notwithstanding their apparent breaking-up in
the nuclear reticulum of the 'resting' nucleus. This new knowledge
throws fresh light in another direction; it proves to us that the
remarkable and complicated processes which go on in the nucleus
during the maturing divisions have really the significance which I
long ago ascribed to them[7], that of effecting the maximum diversity
of intermingling of the paternal and maternal hereditary elements.
For Häcker has shown that during the second maturation-division
the paternal and maternal chromosomes are no longer united each
in a special group, but occur scattered about in the nucleus, and
subsequently come together again to form two differently combined
groups.
[6] Since this was written Häcker has published his results. See
_Anatom. Anzeiger_ xv (1902), p. 440.
[7] See my essay, _Amphimixis_, Jena, 1891.
If this were not so, if the maternal and paternal chromosomes remained
separate, then the reducing division would cause only one of these
groups to reach each of the germ-cells, and thus each mature ovum
or sperm-cell would contain either only paternal or only maternal
hereditary bodies. But this would make a reversion to more than three
generations back impossible, and as such reversions undoubtedly occur,
we must conclude that manifold new combinations of the paternal and
maternal chromosomes take place. This obviously happens during the
maturation-divisions, at least in the Metazoa.
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