We now know that they are cells, and that their origin depends on a
twice repeated division of the egg-cell; but it is a very unequal
division, for these 'directive cells' or 'polar bodies' are always much
smaller than the ovum, and indeed are usually so small that it is easy
to understand why their cellular nature was for so long overlooked.
Yet they have always a cell-body, and in many ova, for instance those
of certain marine Nudibranchs, this is quite considerable; and they
have likewise always a nucleus, which, notwithstanding the smallness of
the cell-body, is in all cases exactly of the same size as the sister
nucleus which remains behind in the ovum after division--a fact which
is in itself enough to indicate that we have here to do essentially
with readjustments and changes in the nucleus of the ovum.
Long before the polar or directive divisions were recognized as
divisions of the egg-cell it was known that the nucleus of the ovum
disappeared as soon as the latter attained to its full size within
the ovary. It was also known that this nucleus--the large so-called
'germinal vesicle' lying in the middle of the ovum--left its central
position and moved to the upper surface of the ovum, there to become
paler and paler, and ultimately to disappear altogether from the
sight of the observer. By many it was believed that it broke up, and
that the 'segmentation nucleus,' which is afterwards obvious, is a
new formation. The truth is that the germinal vesicle, at the time
of its disappearance, is transformed into a division figure which
is invisible without the aid of artificial staining. The nuclear
membrane breaks up; the centrosome of the ovum, which, although hardly
visible, had previously lain beside the germinal vesicle, divides
into two centrosomes and their centrospheres, and these now form the
'mitotic figure' by moving away from each other and sending out their
protoplasmic rays. This nuclear spindle soon ranges itself at right
angles to the surface of the egg, which at the same time arches itself
into a protuberance, and soon two daughter-nuclei are formed, one of
them lying within the protuberance (Fig. 75, _A_, _Rk1_). This soon
separates itself off from the ovum, surrounded by a small quantity
of cell-substance. The other daughter-nucleus remains within the
ovum, but neither of them remains in a state of rest; both are again
transformed into a spindle and divide once more; the minute first
'polar body' dividing into two 'secondary polar bodies' of half the
size (_B_, _Rk1_), while the nuclear spindle within the egg brings
about a second division of the ovum (_B_, _Rk2_) whose unequal products
are the second polar cell and the mature ovum--that is, the ovum ready
for fertilization. The process is now complete; the egg-cell, which
has lost very little plasmic material through the 'polar bodies' and
has not become visibly smaller, has now a nucleus (_B_, _Eik_) which
has become considerably smaller through the two rapidly successive
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