The former problem is comparatively easy, as regards the tendency of a
segmentation cavity to _enlarge_, when once it has been established.
We may then assume that subdivision of the cells is due to the
appearance of a new-formed septum within each cell, that this septum
has a tendency to shrink under surface tension, and that these changes
will be accompanied on the whole by a diminution of surface energy
in the system. This being so, it may be shewn that the volume of the
divided cells must be less than it was prior to division, or in other
words that part of their contents must exude during the process of
segmentation[384]. Accordingly, the case where the segmentation cavity
enlarges and the embryo developes into a hollow blastosphere may, under
the circumstances, be simply described as the case where that outflow
or exudation from the cells of the blastoderm is directed on the whole
inwards.
The physical forces involved in the invagination of the cell-layer to
form the gastrula have been repeatedly discussed[385], but the true
explanation seems as yet to be by no means clear. The case, however,
is probably not a very difficult one, provided that we may assume a
difference of osmotic pressure at the two poles of the blastosphere,
that is to say between the cells which are being differentiated into
outer and inner, into epiblast and hypoblast. It is plain that a
blastosphere, or hollow vesicle bounded by a layer of vesicles, is
under very different physical conditions from a single, simple vesicle
or bubble. The blastosphere has no effective surface tension of its
own, such as to exert pressure on {345} its contents or bring the
whole into a spherical form; nor will local variations of surface
energy be directly capable of affecting the form of the system. But if
the substance of our blastosphere be sufficiently viscous, then osmotic
forces may set up currents which, reacting on the external fluid
pressure, may easily cause modifications of shape; and the particular
case of invagination itself will not be difficult to account for on
this assumption of non-uniform exudation and imbibition.
{346}
CHAPTER VIII
THE FORMS OF TISSUES OR CELL-AGGREGATES (_continued_)
The problems which we have been considering, and especially that of
the bee’s cell, belong to a class of “isoperimetrical” problems, which
deal with figures whose surface is a minimum for a definite content or
volume. Such problems soon become difficult, but we may find many easy
examples which lead us towards the explanation of biological phenomena;
and the particular subject which we shall find most easy of approach
is that of the division, in definite proportions, of some definite
portion of space, by a partition-wall of minimal area. The theoretical
principles so arrived at we shall then attempt to apply, after the
manner of Berthold and Errera, to the actual biological phenomena of
cell-division.
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