This is as much as to say that, supposing each cell tends to divide into two halves when (and not before) its original size is doubled, then, in our flattened disc, the triangular cell T will tend to divide when the radius of the disc has increased by about a third (from 1 to 1·345), but the quadrilateral cell, Q, will not tend to divide until the linear dimensions of the disc have increased by about a half (from 1 to 1·517).
The case here illustrated is of no small general importance. For it shews us that a uniform and symmetrical growth of the organism (symmetrical, that is to say, under the limitations of a plane surface, or plane section) by no means involves a uniform or symmetrical growth of the individual cells, but may, under certain conditions, actually lead to inequality among these; and this inequality may be further emphasised by differences which arise out of it, in regard to the order of frequency of further subdivision. This phenomenon (or to be quite candid, this hypothesis, which is due to Berthold) is entirely independent of any change or variation in individual surface tensions; and accordingly it is essentially different from the phenomenon of unequal segmentation (as studied by Balfour), to which we have referred on p. [348].
In this fashion, we might go on to consider the manner, and the order of succession, in which the subsequent cell-divisions would tend to take place, as governed by the principle of minimal areas. But the calculations would grow more difficult, or the results got by simple methods would grow less and less exact. At the same time, some of these results would be of great interest, and well worth the trouble of obtaining. For instance, the precise manner in which our triangular cell, T, would next divide would be interesting to know, and a general solution of this problem is certainly troublesome to calculate. But in this particular case we can see that the width of the triangular cell near P is so obviously less than that near either of the other two angles, that a circular arc cutting off that angle is bound to be the shortest possible bisecting line; and that, in short, our triangular cell will tend to subdivide, just like the original quadrant, into a triangular and a quadrilateral portion.
But the case will be different next time, because in this new {370} triangle, PRQ, the least width is near the innermost angle, that at Q; and the bisecting circular arc will therefore be opposite to Q, or (approximately) parallel to PR. The importance of this fact is at once evident; for it means to say that there soon comes a time when, whether by the division of triangles or of quadrilaterals, we find only quadrilateral cells adjoining the periphery of our circular disc. In the subsequent division of these quadrilaterals, the partitions will arise transversely to their long axes, that is to say, radially (as U, V); and we shall consequently have a superficial or peripheral layer of quadrilateral cells, with sides approximately parallel, that is to say what we are accustomed to call an epidermis. And this epidermis or superficial layer will be in clear contrast with the more irregularly shaped cells, the products of triangles and quadrilaterals, which make up the deeper, underlying layers of tissue.
Fig. 152.
In following out these theoretic principles and others like to them, in the actual division of living cells, we must always bear in mind certain conditions and qualifications. In the first place, the law of minimal area and the other rules which we have arrived at are not absolute but relative: they are links, and very important links, in a chain of physical causation; they are always at work, but their effects may be overridden and concealed by the operation of other forces. Secondly, we must remember that, in the great majority of cases, the cell-system which we have in view is constantly increasing in magnitude by active growth; and by this means the form and also the proportions of the cells are continually liable to alteration, of which phenomenon we have already had an example. Thirdly, we must carefully remember that, until our cell-walls become absolutely solid and rigid, they are always apt to be modified in form owing to the tension of the adjacent {371} walls; and again, that so long as our partition films are fluid or semifluid, their points and lines of contact with one another may shift, like the shifting outlines of a system of soap-bubbles. This is the physical cause of the movements frequently seen among segmenting cells, like those to which Rauber called attention in the segmenting ovum of the frog, and like those more striking movements or accommodations which give rise to a so-called “spiral” type of segmentation.
Bearing in mind, then, these considerations, let us see what our flattened disc is likely to look like, after a few successive divisions
