The Evolution Theory, Vol. 2 of 2

by August Weismann

He considers that the results of use (that is, increased function) are generally transmitted, because the increase in the organ which is functioning more strongly changes the composition of the blood, by withdrawing from it in a greater degree the specific substances which the organ in question—a muscle, for instance—requires for its activity. All parts of the animal are thereby affected and modified, but especially those smallest vital units or 'fistellæ' (corresponding to biophors) which preside over digestion, and of which there are several sorts. Among them those work most arduously which have to produce the specific substances which serve for the nutrition of the muscles with increased function, because these are needed in larger quantities. This kind of digestive 'fistella' therefore multiplies, while other kinds, whose products are not required and therefore not used up, cease to be so active, diminish in number, and in course of time disappear. In this way the composition of the blood is altered, and with it to a greater or less degree all the characters of the whole organism. Of course the reproductive cells are also under the influence of this change in the composition of the blood, because the different nutritive substances are accumulated within them in an altered proportion corresponding to the changed composition of the blood, the nutritive substances for the muscles with increased function being contained in it in a larger quantity, and thus the greater development of the muscle will repeat itself in the progeny, that is to say, the acquired character is transmitted.

It is obvious that this is precisely the same line of argument as that used in reference to the origin of the worker and soldier ants. The different kinds of 'digestive fistellæ' correspond to the different food-carrying workers, and the blood to the assumed storehouse from which soldiers and workers select the food suitable for their respective needs, while the sex-cells in the one case, the sexual animals in the other, partake of all kinds exactly in the proportion in which they are stored, and thus the organ which functions most vigorously must be stronger in the offspring.

How the minute quantity of nutritive material contained in the ovum, still less in the sperm, is to effect the strengthening of the particular muscles in the descendants is not stated; moreover, such minimal quantities of food must soon be exhausted, and cannot possibly increase. It would seem as if the muscles could not even begin by being stronger, much less that they should remain so, if they were not exercised equally vigorously by the descendants. If the specific nutritive stuffs were 'fistellæ,' that is to say, were living units capable of multiplication, one could understand it. But there can, of course, be no possibility of a production of living units through digestion; that can only give rise to digested substances. Or if the alteration in the composition of the blood produced in the determinant system of the germ-plasm just those variations requisite to bring about a strengthening of the muscular system, it would remain to be shown how this could happen, for the gist of the problem lies in this. For muscles do not lie in the germ-plasm as miniature models of the subsequent muscular system, and even if they did, would not all the muscles, and not merely those which were no longer exercised, decrease hereditarily when a particular group, like the muscles of the ear in man, degenerates? Zehnder replies to this with the hypothesis that the muscles are not all chemically alike, but that each possesses a particular chemical formula, though they may all be very similar, and that, therefore, the nutritive materials required by each must be slightly different. In that case there would require to be, in the ovum and sperm of man, in order that functional modifications might be transmitted, as many special nutritive substances as there are muscles, and, in addition to these, innumerable hosts of other kinds of specific nutritive substances for all the other parts of the body, since all of them can be strengthened by exercise and weakened by disuse. And even if we suppose that all these millions of specific nutritive substances are accommodated within the germ-cells, as Zehnder's theory requires, they could not perform what Zehnder ascribes to them, for, as we have already said, they cannot multiply in the manner of living units, and so control the growing organism. The different specific nutritive materials contained in the blood are just as powerless to perform the task ascribed to them by Zehnder as the specific kinds of food in the hypothetical storehouse of the ants are to give rise to the different persons of the ant-colony.

Zehnder also attempts to overthrow the arguments against the Lamarckian principle which I based on the skeleton of Arthropods.

It does not seem to him probable that the chitinous coat of mail can be an absolutely dead structure, and he supposes that very delicate nerve-fibrils penetrate into all its most minute parts, and so are stimulated by 'every pressure and every strain' exerted on the chitinous skeleton. They 'work' when they are stimulated, and in doing so they use up 'their specific food-stuffs.' At places which are frequently stimulated the corresponding nerves develop more than elsewhere. The necessary specific food-stuffs for these particular nerves therefore increase proportionately within the body, and also in the reproductive cells. Accordingly, in the germ-cells there is an increase of the aforesaid nervous substances, which in the offspring become associated with the relevant part of the chitinous covering, and induce in development the secretion of chitin at this part. At this particular spot, then, the chitin will be specially thick.

This clearly implies that each particular part of the skin has its specific nutritive substances, necessitated by the nerves which traverse it! Thus there must be as many nerve-nutritive substances as there are skin-nerves, specific chemical combinations for every part of the body which is capable of heritable variation. This is so extraordinarily improbable that I need say nothing more about it. If the Lamarckian principle requires this kind of hypothesis to bolster it up, it is undoubtedly doomed.

If we disregard altogether the positive aspect of Zehnder's hypothesis, and assume that the skin-nerves are really stimulated through the chitin by every strain and pressure to which a spot of skin is exposed, and that they cause a correspondingly greater secretion of chitin, which would then, according to the Lamarckian principle, be hereditary, does this harmonize with what actually occurs in the development of the skeleton as we know it in the case of Insects and Crustaceans? Not at all! Can we suppose that the carapace of a crab or the enormously hard wing-covers of a water-beetle are exposed to a continual pounding and pressing and pushing? Exactly the contrary is the case. Every assailant takes care not to grasp the animal where it is so well protected, and seeks out the most vulnerable parts for its attacks. It may be answered that, while this is certainly the case now, the animals were badly protected when the ancestral forms were evolving. But that they could not have become hard by dint of being frequently bitten or otherwise wounded should be obvious from the fact that the whole of the wing-covers and the whole of the carapace is uniformly covered with thick chitin, while each wound would only stimulate particular spots; and we should also have to admit that, since these parts of the skin which are now so well protected are no longer seized and stimulated, they would long ago have become thin again, according to the principle of the degeneration of parts no longer used, or, in this case, no longer stimulated. But there is no need for wasting time over such quibbles, since there is a fact which absolutely contradicts Zehnder's hypothesis. I mean the degeneration of the chitinous skeleton in those Crustaceans and Insects which protect the abdomen within a shelter like the hermit-crabs, the caddis-flies (Phryganidæ), (Fig. 107) and the sack-carrying caterpillars of the Psychidæ among Lepidoptera. The hermit-crabs, as is well known, squeeze their abdomen into a usually spirally-coiled Gasteropod shell, and they always choose houses which are wide enough to conceal the whole body up to the hard claws when necessity arises. In this case there is surely a continual pressure on the abdomen, which, being soft, must be squeezed very tightly every time the animal retreats into its shell. One of my opponents has described the disappearance of the tough integumentary skeleton from the abdomen of these animals as an inherited result of this pressure, and another regards it as the inherited result of the degeneration of the muscles in this part of the body. But, according to Zehnder, this continuous pressure, and the frequent rubbing up and down of the abdomen on the inner surface of the Gasteropod's shell, would undoubtedly have a stimulating effect on the skin-nerves, and would therefore bring about a thickening of the chitinous cuticle. In regard to the larval Phryganidæ and Psychidæ, the case would be the same, though perhaps hardly to the same degree, for while these larvæ make their own houses, and will therefore at least make them big enough to begin with, the pressure and friction must increase with the growth of the animal.

Fig. 107. Larva of a Caddis-fly, after Rösel. A, removed from its case, showing the hooks (h) which attach it thereto, and the whitish abdomen, covered only by a thin cuticle. B, the same larva, moving about with its case.

If the regulation of the strength of the integumentary skeleton be referred to selection, we see at once why carapace and wing-covers should be of equal thickness throughout their whole extent, and why they do not disappear, although they do not function actively, and are less stimulated than any other parts of the skeleton; and we also understand why the abdomen of hermit-crabs and of larval Phryganidæ and Psychidæ has become soft, whether it be exposed to pressure or friction in a greater or a less degree. It no longer requires to be hard, because it is protected by the house, and in the case of the Pagurids it must not be hard because it could not then be readily squeezed into the hard-walled and narrow recesses of the Gasteropod shell; in this case there has therefore been positive selection. I have not yet referred to the fact that the chitinous covering is certainly not living, though it is not exactly dead; it is a secretion of the epidermic cells, not a tissue, and we cannot suppose that there are any nerve-endings in it. It is almost superfluous to say that the fact that the skin is cast is in itself enough to make such an assumption untenable, for the whole of the assumed delicate nervous network would be shed at every moult and torn away from the nerves which lead to it. As far as my knowledge goes, nothing of this kind occurs anywhere in the whole range of the animal kingdom.