The second example of a nascent reflex in dogs which I have to mention is as follows.
Goltz found that his brainless dogs, when wetted with water, would shake themselves as dry as possible, in just the same way as normal dogs will do under similar circumstances. This, of course, proves that the shaking movements may be performed by a reflex mechanism, which can have no other function to perform in the organization of a dog, and which, besides being of a highly elaborate character, will respond only to a very special kind of stimulation. Now, here also I find that the mechanism is congenital, or not acquired by individual experience. For the puppies on which I experimented were kept indoors from the time of their birth—so as never to have had any experience of being wetted by rain, &c.—till they were old enough to run about with a full power of co-ordinating their general movements. If these young animals were suddenly plunged into water, the shock proved too great: they would merely lie and shiver. But if their feet alone were wetted, by being dipped in a basin of water, the puppies would soon afterwards shake their heads in the peculiar manner which is required for shaking water off the ears, and which in adult dogs constitutes the first phase of a general shaking of the whole body.
Here, then, we seem to have good evidence of all the same facts which were presented in the case of the scratching reflex. In the first place, co-adaptation is present in a very high degree, because this shaking reflex in the dog, unlike the skin-twitching reflex in the horse, does not involve only a single muscle, or even a single group of muscles; it involves more or less the co-ordinated activity of many voluntary muscles all over the body. Such, at any rate, is the case when the action is performed by the intelligent volition of an adult dog; and if a brainless dog, or a young puppy, does not perform it so extensively or so vigorously, this only goes to prove that the reflex has not yet been sufficiently developed to serve as a substitute for intelligent volition—i.e. that it is useless, or a mere organic shadow of the really adaptive substance. Again, even if this nascent reflex had been so far developed as to have been capable of superseding voluntary action, still we may fairly doubt whether it could have proved of selective value. For it is questionable whether the immediate riddance of water after a wetting is a matter of life and death to dogs in a state of nature. Moreover, even if it were, every individual dog would always have got rid of the irritation, and so of the danger, by means of a voluntary shake—with the double result that natural selection has never had any opportunity of gradually building up a special reflex mechanism for the purpose of securing a shake, and that the canine race have not had to wait for any such unnecessary process. Lastly, such a process, besides being unnecessary, must surely have been, under any circumstances, impossible. For even if we were to suppose—again for the sake of saving an hypothesis at any cost—that the presence of a fully-formed shaking reflex is of selective value in the struggle for existence, it is perfectly certain that all the stages through which the construction of so elaborate a mechanism must have passed could not have been, under any circumstances, of any such value.
But, it is needless to repeat, according to the hypothesis of use-inheritance, there is no necessity to suppose that these incipient reflex mechanisms are of any value. If function produces structure in the race as it does in the individual, the voluntary and frequently repeated actions of scratching and shaking may very well have led to an organic integration of the neuro-muscular mechanisms concerned. Their various parts having been always co-ordinated for the performance of these actions by the intelligence of innumerable dogs in the past, their co-adapted activity in their now automatic responses to appropriate stimuli presents no difficulty. And the consideration that neither in their prospectively more fully developed condition, nor, a fortiori, in their present and all previous stages of evolution, can these reflex mechanisms be regarded as presenting any selective—or even so much as any adaptive—value, is neither more nor less than the theory of use-inheritance would expect.
Thus, with regard to the phenomena of reflex action in general, all the facts are such as this theory requires, while many of the facts are such as the theory of natural selection alone cannot conceivably explain. Indeed, it is scarcely too much to say, that most of the facts are such as directly contradict the latter theory in its application to them. But, be this as it may, at present there are only two hypotheses in the field whereby to account for the facts of adaptive evolution. One of these hypotheses is universally accepted, and the only question is whether we are to regard it as alone sufficient to explain all the facts. The other hypothesis having been questioned, we can test its validity only by finding cases which it is fully capable of explaining, and which do not admit of being explained by its companion hypothesis. I have endeavoured to show that we have a large class of such cases in the domain of reflex action, and shall next endeavour to show that there is another large class in the domain of instinct.
If instinct be, as Professor Hering, Mr. Samuel Butler, and others have argued, "hereditary habit"—i. e. if it comprises an element of transmitted experience—we at once find a complete explanation of many cases of the display of instinct which otherwise remain inexplicable. For although a large number—or even, as I believe, a large majority—of instincts are explicable by the theory of natural selection alone, or by supposing that they were gradually developed by the survival of fortuitous variations in the way of advantageous psychological peculiarities, this only applies to comparatively simple instincts, such as that of a protectively coloured animal exhibiting a preference for the surroundings which it resembles, or even adopting attitudes in imitation of objects which occur in such surroundings. But in all cases where instincts become complex and refined, we seem almost compelled to accept Darwin's view that their origin is to be sought in consciously intelligent adjustments on the part of ancestors.
Thus, to give only one example, a species of Sphex preys upon caterpillars, which it stings in their nerve-centres for the purpose of paralyzing, without killing them. The victims, when thus rendered motionless, are then buried with the eggs of the Sphex, in order to serve as food for her larvae which subsequently develop from these eggs. Now, in order thus to paralyze a caterpillar, the Sphex has to sting it successively in nine minute and particular points along the ventral surface of the animal—and this the Sphex unerringly does, to the exclusion of all other points of the caterpillar's anatomy. Well, such being the facts—according to M. Fabre, who appears to have observed them carefully—it is conceivable enough, as Darwin supposed[48], that the ancestors of the Sphex, being like many other hymenopterous insects highly intelligent, should have observed that on stinging caterpillars in these particular spots a greater amount of effect was produced than could be produced by stinging them anywhere else; and, therefore, that they habitually stung the caterpillars in these places only, till, in course of time, this originally intelligent habit became by heredity instinctive. But now, on the other hand, if we exclude the possibility of this explanation, it appears to me incredible that such an instinct should ever have been evolved at all; for it appears to me incredible that natural selection, unaided by originally intelligent action, could ever have developed such an instinct out of merely fortuitous variations—there being, by hypothesis, nothing to determine variations of an insect's mind in the direction of stinging caterpillars only in these nine intensely localized spots[49].
Again, there are not a few instincts which appear to be wholly useless to their possessors, and others again which appear to be even deleterious. The dusting over of their excrement by certain freely-roaming carnivora; the choice by certain herbivora of particular places on which to void their urine, or in which to die; the howling of wolves at the moon; purring of cats, &c., under pleasurable emotion; and sundry other hereditary actions of the same apparently unmeaning kind, all admit of being readily accounted for as useless habits originally acquired in various ways, and afterwards perpetuated by heredity, because not sufficiently deleterious to have been stamped out by natural selection[50]. But it does not seem possible to explain them by survival of the fittest in the struggle for existence.
Finally, in the case of our own species, it is self-evident that the aesthetic, moral, and religious instincts admit of a natural and easy explanation on the hypothesis of use-inheritance, while such is by no means the case if that hypothesis is rejected. Our emotions of the ludicrous, of the beautiful, and of the sublime, appear to be of the nature of hereditary instincts; and be this as it may, it would further appear that, whatever else they may be, they are certainly not of a life-preserving character. And although this cannot be said of the moral sense when the theory of natural selection is extended from the individual to the tribe, still, when we remember the extraordinary complexity and refinement to which they have attained in civilized man, we may well doubt whether they can have been due to natural selection alone. But space forbids discussion of this large and important question on the present occasion. Suffice it therefore to say, that I doubt not Weismann himself would be the first to allow that his theory of heredity encounters greater difficulties in the domain of ethics than in any other—unless, indeed, it be that of religion[51].