Chapter IV · INTELLIGENT BEHAVIOUR (continued 2)

Stated in terms of this conception, the familiar phenomena of mimicry are due to the fact that the mimicking form accords sufficiently well with the generic image to carry the same suggested meaning. As is well known, the model has been proved in many cases to be unpalatable or hurtful, while the mimic is in itself neither the one nor the other. The drone-fly, Eristalis, mimics the drone. And it has been urged that this cannot be a true case of mimicry, since the drone is harmless, though the female and “neuter” bees are possessed of stings. But I have satisfied myself by experiments with young birds, that (1) after experience with bees drones are avoided, and (2) that after similar experience drone-flies are also left untouched. Hence it seems that all three fall within the same generic image, the points of resemblance outweighing the differences in detail—as they do, indeed, with many men and women.

Such examples of mimicry belong to what is known as the “Batesian type”—so called after H. W. Bates, who, in 1861, discussed its occurrence among Amazonian insects in the light of the theory of natural selection. There are, however, certain groups of insects which, although themselves “protected,” possess common warning colours, causing them to resemble each other. These are sometimes classed under the head of “Müllerian mimicry”—so called after Fritz Müller, who, in 1879, first offered an explanation of the facts based on the theory of natural selection. He suggested that such mutual resemblance is advantageous to both protected forms, since it lessens the number of those which are killed by young birds and other animals while they are learning by experience what to eat and what to leave. For, as the result of careful observation, Mr. Frank Finn concludes “that each bird has to separately acquire its experience, and well remembers what it has learnt,”—a conclusion with which, as already stated, my own observations are entirely in accord. There is therefore a certain amount of destruction of even well-protected forms by young and inexperienced birds. If, then, two such forms resemble each other, the acquisition of experience is thereby facilitated and the amount of destruction reduced, on the assumption that the two fall within the same generic image. Upholders of natural selection are not, indeed, at one in accepting this explanation, and further observation is unquestionably needed. It is not improbable, however, that common protective coloration, such as the banding of yellow and black, seen in such different forms as the caterpillar of the cinnabar moth and the imago of the wasp, is of mutual utility. The following experiment was made with young chicks. Strips of orange and black paper were pasted beneath glass slips, and on them meal moistened with quinine was placed. On other plain slips meal moistened with water was provided. The young birds soon learnt to avoid the bitter meal, and then would not touch plain meal if it were offered on the banded slip. And these birds, save in two instances, refused to touch cinnabar caterpillars, which were new to their experience. They did not, like other birds, have to learn by particular trials that these caterpillars are unpleasant. Their experience had already been gained through the banded glass slips; or so it seemed. I have also found that young birds who had learnt to avoid cinnabar caterpillars left wasps untouched. Such observations must be repeated and extended. But they seem to show that one aspect of the Müllerian theory is not without some facts in support of it; and, so far as they go, they afford evidence that black and orange banding, irrespective of particular form, may constitute a guiding generic feature in the conscious situation.

It may be said that the generic condensation of experience here indicated implies the formation of general and abstract ideas, and that we cannot in face of the evidence accept Locke’s dictum that abstraction is “an excellency which the faculties of brutes do by no means attain to.” Romanes contended[72] that “all the higher animals have general ideas of ‘good-for-eating’ and ‘not-good-for-eating’ quite apart from any particular objects of which either of these qualities happens to be characteristic,” and he quoted with approval Leroy’s statement,[73] that a fox “will see snares when there are none; his imagination, distorted by fear, will produce deceptive shapes, to which he will attach an abstract notion of danger.” According to such views animals form concepts; and concepts belong to the sphere of rational thought. It is not my intention to enter at length into the refinements of psychological distinction. Many psychologists, however, seek to distinguish between, on the one hand, the predominance by natural emphasis, of certain qualities, such as that of being suitable for food, and, on the other hand, the intentional isolation of these qualities for the purposes of thought and rational explanation. Abstraction they regard as a deliberate process applied with rational intent to the material afforded by experience and reflection. Generalization, too, they regard as deliberate, and carried out with like intent. The result is not merely a composite or generic product, but something more subtle and less dependent on sense. “All trees hitherto seen by me,” said Noiré, “leave in my imagination a mixed image, a kind of ideal presentation of a tree. Quite different is my concept, which is never an image.” The concept “tree” is a deliberate synthesis of abstract qualities intentionally isolated, and recombined in accordance with the general relationships which subsist between them.

If we accept this distinction, if we regard abstraction and generalization as intentional mental processes carried out with the rational intent of discovering the relationships of phenomena with the object of explaining them and recombining their essential features in an ideal scheme of thought, we shall probably admit, with John Locke, that these are excellencies which the faculties of brutes do by no means attain to. But we shall none the less see that the predominance of certain salient features in experience by reiterated emphasis in association with natural needs, and the development of generic in place of merely particular re-presentations will afford the appropriate material for abstraction on the one hand, and generalization on the other. Intelligence supplies the embryonic mental structures from which, under the quickening influence of a rational purpose, abstract and general ideas may be evolved.

The essential features of the evolution of intelligence seem, then, to be, first, the development of controlling nerve-centres, by which the responsive action of reflex automatic or instinctive centres may be checked, augmented, or modified; secondly, the increased differentiation and integration of these control centres with extension of the range and complexity of experience in close touch with practical needs; thirdly, the condensation and concentration of experience by the formation of generic products through the reiterated emphasis begotten of recurrent situations having certain salient features in common, though differing in details; and fourthly, an increased plasticity of behaviour, especially in early life, enabling an animal to deal effectually with an environment far less simple than that to which the more stereotyped instinctive behaviour is fitted by inheritance to respond. And this evolution of intelligent behaviour is working its way up to, though as such it cannot reach, the succeeding phase of mental evolution in which the data, supplied by intelligence, are treated with a new purpose for higher ends in the rational thought which seeks to explain the phenomena, and frame an ideal scheme of their relations and interconnections.

Two further points may be noticed. First, that it is during the early and plastic days or months of life that intelligence is setting its seal on animal behaviour, and stamping it with its distinctive character. Adult life is very much what youth has made it; and old age is stereotyped through habit. In times of progress, the character of the race is determined by plastic possibilities of the young. Among them it is that the incidence of elimination makes itself felt, resulting in the survival of those whose intelligence can mould behaviour in accordance with the new circumstances of a wider life.

Secondly, this selection of the intelligent involves the survival of those in whose higher brain-centres there is room for a greater range and variety of interconnection by means of associating fibres. It involves a selective survival of the larger and more finely organized brains. It is probable, as Professor Ray Lankester has recently indicated, that the ridiculously small-brained mammals and reptiles of the past were creatures of instinct with little capacity for intelligent control. Their lives were simple, and their enemies and competitors no better provided with higher brain-centres than themselves. Stereotyped instinctive behaviour sufficed to enable them to hold their own, and meet the requirements of a life of dull and unprogressive monotony. Strength without cunning made these big-framed animals for a while masters of the situation. But among those existing animals whose skeletons indicate an analogous zoological position, there is none which exhibits a cerebral development so poor. And we may fairly conclude that the fact that these huge creatures have left no lineal descendants may be taken as evidence of the importance and value, in evolution, of that cerebral tissue which is the organic basis of intelligence. The higher brain contains the potentiality of that experience without which the evolution of intelligent behaviour in any race of vertebrate animals is impossible.

V.—The Influence of Intelligence on Instinct

We have seen that the relation of instinct to intelligence is essentially that of congenital to acquired behaviour. We have seen, too, that in the Lamarckian interpretation what is acquired in the course of life may be transmitted through inheritance, and thus the intelligent behaviour of one generation may become instinctive and congenital in the next. But serious biological difficulties stand in the way of the acceptance of this interpretation; there is, moreover, little or no evidence of the assumed transmission to offspring of any acquired modifications of structure or behaviour. We have, therefore, been led to infer that instinctive behaviour has been evolved through the selection of adaptive variations of germinal origin, the influence of intelligence being restricted to the fosterage of co-incident variations, that is to say, of those congenital variations which coincide in direction with the acquired modifications of behaviour due to intelligence. It is clear that on this interpretation the influence of intelligence on instinct is more indirect and less simple than that implied by the Lamarckian hypothesis. Intelligence and instinct are in large degree independent, though there is continual interaction between them. We have now to consider the nature of this interaction, and to this end we must indicate the relation of acquired modifications to the hereditary groundwork of the animal constitution.

The basal fact is, that the bodily tissues are subject to a certain amount of structural change during the course of individual life in accordance with the amount of functional strain put upon them. The labourer’s thickened skin, the enlarged and strengthened muscles of the athlete, the juggler’s acquired suppleness are familiar cases. Less familiar instances are afforded under abnormal conditions. Should one kidney from any cause be slowly destroyed, the other will slowly enlarge to carry on the increased work of elimination of waste products; when the larger shin bone of a dog has been removed after injury, the smaller bone becomes thickened to bear the added strain; new joint surfaces are sometimes formed where bones have been broken and the natural joints injured.

One may say that the normal development of any structure depends upon a due amount of use. But, since in the course of strenuous life any organ is from time to time subject to an abnormal amount of strain, it must be fitted to respond to a super-normal call on its strength and functional activity. Were the heart and the lungs, for example, unable to meet the greatly added drain on their energies, due to unwonted and severe exertion, collapse, perhaps death, would ensue if such exertion were imperatively demanded under special circumstances. And it is clear that many wild animals must be not infrequently placed in such circumstances as will subject their muscular structures and the functional activity of their organs of circulation and respiration to a strain nearly up to their extreme limits of endurance. The carnivorous hunter would often fail to secure his prey if his organization were unequal to a hard and prolonged chase; the hunted prey would not survive to procreate his kind if he fell a victim to the first pursuer through inability to stand the exertion necessary to enable him to make good his escape. It is thus, we may believe, through natural selection that a sufficiently high standard of strength and functional endurance is maintained. The failures in these respects are steadily eliminated. It is difficult to realize the great strain put upon a bird’s organization by the migration flight. Some ten times as many birds leave our shores in the autumn as return to them in the following spring. What proportion of these is weeded out in the act of migration we do not know; but we may be sure that only those fitted to stand a severe test of physical endurance return to rear broods which shall inherit in large degree similar vigour of constitution.

Two factors, then, determine the limits of efficiency in the bodily organs—heredity and use. And these two co-operate in such a way that we may say, either that due use is the essential condition of the effective development of the hereditary powers, or that heredity serves to condition their effective development through use. But though closely related, so that each may be regarded as conditional on the other, they are, if we accept the view that acquired characters are not transmitted as such, so far independent in that use adds nothing to, disuse subtracts nothing from, the hereditary store. It is, indeed, difficult to conceive how, on any view, the absence of the conditioning factor of normal use can be the efficient cause of a positive diminution of the balance at the bank of heredity. And Lamarckian thinkers have not succeeded in placing their conception of the matter in the clear light of a working hypothesis.

The amount of what we may term “modifiability” by use differs a good deal in the several organs and tissues. The teeth of carnivora and the antlers of deer may be cited as structures in which the conditioning effects of use form a relatively unimportant factor. On the other hand, the nervous system, with which we are here primarily concerned, is of all animal structures that in which what is acquired may attain the greatest importance in the successful conduct of life; the nature and the range of behaviour affording an index of the amount of modifiability in this respect.

We have already seen that instinctive behaviour is primarily a matter of the first occasion on which any given action is performed, and that many instinctive acts are subject to subsequent modification in the light of the experience gained during the early performances. The range of such modification varies both in different animals and also with respect to different modes of behaviour in the same animal. The more fixed and deeply rooted an instinct the less readily does intelligence obtain a hold on it, so as to direct the behaviour into new channels of better accommodation to the circumstances. M. Fabre describes how a Sphex, one of the solitary wasps, instinctively draws its prey, a grasshopper, into the burrow by its antennæ. When these were cut off the wasp pulled the grasshopper in by the jaw appendages; but when these were removed she seemed incapable of further accommodation to the unusual circumstances. It would seem an easy and obvious application of intelligence to seize the prey by one of the forelegs. But this was not done; and the grasshopper was then left. Intelligence did not seem equal to meeting the altered conditions presented by the maimed grasshopper. Still, there was some modification of the normal instinctive behaviour; and, as Dr. Peckham has shown, there may be more than Fabre noted. Let us assume the existence of an animal whose every act is instinctive, whose whole behaviour is marked out in strictly hereditary lines, no new departures being acquired in the course of individual life. This extreme case would afford an example of what we may term completely stereotyped behaviour. On the other hand, let us assume the existence of an animal with no hereditary definiteness of reaction, whose every act is intelligent, whose whole behaviour is the result of individual acquisition. This antithetical extreme case would afford an example of what we may term completely plastic behaviour. It is questionable, however, whether either of these extreme types occur in nature. What we find in our study of animal behaviour is some intermediate condition in which both factors co-operate, with a predominance either of stereotyped instinctive response on the one hand, or of plastic intelligent acquisition on the other hand. And in the latter case, as such behaviour approaches its ideal limits, we have modifiability under the circumstances of individual life at its maximum.

The evolution of intelligence as such runs parallel with the evolution of plastic behaviour; and this plasticity is necessitated by the variety and the complexity of the conditions of life—a variety and a complexity requiring many subtle modifications of response to enable the behaviour to reach accommodation to the changeful exigencies of diverse circumstances. To meet constant and relatively fixed conditions stereotyped instinctive responses suffice; and the elimination under natural selection of those individuals which fail to respond in fixed ways by specially adaptive behaviour tends to render definite the hereditary channels of nervous intercommunication. An inherited system of no little complexity may thus be evolved; of which we have seen examples in our study of instinctive behaviour. But the essential condition of the successful working of such a system is constancy in the environment to which the instinctive behaviour is adapted. Completely stereotyped behaviour, in its theoretical perfection, is in exact adaptation to the circumstances. Where instincts are only relatively perfect, further adaptation is secured through congenital variation and the survival of the individuals in which the behaviour is better adapted to the comparatively invariable circumstances. This is one line of evolution. But the evolution of intelligence is along independent lines of progress. Both, however, result from the functional activity of the same nervous system, they jointly determine the behaviour, they interact not only in the course of individual life but in the process of evolution, and they are both subject to the incidence of natural selection, which can determine whether the one line or the other shall preponderate—whether instinct or intelligence shall dominate behaviour.

If an answer must be given to the question whether instinct or intelligence has priority in the course of the evolution of behaviour, it may be urged that, on theoretical grounds, the claims of instinct are the stronger. Taking animals as we actually find them, however, they afford numberless examples of behaviour at first instinctive but subsequently modified, in greater or less degree, in accordance with the teachings of experience. Let us, first, assume that the environment is slowly changing, or has changed, in some definite manner. Such change would, of course, be relative, and might be due, either to new conditions brought to bear on the animal, or to the animal being itself brought, in the expansion of its life, within their influence. The old instinct is no longer quite adapted to the changed circumstances. If the change were sufficient in amount, and occurred somewhat suddenly, variations of instinct might not occur soon enough to enable the animal to reach adaptation by the gradual process of natural selection. If dependent on instinct alone the animal would, under these circumstances, be eliminated. But if intelligence were able to modify the behaviour to meet the new conditions this elimination would be prevented. In successive generations intelligence would constantly modify behaviour in the same manner and in a definite direction. Meanwhile congenital variations in different directions would occur. Those which were in directions antagonistic to that dictated by intelligence would tend to thwart accommodation and render it less effectual; but those which were coincident in direction would conspire with accommodation and render it more effectual. The individuals in which variations of instinct tended to thwart intelligence would be eliminated; while those in which coincident variations assisted and aided intelligent modification would survive. Thus intelligence would lead the way along lines which congenital variations would follow. And in the course of a number of generations the new instinct would reach the fully adaptive level, and further modification by intelligence would become unnecessary unless the environment continued to change yet more. Individual accommodation of behaviour would in this way determine the direction of instinctive variation; and yet throughout the process there would be, strictly speaking, no transmission of the intelligently acquired characters of the behaviour.

But though under constant and uniform changes in the environment the net result would be only a guided variation of the original instinct, under more variable and indefinitely changing circumstances the result would be different. The higher animals exhibit an intelligent plasticity which enables them to meet the requirements of the more complex environment into which their wider life has risen; for evolution lifts the animal from narrower into progressively wider spheres of activity and behaviour, so that its environment becomes relatively more complex. Here stereotyped behaviour would be rather a hindrance than an advantage. The winning animal in life’s struggle would be the one in which behaviour was most rapidly and most surely modified to meet particular needs—the one in which the teachings of experience were most promptly utilized in effective action. The inevitable tendency of the evolution of intelligence must be disintegration of the stereotyped modes of behaviour and the dissolution of instinct. Natural selection, which under a uniform and constant environment leads to the survival of relatively fixed and definite modes of response, under an environment presenting a wide range of possibilities leads to the survival of plastic accommodation through intelligence. It is not that intelligence has any direct influence tending to undermine the hereditary foundations of instinct, for acquired plasticity is not inherited as such; it is rather that when the stereotyped and the plastic are pitted against each other in the struggle for existence in the wider, freer, and more varied life of the higher animals the plastic survives and the stereotyped succumbs.

Imperfect as is our present knowledge of the manner in which the nervous connections implied in psychological associations are established, there can be no question that they are acquired in the course of individual life; they are modifications of nervous structure due to a special mode of use under the conditions of experience. Here, then, in the case of the nervous system, as in that of the bodily organs before mentioned, two co-operating factors determine the limits of efficiency—heredity and use. Just as the heart and lungs must inherit the power of standing abnormal strain if the animal is to avoid elimination in times of unwonted exertion, so must the nervous system inherit some reserve power of dealing effectively with unwonted circumstances by intelligent accommodation, if the animal is not to fall a victim to such circumstances. In other words, at times of heightened competition those animals which can draw on a reserve fund of intelligent accommodation will survive, while the stupid blunderers will be eliminated. We may term this reserve fund of intelligent accommodation, this inherited ability to meet specially difficult circumstances as they arise, innate capacity. From the nature of the case it must be indefinite, for it must carry with it the ability to meet unforeseen combinations of the environing forces by new combinations of the results of experience. Its distinguishing mark is plasticity, in contradistinction to the stereotyped fixity of typical instinct. And accompanying its evolution there is probably, as we have seen, a dissolution of its antithesis, instinct. Thus may we account for the fact that man, with his great store of innate capacity, has so small a number of stereotyped instincts.

But the dissolution of instincts is not complete. Residua are left in the inherited mental constitution. And these we term congenital tendencies and propensities. They differ from the typical instincts in the fact that the definiteness of response has been lost. They dictate a general trend of action, but the particular application in behaviour is due to intelligent accommodation. They are commonly spoken of as instinctive; and their mode of origin justifies the use of the adjective in association with the term “propensities.” But it must be remembered that the behaviour to which they lead is not, as such, wholly instinctive; it is a joint product of instinct and intelligence, the general trend being due to the instinctive propensity, while the mode of application is guided by intelligence.

There is, however, another way in which analogous propensities may be ingrained in the mental constitution, not as residual vestiges of old instincts, but as congenital rudiments fostered by new habits. It is a well-known and familiar fact that the frequent repetition of intelligent accommodation in certain definite lines begets habits, which so far simulate instincts as to be commonly described in popular speech as instinctive. Professor Wundt indeed places them in the category of “acquired instincts”—a usage which we regard as unsatisfactory, seeing that it tends to mask the distinction between the congenital and acquired factors in behaviour, and seeing that we have the well-defined term “habits” for acts rendered to a large extent automatic through repetition. Lamarckian thinkers regard habit as the mother of instinct, assuming that the acquired automatism of one generation may be transmitted to become congenital in the succeeding generation. This conclusion we provisionally reject, regarding the basal assumption as at present unproven. But though we cannot accept the view that habit is the mother of instinct, we regard it as not improbable that habit may be the nurse of congenital propensities. Remembering that similar habits are acquired by animals of the same species throughout a series of succeeding generations, and assuming that congenital variations are constantly occurring in many directions, it seems probable that some of these variations will be coincident in direction with the acquired habits. Thus would arise a congenital propensity to perform the habitual acts; and should they be of sufficient importance in the conduct of life to be subject to the action of natural selection, those animals in which such propensities were congenital would survive, whereas those in which no such propensities existed would be eliminated. It is unnecessary, however, to elaborate this conception further, since it is in line with that already discussed in considering the influence of intelligence in fostering a diversion of instinct under changing circumstances. Then we were considering how habit may lead to a congenital change in an old instinct; here we are dealing with the development of a new propensity.

Sufficient has now been said to illustrate some of the ways in which instinct and intelligence interact in the evolution of behaviour. Such interaction is further exemplified in the social life of animals, which will be dealt with in the next chapter.