Chapter 16

Organic selection has been termed a compromise between the more strictly Darwinian and the Lamarckian principles of interpretation. But it is not in any sense a compromise. The principle of interpretation of that which is instinctive and hereditary is wholly Darwinian. It is true that some of the facts of observation relied upon by Lamarckians are introduced. For Lamarckians however the modifications which are admittedly factors in survival, are regarded as the parents of inherited variations; for believers in organic selection they are only the foster-parents or nurses. It is because organic selection is the direct outcome of and a natural extension of Darwin's cardinal thesis that some reference to it here is justifiable. The matter may be put with the utmost brevity as follows: (1) Variations (V) occur, some of which are in the direction of increased adaptation (+), others in the direction of decreased adaptation (-).

(2) Acquired modifications (M) also occur. Some of these are in the direction of increased accommodation to circumstances (+), while others are in the direction of diminished accommodation (-). Four major combinations are

(_b_) + V with - M, (_c_) - V with + M,

(_a_) + V with + M, (_d_) - V with - M.

Of these (_d_) must inevitably be eliminated while (_a_) are selected. The predominant survival of (_a_) entails the survival of the adaptive variations which are inherited. The contributory acquisitions (+ M) are not inherited; but there are none the less factors in determining the survival of the coincident variations. It is surely abundantly clear that this is Darwinism and has no tincture of Lamarck's essential principle, the inheritance of acquired characters.

Whether Darwin himself would have accepted this interpretation of some at least of the evidence put forward by Lamarckians is unfortunately a matter of conjecture. The fact remains that in his interpretation of instinct and in allied questions he accepted the inheritance of individually acquired modifications of behaviour and structure.

Darwin was chiefly concerned with instinct from the biological rather than from the psychological point of view. Indeed it must be confessed that, from the latter standpoint, his conception of instinct as a "mental faculty" which "impels" an animal to the performance of certain actions, scarcely affords a satisfactory basis for genetic treatment. To carry out the spirit of Darwin's teaching it is necessary to link more closely biological and psychological evolution. The first step towards this is to interpret the phenomena of instinctive behaviour in terms of stimulation and response. It may be well to take a particular case. Swimming on the part of a duckling is, from the biological point of view, a typical example of instinctive behaviour. Gently lower a recently hatched bird into water: coordinated movements of the limbs follow in rhythmical sequence. The behaviour is new to the individual though it is no doubt closely related to that of walking, which is no less instinctive. There is a group of stimuli afforded by the "presentation" which results from partial immersion: upon this there follows as a complex response an application of the functional activities in swimming; the sequence of adaptive application on the appropriate presentation is determined by racial preparation. We know, it is true, but little of the physiological details of what takes place in the central nervous system; but in broad outline the nature of the organic mechanism and the manner of its functioning may at least be provisionally conjectured in the present state of physiological knowledge. Similarly in the case of the pecking of newly-hatched chicks; there is a visual presentation, there is probably a coöperating group of stimuli from the alimentary tract in need of food, there is an adaptive application of the activities in a definite mode of behaviour. Like data are afforded in a great number of cases of instinctive procedure, sometimes occurring very early in life, not infrequently deferred until the organism is more fully developed, but all of them dependent upon racial preparation. No doubt there is some range of variation in the behaviour, just such variation as the theory of natural selection demands. But there can be no question that the higher animals inherit a bodily organisation and a nervous system, the functional working of which gives rise to those inherited modes of behaviour which are termed instinctive.

It is to be noted that the term "instinctive" is here employed in the adjectival form as a descriptive heading under which may be grouped many and various modes of behaviour due to racial preparation. We speak of these as inherited; but in strictness what is transmitted through heredity is the complex of anatomical and physiological conditions under which, in appropriate circumstances, the organism so behaves. So far the term "instinctive" has a restricted biological connotation in terms of behaviour. But the connecting link between biological evolution and psychological evolution is to be sought,--as Darwin fully realised,--in the phenomena of instinct, broadly considered. The term "instinctive" has also a psychological connotation. What is that connotation?

Let us take the case of the swimming duckling or the pecking chick, and fix our attention on the first instinctive performance. Grant that just as there is, strictly speaking, no inherited behaviour, but only the conditions which render such behaviour under appropriate circumstances possible; so too there is no inherited experience, but only the conditions which render such experience possible; then the cerebral conditions in both cases are the same. The biological behaviour-complex, including the total stimulation and the total response with the intervening or resultant processes in the sensorium, is accompanied by an experience-complex including the initial stimulation-consciousness and resulting response-consciousness. In the experience-complex are comprised data which in psychological analysis are grouped under the headings of cognition, affective tone and conation. But the complex is probably experienced as an unanalysed whole. If then we use the term "instinctive" so as to comprise all congenital modes of behaviour which contribute to experience, we are in a position to grasp the view that the net result in consciousness constitutes what we may term the primary tissue of experience. To the development of this experience each instinctive act contributes. The nature and manner of organisation of this primary tissue of experience are dependent on inherited biological aptitudes; but they are from the outset onwards subject to secondary development dependent on acquired aptitudes. Biological values are supplemented by psychological values in terms of satisfaction or the reverse.

In our study of instinct we have to select some particular phase of animal behaviour and isolate it so far as is possible from the life of which it is a part. But the animal is a going concern, restlessly active in many ways. Many instinctive performances, as Darwin pointed out,[166] are serial in their nature. But the whole of active life is a serial and coordinated business. The particular instinctive performance is only an episode in a life-history, and every mode of behaviour is more or less closely correlated with other modes. This coordination of behaviour is accompanied by a correlation of the modes of primary experience. We may classify the instinctive modes of behaviour and their accompanying modes of instinctive experience under as many heads as may be convenient for our purposes of interpretation, and label them instincts of self-preservation, of pugnacity, of acquisition, the reproductive instincts, the parental instincts, and so forth. An instinct, in this sense of the term (for example the parental instinct), may be described as a specialised part of the primary tissue of experience differentiated in relation to some definite biological end. Under such an instinct will fall a large number of particular and often well-defined modes of behaviour, each with its own peculiar mode of experience.

It is no doubt exceedingly difficult as a matter of observation and of inference securely based thereon to distinguish what is primary from what is in part due to secondary acquisition--a fact which Darwin fully appreciated. Animals are educable in different degrees; but where they are educable they begin to profit by experience from the first. Only, therefore, on the occasion of the first instinctive act of a given type can the experience gained be regarded as _wholly_ primary; all subsequent performance is liable to be in some degree, sometimes more, sometimes less, modified by the acquired disposition which the initial behaviour engenders. But the early stages of acquisition are always along the lines predetermined by instinctive differentiation. It is the task of comparative psychology to distinguish the primary tissue of experience from its secondary and acquired modifications. We cannot follow up the matter in further detail. It must here suffice to suggest that this conception of instinct as a primary form of experience lends itself better to natural history treatment than Darwin's conception of an impelling force, and that it is in line with the main trend of Darwin's thought.

In a characteristic work,--characteristic in wealth of detail, in closeness and fidelity of observation, in breadth of outlook, in candour and modesty,--Darwin dealt with _The Expression of the Emotions in Man and Animals_. Sir Charles Bell in his _Anatomy of Expression_ had contended that many of man's facial muscles had been specially created for the sole purpose of being instrumental in the expression of his emotions. Darwin claimed that a natural explanation, consistent with the doctrine of evolution, could in many cases be given and would in other cases be afforded by an extension of the principles he advocated. "No doubt," he said,[167] "as long as man and all other animals are viewed as independent creations, an effectual stop is put to our natural desire to investigate as far as possible the causes of Expression. By this doctrine, anything and everything can be equally well explained.... With mankind, some expressions ... can hardly be understood, except on the belief that man once existed in a much lower and animal-like condition. The community of certain expressions in distinct though allied species ... is rendered somewhat more intelligible, if we believe in their descent from a common progenitor. He who admits on general grounds that the structure and habits of all animals have been gradually evolved, will look at the whole subject of Expression in a new and interesting light."

Darwin relied on three principles of explanation. "The first of these principles is, that movements which are serviceable in gratifying some desire, or in relieving some sensation, if often repeated, become so habitual that they are performed, whether or not of any service, whenever the same desire or sensation is felt, even in a very weak degree."[168] The modes of expression which fall under this head have become instinctive through the hereditary transmission of acquired habit. "As far as we can judge, only a few expressive movements are learnt by each individual; that is, were consciously and voluntarily performed during the early years of life for some definite object, or in imitation of others, and then became habitual. The far greater number of the movements of expression, and all the more important ones, are innate or inherited; and such cannot be said to depend on the will of the individual. Nevertheless, all those included under our first principle were at first voluntarily performed for a definite object,--namely, to escape some danger, to relieve some distress, or to gratify some desire."[169]

"Our second principle is that of antithesis. The habit of voluntarily performing opposite movements under opposite impulses has become firmly established in us by the practice of our whole lives. Hence, if certain actions have been regularly performed, in accordance with our first principle, under a certain frame of mind, there will be a strong and involuntary tendency to the performance of directly opposite actions, whether or not these are of any use, under the excitement of an opposite frame of mind."[170] This principle of antithesis has not been widely accepted. Nor is Darwin's own position easy to grasp.

"Our third principle," he says,[171] "is the direct action of the excited nervous system on the body, independently of the will, and independently, in large part, of habit. Experience shows that nerve-force is generated and set free whenever the cerebro-spinal system is excited. The direction which this nerve-force follows is necessarily determined by the lines of connection between the nerve-cells, with each other and with various parts of the body."

Lack of space prevents our following up the details of Darwin's treatment of expression. Whether we accept or do not accept his three principles of explanation we must regard his work as a masterpiece of descriptive analysis, packed full of observations possessing lasting value. For a further development of the subject it is essential that the instinctive factors in expression should be more fully distinguished from those which are individually acquired--a difficult task--and that the instinctive factors should be rediscussed in the light of modern doctrines of heredity, with a view to determining whether Lamarckian inheritance, on which Darwin so largely relied, is necessary for an interpretation of the facts.

The whole subject as Darwin realised is very complex. Even the term "expression" has a certain amount of ambiguity. When the emotion is in full flood, the animal fights, flees, or faints. Is this full-tide effect to be regarded as expression; or are we to restrict the term to the premonitory or residual effects--the bared canine when the fighting mood is being roused, the ruffled fur when reminiscent representations of the object inducing anger cross the mind? Broadly considered both should be included. The activity of premonitory expression as a means of communication was recognised by Darwin; he might, perhaps, have emphasised it more strongly in dealing with the lower animals. Man so largely relies on a special means of communication, that of language, that he sometimes fails to realise that for animals with their keen powers of perception, and dependent as they are on such means of communication, the more strictly biological means of expression are full of subtle suggestiveness. Many modes of expression, otherwise useless, are signs of behaviour that may be anticipated,--signs which stimulate the appropriate attitude of response. This would not, however, serve to account for the utility of the organic accompaniments--heart-affection, respiratory changes, vaso-motor effects and so forth, together with heightened muscular tone,--on all of which Darwin lays stress[172] under his third principle. The biological value of all this is, however, of great importance, though Darwin was hardly in a position to take it fully into account.

Having regard to the instinctive and hereditary factors of emotional expression we may ask whether Darwin's third principle does not alone suffice as an explanation. Whether we admit or reject Lamarckian inheritance it would appear that all hereditary expression must be due to pre-established connections within the central nervous system and to a transmitted provision for coordinated response under the appropriate stimulation. If this be so, Darwin's first and second principles are subordinate and ancillary to the third, an expression, so far as it is instinctive or heredity, being "the direct result of the constitution of the nervous system."

Darwin accepted the emotions themselves as hereditary or acquired states of mind and devoted his attention to their expression. But these emotions themselves are genetic products and as such dependent on organic conditions. It remained, therefore, for psychologists who accepted evolution and sought to build on biological foundations to trace the genesis of these modes of animal and human experience. The subject has been independently developed by Professors Lange and James;[173] and some modification of their view is regarded by many evolutionists as affording the best explanation of the facts. We must fix our attention on the lower emotions, such as anger or fear, and on their first occurrence in the life of the individual organism. It is a matter of observation that if a group of young birds which have been hatched in an incubator are frightened by an appropriate presentation, auditory or visual, they instinctively respond in special ways. If we speak of this response as the expression, we find that there are many factors. There are certain visible modes of behaviour, crouching at once, scattering and then crouching, remaining motionless, the braced muscles sustaining an attitude of arrest, and so forth, There are also certain visceral or organic effects, such as affections of the heart and respiration. These can be readily observed by taking the young bird in the hand. Other effects cannot be readily observed; vaso-motor changes, affections of the alimentary canal, the skin and so forth. Now the essence of the James-Lange view, as applied to these congenital effects, is that though we are justified in speaking of them as effects of the stimulation, we are not justified, without further evidence, in speaking of them as effects of the emotional state. May it not rather be that the emotion as a primary mode of experience is the concomitant of the net result of the organic situation--the initial presentation, the instinctive mode of behaviour, the visceral disturbances?

According to this interpretation the primary tissue of experience of the emotional order, felt as an unanalysed complex, is generated by the stimulation of the sensorium by afferent or incoming physiological impulses from the special senses, from the organs concerned in the responsive behaviour, from the viscere and vaso-motor system.

Some psychologists, however, contend that the emotional experience is generated in the sensorium prior to, and not subsequent to, the behaviour-response and the visceral disturbances. It is a direct and not an indirect outcome of the presentation to the special senses. Be this as it may, there is a growing tendency to bring into the closest possible relation, or even to identify, instinct and emotion in their primary genesis. The central core of all such interpretations is that instinctive behaviour and experience, its emotional accompaniments, and its expression, are but different aspects of the outcome of the same organic occurrences. Such emotions are, therefore, only a distinguishable aspect of the primary tissue of experience and exhibit a like differentiation. Here again a biological foundation is laid for a psychological doctrine of the mental development of the individual.

The intimate relation between emotion as a psychological mode of experience and expression as a group of organic conditions has an important bearing on biological interpretation. The emotion, as the psychological accompaniment of orderly disturbances in the central nervous system, profoundly influences behaviour and often renders it more vigourous and more effective. The utility of the emotions in the struggle for existence can, therefore, scarcely be over-estimated. Just as keenness of perception has survival-value; just as it is obviously subject to variation; just as it must be enhanced under natural selection, whether individually acquired increments are inherited or not; and just as its value lies not only in this or that special perceptive act but in its importance for life as a whole; so the vigourous effectiveness of activity has survival-value; it is subject to variation; it must be enhanced under natural selection; and its importance lies not only in particular modes of behaviour but in its value for life as a whole. If emotion and its expression as a congenital endowment are but different aspects of the same biological occurrence; and if this is a powerful supplement to vigour effectiveness and persistency of behaviour, it must on Darwin's principles be subject to natural selection.

If we include under the expression of the emotions not only the premonitory symptoms of the initial phases of the organic and mental state, not only the signs or conditions of half-tide emotion, but the full-tide manifestation of an emotion which dominates the situation, we are naturally led on to the consideration of many of the phenomena which are discussed under the head of sexual selection. The subject is difficult and complex, and it was treated by Darwin with all the strength he could summon to the task. It can only be dealt with here from a special point of view--that which may serve to illustrate the influence of certain mental factors on the course of evolution. From this point of view too much stress can scarcely be laid on the dominance of emotion during the period of courtship and pairing in the more highly organised animals. It is a period of maximum vigour, maximum activity, and, correlated with special modes of behaviour and special organic and visceral accompaniments, a period also of maximum emotional excitement. The combats of males, their dances and aerial evolutions, their elaborate behaviour and display, or the flood of song in birds, are emotional expressions which are at any rate coincident in time with sexual periodicity. From the combat of the males there follows on Darwin's principles the elimination of those which are deficient in bodily vigour, deficient in special structures, offensive or protective, which contribute to success, deficient in the emotional supplement of which persistent and whole-hearted fighting is the expression, and deficient in alertness and skill which are the outcome of the psychological development of the powers of perception. Few biologists question that we have here a mode of selection of much importance, though its influence on psychological evolution often fails to receive its due emphasis. Mr. Wallace[174] regards it as "a form of natural selection"; "to it," he says, "we must impute the development of the exceptional strength, size, and activity of the male, together with the possession of special offensive and defensive weapons, and of all other characters which arise from the development of these or are correlated with them." So far there is little disagreement among the followers of Darwin--for Mr. Wallace, with fine magnanimity, has always preferred to be ranked as such, notwithstanding his right, on which a smaller man would have constantly insisted, to the claim of independent originator of the doctrine of natural selection. So far with regard to sexual selection Darwin and Mr. Wallace are agreed; so far and no farther. For Darwin, says Mr. Wallace,[175] "has extended the principle into a totally different field of action, which has none of that character of constancy and of inevitable result that attaches to natural selection, including male rivalry; for by far the larger portion of the phenomena, which he endeavours to explain by the direct action of sexual selection, can only be so explained on the hypothesis that the immediate agency is female choice or preference. It is to this that he imputes the origin of all secondary sexual characters other than weapons of offence and defence.... In this extension of sexual selection to include the action of female choice or preference, and in the attempt to give to that choice such wide-reaching effects, I am unable to follow him more than a very little way."