By C. Lloyd Morgan, LLD., F.R.S

In developing his conception of organic evolution Charles Darwin was of necessity brought into contact with some of the problems of mental evolution. In The Origin of Species he devoted a chapter to "the diversities of instinct and of the other mental faculties in animals of the same class."[153] When he passed to the detailed consideration of The Descent of Man, it was part of his object to show "that there is no fundamental difference between man and the higher mammals in their mental faculties."[154] "If no organic being excepting man," he said, "had possessed any mental power, or if his powers had been of a wholly different nature, from those of the lower animals, then we should never have been able to convince ourselves that our high faculties had been gradually developed."[155] In his discussion of The Expression of the Emotions it was important for his purpose "fully to recognise that actions readily become associated with other actions and with various states of the mind."[156] His hypothesis of sexual selection is largely dependent upon the exercise of choice on the part of the female and her preference for "not only the more attractive but at the same time the more vigourous and vicious males."[157] Mental processes and physiological processes were for Darwin closely correlated; and he accepted the conclusion "that the nervous system not only regulates most of the existing functions of the body, but has indirectly influenced the progressive development of various bodily structures and of certain mental qualities."[158]

Throughout his treatment, mental evolution was for Darwin incidental to and contributory to organic evolution. For specialised research in comparative and genetic psychology, as an independent field of investigation, he had neither the time nor the requisite training. None the less his writings and the spirit of his work have exercised a profound influence on this department of evolutionary thought. And, for those who follow Darwin's lead, mental evolution is still in a measure subservient to organic evolution. Mental processes are the accompaniments or concomitants of the functional activity of specially differentiated parts of the organism. They are in some way dependent on physiological and physical conditions. But though they are not physical in their nature, and though it is difficult or impossible to conceive that they are physical in their origin, they are, for Darwin and his followers, factors in the evolutionary process in its physical or organic aspect. By the physiologist within his special and well-defined universe of discourse they may be properly regarded as epiphenomena; but by the naturalist in his more catholic survey of nature they cannot be so regarded, and were not so regarded by Darwin. Intelligence has contributed to evolution of which it is in a sense a product.

The facts of observation or of inference which Darwin accepted are these: Conscious experience accompanies some of the modes of animal behaviour; it is concomitant with certain physiological processes; these processes are the outcome of development in the individual and evolution in the race; the accompanying mental processes undergo a like development. Into the subtle philosophical questions which arise out of the naÏve acceptance of such a creed it was not Darwin's province to enter; "I have nothing to do," he said,[159] "with the origin of the mental powers, any more than I have with that of life itself." He dealt with the natural history of organisms, including not only their structure but their modes of behaviour; with the natural history of the states of consciousness which accompany some of their actions; and with the relation of behaviour to experience. We will endeavour to follow Darwin in his modesty and candour in making no pretence to give ultimate explanations. But we must note one of the implications of this self-denying ordinance of science. Development and evolution imply continuity. For Darwin and his followers the continuity is organic through physical heredity. Apart from speculative hypothesis, legitimate enough in its proper place but here out of court, we know nothing of continuity of mental evolution as such: consciousness appears afresh in each succeeding generation. Hence it is that for those who follow Darwin's lead, mental evolution is and must ever be, within his universe of discourse, subservient to organic evolution. Only in so far as conscious experience, or its neural correlate, effects some changes in organic structure can it influence the course of heredity; and conversely only in so far as changes in organic structure are transmitted through heredity, is mental evolution rendered possible. Such is the logical outcome of Darwin's teaching.

Those who abide by the cardinal results of this teaching are bound to regard all behaviour as the expression of the functional activities of the living tissues of the organism, and all conscious experience as correlated with such activities. For the purposes of scientific treatment, mental processes are one mode of expression of the same changes of which the physiological processes accompanying behaviour are another mode of expression. This is simply accepted as a fact which others may seek to explain. The behaviour itself is the adaptive application of the energies of the organism; it is called forth by some form of presentation or stimulation brought to bear on the organism by the environment. This presentation is always an individual or personal matter. But in order that the organism may be fitted to respond to the presentation of the environment it must have undergone in some way a suitable preparation. According to the theory of evolution this preparation is primarily racial and is transmitted through heredity. Darwin's main thesis was that the method of preparation is predominantly by natural selection. Subordinate to racial preparation, and always dependent thereon, is individual or personal preparation through some kind of acquisition; of which the guidance of behaviour through individually won experience is a typical example. We here introduce the mental factor because the facts seem to justify the inference. Thus there are some modes of behaviour which are wholly and solely dependent upon inherited racial preparation; there are other modes of behaviour which are also dependent, in part at least, on individual preparation. In the former case the behaviour is adaptive on the first occurrence of the appropriate presentation; in the latter case accommodation to circumstances is only reached after a greater or less amount of acquired organic modification of structure, often accompanied (as we assume) in the higher animals by acquired experience. Logically and biologically the two classes of behaviour are clearly distinguishable: but the analysis of complex cases of behaviour where the two factors coÖperate, is difficult and requires careful and critical study of life-history.

The foundations of the mental life are laid in the conscious experience that accompanies those modes of behaviour, dependent entirely on racial preparation, which may broadly be described as instinctive. In the eighth chapter of The Origin of Species Darwin says,[160] "I will not attempt any definition of instinct.... Every one understands what is meant, when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds' nests. An action, which we ourselves require experience to enable us to perform, when performed by an animal, more especially by a very young one, without experience, and when performed by many individuals in the same way, without their knowing for what purpose it is performed, is usually said to be instinctive." And in the summary at the close of the chapter he says,[161] "I have endeavoured briefly to show that the mental qualities of our domestic animals vary, and that the variations are inherited. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. No one will dispute that instincts are of the highest importance to each animal. Therefore there is no real difficulty, under changing conditions of life, in natural selection accumulating to any extent slight modifications of instinct which are in any way useful. In many cases habit or use and disuse have probably come into play."

Into the details of Darwin's treatment there is neither space nor need to enter. There are some ambiguous passages; but it may be said that for him, as for his followers to-day, instinctive behaviour is wholly the result of racial preparation transmitted through organic heredity. For the performance of the instinctive act no individual preparation under the guidance of personal experience is necessary. It is true that Darwin quotes with approval Huber's saying that "a little dose of judgment or reason often comes into play, even with animals low in the scale of nature."[162] But we may fairly interpret his meaning to be that in behaviour, which is commonly called instinctive, some element of intelligent guidance is often combined. If this be conceded the strictly instinctive performance (or part of the performance) is the outcome of heredity and due to the direct transmission of parental or ancestral aptitudes. Hence the instinctive response as such depends entirely on how the nervous mechanism has been built up through heredity; while intelligent behaviour, or the intelligent factor in behaviour, depends also on how the nervous mechanism has been modified and moulded by use during its development and concurrently with the growth of individual experience in the customary situations of daily life. Of course it is essential to the Darwinian thesis that what Sir E. Ray Lankester has termed "educability," not less than instinct, is hereditary. But it is also essential to the understanding of this thesis that the differentiae of the hereditary factor should be clearly grasped.

For Darwin there were two modes of racial preparation, (1) natural selection, and (2) the establishment of individually acquired habit. He showed that instincts are subject to hereditary variation; he saw that instincts are also subject to modification through acquisition in the course of individual life. He believed that not only the variations but also, to some extent, the modifications are inherited. He therefore held that some instincts (the greater number) are due to natural selection but that others (less numerous) are due, or partly due, to the inheritance of acquired habits. The latter involve Lamarckian inheritance, which of late years has been the centre of so much controversy. It is noteworthy however that Darwin laid especial emphasis on the fact that many of the most typical and also the most complex instincts—those of neuter insects—do not admit of such an interpretation. "I am surprised," he says,[163] "that no one has hitherto advanced this demonstrative case of neuter insects, against the well-known doctrine of inherited habit, as advanced by Lamarck." None the less Darwin admitted this doctrine as supplementary to that which was more distinctively his own—for example in the case of the instincts of domesticated animals. Still, even in such cases, "it may be doubted," he says,[164] "whether any one would have thought of training a dog to point, had not some one dog naturally shown a tendency in this line ... so that habit and some degree of selection have probably concurred in civilising by inheritance our dogs." But in the interpretation of the instincts of domesticated animals, a more recently suggested hypothesis, that of organic selection,[165] may be helpful. According to this hypothesis any intelligent modification of behaviour which is subject to selection is probably coincident in direction with an inherited tendency to behave in this fashion. Hence in such behaviour there are two factors: (1) an incipient variation in the line of such behaviour, and (2) an acquired modification by which the behaviour is carried further along the same line. Under natural selection those organisms in which the two factors coÖperate are likely to survive. Under artificial selection they are deliberately chosen out from among the rest.

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

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."

Into the details of Mr. Wallace's criticisms it is impossible to enter here. We cannot discuss either the mode of origin of the variations in structure which have rendered secondary sexual characters possible or the modes of selection other than sexual which have rendered them, within narrow limits, specifically constant. Mendelism and mutation theories may have something to say on the subject when these theories have been more fully correlated with the basal principles of selection. It is noteworthy that Mr. Wallace says:[176] "Besides the acquisition of weapons by the male for the purpose of fighting with other males, there are some other sexual characters which may have been produced by natural selection. Such are the various sounds and odours which are peculiar to the male, and which serve as a call to the female or as an indication of his presence. These are evidently a valuable addition to the means of recognition of the two sexes, and are a further indication, that the pairing season has arrived; and the production, intensification, and differentiation of these sounds and odours are clearly within the power of natural selection. The same remark will apply to the peculiar calls of birds, and even to the singing of the males." Why the same remark should not apply to their colours and adornments is not obvious. What is obvious is that "means of recognition" and "indication that the pairing season has arrived" are dependent on the perceptive powers of the female who recognises and for whom the indication has meaning. The hypothesis of female preference, stripped of the aesthetic surplusage which is psychologically both unnecessary and unproven, is really only different in degree from that which Mr. Wallace admits in principle when he says that it is probable that the female is pleased or excited by the display.

Let us for our present purpose leave on one side and regard as sub judice the question whether the specific details of secondary sexual characters are the outcome of female choice. For us the question is whether certain psychological accompaniments of the pairing situation have influenced the course of evolution and whether these psychological accompaniments are themselves the outcome of evolution. As a matter of observation, specially differentiated modes of behaviour, often very elaborate, frequently requiring highly developed skill, and apparently highly charged with emotional tone, are the precursors of pairing. They are generally confined to the males, whose fierce combats during the period of sexual activity are part of the emotional manifestation. It is inconceivable that they have no biological meaning; and it is difficult to conceive that they have any other biological end than to evoke in the generally more passive female the pairing impulse. They, are based on instinctive foundations ingrained in the nervous constitution through natural (or may we not say sexual?) selection in virtue of their profound utility. They are called into play by a specialised presentation such as the sight or the scent of the female at, or a little in advance of, a critical period of the physiological rhythm. There is no necessity that the male should have any knowledge of the end to which his strenuous activity leads up. In presence of the female there is an elaborate application of all the energies of behaviour, just because ages of racial preparation have made him biologically and emotionally what he is—a functionally sexual male that must dance or sing or go through hereditary movements of display, when the appropriate stimulation comes. Of course after the first successful courtship his future behaviour will be in some degree modified by his previous experience. No doubt during his first courtship he is gaining the primary data of a peculiarly rich experience, instinctive and emotional. But the biological foundations of the behaviour of courtship are laid in the hereditary coordinations. It would seem that in some cases, not indeed in all, perhaps especially in those cases in which secondary sexual behaviour is most highly evolved,—correlative with the ardour of the male is a certain amount of reluctance in the female. The pairing act on her part only takes place after prolonged stimulation, for affording which the behaviour of male courtship is the requisite presentation. The most vigourous, defiant and mettlesome male is preferred just because he alone affords a contributory stimulation adequate to evoke the pairing impulse with its attendant emotional tone.

It is true that this places female preference or choice on a much lower psychological plane than Darwin in some passages seems to contemplate where, for example, he says that the female appreciates the display of the male and places to her credit a taste for the beautiful. But Darwin himself distinctly states[177] that "it is not probable that she consciously deliberates; but she is most excited or attracted by the most beautiful, or melodious, or gallant males." The view here put forward, which has been developed by Prof. Groos,[178] therefore seems to have Darwin's own sanction. The phenomena are not only biological; there are psychological elements as well. One can hardly suppose that the female is unconscious of the male's presence; the final yielding must surely be accompanied by heightened emotional tone. Whether we call it choice or not is merely a matter of definition of terms. The behaviour is in part determined by supplementary psychological values. Prof. Groos regards the coyness of females as "a most efficient means of preventing the too early and too frequent yielding to the sexual impulse."[179] Be that as it may, it is, in any case, if we grant the facts, a means through which male sexual behaviour with all its biological and psychological implications, is raised to a level otherwise perhaps unattainable by natural means, while in the female it affords opportunities for the development in the individual and evolution in the race of what we may follow Darwin in calling appreciation, if we empty this word of the aesthetic implications which have gathered round it in the mental life of man.

Regarded from this standpoint of sexual selection, broadly considered, has probably been of great importance. The psychological accompaniments of the pairing situation have profoundly influenced the course of biological evolution and are themselves the outcome of that evolution.

Darwin makes only passing reference to those modes of behaviour in animals which go by the name of play. "Nothing," he says,[180] "is more common than for animals to take pleasure in practising whatever instinct they follow at other times for some real good." This is one of the very numerous cases in which a hint of the master has served to stimulate research in his disciples. It was left to Prof. Groos to develop this subject on evolutionary lines and to elaborate in a masterly manner Darwin's suggestion. "The utility of play," he says,[181] "is incalculable. This utility consists in the practice and exercise it affords for some of the more important duties of life,"—that is to say, for the performance of activities which will in adult life be essential to survival. He urges[182] that "the play of young animals has its origin in the fact that certain very important instincts appear at a time when the animal does not seriously need them." It is, however, questionable whether any instincts appear at a time when they are not needed. And it is questionable whether the instinctive and emotional attitude of the play-fight, to take one example, can be identified with those which accompany fighting in earnest, though no doubt they are closely related and have some common factors. It is probable that play, as preparatory behaviour, differs in biological detail (as it almost certainly does in emotional attributes) from the earnest of after-life and that it has been evolved through differentiation and integration of the primary tissue of experience, as a preparation through which certain essential modes of skill may be acquired—those animals in which the preparatory play-propensity was not inherited in due force and requisite amount being subsequently eliminated in the struggle for existence. In any case there is little question that Prof. Groos is right in basing the play-propensity on instinctive foundations.[183] None the less, as he contends, the essential biological value of play is that it is a means of training the educable nerve-tissue, of developing that part of the brain which is modified by experience and which thus acquires new characters, of elaborating the secondary tissue of experience on the predetermined lines of instinctive differentiation and thus furthering the psychological activities which are included under the comprehensive term "intelligent."

In The Descent of Man Darwin dealt at some length with intelligence and the higher mental faculties.[184] His object, he says, is to show that there is no fundamental difference between man and the higher mammals in their mental faculties; that these faculties are variable and the variations tend to be inherited; and that under natural selection beneficial variations of all kinds will have been preserved and injurious ones eliminated.

Darwin was too good an observer and too honest a man to minimise the "enormous difference" between the level of mental attainment of civilised man and that reached by any animal. His contention was that the difference, great as it is, is one of degree and not of kind. He realised that, in the development of the mental faculties of man, new factors in evolution have supervened—factors which play but a subordinate and subsidiary part in animal intelligence. Intercommunication by means of language, approbation and blame, and all that arises out of reflective thought, are but foreshadowed in the mental life of animals. Still he contends that these may be explained on the doctrine of evolution. He urges[185] "that man is variable in body and mind; and that the variations are induced, either directly or indirectly, by the same general causes, and obey the same general laws, as with the lower animals." He correlates mental development with the evolution of the brain.[186] "As the various mental faculties gradually developed themselves, the brain would almost certainly become larger. No one, I presume, doubts that the large proportion which the size of man's brain bears to his body, compared to the same proportion in the gorilla or orang, is closely connected with his higher mental powers." "With respect to the lower animals," he says,[187] "M. E. Lartet,[188] by comparing the crania of tertiary and recent mammals belonging to the same groups, has come to the remarkable conclusion that the brain is generally larger and the convolutions are more complex in the more recent form."

Sir E. Ray Lankester has sought to express in the simplest terms the implications of the increase in size of the cerebrum. "In what," he asks, "does the advantage of a larger cerebral mass consist?" "Man," he replies, "is born with fewer ready-made tricks of the nerve-centres—these performances of an inherited nervous mechanism so often called by the ill-defined term 'instincts'—than are the monkeys or any other animal. Correlated with the absence of inherited ready-made mechanism, man has a greater capacity of developing in the course of his individual growth similar nervous mechanisms (similar to but not identical with those of 'instinct') than any other animal.... The power of being educated—'educability' as we may term it—is what man possesses in excess as compared with the apes. I think we are justified in forming the hypothesis that it is this 'educability' which is the correlative of the increased size of the cerebrum." There has been natural selection of the more educable animals, for "the character which we describe as 'educability' can be transmitted, it is a congenital character. But the results of education can not be transmitted. In each generation they have to be acquired afresh, and with increased 'educability' they are more readily acquired and a larger variety of them.... The fact is that there is no community between the mechanisms of instinct and the mechanisms of intelligence, and that the latter are later in the history of the evolution of the brain than the former and can only develop in proportion as the former become feeble and defective."[189]

In this statement we have a good example of the further development of views which Darwin foreshadowed but did not thoroughly work out. It states the biological case clearly and tersely. Plasticity of behaviour in special accommodation to special circumstances is of survival value; it depends upon acquired characters; it is correlated with increase in size and complexity of the cerebrum; under natural selection therefore the larger and more complex cerebrum as the organ of plastic behaviour has been the outcome of natural selection. We have thus the biological foundations for a further development of genetic psychology.

There are diversities of opinion, as Darwin showed, with regard to the range of instinct in man and the higher animals as contrasted with lower types. Darwin himself said[190] that "Man, perhaps, has somewhat fewer instincts than those possessed by the animals which come next to him in the series." On the other hand, Prof. Wm. James says[191] that man is probably the animal with most instincts. The true position is that man and the higher animals have fewer complete and self-sufficing instincts than those which stand lower in the scale of mental evolution, but that they have an equally large or perhaps larger mass of instinctive raw material which may furnish the stuff to be elaborated by intelligent processes. There is, perhaps, a greater abundance of the primary tissue of experience to be refashioned and integrated by secondary modification; there is probably the same differentiation in relation to the determining biological ends, but there is at the outset less differentiation of the particular and specific modes of behaviour. The specialised instinctive performances and their concomitant experience-complexes are at the outset more indefinite. Only through acquired connections, correlated with experience, do they become definitely organised.

The full working-out of the delicate and subtle relationship of instinct and educability—that is, of the hereditary and the acquired factors in the mental life—is the task which lies before genetic and comparative psychology. They interact throughout the whole of life, and their interactions are very complex. No one can read the chapters of The Descent of Man which Darwin devotes to a consideration of the mental characters of man and animals without noticing, on the one hand, how sedulous he is in his search for hereditary foundations, and, on the other hand, how fully he realises the importance of acquired habits of mind. The fact that educability itself has innate tendencies—is in fact a partially differentiated educability—renders the unravelling of the factors of mental progress all the more difficult.

In his comparison of the mental powers of men and animals it was essential that Darwin should lay stress on points of similarity rather than on points of difference. Seeking to establish a doctrine of evolution, with its basal concept of continuity of process and community of character, he was bound to render clear and to emphasise the contention that the difference in mind between man and the higher animals, great as it is, is one of degree and not of kind. To this end Darwin not only recorded a large number of valuable observations of his own, and collected a considerable body of information from reliable sources, he presented the whole subject in a new light and showed that a natural history of mind might be written and that this method of study offered a wide and rich field for investigation. Of course those who regarded the study of mind only as a branch of metaphysics smiled at the philosophical ineptitude of the mere man of science. But the investigation, on natural history lines, has been prosecuted with a large measure of success. Much indeed still remains to be done; for special training is required, and the workers are still few. Promise for the future is however afforded by the fact that investigation is prosecuted on experimental lines and that something like organised methods of research are taking form. There is now but little reliance on casual observations recorded by those who have not undergone the necessary discipline in these methods. There is also some change of emphasis in formulating conclusions. Now that the general evolutionary thesis is fully and freely accepted by those who carry on such researches, more stress is laid on the differentiation of the stages of evolutionary advance than on the fact of their underlying community of nature. The conceptual intelligence which is especially characteristic of the higher mental procedure of man is more firmly distinguished from the perceptual intelligence which he shares with the lower animals—distinguished now as a higher product of evolution, no longer as differing in origin or different in kind. Some progress has been made, on the one hand in rendering an account of intelligent profiting by experience under the guidance of pleasure and pain in the perceptual field, on lines predetermined by instinctive differentiation for biological ends, and on the other hand in elucidating the method of conceptual thought employed, for example, by the investigator himself in interpreting the perceptual experience of the lower animals.

Thus there is a growing tendency to realise more fully that there are two orders of educability—first an educability of the perceptual intelligence based on the biological foundation of instinct, and secondly an educability of the conceptual intelligence which refashions and rearranges the data afforded by previous inheritance and acquisition. It is in relation to this second and higher order of educability that the cerebrum of man shows so large an increase of mass and a yet larger increase of effective surface through its rich convolutions. It is through educability of this order that the human child is brought intellectually and affectively into touch with the ideal constructions by means of which man has endeavoured, with more or less success, to reach an interpretation of nature, and to guide the course of the further evolution of his race—ideal constructions which form part of man's environment.

It formed no part of Darwin's purpose to consider, save in broad outline, the methods, or to discuss in any fulness of detail the results of the process by which a differentiation of the mental faculties of man from those of the lower animals has been brought about—a differentiation the existence of which he again and again acknowledges. His purpose was rather to show that, notwithstanding this differentiation, there is basal community in kind. This must be remembered in considering his treatment of the biological foundations on which man's systems of ethics are built. He definitely stated that he approached the subject "exclusively from the side of natural history."[192] His general conclusion is that the moral sense is fundamentally identical with the social instincts, which have been developed for the good of the community; and he suggests that the concept which thus enables us to interpret the biological ground-plan of morals also enables us to frame a rational ideal of the moral end. "As the social instincts," he says,[193] "both of man and the lower animals have no doubt been developed by nearly the same steps, it would be advisable, if found practicable, to use the same definition in both cases, and to take as the standard of morality, the general good or welfare of the community, rather than the general happiness." But the kind of community for the good of which the social instincts of animals and primitive men were biologically developed may be different from that which is the product of civilisation, as Darwin no doubt realised. Darwin's contention was that conscience is a social instinct and has been evolved because it is useful to the tribe in the struggle for existence against other tribes. One the other hand J. S. Mill urged that the moral feelings are not innate but acquired, and Bain held the same view, believing that the moral sense is acquired by each individual during his life-time. Darwin, who notes[194] their opinion with his usual candour, adds that "on the general theory of evolution this is at least extremely improbable." It is impossible to enter into the question here: much turns on the exact connotation of the terms "conscience" and "moral sense," and on the meaning we attach to the statement that the moral sense is fundamentally identical with the social instincts.

Presumably the majority of those who approach the subjects discussed in the third, fourth, and fifth chapters of The Descent of Man in the full conviction that mental phenomena, not less than organic phenomena, have a natural genesis, would, without hesitation, admit that the intellectual and moral systems of civilised man are ideal constructions, the products of conceptual thought, and that as such they are, in their developed form, acquired. The moral sentiments are the emotional analogues of highly developed concepts. This does not however imply that they are outside the range of natural history treatment. Even though it may be desirable to differentiate the moral conduct of men from the social behaviour of animals (to which some such term as "pre-moral" or "quasi-moral" may be applied), still the fact remains that, as Darwin showed, there is abundant evidence of the occurrence of such social behaviour—social behaviour which, even granted that it is in large part intelligently acquired, and is itself so far a product of educability, is of survival value. It makes for that integration without which no social group could hold together and escape elimination. Furthermore, even if we grant that such behaviour is intelligently acquired, that is to say arises through the modification of hereditary instincts and emotions, the fact remains that only through these instinctive and emotional data is afforded the primary tissue of the experience which is susceptible of such modification.

Darwin sought to show, and succeeded in showing, that for the intellectual and moral life there are instinctive foundations which a biological treatment alone can disclose. It is true that he did not in all cases analytically distinguish the foundations from the superstructure. Even to-day we are scarcely in a position to do so adequately. But his treatment was of great value in giving an impetus to further research. This value indeed can scarcely be over-estimated. And when the natural history of the mental operations shall have been written, the cardinal fact will stand forth, that the instinctive and emotional foundations are the outcome of biological evolution and have been ingrained in the race through natural selection. We shall more clearly realise that educability itself is a product of natural selection, though the specific results acquired through cerebral modifications are not transmitted through heredity. It will, perhaps, also be realised that the instinctive foundations of social behaviour are, for us, somewhat out of date and have undergone but little change throughout the progress of civilisation, because natural selection has long since ceased to be the dominant factor in human progress. The history of human progress has been mainly the history of man's higher educability, the products of which he has projected on to his environment. This educability remains on the average what it was a dozen generations ago; but the thought-woven tapestry of his surroundings is refashioned and improved by each succeeding generation. Few men have in greater measure enriched the thought-environment with which it is the aim of education to bring educable human beings into vital contact, than has Charles Darwin. His special field of work was the wide province of biology; but he did much to help us to realise that mental factors have contributed to organic evolution and that in man, the highest product of Evolution, they have reached a position of unquestioned supremacy.

[153] Origin of Species (6th edit.), p. 205.