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 ("Origin of Species" (6th edition), page 206.), 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 weighed 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 ("Expression of the Emotions", page 13. The passage is here somewhat condensed.), "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." (Ibid. page 368.) 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." (Ibid. pages 373, 374.)
"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." ("Expression of the Emotions", page 368.) This principle of antithesis has not been widely accepted. Nor is Darwin's own position easy to grasp.
"Our third principle," he says (Ibid. page 369.), "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 ("Expression of the Emotions", pages 65 ff.) 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.