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.