XXI. MENTAL FACTORS IN EVOLUTION. By C. Lloyd Morgan, LL.D., 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." ("Origin of Species" (6th edition), page 205.) 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." ("Descent

of Man" (2nd edition 1888), Vol. I. page 99; Popular edition page 99.)

"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." (Ibid.

page 99.) 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." ("The Expression of the Emotions" (2nd edition), page 32.) 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 vigorous and victorious

males." ("Descent of Man", Vol. II. page 435.) 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." (Ibid. pages 437, 438.)

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 naive acceptance of such a creed it was not Darwin's province

to enter; "I have nothing to do," he said ("Origin of Species" (6th

edition), page 205.), "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 cooperate,

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 ("Origin of

Species" (6th edition), page 205.), "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 ("Origin of Species" (6th edition), page 233.), "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." (Ibid. page 205.) 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 factors 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 ("Origin of Species"

(6th edition), page 233.), "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 (Ibid. pages

210, 211.), "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 (Independently suggested, on somewhat

different lines, by Profs. J. Mark Baldwin, Henry F. Osborn and the

writer.), 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

cooperate 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

(a) + V with + M,

(b) + V with - M,

(c) - 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 they 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 cooperating 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

("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.

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 hereditary, 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

(Cf. William James, "Principles of Psychology", Vol. II. Chap. XXV,

London, 1890.); 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 viscera 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

vigorous 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

vigorous 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 ("Darwinism", pages 282, 283, London,

1889.) 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

(Ibid. page 283.), "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 ("Darwinism", pages 283, 284.):

"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, but 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 vigorous, 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 ("Descent of Man" (2nd edition), Vol. II.

pages 136, 137; (Popular edition), pages 642, 643.) 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 ("The

Play of Animals", page 244, London, 1898.), 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." (Ibid. page

283.) 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 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 ("Descent of

Man", Vol. II. page 60; (Popular edition), page 566.), "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 ("The Play of

Animals", page 76.), "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 (Ibid. page 75.) 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. ("The Play of Animals" page 24.) 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. ("Descent of Man" (1st edition),

Chapters II, III, V; (2nd edition), Chapters III, IV, V.) 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 (Ibid. Vol. I. pages 70, 71;

(Popular edition), pages 70, 71.)" 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. (Ibid. page 81.) "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

("Descent of Man" (Popular edition), page 82.), "M.E. Lartet ("Comptes

Rendus des Sciences", June 1, 1868.), 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." ("Nature", Vol.

LXI. pages 624, 625 (1900).)

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 ("Descent of Man", Vol. I. page 100.) 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 ("Principles of Psychology," Vol. II. page 289.) 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."

("Descent of Man", Vol. I. page 149.) 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 (Ibid. page 185.), "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. On 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 (Ibid. page 150

(footnote).) 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 overestimated.

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 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.

XXII. THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY.

By H. Hoffding.

Professor of Philosophy in the University of Copenhagen.

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