Physiological Foundations of Behavior

of the more stable features of the polar-symmetrical excito-motor — patterns of the lower levels of the gradients, e. g., the reflex and instinctive complexes, may undergo more or less modification under the superdominance of intelligence, of which the physiological pattern exists primarily in the cortex.! 1 Compare Herrick’s discussion of the physiological characteristics of the cerebral cortex (’24, at the close of Chap. XVII). In the preceding chapters I have tried to show that the individual organism represents fundamentally a behavior pattern, 7. e., the behavior of a specific protoplasm in a certain environment. The development of the individual from the stage of the simple quantitative gradient to the mature organism, of whatever species, represents the modification of behavior resulting from establishment of the primary pattern and the changes which this brings about in the protoplasm. This conception of the organism as fundamentally a dynamic integration of behavior pattern in a specific protoplasm seems to throw light upon various fields of biology, which from other viewpoints have remained more or less obscure. Probably no one doubts that social integrations, patterns, institutions are fundamentally integrations of the behavior of living organisms in certain environments. If both the individual organism and human society represent behavior integrations of living systems, a comparison between them from this viewpoint should prove of interest.

Sociology and its problems are very often outlined in terms of human beings solely, but as a matter of fact various social patterns are found among various organisms in various degrees of development, from rudimentary and evanescent integration to highly stable and complex organizations. The fact that these integrations are based wholly or largely on instinctive, rather than on intelligent behavior does not make them any the less social in character, at least to the biologist. Social integration in man could not have been primitively an intelligent, fully self-conscious integration, but was unquestionably, in large measure, instinctive. Indeed it is a pertinent question whether many of the more primitive and evanescent types of social integration among human beings at the present day, @. g., crowds, mobs, boys’ gangs, etc., are not to a considerable extent instinctive rather than intelligent. If instinctive behavior does play any considerable part in them, they appear to be very similar or essentially identical with many social integrations among animals. It appears to the biologist that the attempts so often made by sociol-

ogists to define and consider social problems in terms of man alone are responsible for at least certain features of the vagueness, uncertainty and difference of opinion, particularly as regards social origins. The delimitation of the subject to the human species makes any adequate interpretation of social origins practically impossible. If man is a product of evolution, the foundations of human society lie, not in the human race, but in other organisms. It is certain that human society represents the highest order or plane of integration upon our planet. If man is a part of organic nature it appears a difficult and uncertain task to attempt to interpret this highest order of integration without attempting to determine the relations between it and the lower and simpler orders of integration on which it must be in part based.

Some sociologists, e. g., Comte, Spencer, Schiffle, have interpreted human society more or less in terms of biological conceptions. Spencer emphasizes the morphological aspects, the mechanisms, of society, while Schiiffle is interested not merely in the structure but in the function of the mechanisms. Both authors carry the biological analogy into minute detail. Of course the question whether or not we shall call human society an organism must depend on our definition of an organism. The objections made by various sociologists to biological forms of interpretation in sociology are of some interest, and some of them suggest that some sociologists have not considered what organisms are. For example, the objection to the biological analogy has been based on the ground that there is no protoplasmic or physical continuity between human individuals. I¢ may be pointed out that the blood cells, though parts, or organs of the organisms, are not continuous and that in the simpler forms cells may lose their continuity with other cells, migrate and establish continuity with a new group. On the other hand, many sociologists maintain that the tool in use becomes essentially an organ, or that in working with different tools the hand plus the tool become essentially different organs. If we grant this, human society consists not simply of human bodies, but of these bodies plus environmental factors, whether natural or in the form of man-made tools which permit communication. Are not two human beings conversing over the telephone or even by means of atmospheric sound waves brought into a dynamic correlation very similar to the dynamic correlation in the organism?

Again, it has been pointed out repeatedly that human society has no permanent organs, but essentially the same thing is true of many: of the simpler organisms. In this connection it may also be pointed out that the tools, the fabrications, are the permanent organs of society. The human individual may be regarded as the dynamic, the metabolic factor in the pattern. The objection has also been raised that the basis of human association is psychic, but unless our definition of the word is very narrow, the psychic factor exists in many forms of animal association as well as in human society. Certainly not all human association is intelligent and self-conscious in character, while many integrations among

animals are more or less psychic, perhaps conscious in character. Objections such as those that human society is not a big animal, that it has for example no stomach, no muscles, etc., are Just as true for many organisms as for society. It has been said that the social mind has no sensorium. But do not the individuals in relation to each other and to environment constitute the sensorium of the social mind just as truly as cells and cell groups in relation to each other and to the external world constitute the sensorium of the individual mind?

And finally many sociologists who admit that the biological analogy may have a certain value are inclined to regard it as a crude and merely preliminary form of interpretation. Undoubtedly human behavior and its integration patterns differ in many respects from anything recognizable in other organisms, but the foundations of human behavior include all of biology, not simply the social behavior of animals, but the behavior of individuals and its mechanisms, and even the behavior of living protoplasm which constitutes the organism. How can we hope to analyze and interpret adequately the more advanced forms of integration in human society without knowledge of the foundations on which they rest?

It is perhaps not of great importance whether or not we call human society an organism but it is a highly significant fact that biological terms apply better than any others. Even though we decline to speak of the social organism we find it very difficult to dispense with the term organization, simply because the term expresses better than any other certain processes and phenomena in society as well as in the organism. The term “ differentiation” is also used in the biological sense in sociology, likewise the term “function,” and so on. The fact that these terms are universally recognized as practically indispensable must mean that the social process is in certain respects, at least, very similar to the organismic process.

The biologist, likewise, has found that certain sociological terms afford more satisfactory analogies for purposes of interpretation than any others. For example, the conception of the multicellular organism as a cell state, with division of labor among its constitutent cells is familiar to every student of elementary biology, and it would be difficult to express certain aspects of the pattern of organisms so clearly in any other terms. In preceding chapters (pp. 146, 258) I have called the more primitive axiate type of pattern an autocratic form of government and have pointed out that evolution has progressed toward democracy and representative government. Various neurologists and psychologists have found the conception of a parliament or deliberative assembly of very great value in interpreting the rdéle of the cerebral cortex.

But there seems to be no good reason for stopping here. The sociologist is accustomed to call the relatively fixed and permanent social patterns institutions. Are there not patterns or orders within the organism fundamentally similar to social institutions in that they represent integrations of living units? Is not the organism itself an institution of some sort in the living protoplasm of one or more cells? If the physiological conception of axiate pattern developed in earlier chapters and elsewhere (Child, ’15c, ’21a) is correct the axiate organism is an institution resembling the state. It represents fundamentally a relation of dominance and subordination, 7. e., of government and governed.

In short, whether we are primarily concerned with the organism or with human society, we cannot help but see the fundamental similarities in the processes of integration in the two patterns, and even though sociologists may continue to insist that sociology remain distinct from biology, the development in biology of the dynamic, in place of the older morphological viewpoint, will make this more and more difficult. In fact the definition of the organism to which the strictly physiological viewpoint of the preceding chapters leads us will serve almost equally well as a definition of society. The organism is a dynamic order, pattern, or integration among living systems or units. -A social organization is exactly the same thing. The fundamental difference between the organism and social integrations among human beings is apparently one of degree or order of magnitude. So- cial integration is a dynamic integration of human organisms, and the human organism is an integration of cells, tissues and organs. Unquestionably these differences in the order of magnitude of the integration determine differences in detail in the processes concerned,

- : pokes but both cell and human being are living systems, and we believe - that the human being is a product of evolution from the cell. We may expect therefore, to discover a fundamental similarity or identity in the more general laws and processes of integration from the one extreme of the cells or the simplest organism to the other of the great modern state or nation. The overlapping of biology and sociology and their use of each other’s terminology indicate that this is the case, but from the dynamic viewpoint this fundamental similarity appears even more clearly than from any other. The morphological analogy between organism and society is at best very general and may readily be overworked. As the morphological mechanisms of the human body are different from those of a bacterium or an Ameba, so the morphological mechanisms of human society differ from those of the human body. But when we come to the problem of the nature of the integration process which makes the micro- érganism and the human being organisms and human society a “superorganismic” pattern, we find certain highly significant similarities. To some of these attention is briefly called in following sections.

Among animals and even among plants we find not only various degrees of organismic integration, from the simple cell to the multiaxiate or “colonial” plant and animal, but we also find certain superorganismic integrations. Some are unquestionably social, while others are near the border line between social relation and relation to environment in general. Biologically speaking; social integration represents for the individual a reaction to a particular factor in his environment, viz., another individual or group of individuals. We must search, therefore, for the foundations of social integration in the relations of the individual to his environment and more particularly his living environment.

The most general relations of this sort are the ecological relations between different species. Ecological communities represent integrations of various species of animals and plants based on the relations of each species to its environment, including both inorganic factors and the other species of the community. As regards the living environment, these relations are of various sorts. The one species or some of its products of metabolism or its body may be merely a source of food in the environment of the other. Even this relation may differ widely in character. The relation between the herbivorous

animal and its food, for example, is passive and accidental on the side _ of the plant, but a definite feature of excito-motor behavior on the part of the animal. The relation of the parasite and its host is likewise usually based on a onesided reaction. The relation between the predatory animal and its prey, however, may involve reaction on both sides. The various forms of symbiosis, mutualism, inquilinism, “guest”’ relations, range from cases like the symbiosis of hydra and other simple animals with green algae in which both species are apparently passive and the relation a matter of chance, to cases among insect species in which each shows definite behavior with respect to the other. This is particularly the case as regards the ants and some of their “guests.”’ Certain species of ants raid the nests of other species, capture and carry home the larve or pup, which are brought up to be “‘slaves.”’ In some relations of this sort the one species is wholly dependent on the other for continued existence. For example, the eggs of certain species of plant lice from which ants obtain nutritive secretions during the summer aré carried by the ants to their nests, cared for during the winter and in the spring the young aphids are carried to their food plants. Certain species of ant aphids are carried out to pasture in good weather and back to the ants’ nests in bad weather, cold nights, etc. Also the ants move them from one pasture to another. The species of plant lice in which these mutualistic relations with ants occur react differently to the ants from other independent species, and their behavior shows adaptation in various ways to the ants. Students of ants have described many other mutualistic relations between ants and other insects (see, for example, Wheeler, ’10, Chap. XIX, also 2)

Discussion of the question whether any of these relations between species, or which of them are to be called social is unprofitable. It may merely be noted that at least some of them exhibit features which play a part in true social integrations. The more general ecological relations between different species of living organisms are largely or wholly material in character and as such they constitute the material basis of a relation of exchange, a commercial relation, but unregulated by communication between individuals. In the more intimate relations between species, e. g., some of those between ants and aphids, communication, excito-motor reaction, and often complex instinctive behavior of one or both species may occur.

It is perhaps justifiable to characterize as fundamentally nonsocial the general ecological relations existing between ‘ differerit species and having to do with material factors such as food, oxygett, or other factors concerned in vegetative life. Social relations are then theoretically distinguishable from such ecological relations in that they are fundamentally relations between members of the same species and involve communication of some sort. But it is probably impossible to make this distinction sharp, for in certain cases the ecological relations obviously approach a social character.

Certain sorts of integration may occur more or less indifferently between individuals of different but closely related species, and between members of the same species. For example, Allee (’20) has described aggregations among ophiurid starfish and among isopod crustacea which may include one or more species. The animals concerned tend to keep their bodies in contact with solid surfaces (positive thigmotaxis) and when such surfaces are reduced to a minimum in their environment they react positively to each other and form masses or bunches of individuals. Dr. Allee informs me that while such aggregations may consist of members of more than one species they form more readily and to a greater extent between individuals of the same species. This being the case, the aggregating individuals must be able to distinguish in some way and to some degree between other individuals of the same and those of other species. Apparently even in these physical aggregations of individuals there is a factor remotely akin to the social factor.

In many other relations between individuals of the same species the social character of the integration appears more clearly. Many animal species aggregate into herds, flocks, packs, schools, ete. The simplest forms of aggregation are temporary and shifting, consisting of individuals which happen to be near enough together to react te each other. Concerning many such aggregations we can say little more than that they originate in a gregarious instinct, but this is no more than saying that they exist because members of the species behave in a certain way. Aggregations of this kind may become more definite and distinct under certain special conditions, e. g., the integration of the herd of cattle through fear, the flocking of various species of birds and of insects, at the migratory season. Again, such ageregations may be based on the brood, the family, or they may have some sexual significance. The flying or “dancing” groups of various species of the gnats and midges which appear in spring and early summer apparently represent a rudimentary and temporary grouping, but one apparently involving some sort of reaction of the individuals to each other. For the formation of such groups a certain atmospheric humidity and absence of wind are obviously necessary. Such a

group may divide or join with others, and individual insects may enter it or leave it. We can distinguish no indications of leadership and in some forms the group can be temporarily or permanently disintegrated by a sound of certain pitch and intensity. But the fact that such groups are at least often, perhaps always, composed of males only is evidence of their relation in some way to sex. We also find animal groups with definite leadership; either temporary and shifting or more or less permanent. The migrating flocks of wild geese are led by a gander and assume a more or less definite arrangement of individuals in two lines diverging from the position of the leader. A more or less definite and fixed relation of leadership and led is commonly believed to exist in the wolf pack and in the aggregations of various other mammalian species. Such groups appear to be truly social. The behavior of the individuals is regulated to some extent by their relation to the group and this relation is dynamic, not material, 7. e., it rests on communication of some sort, not necessarily volitional, but none the less real. Some such groups seem to show something approaching consciousness of the group, however rudimentary. In groups of various birds and mammals certain individuals function as sentinels while the others feed.

The family in one form or another is of wide occurrence among animals and the fundamental factor underlying the origin of the family, viz., sex, is a general biological phenomenon. The parental instinct appears in one form or another, and as a temporary and evanescent, or periodic phenomenon far down among the invertebrates, and the true family appears, at least as a temporary or periodic integration even in certain fishes and in many species among the higher vertebrates. Often the relations of the male to the family are little or nothing more than the primary sexual relation, but there are various species among both birds and mammals in which it is much more than this, and in some animals the male is permanently a member of the family.

The social integration among ants, bees, wasps and termites appears to represent in large measure modifications of. the family. The various castes represent different degrees of development and differentiation of the sexes, in bees, wasps and ants primarily of the females, in termites of both sexes. In the differentiation of the castes heredity, the occurrence or non-occurrence of fertilization, at least in the bees, nutritive conditions during development and perhaps other factors are concerned. The bee and ant communities -are social systems centered about the queen mother, matriarchal systems, with various

complications associated with the differentiation of castes. With the development of the different castes within one sex, or as in the termites, in both sexes, the division of labor has gone further than the primary sexual division and so the community has come to resemble in certain respects a state rather than a family, but its relation to the family seems evident. - Wheeler (’18) has suggested the possibility of a very different origin of social life in the insects, viz., the exchange of nutritive substances, including secretions and excretions, between individuals, a relation which he calls trophallaxis. Unquestionably such exchange plays a part of great importance in many insect communities, but the interpretation of the relation of the community to sex, to the queen mother and to the offspring in terms of trophallaxis appears to offer some difficulties.

Within the communities of different species of ants and other social insects various institutions of more or less definite social character exist in addition to the nutritive exchange. Among the ants for example, we find agriculture, slavery, social care of the young, war, to mention only a few. As a predominantly instinctive, rather than an intelligent integration, the insect community is relatively rigid, but it involves dynamic communication, direct or indirect, between individuals and recognition of, and distinction between, individuals of different sex, caste, stage of development, nest, species, etc. Moreover, the fixity of the integrative behavior is by no means absolute, but modification does occur through learning. In the ordinary life of most social insects there is little indication of anything approaching real leadership, though some vestiges of it do perhaps appear at times. In general the integration of behavior appears to be largely the result of the uniformity of instinctive response of the individuals of each caste to any particular factor. The absence of leadership and the apparent dead level of individuality in each caste make social progress in any strict sense impossible. Probably progress in the insect community can occur only with evolution of castes and instincts, in whatever way this takes place.

That social integration among human beings is fundamentally psychic in character needs no proof. In its simpler forms it may be largely instinctive, but in its higher development it’ becomes conscious and intelligent and in modern society group self-consciousness has developed rapidly and is obviously of fundamental importance for the future progress of society.! Unquestionably human social integration is a matter of the behavior of human beings, particularly with respect to each other, and at the foundation of that behavior as of the individual organism, lie the physicochemical processes of excitation and transmission in human protoplasms and their significance as correlative factors in the organism. In human integration transmissive correlation within the cerebral cortex has become the chief factor, or to speak in psychological terms, the development of human society represents the progressive dominance of the idea and its effects upon behavior.

This brief and fragmentary consideration of relation and integration among organisms is sufficient to show that the fundamental factors in such integration are the excitatory or sensory impressions determined in individual organisms by other organisms and the effects of such impressions upon behavior. In its simpler forms such integration may be little or nothing more than a tropism or a trial- and-error reaction in which individuals react to each other simply as a part of their environment. In somewhat more highly developed forms of animal integration more or less blind and stereotyped instincts are brought into play by the sensory impressions, and the integration becomes more distinctly social in character, though not necessarily involving any conscious factor. Some forms of animal integration, particularly among the higher vertebrates, appear to involve some degree of consciousness of the group, doubtless vague and rudimentary. And finally, the higher forms of integration among human beings involve not only consciousness but. self-consciousness of the group. With evolution of the nervous system and its behavior mechanisms the fundamentally psychic character of integration among individual organisms becomes always more distinct.

In preceding chapters I have attempted to show that the individual organism as an integration or pattern represents the behavior of a specific protoplasm with certain hereditary potentialities in a certain physical environment. The nature of such behavior depends of course upon the behavior mechanisms present in the specific protoplasm. That the integration of individual organisms into groups or patterns of a higher order of magnitude is fundamentally dependent on behavior is a self-evident fact. As the character of the particular organism depends primarily upon the specific constitution of its protoplasm and the behavior mechanisms resulting from this constitution, so the character of a particular integration among orguisms depends primarily upon the sort of organisms concerned

and the behavior mechanisms which they possess. With the development of excito-motor behavior and its mechanisms in the individual organism new potentialities of physiological integration arise and similarly with the evolution of excito-motor behavior and its mechanisms new potentialities of integration among organisms arise. We may analyze these into social and non-social, conscious and unconscious integration, etc., but the point which I wish to emphasize is that, from the viewpoint of behavior, synthesis is also possible. From the individual organism onward through various forms of integration of organisms, we are dealing with the reactions of specific living systems to their environments, including other living systems, and with the laws which govern those reactions. The degree of complication of mechanism differs enormously, but, if we admit that man is a part of organic nature, then we must also admit physiological continuity from the simple excitation-transmission gradient to the complex workings of the human mind in its social relations. And at this point the question arises whether the synthesis is completed with the recognition of such continuity. This question is dealt with in following sections.

Association.—In the organism the association of cells or other parts in normal development usually represents rather an incomplete dissociation or separation than an active coming together of independent individuals. Under experimental conditions, however, isolated cells not too far distant from each other do move toward each other and aggregate into masses (Roux, ’94; H. V. Wilson, 07, ’11). In certain cases, ¢. 9., cells of sponges, these aggregates, if exposed to an environmental differential may become new individual organisms. In such cases active association does occur, supposedly through the action of chemical substances given off by the cells. The swarm spores of certain simple plants also undergo normally an active association to form multicellular individuals of more or less definite form and cell-configuration (R. A. Harper, 418°a, b).

The basis of social integration is likewise association of individuals and such association may be predetermined and passive, so far as the individual is concerned, or it may be active. For example, the individual is born into @ group as a new cell arises by division in an organ, but he associates actively with other individuais, whether it be in a conflict of interests or under the compulsion of a common instinct, the stress of a common emotion, or the influence of a common purpose.