Henry W. Nissen (1901-1958)
Biographical Reference:
Dewsbury, D. A.
(1998). Henry W. Nissen: Quiet comparative psychologist. In G.A. Kimble, C.A.
Boneau, & M. Wertheimer (Eds.) Portraits of Pioneers in Psychology: Volume
III. (pp. 228-242). Washington, DC: APA.
Phylogenetic Comparison
HENRY W. NISSEN
Yerkes Laboratories of Primate Biology and Yale University
In this chapter we shall examine the role of comparative psychology in general biology, and its contribution towards the solution of basic problems of experimental and theoretical psychology. A search will be made for concepts or behavioral mechanisms in terms of which the behaviors of different animals may be compared. This treatment does not promote one or another of the currently active schools; it tries rather to synthesize a systematic, theoretical approach with the naturalistic point of view which looks at behavior in its wider setting and which refuses to exclude from consideration phenomena lying outside an arbitrarily circumscribed area of interest.
FUNCTIONS AND METHODS
Comparative psychology may be defined broadly as the science concerned with similarities and differences of behavior at various phylogenetic levels. Its aims and functions (1) as a branch of biology and (2) as part of experimental and theoretical psychology are closely interrelated.
Comparative Psychology and Evolution
As a biological science, comparative psychology supplements morphological and physiological evidence concerning phylogenetic development and the interrelations of species and phyla. The fragmentary story of evolution is interrupted most conspicuously and exasperatingly at the critical points of transition from one main grouping to another. One reason for this may be, as Lecomte du Noüy (1947, p. 79) suggests, that the transitional forms evolved quickly and survived only briefly. "Each group, order, or family," he says, "seems to be born suddenly and we hardly ever find the forms which link them to the preceding strain. When we discover them they are already completely differentiated." The present status of our knowledge about phylogenetic origins is admirably summarized by [D'Arcy W.] Thompson:
We have long known, in more or less satisfactory detail, the pedigree of horses, elephants, turtles, crocodiles and some few more; and our conclusions tally as to these, again more or less to our satisfaction, with the direct evidence of paleontological succession. But the larger and at first sight simpler questions remain unanswered; for eighty years' study of Darwinian evolution has not taught us how birds descend from reptiles, mammals from earlier quadrupeds, quadrupeds from fishes, nor vertebrates from the invertebrate stock.... We may fail to find the actual links between the vertebrate groups, but yet their resemblance and their relationship, real though indefinable, are plain to see; there are gaps between the groups, but we can see, so to speak, across the gap. On the other hand, the breach between vertebrate and invertebrate, worm and coelenterate, coelenterate and protozoan, is in each case of another order, and is so wide that we cannot see across the intervening gap at all (1942, p. 1093). [p. 348]
The conclusion is already implicit in these words that evolution, as manifested in structural characteristics, has been a discontinuous affair rather than the continuous process envisioned by Darwin. Thompson proceeds to draw an analogy with mathematics and phenomena of the inorganic world:
An algebraic curve has its fundamental formula, which defines the family to which it belongs; and its parameters, whose quantitative variation admits of infinite variety within the limits which the formula prescribes. With some extension of the meaning of parameters, we may say the same of the families, or genera, or other classificatory groups of plants and animals.... We never think of "transforming" a helicoid into an ellipsoid, or a circle into a frequency curve. So it is with the forms of animals. We cannot transform an invertebrate into a vertebrate, nor a coelenterate into a worm, by any simple and legitimate deformation, nor by anything short of reduction to elementary principles.... A "principle of discontinuity," then, is inherent in all our classifications, whether mathematical, physical or biological.... This is no argument against the theory of evolutionary descent.... Our argument indicates, if it does not prove, that such mutations, occurring on a comparatively few definite lines, or plain alternatives, of physico-mathematical possibility, are likely to repeat themselves; that the "higher" protozoa, for instance, may have sprung not from or through one another, but severally from the simpler forms; or that the worm-type, to take another example, may have come into being again and again (1942, pp. 1094-95).
This emphasis on large and abrupt variations in evolution is representative of most modern biological thinking. Its implication for comparative psychology is that in behavior also we may expect to find discontinuity -qualitative rather than merely quantitative changes- as we pass from the lower to the higher animal forms. Unlike the early post-Darwinians, who looked for a gradual development of behavioral capacity in the animal kingdom, the comparative psychologist is now relatively free to postulate new capacities or new behavioral mechanisms at various points along the phylogenetic scale, without doing violence to general biological theory. It would be expected, however, that such major psychological emergents would coincide with the major and abrupt changes in structural characteristics differentiating the larger taxonomic groups, especially the phyla. This expectation, we may say now, is not substantiated by the considerations that follow. [fn-1]
Taxonomy and evolutionary theory in general have been based almost entirely on "formal resemblance" among structural criteria, which often are of a most superficial character. Now there is no a priori reason, and certainly no compulsion, to make the study of mental (i.e. behavioral) evolution subsidiary to the known or inferred facts of structural evolution. A comparative evolutionary psychology could be developed in vacuo, strictly and completely "on the behavioral level," a la Skinner (1938). This however, would be unnecessarily restrictive and would repeat in principle the error of the taxonomists who confine themselves entirely to the structural aspects of evolution.
As Zuckerman (1933) suggested, behavioral data should contribute to an understanding of the historical relations among animals and to a more rational basis for their classification. Zuckerman has already demonstrated how the inclusion of functional [p. 349] (i.e. physiological and behavioral) data can help to disentangle the confused relations within the order of primates. Lorenz (1937) points out that, although the family of pigeons, Columbidae, form a definite taxonomic group, there is no one distinctive morphological character that is common to all species and that distinguishes them all from other groups. All pigeons, however, are distinguished from other birds by a behavioral criterion, namely, the manner of drinking. Pigeons pump water, whereas all other birds scoop it up. Petrunkevitsch (1926) has pointed out "the value of instinct as a taxonomic character in spiders," and other examples are given by Heinroth (1910), and Adriaanse (1947).
The contribution that comparative psychology can make to the problems of phylogeny is a function of (1) the scope and precision of its data, (2) the adequacy of the behavioral categories under which the data are organized, and (3) the comparability of the data. The second of these problems is especially critical. What are the "dimensions of behavioral evolution," that behavioral units, mechanisms, processes, or capacities that differentiate, quantitatively or qualitatively, among different levels on the phylogenetic scale? Some of the simpler ones, such as sensory acuity, are relatively easy to identify. But it is the more involved patterning of behavior that is most significant in differentiating the behavior of various animal groups, and it is here that the problem of identification is most difficult.
Given that the aspect of behavior in terms of which two species are to be compared has been satisfactorily defined, problems of technique remain to tax the ingenuity of the experimenter. If an intellective function is to be tested, all those factors that affect the expression of the function, but that are no part of it, must be experimentally controlled. Conditions of motivation, past experience, and sensory and motor characteristics are among the determinants of any performance, and, unless this entire "assisting cast" is rendered functionally equivalent, the differences in performance in the testing situation cannot be interpreted as reflecting an difference in the function under investigation. In addition, the possibility that a given problem may be solved in different ways by different species is a source of difficulty.
Animal Psychology
Psychological studies using animal subjects [fn-2] may be grouped according to the purpose of the investigator. Those experiments in which interest is focused on a special problem, or on a particular animal form, are here considered under the heading "animal psychology."
Animal experimentation. In a large proportion of animal experiments the use of this or that species is incidental or even accidental; a particular animal is chosen because it happens to be convenient. The fact that insects, fish, and birds have been used more that other forms for studies of instinct is attributable to the fact that these animals display instinctive behavior with particular clarity, not because it is absent in other animals. All such animal experimentation is simply an extension of general (human) experimental psychology; the phylogenetic status of the species used is more or less irrelevant.
For certain types of investigation animal have definite advantages over human subjects. Their life span is usually shorter, their entire life history prior to experimentation can be known and controlled, and they may be subjected to operative and other procedures that could not otherwise be used. Most important of all, basic mechanisms that in man are obscured by the complexity of his behavior, as well as by cultural (especially language) factors, may often be [p. 350] clearly discernible in simpler organism. These points have been elaborated by Washburn (1936), Katz (1937), Thorndike (1942), Harlow (1942) and others.
Studies of the animal-as-a-whole. A second class of studies is concerned more with a rounded-out delineation of a given kind of animal than with a specific problem in behavior analysis. The method is more often observational than experimental. Such studies are in the tradition of the older naturalist who set about to give complete accounts of every form of animal life. The early emphasis was on morphological characteristics, but ecological and behavioral features are being given increasing attention.[fn-3]
Most contributions of the so-called anecdotal school (roughly 1852-1890) belong to this category, although the anecdotalists were motivated largely by the desire to produce "evidence of reasoning and other characteristically human mental life in the higher animals" (Warden, 1927, p. 147). Field studies by trained observers, casual or incidental observations by ecologists, hunters, and explorers, the reports of animal trainers, zoo caretakers, and veterinarians, and the books written about pets of various kinds also belong here. Some of these are valuable contributions to our knowledge; others are useless or worse. Many of them are guilty of anthropomorphism -or of its opposite extreme (a delinquency for which there seems to be no good term: "brutalization" and "mechanomophism" say to much).
The importance of good descriptive studies of animal behavior must not be minimized. The value of careful observation under natural conditions is indicated by Tinbergen (1942). He points out that the controversy between von Hess and von Frisch regarding "knowledge of the whole behavior-pattern" of the species: in its food-gathering activities the honeybee responds to the colors of flowers, whereas in other situations its visual behavior is guided entirely by relative brightness, as if it were color-blind. In the case of the chimpanzee, Nissen (1931) has suggested that intellectual capacities will be revealed more clearly in the laboratory, where artificially imposed obstacles may be used to encourage that exploitation of those capacities, than in their natural habitat, where the necessities of life are obtained with little effort. The emotional and social aspects of anthropoid behavior, on the other hand, are greatly distorted in captivity and are more validly observed in the field.
Comparative Psychology
We wish to consider now the bearing of all animal behavior work on the central problems of psychology. How may phylogenetic differences and similarities of behavior be used to clarify the important principles of behavior? This, as I see it, is the raison d'être of what is called comparative psychology.
The genetic method. The genetic or historical method attempts to find explanations in origins. With the aid of temporal and causal continuity the present is explained in terms of the past. Ontology, phylogeny, and cultural are the three areas within which behavioral development may occur. These three areas of development are, of course, interdependent: the accomplishments of ontogeny and cultural depend, in the first place, on phylogenetic heritage. What the individual contributes to the culture is a function of complex interactions between phylogenetic status, ontogenetic development, and prior cultural accumulation. And some writers such as Hooton (1940) are perturbed about the debilitating effects of cultural influences on the future biological status of the human race.
The credo of the comparative psychologist is well stated by Tolman (1945):
... whereas man's successes, persistences and socially unacceptable divagations -that is, [p. 351] his intelligences, his motivations and his instabilities- are all ultimately shaped and materialized by specific cultures, it is still true that most of the formal underlying laws of intelligence, motivation and instability can still be studied in rats well as, and more easily than, in men.
Unless there is a continuity or homology of behavioral mechanisms from the lower to the higher animals (including man), there would be no rationale for the comparative method. This does not mean, necessarily, that the higher (later) is merely an extension or elaboration of the lower (earlier), although this is and has been a most provocative and fruitful working hypothesis. Quantitative complication may become so great that it produces, in effect, qualitative differences with new, "emergent" properties.
One further point deserves mention: although the laws that hold on one level may not be adequate to explain all phenomena on the next higher level, the former are never contradicted, reversed, or negated by the later. "No physical law, any more than gravity itself, not even among the puzzles of stereochemistry or of physiological surface-action and osmosis, is known to be transgressed by the bodily mechanism" (Thompson, 1942, p. 13).
When interest is focused on individual differences within the higher group, it may be that these later emergent properties have primary significance in distinguishing between this and that individual or class. In this sense and to this extent only, we may agree with [Gordon] Allport (1947) in his objection to the "phylogenetic model" and "the animal paradigm for personality and for social psychology." We can agree that differences of intention among people are of great practical significance. But these intentions are superstructures erected on a phylogenetically inherited foundation. Would the superstructure remain the same, or ever have become what it is, without the foundation? Can anyone conceive of what our "intentional morality" would be like in the absence of food, water, and sex hungers, without competition, without frustration or conflict? Intentions will never be understood without reference to the phylogenetic matrix from which they have arisen. They are the product of a long and tortuous development.
Structure-function correlations and homologies. There is a school of thought in psychology that holds that a psychological (behavioral) event is explained when, and only when, its underlying physiological-morphological basis has been identified. Whether or not we concur with this extreme position, it is undeniable that the demonstrated correlation between "physical" and behavioral phenomena is a long step towards explanation. A psychological theory that is at variance with established principles of physiological (e.g. neurological) action is thereby summarily ruled out, whereas one that does mesh with such principles is ipso facto given increased probability.
Within any one species the expression of a given mechanism or process is almost certain to be obscured by other mechanisms, the problem of isolation (experimental or otherwise) being often insuperably difficult. Within the one species it is often impossible to abstract what is essential from what is incidental. But, when the process is observed in various contexts (i.e. species), the irreducible minimum, the essence of the process under consideration, gradually becomes clear. Zuckerman has stated it as follows: "What is obscure in a functional activity of one species, for example man, may be clarified by some peculiarity in the working of the [p. 352] homologous process in a related species, and problems whose existence had not previously been realized might be revealed by this method of approach" (1933, p. xvi).
The comparison of the behavior of species and of phyla is one of the more important ways in which we may exploit naturally given, ready-made variations. Nature has provided us with well over half a million species that vary, greatly or minutely, in structure and behavior. Also, they show greater or lesser similarities, resemblances, homologies, in both aspects. This wealth of materials is ready to be used (1) to help in the definition of fundamental mechanisms or principles of behavior, and (2) in explaining behavior in terms of its underlying physical bases.
Dimensions of behavioral evolution. The literature contains an amazingly small number of studies in comparative psychology -investigations in which two or more species have been compared directly. Comparisons are mostly attempts to relate the findings of different workers, using various organisms in more or less similar experimental or naturalistic situations. [fn-4]
Whether we are impressed more by the similarities or by the differences of behavior within the animal kingdom depends on the scale of observation used. The broader the category, the more do the similarities stand out, and vice versa. All animals reproduce and maintain a relatively constant internal environment, but the specific behaviors associated with reproduction and with homeostasis differ greatly among species. All animals are sensitive to stimuli, and almost all move from place to place, but the mechanisms of receptivity and of locomotion vary widely. In general the similarities are expressed in terms of function or "purpose"; the differences in terms of mechanisms or efficient causes.
Many criteria for differentiation among the behaviors exhibited at various stages of the phyletic scale have been suggested. Some of these are proposed as continua of quantitative variation from the most primitive to the highest forms. Others refer to emergent characteristics, absent at one level, present full-blown at the next.... [p. 353]
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RECAPITULATION
The present section, being, so to speak, a summary of summaries, must be presented in condensed and dogmatic form, without the development of implications or the consideration of alternative points of view.
Motivational versus Cognitive Differences in Evolution
For psychology, differences among animals with respect to what they want are trifling by comparison with differences in the means by which they satisfy their wants. The fact that one organism feeds itself by ingesting algae, others by eating crabs, insects, grass, or eggs, is of only incidental interest in contrast to the various behaviors by which algae or crabs, insects, grass, or eggs are obtained. It is in the cognitive rather than in the motivational aspects of behavior that we find the significant axes of behavioral evolution. Differences in motivation consist largely in an increasing complexity or degree of indirection of drive sequences terminating in goal attainment. These motivational differences are derivative, depending on the cognitive equipment of the animal. Differences in forms of social interaction, including those aspects we call culture, also derive from cognitive factors. In an earlier section we found only one type of behavior modification, namely, "transformation of mechanisms into drives," for which there is any convincing evidence of phylogenetic difference, and this distinction applies mainly to man as contrasted with other animals. This emphasis on cognitive factors does not mean, of course, that the student of comparative behavior may disregard its motivational aspects. Such neglect would be as inexcusable as disregarding the animal's sensory equipment or mode of locomotion, as if its receptor capacities and whether it walks, crawls, flies, or swims were irrelevant to the problem.
Axes of Cognitive Development
Sensory elements and response units may be thought of as the raw materials of behavior. The range and differentiation of environmental energies to which the organism is sensitive depend on the sense organs [p. 381] and their central connections. With increasing sensory differentiation, the possibilities of central (perceptual) organization into unique patterns are rapidly multiplied. Sensory differentiation is, therefore, a necessary, but not a sufficient, condition for breadth of perceptual scope. Behavioral complexity is likewise increased as the organism is able to make discrete and independent movements of all parts of its body. Phylogenetically the locomotor coordinations develop earlier than those involved in manipulation. Variety and precision of motor patterning culminates in the manual dexterities and speech of man.
Perceptual organization (sensory correlation) in evolutionary development proceeds along two axes: (1) inherited perceptual patterns and (2) individual acquisition of perceptions. Acquired perceptions are organized in the course of experience by the postulated mechanisms of pattern identification, the selective process, and symbolization. As regards (1), an initial peak, phylogenetically, is seen in the insects, and a second one in the birds. In the higher primates this trend takes a pronounced drop. Experiential perceptual organization (2), on the other hand, seems to play only a minor part until we come to some of the birds and especially the mammals. Among the monkeys and apes (2) rapidly displaces (1) and leads to a wider perceptual scope than is found among any of the lower organisms. Somewhere among the higher mammals, perhaps not before the primate level, there emerges a new instrumentality for perceptual and conceptual organization, namely, symbolization. Although we have suggested a gradual development of the symbolic process, the evidence is overwhelming that the curves representing this dimension of behavioral evolution rises sharply upwards with the appearance of man.
The effectiveness of the S-R connecting process rises from the lower to the higher invertebrates; and no further rise is apparent until we come to the higher primates. It is probable that symbolization helps not only in perceptual organization but also in making possible prompt and efficient connection of percepts and concepts to specific responses. One-trial learning becomes the rule rather than the exception among the highest primates. There the connecting of the central symbolic processes with one another, and with overt responses, seems to occur with unique facility.
A Phylogenetic Comparison of Intelligence
Behavioral evolution is characterized by increasing complexity. With ascent in the phyletic scale, the effective environment expands spatially and temporally and provides an ever-wider range of behavior determinants. This expansion follows upon the organism's ability to integrate and organize. It is only with the advent of "culture" that the effective environment itself expands rapidly from generation to generation.
We have now discussed and summarized what appear to be important axes along which phylogenetic changes in behavioral complexity may be traced. We have also seen, usually rather vaguely, the points of the phyletic scale at which inflections or origins of the curves representing these several axes occur. The accompanying figure is a rash venture, presenting a tentative estimate of the composite of those curves.
.jpg)
The
ordinate of Fig. 2, it will be noted, is labeled "Intelligence level."
It could as well have been called "Behavioral complexity" or "Spatio-temporal
span of behavior determination." Or it may be taken to represent the sum
total of the organism's cognitions and adjustments. [fn-5] [p. 382]
The particular divisions of the animal kingdom indicated in Fig. 2, where one division contains a single species and another includes several phyla, represent a compromise between conventional classification and present behavioral evidence. (A single continuous curve could have been drawn by reordering the usual taxonomic sequence.) Within the division of protista the Rhizopoda, Mastigophora, and Infusoria are represented by the upper (right-hand) portion of the curve; among the lower invertebrates the Echinodermata are displace to the left of their usual taxonomic position; within the lower vertebrates and the primates the conventional order is maintained; for the division of lower mammals behavioral data are inadequate to permit an estimate of the relative positions of the various orders.
It is quite possible that the continuous curves drawn for each division should be broken up into numerous discrete curves, each representing a subgroup. Further refinements in behavioral measurement might indicate, for instance, that the higher amphibians are more advanced than the most primitive reptiles. Needless to say, distance along the abscissa is not proportional to the number of species or individuals included. The fact that the curve represents an average, a pooling of attainments along each of several dimensions, means that, with respect to one or another of these, a given species may stand considerably higher or lower than its position on the curve indicates. There are, obviously, no absolute units of measurement for the ordinate; a given distance on one curve cannot be presumed to be equal to the same distance on another curve or at another level of the same curve. The vertical axis thus represents only order, or direction [p. 383] of difference. The top of the final curse suggests the debatable position of modern natural man. Because we know so little about the behavior of earlier forms of Homo Sapiens, this curve is dotted.
[SELECTED] REFERENCES
Allport, G. W. Scientific models and human morals. Psychol. Rev., 1947, 54, 182-192.
Harlow, H. F. Animal behavior. Chapter 12 in R. H. Seashore (Ed.). Fields of psychology. New York: Holt, 1942.
Hobhouse, L. T. Mind in evolution. New York: Macmillan, 1901. (2nd ed., 1915, 3rd ed., 1926).
Hooton, E. A. Why men behave like apes and vice versa. Princeton: Princeton University Press, 1940.
Hull, C. L. Principles of behavior. An introduction to behavior theory. New York: Appleton-Century-Crofts, 1943.
Hull, C. L. The place of innate individual and species differences in a natural-science theory of behavior. Psychol. Rev., 1945, 52, 55-60.
Hunter, W. S. The delayed reaction in animals and children. Behav. Monogr., 1913, 2.
Katz, David. Animals and men. Studies in comparative psychology. New York: Longmans, Green, 1937.
Lecomte du Noüy, P. Human destiny. New York: Longmans, Green, 1947.
Lorenz, K. über die Bildung des Instinktbegriffs. Naturwissenschaften, 1937, 25, 289-300., 307-318, 324-331.
Maier, N. R. F. Reasoning in white rats. Comp. Psychol. Monogr., 1929, 6.
Maier, N. R. F. and T. C. Schneirla. Principles of animal psychology. New York: McGraw-Hill, 1935.
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Munn, N. L. Psychological development. New York: Houghton Mifflin, 1938.
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Morgan, C. L. Animal behavior. London: Arnold, 1900.
Morgan, C. L. Emergent evolution. New York: Holt, 1927.
Nissen, H. W., A field study of the chimpanzee. Comp. Psychol. Monogr., 1931 S.
Nissen, H. W., J. S. Blum, and R. A. Blum. Conditional matching behavior of chimpanzees. J. Comp. Psychol., 1949, 42, 339-356.
Nissen, H. W., A. H. Riesen, and V. Nowlis. Delayed response and discrimination learning by chimpanzees. J. Comp. Psychol., 1938, 26, 361-386.
Novikoff, A. B. The concept of integrative levels and biology. Science, 1945, 101, 209-215.
Petrunkevitsch, A. The value of instinct as a taxonomic character in spiders. Biol. Bull. Woods Hole, 1926, 50, 427-432.
Skinner, B. F. The behavior of organisms. An experimental analysis. New York: Appleton Century-Crofts, 1938.
Thompson, D. W. On Growth and form. Cambridge: Cambridge University Press, 1942.
Thorndike, E. L. Why study animal psychology? Chapter 1 in F. A. Moss (Ed.) Comparative psychology. New York: Prentice-Hall, 1942.
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Tolman, E. C. A stimulus-expectancy need-cathexis psychology. Science, 1945, 101, 160-166.
Warden, C. J. The historical development of comparative psychology. Psychol. Rev., 1927, 34, 57-85, 135-168.
Warden, C. J., T. N. Jenkins, and L. H. Warner. Comparative psychology. Volume I. Principles and Methods. New York: Ronald, 1935; Volume III. Vertebrates, 1936; Volume II. Plants and invertebrates, 1940.
Waters, R. H. The historical background of comparative psychology. Chapter 2 in F. A. Moss (Ed.), Comparative psychology. New York: Prentice-Hall, 1942.
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[Extracted Footnotes]
[fn-1] Thompson's "principle of discontinuity" would suggest also that, in a systematic treatemnt such as Hull's (1943), species differences could be accounted for merely by changing the "empirical constants" of the equations (Hull, 1945) only within limits; at other points the forms of the equations would need to be changed, or new equations would have to be added.
[fn-2] Except where otherwise indicated, the term "animal" will be used in this chapter to refer to infrahuman organisms.
[fn-3] For brief histories of comparative psychology the reader may consult Warden (1927), Warden, Jenkins, and Warner (1935), or Waters (1942).
[fn-4] Among the major American contributions which cover the entire range of the animal kingdom are the books of Washburn (1936) and of Maier and Schneirla (1935). The three-volume work of Warden, Jenkins, and Warner (1935, 1936, 1940) is an invaluable handbook for the field of animal behavior; these volumes contain also comprehensive bibliographies of the literature. Among briefer or more specialized treatments that have appeared since 1930 may be mentioned Lashley (1934), Warden, Jenkins and Warner (1934), Russell (1934), Moss (1934, 1942), Katz (1937), Munn (1938), Fischel (1938), Bierens de Haan (1940), Werner (1940), Moore (1941), C. T. Morgan (1943). Earlier classics in the field include Romanes (1881), C. L. Morgan (1894, 1900, 1927), Loeb (1900, 1918), Hobhouse (1901), Jennings (1906), Yerkes (1907), Thorndike (1911), Watson (1914), Kafka (1914, 1922), Holmes (1916), Uexküll (1921, 1926), Child (1924), Herrick (1924), Köhler (1925), Hemplemann (1926), Hingston (1928), Pavlov (1927), and Coghill (1929). The present chapter owes much to these earlier compilations; as a rule, studies already described in them will be treated as "general knowledge," and specific reference to those studies and their authors will usually be omitted in our subsequent discussions.
[fn-5] By thus including those behavior integrations strictly determined by the inherited structure, as well as those whose development is contingent upon individual interaction with the environment, we depart from the traditional instinct-intelligence dichotomy. The distinction between the innate and the acquired is certainly an important one. But the usual connotation of "intelligence" is a measurable, or at least describable, performance. Intelligence that does not express itself in behavior is a contradiction of terms. Performance reflects neural organization, and this comprises structurally determined patterns as well as patterns formed in the course of experience. The fact that there will be organizing activity is determined structurally as much as is the fact that there are certain species-constant organizations. The variability of environmental factors that enter into individually acquired organizations provides for a greater indication in the final test of survival, however, that nature is prejudiced in favor of one or the other mode of intelligence [(see Munn, 1971; and Leontyev, 1981 -chapters 1 & 2- for more on these points].