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Animal models of human behavior: Conceptual, evolutionary and neurobiological perspectives
Book Review Animal Models of Human Behavior: Conceptual, Evolutionary and Neurobiological Perspectives. Edited by Graham C. L, Davey. Chichester: Wiley, 1983, f23.75
This volume collects together in one place a number of papers dealing with the problems of extrapolation from animals to man. It is organized in three sections, each rather different in character. Although the first section is entitled “Conceptual and Historical Issues,” conceptual problems are discussed throughout the book, and this section is in fact a set of six papers dealing mainly with the usefulness or otherwise of the rat as a model of human associative learning. Most authors are rather critical of the assumption of ready extrapolation from this or other species to man. Thus, Miles notes that the language we use for many human concepts, such as ‘responsibility’, is applied only reluctantly if at all to animals. He does not, however, offer any criterion by which we should judge when to accept and when to reject the promptings of linguistic propriety as a guide to real species differences. Boakes suggests that although J. B. Watson thought the environment was important, he was reacting against an unthinking hereditarianism rather than making a strong claim for environmentalism as such. He then argues that nevertheless a consequence of this may have been the diversion of psychology into an excessive concentration on conditioning in animals at the expense of what might actually be important environmental influences on behavior. Boakes might also have said, but has left it to the contributors in section two, that another consequence has been for psychology to disregard the natural history and evolutionary significance of behavior. Blackman offers a spirited defense of the conventional behavior analyst’s argument that animal behavior be explained without re-
course to cognitive terms extrapolated from humans. As his examples make clear, this amounts to asserting that descriptions of necessary and sufficient environmental conditions for behavior, if found, constitute an explanation and are not themselves in need of some further level of explanation. This approach allows explanation inductively, but it severely limits the applicability of animal models if it is accepted that human behavior may actually not be wholly explicable in similar terms. Blackman argues against extrapolating explanatory cognitive constructs from man to animals, but his view on extrapolation the other way is not clear. Catania, too, seems to think that the important thing is to describe behavior in terms of antecedentconsequent relationships but leaves the nature of explanation rather obscure. Lowe and Davey both argue that there are qualitative differences between human and animal learning, claiming that the way humans “label” or “describe” environmental events (Lowe) or “restructure knowledge about the UCS” (Davey) affects their responses. The second section is entitled “Evolutionary Aspects” and comprises live chapters discussing the implications of evolutionary theory for attempts to extrapolate from animals to man, and also the way in which it informs the sorts of inquiry that are appropriate. Plotkin argues at length that comparative studies of learning are of limited value without a conceptual framework, by which he means a theory of the biological function of the behavior under consideration. If correct, his thesis preempts much of the preceding section. Odling-Smee offers an 123
Ethology and Sociobiology 6: 123-125 (1985) 0 Elsevier Science Publishing Co., Inc., 1985 52 Vanderbilt Ave., New York, New York 10017
0162-3095/85/$03.30
124
elaboration of his theory of multiple levels in evolution as a conceptual resolution of the naturenurture issue in behavior. His work is dealing with evolutionary models rather than animal models as such. Smith, in a review of the sociobiology of reproduction and child care in man makes it pretty clear that whereas the ideas derived from animal studies in this field have been useful in generating research, the predictions from animal models are only weakly supported, and he concludes that evolutionary biology cannot account for behavior in contemporary societies without additional assumptions about cultural evolution. He does, however, feel that some aspects at least of parent child behavior do tend to maximize inclusive fitness. Evidently the development of distinct cultural systems may not be independent of selection pressures. Blurton Jones shows even more clearly how assumption from evolutionary biology serve to structure questions about human behavior and illustrates the point with accounts of two diverse studies. He is concerned with the question of whether cultural activities and learning mechanisms actually do tend to improve inclusive litness, and concludes that they may. He also tackles head on the objections raised against applying sociobiology to man. He further provides an object lesson in the principle that it is not the stability of a behavioral measure but what it means that is important. Blurton Jones writes with great clarity, though with perhaps a hint of irritable defensiveness. The net conclusion from this section seems to be that as soon as the adaptive function of behavior is considered, the validity of particular cross species comparisons declines sharply and has to be qualified by a grasp of the function, but that the use of animal behavior in an evolutionary framework to suggest functional hypotheses is valuable. “Neurobiological AsThe final section, pects,” contains a further seven chapters examining the nature of extrapolation in neurobiology, including several dealing with aspects of the neurochemistry of behavior, together with a scathing review of the way simple analogies have apparently inspired a number of psychosurgical procedures (Carroll and O’Callaghan). It is difficult to disagree with the latter authors when they suggest that without convergent indications from human studies, animal studies alone should not stand as the basis for psychiatric intervention (especially when irreversible).
Green, in the next chapter, is principally concerned with schizophrenia and comes to the unsurprising conclusion that it is not much use looking in animals for behavioral analogues to a condition primarily characterized by disorders of thought and affect. However, he thinks it important to understand the dopamine system, and argues that animals may be useful here. In so doing he implicitly follows a logic of extrapolation explicitly laid out for neurochemical work by Warburton. Green also discusses the psychosurgical procedures once used for schizophrenia, an overlap with the’preceding chapter that could properly have been edited out. One really does not need to be told twice that the founder of prefrontal lobotomy was shot by one of his own patients in 1944. The editor might also have intervened in the chapters by Legg and Oakley. Legg argues at some length that neither the principle of encephalization-translation of brainstem functions to the neocortex with advancing phylogenetic status-nor that of “conservation,” which in this context means the retention by homologous brain structures of their functions in the face of phylogenetic advances, can meet the data on cross species comparisons of visual function without circularity. Inter alia he notes that, “Since the structures implicated in residual [destriatel vision . . . . are all brain stem regions, the principle of conservation would lead us to expect that they should operate similarly in all mammals, but this assumption is directly contradicted by the data” (p. 229). Oakley, immediately following, seems rather more taken with the principle of conservation: “It now seems evident from the use of comparable testing procedures across species that all mammals with striate cortex removed show similar visual abilities” (p. 248). Legg and Oakley, it may be noted, do not cite a single reference in common. This is probably because the main substance of Oakley’s chapter is a claim that subcortical mechanisms can be important in associative learning, and that the study of mammalian subcortical function is useful in establishing this conclusion. The chapters by Rick and Benton deal with fairly specific areas of neurochemical research. Rick argues that rats injected with anti-ganglioside antibody may show some structural and behavioral analogies with human mental retardates, and Benton argues on conceptual and methodological grounds that the relationship between testosterone and aggres-
125
sion has been exaggerated in humans despite animal studies suggesting a rather strong effect. It is certainly valuable to have this material collected together in one place, but this is, on the whole, a collection of discrete and self-contained contributions. There are some omissions. Almost nothing is said of the use of primates as models, for example in studies of communication or of social development, nor is there any discussion of classical ethology and attempts to extend it to man. Some authors, notably Catania, Plotkin, Benton, Legg, and Warburton have made the general relevance of animal models a relatively central concern of their chapter, but others have raised questions of animal models in the context of rather specific pieces of research. Authors generally give an impression of not having talked much to one another. Consequently, some issues arise as it were interstitially, and are only apparent to the reader by a comparison of the various articles. For example, the idea of animal models evidently means quite different things in different sections, and the criteria by which it should be evaluated vary accordingly. Thus evolutionary theory is now concerned with ideas of inclusive fitness, and uses various species to illustrate principles rather than to extrapolate behavior directly (a strategy open to the charge of selectivity in the material), whereas neurochemists are happy to extrapolate, provided the biochemical, neurological, and behavioral identities seem sufficient
when allowance is made for species differences (but assuming such allowance can be made, see Legg). Similarly, the first section conveys an impression of researchers rather narrowly concerned with details of species similarities and differences in various types of associative learning, but reading the book as a whole left this reader in some doubt as to the extent to which associative learning in any species can properly be treated without regard to its evolutionary function and the actual neurobiological mechanisms involved. A discussant chapter directed at these and similar issues would have been extremely helpful. The intended readership of the book is not explicitly stated, but the summary of the contents offered above should indicate the scope of readership interest. It would be a handy volume for researchers at postgraduate level and above to dip into. Some of the more wide-ranging chapters could be recommended to honours undergraduates. The standard of production is relatively high, with few misprints. All but three of the contributors are at British University Departments; but “behaviour” is spelled “behavior” throughout. J. M. Elliott Department of Psychology University of Sheffield Shefleld SIO 2TN England
This volume collects together in one place a number of papers dealing with the problems of extrapolation from animals to man. It is organized in three sections, each rather different in character. Although the first section is entitled “Conceptual and Historical Issues,” conceptual problems are discussed throughout the book, and this section is in fact a set of six papers dealing mainly with the usefulness or otherwise of the rat as a model of human associative learning. Most authors are rather critical of the assumption of ready extrapolation from this or other species to man. Thus, Miles notes that the language we use for many human concepts, such as ‘responsibility’, is applied only reluctantly if at all to animals. He does not, however, offer any criterion by which we should judge when to accept and when to reject the promptings of linguistic propriety as a guide to real species differences. Boakes suggests that although J. B. Watson thought the environment was important, he was reacting against an unthinking hereditarianism rather than making a strong claim for environmentalism as such. He then argues that nevertheless a consequence of this may have been the diversion of psychology into an excessive concentration on conditioning in animals at the expense of what might actually be important environmental influences on behavior. Boakes might also have said, but has left it to the contributors in section two, that another consequence has been for psychology to disregard the natural history and evolutionary significance of behavior. Blackman offers a spirited defense of the conventional behavior analyst’s argument that animal behavior be explained without re-
course to cognitive terms extrapolated from humans. As his examples make clear, this amounts to asserting that descriptions of necessary and sufficient environmental conditions for behavior, if found, constitute an explanation and are not themselves in need of some further level of explanation. This approach allows explanation inductively, but it severely limits the applicability of animal models if it is accepted that human behavior may actually not be wholly explicable in similar terms. Blackman argues against extrapolating explanatory cognitive constructs from man to animals, but his view on extrapolation the other way is not clear. Catania, too, seems to think that the important thing is to describe behavior in terms of antecedentconsequent relationships but leaves the nature of explanation rather obscure. Lowe and Davey both argue that there are qualitative differences between human and animal learning, claiming that the way humans “label” or “describe” environmental events (Lowe) or “restructure knowledge about the UCS” (Davey) affects their responses. The second section is entitled “Evolutionary Aspects” and comprises live chapters discussing the implications of evolutionary theory for attempts to extrapolate from animals to man, and also the way in which it informs the sorts of inquiry that are appropriate. Plotkin argues at length that comparative studies of learning are of limited value without a conceptual framework, by which he means a theory of the biological function of the behavior under consideration. If correct, his thesis preempts much of the preceding section. Odling-Smee offers an 123
Ethology and Sociobiology 6: 123-125 (1985) 0 Elsevier Science Publishing Co., Inc., 1985 52 Vanderbilt Ave., New York, New York 10017
0162-3095/85/$03.30
124
elaboration of his theory of multiple levels in evolution as a conceptual resolution of the naturenurture issue in behavior. His work is dealing with evolutionary models rather than animal models as such. Smith, in a review of the sociobiology of reproduction and child care in man makes it pretty clear that whereas the ideas derived from animal studies in this field have been useful in generating research, the predictions from animal models are only weakly supported, and he concludes that evolutionary biology cannot account for behavior in contemporary societies without additional assumptions about cultural evolution. He does, however, feel that some aspects at least of parent child behavior do tend to maximize inclusive fitness. Evidently the development of distinct cultural systems may not be independent of selection pressures. Blurton Jones shows even more clearly how assumption from evolutionary biology serve to structure questions about human behavior and illustrates the point with accounts of two diverse studies. He is concerned with the question of whether cultural activities and learning mechanisms actually do tend to improve inclusive litness, and concludes that they may. He also tackles head on the objections raised against applying sociobiology to man. He further provides an object lesson in the principle that it is not the stability of a behavioral measure but what it means that is important. Blurton Jones writes with great clarity, though with perhaps a hint of irritable defensiveness. The net conclusion from this section seems to be that as soon as the adaptive function of behavior is considered, the validity of particular cross species comparisons declines sharply and has to be qualified by a grasp of the function, but that the use of animal behavior in an evolutionary framework to suggest functional hypotheses is valuable. “Neurobiological AsThe final section, pects,” contains a further seven chapters examining the nature of extrapolation in neurobiology, including several dealing with aspects of the neurochemistry of behavior, together with a scathing review of the way simple analogies have apparently inspired a number of psychosurgical procedures (Carroll and O’Callaghan). It is difficult to disagree with the latter authors when they suggest that without convergent indications from human studies, animal studies alone should not stand as the basis for psychiatric intervention (especially when irreversible).
Green, in the next chapter, is principally concerned with schizophrenia and comes to the unsurprising conclusion that it is not much use looking in animals for behavioral analogues to a condition primarily characterized by disorders of thought and affect. However, he thinks it important to understand the dopamine system, and argues that animals may be useful here. In so doing he implicitly follows a logic of extrapolation explicitly laid out for neurochemical work by Warburton. Green also discusses the psychosurgical procedures once used for schizophrenia, an overlap with the’preceding chapter that could properly have been edited out. One really does not need to be told twice that the founder of prefrontal lobotomy was shot by one of his own patients in 1944. The editor might also have intervened in the chapters by Legg and Oakley. Legg argues at some length that neither the principle of encephalization-translation of brainstem functions to the neocortex with advancing phylogenetic status-nor that of “conservation,” which in this context means the retention by homologous brain structures of their functions in the face of phylogenetic advances, can meet the data on cross species comparisons of visual function without circularity. Inter alia he notes that, “Since the structures implicated in residual [destriatel vision . . . . are all brain stem regions, the principle of conservation would lead us to expect that they should operate similarly in all mammals, but this assumption is directly contradicted by the data” (p. 229). Oakley, immediately following, seems rather more taken with the principle of conservation: “It now seems evident from the use of comparable testing procedures across species that all mammals with striate cortex removed show similar visual abilities” (p. 248). Legg and Oakley, it may be noted, do not cite a single reference in common. This is probably because the main substance of Oakley’s chapter is a claim that subcortical mechanisms can be important in associative learning, and that the study of mammalian subcortical function is useful in establishing this conclusion. The chapters by Rick and Benton deal with fairly specific areas of neurochemical research. Rick argues that rats injected with anti-ganglioside antibody may show some structural and behavioral analogies with human mental retardates, and Benton argues on conceptual and methodological grounds that the relationship between testosterone and aggres-
125
sion has been exaggerated in humans despite animal studies suggesting a rather strong effect. It is certainly valuable to have this material collected together in one place, but this is, on the whole, a collection of discrete and self-contained contributions. There are some omissions. Almost nothing is said of the use of primates as models, for example in studies of communication or of social development, nor is there any discussion of classical ethology and attempts to extend it to man. Some authors, notably Catania, Plotkin, Benton, Legg, and Warburton have made the general relevance of animal models a relatively central concern of their chapter, but others have raised questions of animal models in the context of rather specific pieces of research. Authors generally give an impression of not having talked much to one another. Consequently, some issues arise as it were interstitially, and are only apparent to the reader by a comparison of the various articles. For example, the idea of animal models evidently means quite different things in different sections, and the criteria by which it should be evaluated vary accordingly. Thus evolutionary theory is now concerned with ideas of inclusive fitness, and uses various species to illustrate principles rather than to extrapolate behavior directly (a strategy open to the charge of selectivity in the material), whereas neurochemists are happy to extrapolate, provided the biochemical, neurological, and behavioral identities seem sufficient
when allowance is made for species differences (but assuming such allowance can be made, see Legg). Similarly, the first section conveys an impression of researchers rather narrowly concerned with details of species similarities and differences in various types of associative learning, but reading the book as a whole left this reader in some doubt as to the extent to which associative learning in any species can properly be treated without regard to its evolutionary function and the actual neurobiological mechanisms involved. A discussant chapter directed at these and similar issues would have been extremely helpful. The intended readership of the book is not explicitly stated, but the summary of the contents offered above should indicate the scope of readership interest. It would be a handy volume for researchers at postgraduate level and above to dip into. Some of the more wide-ranging chapters could be recommended to honours undergraduates. The standard of production is relatively high, with few misprints. All but three of the contributors are at British University Departments; but “behaviour” is spelled “behavior” throughout. J. M. Elliott Department of Psychology University of Sheffield Shefleld SIO 2TN England