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The Cognitive Animal Empirical and Theoretical Perspectives on Animal Cognition

© 2002 Massachusetts Institute of Technology

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There are as many approaches to studying ani- and The Expression of the Emotions in Man and mal cognition as there are definitions of cogni- Animals (1872). Consequently, both disciplines tion itself. This diversity is reflected in the essays are almost inextricably linked to the concept of that follow, to a degree that we believe is un- instinct. Darwin viewed instinct primarily in be- paralleled in any other volume that has been havioral terms and considered his ability to ex- produced on this subject. This diversity is philo- plain instinct through natural selection to be one sophical and methodological, with contributors of the most critical tests of his theories. Thus he demonstrating various degrees of acceptance or compared closely related species of bees to ex- disdain for terms such as ‘‘consciousness’’ and plain the evolution of hive building and closely various degrees of concern for the rigors of lab- related species of ants to explain the origins of oratory experimentation versus the validity of slave making. He also focused on domesti- naturalistic research. The diversity is also appar- cated species to show how human intervention ent in the wide range of species to be found be- and selection could have formed variants. His tween these covers. Of course there are many prime examples here were di¤erent breeds of chapters on primates, and especially the great dogs and pigeons. Soon, zoologists such as C. O. apes, reflecting our human-centered interest in Whitman, E. C. Poulton, Oskar Heinroth, Julian our closest relatives, but there are also chapters Huxley, and others began to exploit the com- that touch on cognition in animals as diverse as parative method to trace the evolution of even earthworms, antelopes, dogs, spiders, dolphins, the most complex social displays of animals. bees, fish, hyenas, snakes, sea lions, prairie dogs, This work inspired the development of ethology virtual organisms, parrots, rats, ravens, and primarily through the research and promotional squirrels, to name but a few. We think we have e¤orts of the Nobel laureates Konrad Lorenz produced one of the most wide-ranging menag- and Niko Tinbergen. eries of scientific studies of animal cognition ever Although when the Origin appeared in 1859 assembled. Yet it is humbling to realize that in Darwin deliberately discussed neither the evolu- terms of the diversity of life itself we cannot tion nor the behavior of human beings, he did, claim to have even scratched the surface. near the very end of the book, include this intri- One of our objectives in bringing such a di- guing passage: verse collection of research studies together is to show that whatever the ideological di¤erences, In the distant future I see open fields for far more im- portant researches. Psychology will be based on a new behavioristic psychologists and cognitive etholo- foundation, that of the necessary acquirement of each gists have a lot to learn from each other and mental power and capacity by gradation. Light will be from the neurosciences. Yes there are di¤erences thrown on the origin of man and his history. (Darwin of opinion about how to pursue the study of an- 1859, p. 488) imal cognition, but all sides stand to gain from listening carefully to the concerns of others. De- Note that this passage considers the psychologi- spite the di¤erences, there is in fact a great deal cal aspects of evolution as far more important, in of similarity among the di¤erent approaches, for the long run, than the morphological and taxo- they have, after all, evolved from the same start- nomic issues to which the Origin was largely de- ing point in the theory of evolution by natural voted. Most biologists have ignored the clear selection. evidence, found in Darwin’s early notebooks, The roots of both comparative ethology and that while Darwin may have loved natural his- comparative psychology are found in the writ- tory, his ultimate agenda was to explain the ings of Charles Darwin, particularly in the Ori- origin of perhaps the strangest species of all, gin of Species (1859), The Descent of Man (1871), ourselves. Thus, in his later writings Darwin

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applied the comparative method to show the 1984). This focus on parsimonious and fairly possible origins of facial expressions in human simple mechanisms to explain diverse abilities beings and speculated about many other aspects had salutary e¤ects in terms of systematizing of abilities typically considered to be uniquely quantitative research methodology and inter- human, such as conceptual thought, language, pretation when the field was young (see Boakes loyalty, love, and morality (the latter was to 1984). Darwin’s cousin, Francis Galton, was Darwin a social instinct). In this way Darwin among the first to develop quantitative and sta- sought to show that our di¤erences from other tistical means to study behavior. Nevertheless, species were not major qualitative leaps, but by the 1940s, the study of comparative cogni- were based on quantitative change that was due tion, especially in psychology in the United to basic evolutionary processes. Furthermore, States, had narrowed greatly; field and natural- these di¤erences in degree, not kind, were not istic research was disparaged and many impor- due to some external supernatural intervention or tant problems were ignored. The seminal volume a nonmaterial mind. Darwin thus tried to docu- of this period is useful in showing the stated ment that human behavior contained instinctive breadth and e¤ective narrowing of comparative roots that could be traced to our animal ances- psychology (Warden et al. 1935). These authors tors while at the same time arguing that the con- thoroughly criticized early work that was not ceptual, communicative, intellectual, emotional, based on laboratory experimental paradigms and social, and moral aspects of our behavior also concluded that most of it was virtually worthless, had roots in the behavior and psychology of being contaminated by anecdotal, anthropomor- other species. phic, and introspectionist errors. These later writings by Darwin greatly influ- Furthermore, few biology or zoology depart- enced the comparative psychology of the nine- ments incorporated behavior as an important teenth century. Many early writers, going back element in their scientific training and research. to Aristotle, Pliny, and even earlier, had com- European ethologists, with the primary excep- pared the abilities of people with those of other tion of von Uexku¨ll, von Frisch, and Lorenz, animals and pointed out the surprising abilities ignored the study of the higher levels of cogni- of the latter. However, it was Darwin who sys- tion and were especially suspicious of studying tematically set out to show that the gap between consciousness and subjective states. Tinbergen humans and other animals was smaller than pre- (1951) was adamant on this point, almost cer- viously thought, and even more important, how tainly because of the lingering vitalism con- the gap could be bridged by natural selection cerning instinct and mind found in so many and su‰cient time. Darwin’s notebooks from the scientists, including his countryman Birrens de 40 years preceding The Descent of Man showed Haan. Concerning play behavior, Tinbergen how influenced he was by the similarities of mon- (1963, p. 413) wrote: ‘‘Concepts such as play and keys and apes to human beings. learning have not yet been purged completely Darwin and his prote´ge´ in comparative from their subjectivist, anthropomorphic under- psychology, George John Romanes (see, e.g., tones. Both terms have not yet been satisfactorily Romanes 1883, 1892) often relied on casually defined objectively, and this might well prove collected anecdotes and an uncritical anthropo- impossible. . . .’’ morphism that troubled more experimentally However, the seeds of the conceptual and sophisticated scientists. Soon, the study of ani- methodological tools necessary to reexplore the mal intelligence and abilities was shaped by complex lives of animals were already in place. the writings and work of C. Lloyd Morgan and Wallace Craig (1918) had shown how to separate E. L. Thorndike (Burghardt 1985a,b; Dewsbury the motivational and cognitive aspects of instinct

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as well as the importance of sequential analysis, animal learning and comparative psychology primarily in his distinction between appetitive were ignoring the really interesting phenomena behavior and consummatory acts, a distinction presented by animals as a result of their narrowly now finally being formally recognized and ex- focused laboratory studies of a few selected tended in comparative cognition, as many chap- domesticated species. ters in this book acknowledge. Von Uexku¨ll Ethology, on the other hand, was falling under (1909/1985) gave us the concepts of Umwelt, the influence of behaviorists and ignoring the Innenwelt, counterworld, sign stimulus, search cognitive implications of the flexible and diverse image, and other useful means for approaching accomplishments their studies of animals were the behavior of other species from their per- uncovering. For Gri‰n, cognitive ethology was ceptual worlds and not ours. Von Frisch was a not only the study of the complex natural be- brilliant experimenter who, among other accom- haviors of diverse species, it was also the study of plishments, showed that fish could hear, honey- taboo subjects such as consciousness and aware- bees could see colors, and, most amazingly, that ness in other species. A cognitive ethology that bees could communicate information about dis- ignored consciousness, choice, deliberation, plan- tant resources, which virtually no other species, ning, intentions, and other mental processes was, except people, could accomplish. for Gri‰n, not possible. Then, at around the beginning of World War The Question of Animal Awareness produced II, other discoveries were being made that would ‘‘Amens’’ in some quarters and outrage in have consequences. One of these was Baerends’ others. Experimental psychologists in particular demonstration that wasps could assess and hold felt that they had developed methods that could in memory for a day or more how many cat- answer the important questions without encour- erpillars they needed to provision a nest. This aging unbridled speculation that they, like Tin- study, presented in a dissertation, was made bergen, felt could never be assessed through the widely known by Baerends’ professor, Niko Tin- methods of normal science. Nonetheless, in psy- bergen, in the seminal volume that brought Eu- chology a new cognitivism was displacing be- ropean ethology to English-speaking scientists haviorism as the dominant approach and already (Tinbergen 1951). The tidy view that vertebrates, psychologists studying animal learning were in- especially mammals, were the most cognitively corporating cognitive approaches while still using advanced animals was in e¤ect being challenged standard laboratory methods, although they were by the bees and wasps. applied to more cognitive topics such as concept Also around 1940, Donald Gri‰n, as a stu- formation, information retrieval, and memory. dent, co-authored a remarkable paper proving The rich history of behaviorism in fact in- that bats used echolocation to navigate in their cluded such concepts put forth by E. C. Tolman environment and locate prey. Decades after this in the 1930s as cognitive maps and latent learn- work, Gri‰n finally integrated much ethological ing, which were eventually applied to in animal work, together with the implications of the abili- learning (e.g., the radial arm maze) as well as ties uncovered by experimental work on bats cognitive psychology and ethology. By the 1980s and insects, the results of studies on chimpanzee the behavioral concepts and methods of Hull communication, and other diverse information and especially Skinner, which were largely codi- in a remarkable small book, The Question of An- fied in the 1940s, were increasingly applied by imal Awareness: Evolutionary Continuity of Men- psychologists to issues of interest to ethologists, tal Experience (Gri‰n 1976). Gri‰n advocated a behavioral ecologists, and neuroscientists. new field, to be called ‘‘cognitive ethology’’ (see Computer scientists, robotics engineers, and Gri‰n 2001), because he thought that traditional modelers also became interested in the problems

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faced by those working with real animals, and philosophers such as Peter Singer and Tom neuroscientists began exploring more naturalistic Regan, among others, raised issues concerning phenomena, such as face recognition, as well. our treatment of animals, increasing the stakes The results of all these broadened perspectives and relevance of obtaining accurate information are on display in this volume. Nevertheless, there on the emotional and cognitive lives of other remain real di¤erences, many of them high- species at the individual level and challenging lighted in this volume. Some authors are willing, philosophers to pay more attention to evolu- even anxious, to enter the cognitive world and tionary relationships among species. At the same conscious life of their animal subjects. Others time, ecological and environmental concerns explicitly or by omission are clearly uncomfort- arose that involved extinction and habitat de- able with a cognitive study of nonhuman ani- struction. Accurate knowledge of the complex mals that attempts to explore subjective states, behavior and needs of animals became seen as intentionality, consciousness, or self-awareness. imperative for their very survival in nature or in And, if the science holds, such diversity is both captivity (Beko¤ 1998, 2002). appropriate and defendable. Today, all the issues and concerns outlined In addition, philosophers became interested above swirl over, within, and below seemingly in developing naturalistic approaches to age-old straightforward questions about the cognitive questions about the nature of mind and thereby lives of animals. The way we answer such ques- came to face the challenge that nonhuman ani- tions and apply the findings may tell us much mals presented to the traditional anthropocentric about ourselves as a species as well as having philosophy of mind. Ethology, animal learning serious consequences for the other inhabitants of and communication, and improved understand- this planet. Thus, the way in which the research ing of the workings of the brain promised some issues addressed in this volume are eventually advances in understanding everything from resolved (or not resolved) has enormous political mind–body relationships to the nature of lan- implications at many levels. This is probably guage, intentionality, ethics, and knowledge it- something most of those working on cognitive self. Tom Nagel, Daniel Dennett, Steven Stich, aspects of animal behavior are aware of at some John Searle, Ruth Millikan, Jerry Fodor, Dale level, but typically do not address in their formal Jamieson, Fred Dretske, and many other philos- writings. ophers, despite often remaining wed to tradi- With the above thoughts as a backdrop, we tional philosophical methods of reflection on now can turn to the book at hand. The editors thought experiments, became fellow travelers bring considerable and diverse research back- with scientists who were studying animal cogni- grounds and perspectives to this project. Marc tion in both the laboratory and the field. Beko¤ was trained in neurobiology and be- For many years, the primary philosophers havior, later as an ethologist, and has worked to whom experimental psychologists were in- on comparative aspects of behavioral develop- troduced were the positivists and Ludwig Witt- ment, social communication, quantitative meth- genstein, while ethologists typically ignored all ods, play, animal ethics, and cognitive ethology philosophers (except for Lorenz and von Uex- for many years, with a particular focus on mam- ku¨ll, who were very partial to Kant). Evolution- malian carnivores. In 1978 he and Gordon ary biologists were largely influenced by Karl Burghardt edited a volume on behavioral devel- Popper. opment and evolution (Burghardt and Beko¤ These trends changed as both cognitive ethol- 1978). Colin Allen was trained as a philosopher ogy and sociobiology became popular in the and has written extensively on topics in lan- mid-1970s. Also, at this time the writings of guage, communication, play, and mental evolu-

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tion. He has conducted fieldwork on the behavior voices and not the tight control or limited cov- of birds with Marc Beko¤ and has studied erage found in many edited books. learning in domestic pigs. Gordon Burghardt Because this volume did not begin as part of a was trained in an interdisciplinary biopsychology conference or symposium, we were able to invite program and has focused on the comparative a far larger and broader set of authors, both geo- behavioral development of squamate reptiles and graphically and disciplinarily, than is typically the complex interactions between genetic and found in a single book, certainly in a volume on environmental factors, as well as on play, com- this topic. The result is a book about which we munication, and the intertwined history of ethol- are very excited, a volume that could easily be ogy and psychology. used in advanced undergraduate and graduate The Cognitive Animal contains fifty-seven courses. It can serve as a useful and fascinating compact essays dealing with numerous di¤erent introduction to modern studies and provides topics on a wide range of organisms by re- examples for those with a general interest in an- searchers in many disciplines. The contributors imal cognition. For the professional, it provides were asked to consider five questions in their an update to work from approaches other than essays. These were (1) What are the central re- that in which he or she works. Thus there are search questions in your study of animal cogni- chapters by cognitive ethologists, behavioral ecol- tion? (2) What theoretical or empirical methods ogists, experimental psychologists, behaviorists, have enabled you to address those questions? (3) philosophers, neuroscientists, developmental psy- What, if anything, do your results reveal about chologists, computer scientists and modelers, field the internal psychological states of animals? (4) biologists, and others. What future work is suggested by your inves- The potpourri of topics covered in this book tigations? (5) What phylogenetic and method- resists an easy or concise summary. We have ological limits are there to the study of animal chosen to keep this introduction brief rather cognition? Some authors stuck to these ques- than attempting to provide a beginner’s guide, tions more explicitly than others, but generally for our best advice to beginners is to just dive the essays address these themes. Also, when in. Each chapter is short enough that it may taken together, the essays reflect evolutionary, be read quickly and easily reread when con- ecological, and comparative approaches to the trasting views are encountered in other chapters; problems at hand. The importance of careful in this way understanding can be progressively observation, description, and experiments is em- deepened. The topics include (but are not limited phasized; all are important. to) definitions of cognition, the role of anecdotes Since our contributors are all well-respected in the study of animal cognition, naturalizing the researchers, we allowed them considerable lati- study of animal cognition, anthropomorphism, tude in their presentations, realizing that some attention, perception, learning, memory, ecology, readers might prefer some of the chapters to evolution (including discussions of di¤erent lev- provide more citations to the available literature els of selection), communication, reproduction, and to be written in a more formal tone, while thinking, consciousness, intentionality, rational- other readers might prefer the opposite. As ity, play, aggression, dominance, predation, par- editors, we read each essay carefully and often ent–young interactions and care giving, the role o¤ered organizational, stylistic, conceptual, and of models, planning, anticipation, kin selection, historical suggestions, especially those that would cooperation, recognition, assessment of self and aid readers with di¤erent perspectives. However, others, neuroethology, choice, social knowledge, mindful of our own involvement in the issues the role of touch, empathy, social symmetries covered in this book, we encouraged diverse and asymmetries, and conflict resolution.

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Even a cursory glance at the essays makes it 4. Self and other: the evolution of cognitive obvious that in some cases there are di¤erent cooperators (from self-recognition to social views on the same or similar topics, an aspect of cognition) The Cognitive Animal with which we are very pleased. Pluralism is a good route to take, given Narrower categories would have separated con- the state of the art. Behaviorism is alive and well, tributions that are clearly relevant to one an- as are rich cognitivism and more middle-of-the- other. But even within the broad categories we road perspectives, and all of these approaches ended up with, there are chapters that might need to be given serious attention in future re- reasonably be paired with others in di¤erent search. We also hope that these chapters will aid categories. For instance, many approaches to in the development of more understanding and animal concepts or to social cognition involve reduce the polemics among all those who strive language or communication, and many of the to elucidate the minds of animals. chapters on communication touch upon catego- Given the rich and wide variety of these rization or concept formation. Furthermore, all essays, the task of organizing them in a linear the chapters provide a theoretical perspective, sequence necessarily involved compromises. Two but those in the first section tend to be more ex- main lines of organization suggested themselves: plicit in raising basic and general theoretical topical and methodological. We opted for a questions. The fact that this organizational task topical organization because we found that re- was so di‰cult, and that there are so many over- searchers working on similar topics from a va- lapping themes even among researchers with riety of di¤erent methodological backgrounds very di¤erent approaches, speaks to the interdis- have many interesting things to say to each ciplinary nature of cognitive studies of animals. other. Ultimately, people (particularly potential We hope that these essays stimulate us all to be students) are interested in the field of animal be- more interdisciplinary and to rise to challenges havior because of what animals do, not because to try to answer, and not to dismiss, di‰cult of the methods that we use to study them. How- questions. Di‰cult does not mean impossible. ever, we also believe that the importance of Learning more about the cognitive capacities of thinking about those methods is highlighted by other animals will inform not only the general putting contrasting approaches to similar ques- topic of animal cognition, but also how we in- tions alongside each other. teract with and treat other animals. The degree We also chose to organize the essays along of development of cognitive skills and assess- broad rather than narrow topical lines. We ments of intelligence, along with information found that the chapters could be grouped into about the emotional lives of animals, are being four categories under the following headings increasingly used to make informed decisions (with brief explanations): about the use of animals in research and educa- tion, and for entertainment and food. 1. The diversity of cognition (taxonomic, meth- We thank all of our contributors and The odological, and theoretical) MIT Press for working with us with enthusiasm 2. Concepts and categories (ways in which or- and dedication on this project. We are grateful to ganisms divide up the world) Mrs. Rosalie Glenn of the Department of Phi- losophy at Texas A&M University for her help 3. Communication, language, and meaning (a with printing and mailing. Don Gri‰n gra- possible window on animal minds, to borrow ciously agreed to contribute an afterword. Marc Gri‰n’s metaphor) Beko¤ thanks the University of Colorado for a

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sabbatical leave for the academic year 2000– Romanes, G. J. (1892). Animal Intelligence. 5th ed. 2001. Colin Allen acknowledges support in the London: Kegan, Paul, Trench, Tru¨bner. form of a Texas A&M Faculty Development Tinbergen, N. (1951). The Study of Instinct. Oxford: Leave during 2000–2001, and a Big 12 Faculty Clarendon Press. Fellowship in 1999 that enabled a trip to the Tinbergen, N. (1963). On aims and methods in ethol- University of Colorado when this book was ogy. Zeitschrift fu¨r Tierpsychologie 20: 410–433. conceived. Uexku¨ll, J. von (1909/1985). Environment (Umwelt) and the inner world of animals (C. J. Mellor and D. Gove, trans.). In The Foundations of Comparative References Ethology, G. M. Burghardt, ed., pp. 222–245. New York: Van Nostrand Reinhold. (Reprinted from J. von Beko¤, M. (ed.) (1998). Encyclopedia of Animal Rights Uexku¨ll, 1909, Umwelt and Innenwelt der Tiere. Berlin: and Animal Welfare. Westport, Conn.: Greenwood. Jena.) Beko¤, M. (2002). Minding Animals: Awareness, Emo- Warden, C. J., Jenkins, T. N., and Warner, L. H. tions, and Heart. New York, Oxford University Press. (1935). Comparative Psychology: A Comprehensive Boakes, R. (1984). From Darwin to Behaviorism: Psy- Treatise. Vol. 1, Principles and Methods. New York: chology and the Minds of Animals. Cambridge: Cam- Ronald Press. bridge University Press. Burghardt, G. M. (1985a). Animal awareness: Current perceptions and historical perspective. American Psy- chologist 40: 905–919. Burghardt, G. M. (ed.) (1985b). Foundations of Comparative Ethology. New York: Van Nostrand Reinhold. Burghardt, G. M. and Beko¤, M. (eds.) (1978). The Development of Behavior: Comparative and Evolution- ary Aspects. New York: Garland. Craig, W. (1918). Appetites and aversions as con- stituents of instincts. Biological Bulletin 34: 91–107. Darwin, C. (1859). On the Origin of Species. London: Murray. Darwin, C. (1871). The Descent of Man and Selection in Relation to Sex. London: Murray. Darwin, C. (1872). The Expression of the Emotions in Man and Animals. London: Murray. Dewsbury, D. A. (1984). Comparative Psychology in the Twentieth Century. Stroudsburg, Pa.: Hutchinson Ross. Gri‰n, D. R. (1976). The Question of Animal Aware- ness: Evolutionary Continuity of Mental Experience. New York: Rockefeller University Press. Gri‰n, D. R. (2001). Animal Minds: Beyond Cognition to Consciousness. 2nd ed. Chicago: University of Chi- cago Press. Romanes, G. J. (1883). Mental Life of Animals. Lon- don: Kegan, Paul, Trench, Tru¨bner.

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