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A Behavioral Biology for the Future

2012, Ethology

Some comments are appended to Patrick Bateson's reflections on the future of behavioral biology that were triggered by remembering the contributions of Gu ¨nter Tembrock to ethology. While the suggestions made are valid and insightful, a few specific areas where exciting research possibilities may reside are added including those involving communication and language, culture and ritual, brain imaging, genetics, animal tracking, and aesthetics. The need for the historical grounding of graduate education in behavior studies is also emphasized as an important possible factor if the future will effectively fulfill its mission and recognize its legacy.

Ethology COMMENTARY A Behavioral Biology for the Future (Commentary on Pat Bateson – Behavioural Biology: The Past and a Future) Gordon M. Burghardt Departments of Psychology and Ecology & Evolutionary Biology, University of Tennessee, 1404 Circle Drive, Austin Peay Building, Knoxville, TN 37996, USA Correspondence: E-mail: [email protected] doi: 10.1111/j.1439-0310.2012.02024.x Abstract Some comments are appended to Patrick Bateson’s reflections on the future of behavioral biology that were triggered by remembering the contributions of Günter Tembrock to ethology. While the suggestions made are valid and insightful, a few specific areas where exciting research possibilities may reside are added including those involving communication and language, culture and ritual, brain imaging, genetics, animal tracking, and aesthetics. The need for the historical grounding of graduate education in behavior studies is also emphasized as an important possible factor if the future will effectively fulfill its mission and recognize its legacy. Pat Bateson (2012) has written a moving tribute to the often neglected writings of Günther Tembrock, who died in 2011. While not having on my bookshelf the volumes cited by Pat, I do have his short textbook in German (Tembrock 1987). This book is a schematic, annotated outline that covers much contemporary research and is enlivened with many small, elegant drawings as well as copious graphs, charts, and quantitative analysis. Pat then devotes the first major part of his essay to the relationships among various fields including psychology, behavioral ecology, sociobiology, and ethology that seem to constitute his characterization of behavioral biology. This is well-worked ground that many have trod (e.g., Burghardt 1973; Dewsbury 1984). Tinbergen’s aims are described, which, perhaps unnecessary for Ethology’s readership, need to be continually reinforced as the pressures leading to splintering, specializing, and balkanization of scientific fields are ever present, and integrative and interdisciplinary approaches are too often honored more in words than in practice. It is on the prospects for the future that I want to focus on in my brief comments. Bateson mentions several issues including the need for a functional 222 approach, reciprocity between how and why questions, appreciating the increasingly sophisticated and complex epigenetic processes underlying behavioral plasticity as instruments of evolution, taking seriously three heterodox principles for studying behavior, understanding interactions among levels of analysis, and the need for interdisciplinary approaches that break out of intellectual silos. These are all important considerations for the future, clearly delineated, and I comment on a few of them as I propose some additional features that may characterize the future of behavioral biology. Bateson refers to the field of animal behavior as behavioral biology. This is a common term and is certainly broad and inclusive. The impetus seems to be that as animal behavior ⁄ ethology incorporated more explicit ecological, evolutionary, adaptationist, and other perspectives, behavioral biology was born. Actually, the term ‘behavioral biology’ can also include behavioral neuroscience, experimental analysis of behavior, psychopharmacology, neuroendocrinology, psychoimmunology, toxicology, psychobiology, biopsychology, cognitive neuroscience, behavioral medicine, evolutionary psychology, Darwinian anthropology, mathematical modeling, computer simulations, Ethology 118 (2012) 222–225 ª 2012 Blackwell Verlag GmbH G. M. Burghardt robotics, and many other often overlapping labels. While behavior questions are addressed in all these areas, we need to be careful as to how wide a net to cast. In my mind, what Pat seems most to want to include is Tinbergian ethology writ large: the naturalistic understanding of behavior from an evolutionary and developmental perspective using all available tools. Pat also points out, as have many others, that the simplistic notion that psychologists do experiments on behavior in often highly artificial settings while ethologists focus on observational studies of animals in naturalistic settings is misleading and outdated (c.f., Burghardt 1973; Dewsbury 1984; Hirsch 1987). Given that, where is the field going?, Pat wisely provides ‘a’ future rather than ‘the’ future and I will do likewise. Pat sees a greater reliance on a functional analysis, particularly one dealing with behavioral development and the ‘scaffolding’ approach that Pat has championed over the years. Here, we see a combination of the ethological aims of causation ⁄ mechanism, adaptive function, and ontogeny, all in the service of understanding the process and the pattern in evolution. Actually, I may have expanded this a bit beyond Pat’s claim, but I think it works. Understanding the reciprocal relationship between how and why (proximal and distal ⁄ ultimate) is essential. Epigenetics and the role of behavioral plasticity are certainly at the forefront of many biological studies these days. While Bateson’s ‘a future’ lists important components that I am sure will animate us for many years, I think that there are additional fields and questions that ethologists should ponder and be prepared to study. I discussed some of these in a retrospective paper on Charles Darwin’s legacy in psychology and ethology (Burghardt 2009). Besides enumerating ten major influences Darwin had on modern studies of animal behavior, I listed several additional ones that seem to highlight how far the students of animal behavior have come in the range of phenomena that are considered amenable to scientific analysis. As such, they are controversial, but so was the initial resurgence of interest in instinct and then in comparative animal cognition (i.e., ‘intelligence’). One of these is language and its characterization, origins, and evolution. Darwin pioneered communication research and early ethologists, and then birdsong researchers, developed the field with conceptual and empirical work. Although some links with human style language were explored in the 70s by the Gardners and others working on chimpanzees and by Peter Marler and others working on birdsong (e.g., Marler 1970), it is only in recent years that the Ethology 118 (2012) 222–225 ª 2012 Blackwell Verlag GmbH Future of Behavioral Biology animal communication ⁄ human language nexus has fledged into a vibrant area with Alex and other gray parrots (cited by Bateson), the extensive research emanating from the Hailman and Ficken studies on chickadee song complexity and flexibility (Hailman et al. 1987), and the re-engagement of linguistics and animal communication scholars (e.g., Hauser et al. 2002). Culture in human and non-human animals is also a vibrant growing field fertilized by the studies on the dialect learning in birds and tool-use diversity in apes along with the seminal writings on geneenvironment aspects of cultural evolution (Boyd & Richerson 1985). The parameters and limits of social learning, imitation, copying, and other phenomena in diverse species are uncovering examples from many species including reptiles, fish, etc. Although cited in a different context by Pat, the edited book by Laland & Galef (2009) highlights many of the current contributions and controversies on animal culture. These will stimulate our field for some time. Some years ago, I suggested that we need to amend Tinbergen’s four aims with a fifth one, the aim of assessing the experiential life of the animal or what I termed ‘private experience’ (Burghardt 1997). There were several reasons for such a move. One of the limitations of Tinbergen (1963) was that he was so reluctant to being associated with anything ‘subjective’ that he even disparaged the study of play, although Pat himself has been a major contributor to its study! Another incentive for this move was the increasing sophistication of brain imaging in alert functioning animals as well as the need to address the issues of animal welfare and treatment. Influenced by Jakob von Uexküll’s concepts of the Umwelt and Innenwelt, I argued that if ethologists and other behavioral biologists did not consider such methods and related questions, incorporating the perspective of ‘the other’ and not just their overt behavior, we would be missing out on exciting research opportunities and questions. Work integrating brain activity, including mirror neurons, gene expression, and chemical transmitters is transforming our understanding of brain–behavior interactions underlying the normal behavior and the social emotions as well (Panksepp 1998). And there is more! Morality, fairness, ethics, social reciprocity, and other questions are now being asked of non-human animals in many ingenious (and sometimes questionable) ways (de Waal 2006). The book by Bekoff & Pierce (2009) discusses much of this research carefully compiled from field and captive studies in numerous species, not just non223 Future of Behavioral Biology human primates. Similarly, aesthetics and beauty including the arts, music, etc. are being explored, and not just from the sexual selection angle. I also suggest that the field of ritualization, so dear to many of the early ethologists, is ripe for reinvigorating links to the processes underlying religion, play, sexual selection, and behavioral development (e.g., Feierman 2009). Besides addressing the neuroscience aspects of behavior, the increasing use of genetic methodology for everything from paternity analysis to individual recognition to analyzing the diet of the predators to remotely triggered or monitored webcams, etc. offers loads of opportunities for path-breaking findings in understanding hitherto enigmatic behaviors and phenomena. The miniaturizing of tracking devices, camcorders, and use of satellites are changing our perception of animal behavior, dispersal, and migrations. In short, the widespread availability and the ease of use of modern technology are truly exciting, which helps to gather knowledge on the private lives of animals in hitherto unthinkable ways. Used carefully, these methods can do lots of ‘gee whiz’ work with YouTube clips the mark of success. And yet, I think that these studies, and the people who carry them out, need to relate back to larger questions in understanding behavior as well as contribute to conserving the increasingly endangered subjects of our love and scientific interest. Finally, I will comment on the Bateson’s ‘three principles,’ put-ons with a grain of truth. Certainly, the first, about treating animals as humans, is traditionally one of the first sins students are taught about in most animal-behavior courses. Anthropomorphism is a sin, but not all anthropomorphic thinking is equivalent. In fact, critical anthropomorphism is most useful in moving into the private experience realm and in devising experimental questions. Many insightful and creative students of animal behavior have done this, although they rarely acknowledged doing so in published papers (Burghardt 1985, 1997). Nevertheless, anthropomorphism is the focus of many recent studies and books. The second principle, being suspicious of simple explanations, was certainly shown by the attempts of experimental psychology theorists to squeeze all learning, for example, into Pavlovian and ⁄ or operant conditioning or even old line associationism. Thus, illness-induced aversions, imprinting, song learning, and other phenomena were so treated, often harshly in the literature. Many of Don Griffin’s claims about animal awareness met the same fate. Many did not jump on the optimal foraging theory way of looking 224 G. M. Burghardt at animal feeding behavior, on game theory (hawks and doves) for understanding animal fighting, or on kin selection for understanding mating systems because the initial predictions, while elegant and even remarkably accurate, seemingly failed to capture the complexity of behavior. Nonetheless, all these efforts, by forcing quantification and conceptual explication of terms, were extremely useful in advancing the field, and the early simple models and the conceptions could be (sometimes reluctantly) discarded. However, the third principle, the Oxford Principle, about using teleological, anthropomorphic, and folk psychology labels for the actions of genes, bacteria, and animals is, I believe, far more dangerous than Pat allows (Burghardt & Bekoff 2009). This is because speaking about insulting the flag or thermostats seeking their set point implies intentions that no one takes seriously. But when one refers to animals and the need to understand mechanisms that may indeed be intentional, deliberate, emotional, or conscious, the misuse of labels becomes a hindrance to good science. The old idea of teleonomic vs. teleological thinking should be more transparently employed as new generations of researchers come on line. In his last paragraph, Bateson refers to the excitement of behavioral biology, its essential links with the other fields, and the competitive nature of science today in fashioning the future. These characteristics, combined with the richness and the wealth of opportunities now often readily available to amateurs and professionals alike, suggest the need to ponder carefully the education that new graduate students going into this field receive. Will they be trained in at least some aspects of all four of Tinbergen’s aims, let alone the fifth aim? Will they, besides learning the latest findings in ecology, molecular genetics, and phylogenetic programming, also learn the details of experimental design, reliability testing, how the receptors and the central nervous system work, the expanding purview of behavior genetics, the intricacies of animal communication, and so forth? More importantly, will they have the opportunity to study and think through the seminal contributions of the writers in our field since the time of Darwin – including the writings by Bateson himself? If students focus only on the latest research studies, read only work from the last decade or so, and are insulated from the larger issues that motivated the early workers and seem to constantly cycle back, then the future will be composed of fits and starts, rediscoveries and priority claims, overlooked Ethology 118 (2012) 222–225 ª 2012 Blackwell Verlag GmbH G. M. Burghardt opportunities, and other misadventures. I fear that it will not only be Tembrock whom no one will read and may be forgotten, but the entire larger history of research and ideas out of which our current understanding has evolved. Fortunately, the existence of ‘virtual’ libraries of books and journals going back hundreds of years makes recovering the past more accessible than ever for those with the desire to do so. Literature Cited Bateson, P. 2012: Behavioural biology: the past and a future. Ethology 118, 216—221. Bekoff, M. & Pierce, J. 2009: Wild Justice. Univ. of Chicago Press, Chicago, IL. Boyd, R. & Richerson, P. J. 1985: Culture and the Evolutionary Process. Univ. of Chicago Press, Chicago, IL. Burghardt, G. M. 1973: Instinct and innate behavior: toward an ethological psychology. In: The Study of Behavior: Learning, Motivation, Emotion, and Instinct (Nevin, J. A. & Reynolds, G. S., eds). Scott, Foresman, Glenview, IL, pp. 322—400. Burghardt, G. M. 1985: Animal awareness: current perceptions and historical perspective. Am. Psychol. 40, 905—919. Burghardt, G. M. 1997: Amending Tinbergen: a fifth aim for ethology. In: Anthropomorphism, Anecdotes, and Animals (Mitchell, R. W., Thompson, N. S. & Miles, H. L., eds). SUNY Press, Albany, NY, pp. 254—276. Ethology 118 (2012) 222–225 ª 2012 Blackwell Verlag GmbH Future of Behavioral Biology Burghardt, G. M. 2009: Darwin’s legacy to comparative psychology and ethology. Am. Psychol. 64, 102—110. Burghardt, G. M. & Bekoff, M. 2009: Animal consciousness. In: Oxford Companion to Consciousness (Bayne, T., Cleeremans, A. & Wilken, P., eds). Oxford Univ. Press, Oxford, UK, pp. 39—43. Dewsbury, D. 1984: Comparative Psychology in the Twentieth Century. Hutchinson Ross, Stroudsburg, PA. Feierman, J. R. (ed.) 2009: The Biology of Religious Behavior. Praeger, Santa Barbara, CA. Hailman, J. P., Ficken, M. S. & Ficken, R. W. 1987: Constraints on the structure of combinatorial ‘‘chick-adee’’ calls. Ethology 75, 62—80. Hauser, M. D., Chomsky, N. & Fitch, T. S. 2002: The faculty of language: what is it, who has it, and how did it evolve? Science 298, 1569—1579. Hirsch, J. (ed.) 1987: Comparative Psychology – Past, Present, and Future. (special issue). J. Comp. Psychol. 101, 219—291. Laland, K. N. & Galef, B. G. (eds) 2009: The Question of Animal Culture. Harvard Univ. Press, Cambridge, MA. Marler, P. 1970: Birdsong and speech development: could there be parallels? Am. Sci. 58, 669—673. Panksepp, J. 1998: Affective Neuroscience. Oxford Univ. Press, New York. Tembrock, G. 1987: Verhaltensbiologie. Gustav Fischer, Jena. Tinbergen, N. 1963: On the aims and methods of ethology. Z. Tierpsychol. 20, 410—433. de Waal, F. 2006: Primates and Philosophers – How Morality Evolved. Princeton Univ. Press, Princeton, NJ. 225