information should be heavily influenced by its ecology. Neuroecology refers to the study of the neural mechanisms of behavior guided by functional and evolutionary principles. How do the evolutionary pressures for complex birdsong affect the evolution of the underlying neural substrate? How does having a large home range affect one’s ability to navigate? Does having to store food place selective pressure on spatial memory and its underlying brain regions? These functional and evolutionary approaches to the study of brain and cognition have come under considerable criticism from authors who claim that they are flawed, because Tinbergen’s four whys are being confounded (Bolhuis & Macphail 2001; Macphail & Bolhuis 2001; Bolhuis & Wynne 2009; Bolhuis 2015). For example, food storing in certain species of titmice and corvids has been interpreted as a case of adaptive specialization (Healy & Braithwaite 2000; Shettleworth 2010). According to this hypothesis, food-storing birds would have evolved superior spatial memory as well as a larger hippocampus (thought to be its neural substrate) compared to their nonstoring counterparts. Both parts of this hypothesis have not been confirmed by the data (see Chapter 8 for further discussion).
Animal welfare and human nature
Many people are interested in animal behavior out of mere curiosity, the need to know more about something. This is all very fine but there always comes a time when someone will ask “what is the purpose of studying animal behavior?” This question, whether from a research colleague, a friend, or a granting agency requires an answer expressed in terms of benefits to society. We see two areas in which animal behavior research can contribute to human society: animal welfare and understanding human nature.
Animals are important contributors to wealth and quality of life. They provide us with nourishment, the means to find cures and treatments for our illnesses, as well as invaluable companionship. Almost all of the information contained in this book relies on experiments and research conducted with animals. There is a growing concern that animals used for human benefit, however, be exposed to as little unpleasantness as necessary. Are housing cages too small, or densities of individuals too high? Is the knowledge acquired from experiments sufficiently important to authorize animal experimentation? The answer to such, often difficult, questions depends in many ways on knowing something about an animal’s behavior (Chapter 10).
People are endlessly curious about people and the sheer number of disciplines devoted uniquely to the study of human beings is eloquent testimony to this fact (e.g., medicine, anthropology, psychology, sociology, criminology). Animal behavior can provide insight into human behavior in two ways. More conventionally, phenomena observed in animals can be generalized, although often in some modified way, to humans. For example, just as a new antibiotic drug that cures an infection in some nonhuman primate can also be used, perhaps in a slightly modified way, to cure infections in humans, so can knowledge about how an animal learns be extended and applied to human learning. The second way, however, involves generalizing an approach rather than a result. For instance, can we learn anything new about human behavior by applying an evolutionary cost–benefit analysis to the things we do? This is what an area known as evolutionary psychology does (Chapter 17).
SUMMARY AND CONCLUSIONS
The study of the behavior of animals has grown into a highly diverse set of approaches and disciplines. Its subject area ranges from molecules and neurons to individuals and populations. One of Tinbergen’s major contributions to the study of animal behavior has been to make its goals explicit and clarify four types of questions that can be asked of behavior: causation, development, survival value, and evolution. In this book we strongly advocate Tinbergen’s position that behavior can only be understood through research on all four questions. In addition, we suggest that it should be made clear which of Tinbergen’s questions is addressed when a behavioral problem is investigated: a problem in one domain should not be investigated with concepts from another. The early chapters in this book examine primarily causal, mechanistic, and developmental questions, while the latter chapters examine survival value and evolution issues. But examples in many chapters also illustrate that multiple approaches are necessary for understanding a problem, affirming Tinbergen’s view.
FURTHER READING
Tinbergen’s (1963) paper on the four whys is essential reading for any serious student of animal behavior. It was reprinted in Houck and Drickamer (1996), which is a collection of classic papers on all aspects of animal behavior, and in Tinbergen’s Legacy (Bolhuis & Verhulst 2009), which is a collection of contemporary essays reflecting on Tinbergen’s classic paper. The four-part reader by Bolhuis and Giraldeau (2010) is organized on the basis of Tinbergen’s four whys and has a collection of classic and contemporary papers on the evolution, function, development, and causation of animal behavior. Tinbergen’s (1951) classic book is still very much worthwhile. It was reprinted in 1992 and is still available. The British ethologist William Thorpe (1979) has written a brief history of ethology, viewed from the inside, while Dewsbury (1989) provides a more recent account from the North American perspective. Boakes (1984) is an excellent review of the history of the study of animal behavior by psychologists, while Laland and Brown (2011) provide a very clear account of the different ways in which the behavior of animals (including humans) can be studied from an evolutionary perspective. Functional and evolutionary aspects of behavior are also discussed in Davies et al. (2012). Hogan’s (2017) book provides an exposition and critique of behavioral concepts and of historical and contemporary studies of behavior.
REFERENCES
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17 Hogan,