Ethology, Natural History, the Life Sciences, and the Problem of Place

Investigators of animal behavior since the eighteenth century have sought to make their work integral to the enterprises of natural history and/or the life sciences. In their efforts to do so, they have frequently based their claims of authority on the advantages offered by the special places where they have conducted their research. The zoo, the laboratory, and the field have been major settings for animal behavior studies. The issue of the relative advantages of these different sites has been a persistent one in the history of animal behavior studies up to and including the work of the ethologists of the twentieth century. This revised version was published online in July 2006 with corrections to the Cover Date.     

Tribute to Tinbergen: Putting Niko Tinbergen's ‘Four Questions’ in Historical Context

Niko Tinbergen's (Zeit. Tier. 20, 1963, 410) paper ‘On aims and methods of ethology’ is appropriately remembered as the paper in which Tinbergen characterized ethology as ‘the biological study of behavior’ and went on to explain that to study behavior biologically is to ask four distinct questions about it: (1) How is it caused physiologically? (2) What is its survival value? (3) How has it evolved? and (4) How does it develop in the individual? Here, we consider Tinbergen's paper in its historical context by looking at it from three different perspectives: (1) a comparison of Tinbergen's formulation of ‘ethology's four questions’ with similar, but different formulations of biology's basic problems offered by Julian Huxley, Konrad Lorenz, and Ernst Mayr; (2) a survey of the roles that the four questions played in Tinbergen's own work over the course of his career; and (3) a consideration of the two explicit goals of Tinbergen's (Zeit. Tier., 20, 1963, 410) paper, namely (a.) to honor Tinbergen's friend and colleague Konrad Lorenz (as part of a Festschrift for Lorenz on the occasion of his sixtieth birthday) and (b.) to provide a sketch of ethology's scope and an evaluation of the ways the field needed to develop in the future. We suggest that just as the work of Tinbergen's Oxford research team revealed how the behavior of gulls reflected compromises worked out in the face of the diverse selective pressures of particular environments, we can identify certain conflicts that arose for Tinbergen in trying to write something that his friend Lorenz would like while also assessing ethology's current state and future prospects. That said, however, Tinbergen's enduring concern was to do all he could to ensure that ethology thrive as a field and develop a scientific understanding of animal (and human) behavior. For this to happen, he insisted, the four questions of ethology needed to be pursued in a balanced, comprehensive, and integrated fashion.     

What can whiskers tell us about mammalian evolution,behaviour, and ecology?

1. Most mammals have whiskers; however, nearly everything we know about whiskers derives from just a handful of species, including laboratory rats Rattus norvegicus and mice Mus musculus, as well as some species of pinniped and marsupial.
2. We explore the extent to which the knowledge of the whisker system from a handful of species applies to mammals generally. This will help us understand whisker evolution and function, in order to gain more insights into mammalian behaviour and ecology.
3. This review is structured around Tinbergen’s four questions, since this method is an established, comprehensive, and logical approach to studying behaviour. We ask: how do whiskers work, develop, and evolve? And what are they for?
4. While whiskers are all slender, curved, tapered, keratinised hairs that transmit vibrotactile information, we show that there are marked differences between species with respect to whisker arrangement, numbers, length, musculature, development, and growth cycles.
5. The conservation of form and a common muscle architecture in mammals suggests that early mammals had whiskers. Whiskers may have been functional even in therapsids.
6. However, certain extant mammalian species are equipped with especially long and sensitive whiskers, in particular nocturnal, arboreal species, and aquatic species, which live in complex environments and hunt moving prey.
7. Knowledge of whiskers and whisker use can guide us in developing conservation protocols and designing enriched enclosures for captive mammals.
8. We suggest that further comparative studies, embracing a wider variety of mammalian species, are required before one can make large-scale predictions relating to evolution and function of whiskers. More research is needed to develop robust techniques to enhance the welfare and conservation of mammals.

     

Tribute to Tinbergen: The Place of Animal Behavior in Biology

Tinbergen is famous for emphasizing behavioral fieldwork and experimentation under natural circumstances, for founding the field of ethology, for getting a Nobel Prize, and for mentoring Richard Dawkins. He is known for dividing behavior studies into physiology, development, natural selection, and evolutionary history. In the decades since Tinbergen was active, some of the best research in animal behavior fuses Tinbergen's questions, connecting genes to behavioral phenotypes, for example. Behavior is the most synthetic of the life sciences, because observing the actions of an organism can tell us what all those physical and physiological traits are for. Insights from behavior tell us how traits in one individual impact those in another in ways that challenge our definition of an organism. Behavioral conflict and cooperation among animals has led to theory that explains within‐organism conflict and cooperation and human malfunctions of many kinds. Darwin certainly began the evolutionary study of behavior, but Tinbergen brought it forward to the heart of biology. The challenge for the future is to apply concepts from animal behavior across biology with tools that would have amazed Tinbergen.