Histories of Ethology: Methods, Sites, and Dynamics of an Unbound Discipline

Abstract

Ethology is considered the leading biological discipline within behavioral research in the 20th century. Its history is told as a seemingly straightforward narrative: Ethology has its roots in the 1930s in German-speaking countries, a disciplinary heyday in the 1950s and 1960s, after which it slowly lost relevance. It employs a distinct approach to the comparative study of animal behavior, which is characterized by a physiological method of non-invasive, often observational studies of natural behavioral patterns, which were conceived of as shaped by evolution. Ethology contains stories of charismatic research animals such as the jackdaw Tschock or the goose Martina,¹ draws on academic disciplines such as ornithology, ichthyology, and entomology,² and also incorporates contexts and practices of animal lovers, bird watchers, and hunters, as well as those involved in animal husbandry, wildlife preservation, and livestock farming, or who work in nature reserves or zoological gardens.³ Ethology is further connected to the development of certain visual media, such as chronophotography and the film loop,⁴ and corresponding forms of perception, such as pattern recognition⁵ or comparative visual analysis.⁶ Other methodical highlights include the ethogram,⁷ dummies,⁸ and the Kaspar Hauser experiment.⁹ The history of ethology conventionally focuses on several elements: an illustrious circle of founding figures, indeed founding “fathers,” such as Konrad Lorenz (1903–1989), Nikolaas Tinbergen (1907–1988), Karl von Frisch (1886–1982), Jakob von Uexküll (1864–1944),¹⁰ Oskar Heinroth (1871–1945),¹¹ Erwin Stresemann (1889–1972),¹² and Otto Koehler (1889–1974),¹³ the importance of the Zeitschrift für Tierpsychologie (later renamed Ethology),¹⁴ legendary encounters of individual scholars with their research subjects¹⁵ and colleagues during conferences¹⁶, and towering intellectual achievements such as famous talks¹⁷ or foundational monographs such as Tinbergen's The Study of Instinct, published in 1951¹⁸. Further markers of ethology's disciplinary history are the recognition of its achievement and disciplinary status with the Nobel Prize in Physiology or Medicine awarded to Lorenz, Tinbergen and von Frisch in 1973 and the ensuing controversy about Lorenz's political background.¹⁹

All scientific disciplines, of course, are outlined by a set of protagonists, places, publications, and practices.²⁰ However, ethology is characterized by an unusual preoccupation with its own disciplinary status, demarcating its core and periphery, its borders and boundaries as well as delineating its historical lineage and possible trajectories—all adding to the sense that ethologists’ historical accounts are full of strategic self-historization and purposeful narratives. Consequently, historians of science have engaged with this core question of disciplinary identity, unveiling the layers of strategic narratives employed by the actors and adding complexity to their presentations of disciplinary success stories. Historians of science, ethologists turned historians and ethologists themselves offered narratives of ethology, some motivated by scholarly curiosity, others by a desire for commemoration, historical legitimacy, or strategic considerations.

This special issue contributes to the historiography of ethology beyond the classical phase and reflects ethology's nature as an unbound discipline that is not as uniform and cohesive as the historical efforts of Lorenz and other early ethologists made it out to be.²¹ It invites an interdisciplinary audience to rediscover the history of ethology from the perspectives of epistemology, animal studies, media studies as well as publication and data history. Contributions in this issue focus on novel methods, hidden sites, and overlooked dynamics and exemplify a wide range of approaches, including archival work, multi-media analysis, oral history interviews, and publication tracing for a multi-focal history of ethology. We engage with central countries in the history and historiography of ethology: German-speaking countries, the United Kingdom, and USA. While we acknowledge the limits of this choice, the historiography of ethology is shaped by the Lorenz-Tinbergen axis in Germany and the United Kingdom, with some of the most important ethological and historical research carried out in connection or succession to the scientific impact they had in each country.²² Therefore, writing the history of ethology includes dealing with the historical contexts and social motives of these localized disciplinary histories. We will recount the central authors, phases, and genres in the historiography of ethology, ending with a reflection on current themes.

The work of historians and ethologists over the last decades has shown that ethology was never a monolithic, cohesive framework, but instead the result of social dynamics, strategic agendas, and contingent developments, containing diverse approaches and diverging theoretical premises. Its disciplinary boundaries are porous and permeable. We therefore suggest understanding ethology as an unbound discipline from its very inception. This allows us to add new perspectives of its further unbounding, especially by looking at researchers that were not considered to be in the mainstream of ethology and after what is considered its classical stage. Our contributions explore different dynamics of ethology's development. Novel methods allowed an unbounding from the constraints of time and life span: photography, card indexing, and long term data collection moved the object of study from individual animal behaviors to synthesized behavior repertoires over ever-increasing time spans. Similarly, an unbounding from limits placed on research subjects and questions took place, such as the inclusion of human behavior or animal cognition into the scope of ethological research. These changes also imply an unbounding from classic sites of knowledge production in ethology and our contributions explore new sites such as long-term field stations in Kenya, a research compound in the East German part of Berlin, a eugenically minded dog research station in the US as well as Munich kindergartens. These dynamics add complexity to the perception of ethology as a disciplinary monolith and, instead, pose the question of how ethology is kept together as a discipline. Our contributions point to the essentially social nature of ethology as a discipline through shared institutions and publications organs, academic genealogies and lineages, and communal research stations and vibrant university departments.

Kelle Dhein sheds light on Karl von Frisch's hitherto unclarified relationship with ethology regarding experimental methods. The focus of Dhein's article is the grey card experimental method developed by von Frisch in the 1910s. Combining a holistic view of the behavior of his research subjects with a controlled, at times reductionist experimental design, von Frisch succeeded in producing quantifiable research results which provided information about the sensory perception of the bees in his experiments. Among other aspects, the conditioning of the bees, which was necessary for his experiments at the time, represented a clear departure from the methodologies of classical ethology, even leading to von Frisch's exclusion from ethologist circles. It is one of the paradoxes of the history of ethology that precisely this experimental design would become a key component in later phases of the discipline's history and, from the 1970s onwards, play a significant role in the success of research projects associated with the field of neuroethology. In examining how an experiment “has a life on its own,”⁷² thus traversing different premises, questions and directions of research, Dhein succeeds in tracing a new line in the history of ethology.

Sophia Gräfe’s contribution also adopts a praxeological approach. Her study of visual media in classical ethology adds archives to the existing catalogue of significant sites in the history of the discipline. Taking as her subject the ethological image collection established by Günter Tembrock at the Forschungsstätte für Tierpsychologie in Berlin, she demonstrates the extent to which these images, along with other visual media such as drawings and film recordings, significantly advanced the ethological search for innate forms of behavior. Images were used to assist Gestalt perception and pattern recognition. They also served as a memory aid, as well as to display the scientists’ detailed knowledge and research experience to their colleagues. In addition, collections of images provided a basis for developing the clearest possible nomenclature of behavior, which, as an ephemeral phenomenon, required new forms of representation. Since the images in ethological collections generally come from various sources, and their motifs span lengthy periods of time, Gräfe's analysis underlines the predominantly collective character of comparative behavioral research. Finally, an examination of the history of ethology from the perspective of its visual media addresses the synthetic character of the knowledge acquired about the behavior of living beings. Tembrock's seminal research on the behavior of red foxes was the result of both his experiments with the animals in the behavioral laboratory and the expert analysis of archive images.

Brad Bolman achieves a multi-layered portrait of canine behavioral studies in the US by using approaches from model organism research. Previous historiographies had so far excluded these studies from the history of ethology and thus overlooked the canine history of this line of behavioral research. The projects reconstructed by Bolman at the intersection of eugenics, genetics, behavioral research, and animal breeding are situated on the periphery of classical ethology and form the starting point for a historiographical intervention: Charles Rupert Stockard, Clarence Cook Little, Raymond C. and Lorna Coppinger and Harry and Martha G. Frank, among others, all studied canine behavior based on the hypothesis of a critical period, in other words certain phases in the early development of living beings in which the influence of specific environmental conditions determines the development of specific behaviors. The gradual incorporation of the critical period concept into everyday usage since 1950 and its more recent establishment as an analytical category in the history of science evokes teleological narratives. Rather than portraying the birth of the discipline as the result of favorable conditions, Bolman's study highlights the winding paths of its history, marked by interruptions and detours.

By the time Lorenz, Tinbergen and Frisch received the Nobel Prize in 1973, the heyday of classical ethology may already have passed. Jakob Odenwald shows in his contribution how from the late 1960s onwards a new field emerged in comparative behavioral research, when biologist Barbara Hold initiated the innovative “Kindergarten Project,” which served as an excellent example for the developing field of human ethology. Following the principle of Irenäus Eibl-Eibesfeldt's “Cross-Cultural Long-Term Study” (Kulturvergleichende Langzeitstudie), children at several kindergartens in the Munich area became the protagonists of a behavioral study. The idea was that as research subjects the children would provide undistorted access to the natural foundations of social behavior. Odenwald's contribution decisively highlights the political background of the Kindergarten Project, in which the focus of the research was social ranking behavior. The educational approach varied among the groups in the study—a group with an anti-authoritarian approach was also included. Against a backdrop of social debate about class and social justice, Hold's theses on the biological foundations of social ranking provided ample fuel for discussion, as Odenwald illustrates using samples of the surviving correspondence between the scientific, public, and private actors.

The long-term behavioral biology projects gathered together in Erika Milam’s contribution also colonized new spaces. In their search for cross-generational mechanisms underlying the development of social behavior, population biologists, primatologists, and sociobiologists shifted their research from the laboratory to the field and focused on long-term studies from the 1960s and 1970s onwards. While research locations are already a common theme in the history of ethology, Milam's study focuses on the time factor and correlates specific questions about behavior with the timelines of the research. One effect of adopting this perspective is the emergence of new protagonists in the history of ethology, including local researchers who continued collecting data in the absence of principal investigators, and data analysts who used their skills to analyze vast amounts of behavioral data. Milam's account also draws attention to the biographical side-effects of this new long-term perspective on behavior. In interviews with Kenneth Armitage, Jeanne Altmann, Timothy Hugh Clutton-Brock, and others, she outlines how the professional environment was transformed, turning some researchers into migrating project workers and others into decades-long companions of “their” animal population.

Finally, Cora Stuhrmann examines the development of two disciplines that followed ethology. By the 1970s, ethology had lost its methodological coherence. The growing emphasis on mathematical methods was undermining the leading status of its key figures, some of whom had by then reached old age. Stuhrmann's examination of the fate of ethology as a discipline explores the dynamics between ethology, behavioral ecology and sociobiology at the inter-disciplinary level and adds the analytical framework of competition to the history of the discipline. She identifies the genre of edited volumes, textbooks, and their respective reviews as the forum of this driving social force, interpreting these publications as strategic components of scientific negotiation processes. The key finding of her study is the negation of the frequently raised question of whether ethology has died. From the loss of relevance of its central theoretical foundations and the rise of a new generation of researchers we can conclude that although ethology has lost relevance as a discipline, it has expanded terminologically to include all studies on animal behavior with a biological foundation.