In the German-speaking world, ethology is traditionally referred to as “classical” comparative behavioral research, which established itself as an independent research area from the 1930s, but occasionally also behavioral biology in general . Ethology is consequently a branch of zoology and a neighboring discipline of psychology , but within zoology it is also a supplement to the comparative approaches of morphology , anatomy and physiology in the service of systematic kinship research.
The term ethology is derived from the Greek ἔθος ethos ("habit, custom, custom") and λόγος lógos (including "philosophical doctrine" or in the plural "sciences", cf. logic ). Ethology literally means "the science of habits ".
The made-up word ethology was coined in France in the middle of the 19th century by Isidore Geoffroy Saint-Hilaire , and Friedrich Dahl had already proposed in 1898 that the French word for the habits of animals should be translated into German. But only after William Morton Wheeler introduced ethology to the English-speaking world in 1902 and this term established itself there, did it return to Germany via this detour.
The classic ethological research on instinct
The ethological research is closely linked to the work of Oskar Heinroth , Erich von Holst , Konrad Lorenz , Günter Tembrock , Nikolaas Tinbergen and Irenäus Eibl-Eibesfeldt , the draft of a theory of instinct and the former Max Planck Institute for Behavioral Physiology . Jean-Henri Fabre , who studied the instinct in insects, can be regarded as an important precursor .
The field of ethology integrated elements from several disciplines. According to Ilse Jahn and Ulrich Sucker, the ethologists initially took over "elements from the scientifically oriented human psychology of Wilhelm Wundt ", furthermore suggestions "from the evolutionist comparative morphology and development history", and they finally integrated "methods of experimental animal physiology and taxonomic field research ."
Even Hermann Samuel Reimarus (1694-1768) had in 1760 in his book General observations on the instincts of animals described "instincts, impulses, arts" that "without and prior to all experience, right after their birth" are suitable for their "self-preservation " to serve. A hundred years later, Charles Darwin was able to make plausible, based on years of his own breeding experiments (including on domestic pigeons ), that the animals' often very complex behaviors must have arisen on the basis of the same regularities as their anatomical features: i.e. because of the random variability of the individual features and their meaning in the "Struggle for survival" of their porters. William James , who as the founder of psychology in the United States and as a prominent representative of the so-called instinct psychology applies, went in 1890 in his major work, The Principles of Psychology from evolutionary assumptions example, assume that the man possesses several dozen instincts, and William McDougall , the Co-founder of the British Psychological Society , attributed numerous "primary instincts" to humans in 1908, including: a. Flight instinct, disgust instinct, curiosity instinct, aggression instinct, self-assertion instinct, sexual instinct, parental instinct as well as one instinct each for self-humiliation by showing oneself and for sociability.
So-called vitalistic views held up well into the early 20th century, although they did not deny innate instinctive behavior and even showed its usefulness. However, they answered the question about the emergence of this expediency with the assumption of a life force (Latin vis vitalis , hence: vitalism), a natural force or divine guidance . For a long time, this assumption of supernatural forces blocked any scientific research into causes. A prominent representative of this direction was Alfred Russel Wallace . Next to Darwin, Wallace is considered to be the founder of the modern theory of evolution ; but he distanced himself far from all evolutionary biological ways of thinking as soon as it came to the emergence of instincts .
In sharp contrast to the vitalistic directions stood the so-called mechanists , who interpreted all behavior as the quasi passive reaction to external stimuli, as a chain of reflexes (" reflex chain theory "). Their views were based primarily on the research results of Nobel Prize winner Ivan Pavlov and denied internal drives or excluded them as an object of scientific investigation due to lack of accessibility. Prominent representatives of this direction were, besides Pavlov, the American psychologists John B. Watson , the founder of classical behaviorism , and BF Skinner . In the German-speaking area, an independent research direction developed after the First World War , which on the one hand - in contrast to behaviorism - emphasized the spontaneous occurrence of innate behavior due to internal, central nervous causes ("drives"). On the other hand, due to the assumed inheritance of such behavior, their representatives compared the behavior of related species with one another in a manner similar to how anatomists compare anatomical features with one another (hence also comparative behavioral research ). In 1937 this line of research created its own publication organ through the Zeitschrift für Tierpsychologie . It was only after the Second World War that the term animal psychology was replaced by the term ethology , since animal psychology was now reputed to be a mere hobby.
In the 1970s, the terms ethology , instinct research and comparative behavioral research were still being used as synonyms by researchers in this field .
To the extent that the instinct theory that emerged from “classical” comparative behavioral research was viewed as outdated due to more recent behavioral ecological and neurobiological findings, many behavioral researchers have also been using the term ethology less and less since the early 1980s and replaced it with the more neutral one perceived term behavioral biology .
Outside of the German-speaking area, however, ethology (English), éthologie (French), etología (Spanish), etologia (Italian), etoloji (Turkish) and etologi (Danish) generally stand for behavioral biology . For this reason, the magazine for animal psychology co-founded by Konrad Lorenz in 1937 , alongside Behavior and Animal Behavior for decades the most important specialist publication in behavioral biology, was renamed Ethology from 1986 by Wolfgang Wickler - following international usage .
The concept of instinct
Behavioral research took a fundamental turn by Oskar Heinroth , in whose lecture published in 1911 before the 5th International Ornithological Congress, the word ethology was used for the first time in today's sense in Germany, also in front of a large specialist audience. Heinroth had first studied the behavior of various types of geese and ducks and discovered that certain movements (for example during courtship) are always carried out by animals of the same sex and the same type with the same gestures and postures. Heinroth called such constant-form movements specific instinctual actions and was able to show that related species have more or less strong modifications of such behavior. From these precise behavioral observations to an evolutionary interpretation of their emergence, it was not a big step either for Heinroth or for his later pupil Konrad Lorenz . Lorenz took up the term ethology for the first time in 1931 when he published an extensive essay on the "ethology of social corvids ".
The ethological instinct theory states that instinctive behavior is anchored in the genome and can be triggered by key stimuli , as long as an inner action-specific energy is present. The usefulness of this interlocking of external trigger, willingness to act and specific behavior developed in the process of evolution and ultimately serves to pass on the genes to the next generation.
A frequently cited example of such an instinctive movement is the egg rolling movement of the greylag goose : When an egg (the key stimulus ) gets outside of the nest, the goose stretches its beak over the egg and uses its beak to roll the egg back into the nest. This movement always proceeds in the same way and is completed even if the egg is removed by an experimenter during the process. This rigid, innate form of behavior is considered to be a species-specific instinctual act in the sense of Oskar Heinroth and was described by Konrad Lorenz as hereditary coordination .
Characteristic of ethological instinct research is, on the one hand, the emphasis on field research, i.e. the observation and explanation of behavior under natural environmental conditions, and, on the other hand, so-called ethograms : These are exact descriptions of all behavioral patterns that can be observed in an animal species. On the basis of these ethograms, behavioral protocols can be drawn up in which the frequency of behavior and their chronological sequence are listed (e.g. eating, sleeping, cleaning oneself, running away quickly, bringing young animals to the nest). This makes it possible to qualitatively and quantitatively describe both the frequency and the sequence of behaviors.
Until the end of the 1960s, the term instinctive movement or hereditary coordination was associated with the view that the behaviors interpreted in this way are purely innate activities that are “triggered by external stimuli and influenced in their intensity and orientation”; their sequence, however, "ie the type of movement, is independent of external stimuli and is determined specifically for each species." In the meantime, however, research has found more and more indications that such rigid reactions to external stimuli are an exception , that heritage and the environment are also related are closely linked to individual behaviors (see reaction norm ).
Central concepts of classical ethology were criticized in 1990 by Wolfgang Wickler , a student of Konrad Lorenz , and in 1992 by Hanna-Maria Zippelius , a student of Karl von Frisch (compare, among other things, skipping and idling ). Their criticism, however, was preceded by a detailed analysis in a review article in the USA almost 30 years earlier , in which the instinct-based ethology was described as “ preformationistic ” and “biased” with regard to its “rigid concepts” of innate behavior and maturation Perfection in the course of individual development without practice - was designated.
The turning away from the instinct theory was formulated most clearly in 1990 by Wolfgang Wickler using the example of jumping and idling : "The action-specific energy turned out to be a modern phlogiston and the psychohydraulic model, despite ingenious changes, was unsuitable for adequately depicting the changes in readiness and state in the animal." Modeling were u. a. young sub-areas of ecology such as population ecology and behavioral ecology , which describe, for example, the search for food and other decision-making in conflict situations with the help of the concept of cost-benefit analysis as optimal foraging . For the investigation of social behavior, sociobiology and bioacoustics have established themselves as independent disciplines, and game theory , biolinguistics and evolutionary psychology also provide fruitful approaches for the investigation of the evolution of behavior .
Farm animal ethology
As ethology by a branch of is Agricultural Sciences called, devoted to the study of the behavior of animals is concerned. The aim is to optimize the keeping conditions in livestock keeping in the sense of an almost species-appropriate keeping .
- Irenäus Eibl-Eibesfeldt : Outline of Comparative Behavioral Research. 7th completely revised and expanded edition, Piper, Munich 1987.
- Ilse Jahn and Ulrich Sucker: The Development of Behavioral Biology. In: Ilse Jahn (Ed.): History of Biology. 2nd, corrected edition of the 3rd edition. Spektrum Akademischer Verlag, Heidelberg and Berlin 2002, pp. 580–600, ISBN 3-8274-1023-1 .
- Konrad Lorenz : About animal and human behavior. From the development of the theory of behavior. Collected Treatises. Volume 1 and 2, Piper, Munich 1965.
- Konrad Lorenz: Comparative behavior research. Basics of ethology. Springer Verlag, Vienna 1978.
- Konrad Lorenz: The natural science of humans. An introduction to comparative behavioral research. The "Russian Manuscript ." Piper, Munich 1992.
- Volker Schurig : One hundred years of the scientific term 'ethology': a critical balance sheet. In: Biology in our Time , Volume 41, No. 2, 2011, pp. 92–94.
- Uta Seibt and Wolfgang Wickler : History of behavior research. In: Lexicon of Biology. Volume 10, 1992, pp. 353-358.
- Keyword ethology on biological-concepts.com
- "its usage in the sense of habit, manners, etc. (...) expresses what we mean by animal behavior". William Morton Wheeler : Natural history, 'oecology' or 'ethology'? In: Science . Volume 15, No. 390, 1902, p. 975, doi: 10.1126 / science.15.390.971 .
- Isidore Geoffroy Saint-Hilaire: Histoire naturelle des règnes organiques. Volume 2, 1854, p. 285, access page to the full text .
- Friedrich Dahl : Experimental statistical ethology. In: Negotiations of the German Zoological Society. 1898, pp. 121-1131. For the delimitation of biology , ecology and ethology around 1900 see: Ulrich Sucker: The Kaiser Wilhelm Institute for Biology. Its founding history, its problem-historical and scientific-theoretical prerequisites (1911–1916). Franz Steiner Verlag, Stuttgart 2002, pp. 59 ff., ISBN 3-515-07912-2 .
- William Morton Wheeler: Natural history, 'oecology' or 'ethology'? In: Science . Volume 15, No. 390, 1902, pp. 971-976, doi: 10.1126 / science.15.390.971 .
- Ilse Jahn and Ulrich Sucker: The development of behavioral biology. In: Ilse Jahn (Ed.): History of Biology. 2nd, corrected edition of the 3rd edition. Spektrum Akademischer Verlag, Heidelberg and Berlin 2002, p. 593, ISBN 3-8274-1023-1 .
- Hermann Samuel Reimarus : General considerations about the instincts of animals, mainly about their artistic instincts. To the knowledge of the connection of the world, the Creator and ourselves. Johann Carl Bohn, Hamburg 1760, S. 92 and 96, full text .
- Charles Darwin : On the origin of species by means of natural selection, or the preservation of favored races in the struggle for life . John Murray, London 1859, chapter 1; digitized version .
- William James : The Principles of Psychology. 2 volumes. Henry Holt and Company, New York 1890.
- William McDougall : The Principal Instincts and the Primary Emotions. Chapter 3 in: The Same: An Introduction to Social Psychology. Methuen & Co. Ltd., London 1919, full text
Two early studies are:
Günther Schlesinger : Zur Ethologie der Mormyriden . In: Annals of the Natural History Museum in Vienna. Volume 23, No. 3/4, 1909, pp. 282-311, introduction .
Oskar Heinroth : Contributions to biology, especially ethology and psychology of the anatids. In: Negotiations of the 5th International Ornithological Congress in Berlin, May 30 to June 4, 1910 . German Ornithological Society, Berlin 1911, pp. 559–702, full text .
- See e.g. B. Katharina Heinroth in Grzimek's Animal Life , Complementary Volume Behavioral Research, Chap. 1, Kindler Verlag, Zurich 1974.
- Volker Schurig: A hundred years of the scientific term 'ethology': a critical balance sheet. In: Biology in Our Time. No. 2/2011, p. 92. Here it says literally: "The scientific term 'ethology' has been disappearing increasingly from scientific linguistic usage since 1980."
- Oskar Heinroth: Contributions to biology, namely ethology and psychology of the anatids. In: Reports of the V. Int. Ornithologists Congress. Berlin 1910, p. 559 ff.
- Konrad Lorenz: Contributions to the ethology of social corvids. In: Journal of Ornithology. Volume 79, No. 1, 1931, pp. 67–127, full text (PDF) .
- Konrad Lorenz, Nikolaas Tinbergen: Taxis and instinctive action in the egg rolling movement of the gray goose. In: Zeitschrift für Tierpsychologie. Volume 2, No. 1-3, pp. 1-29, 1939, doi: 10.1111 / j.1439-0310.1939.tb01558.x .
- Gerard Baerends: Structure of animal behavior. In: J.-G. Helmcke, H. von Lengerken and D. Starck (Hrsg.): Handbuch der Zoologie. Volume VIII: Mammalia, 10th part, 1st half. Walter de Gruyter, Berlin 1956, p. 1.
- Entry in hereditary coordination in: Klaus Immelmann : Grzimeks Tierleben , supplementary volume behavior research. Kindler Verlag, Zurich 1974, p. 626.
- Wolfgang Wickler : From ethology to sociobiology. In: Jost Herbig, Rainer Hohlfeld (ed.): The second creation. Spirit and Demon in 20th Century Biology. Carl Hanser, Munich 1990, p. 176, ISBN 978-3-446-15293-9 .
- Hanna-Maria Zippelius : The measured theory. A critical examination of the instinct theory of Konrad Lorenz and behavioral research practice. Vieweg, Braunschweig 1992, ISBN 3-528-06458-7 .
- Daniel S. Lehrman: A Critique of Konrad Lorenz's Theory of Instinctive Behavior. In: The Quarterly Review of Biology: Volume 28, No. 4, 1953, pp. 337–363, doi: 10.1086 / 399858 , full text (PDF)
- Steffen Hoy: farm animal ethology. Eugen Ulmer, Stuttgart 2009, ISBN 978-3-8252-3312-9 .