Skipping movement

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Skipping movement (also: skipping action , skipping behavior ; English : displacement activity , occasionally also: substitute activity or behavior out of context ) is a technical term of the instinct theory of classical comparative behavioral research (ethology) developed primarily by Konrad Lorenz and Nikolaas Tinbergen . He was Nikolaas Tinbergen and Adriaan Kortlandt in the ethology introduced and referred to certain behavior patterns that are perceived "unexpected" by the observer than as they occur within a behavioral sequence in which they appear to serve no immediate purpose. Nikolaas Tinbergen described them as follows: "These movements seem irrelevant in the sense that they occur regardless of the context of the behavior immediately preceding or following."

Such behavior , which appears to be “inappropriate” to the observer, without a comprehensible reference to the given situation, was interpreted as an expression of “a conflict between two instincts ”, which is why the continuation of the previously observable instinctual behavior is not possible - at least temporarily - and instead a behavior is shown that (according to the theory of instinct) comes from a completely different - third - functional circle of the behavioral repertoire.

Later research in behavioral biology interpreted behaviors that were originally interpreted as skipping movements as social signals and thus by no means irrelevant in the respective context.

Models of skipping behavior

The concept of jumping movements was based on the assumption that two opposing instinctual actions (for example attack and flight) mutually inhibit each other and the "drive energy" released for both in this situation jumps to a third behavior , so that this third behavior is carried out - that is “Movements that belong to a different instinct than the currently activated instinct or the currently activated instinct.” Klaus Immelmann explained that the behavior pattern classified by the representatives of classical comparative behavioral research as jumping behavior is “unexpected in the sense that it is in the situation in which it occurs does not fulfill the normal biological function for which it was developed in the course of tribal history. ”In other words: If the course of an instinctual act is caused by deficiencies in the triggering situation or the occurrence of a conflict between unerei If the instincts are disturbed, the pent-up drive energy can be reacted to via behavior that is apparently irrelevant in the situation and that belongs to another instinct.

Skip hypothesis

The skipping hypothesis developed by Tinbergen around 1940 assumes that whenever the "discharge" of an action-specific excitation, i.e. the activated instinct, is not possible, another - but always the same - movement pattern is produced. According to Tinbergen's "Instinctive Doctrine" from 1952, an animal then shows jumping behavior "when with a very strong drive ... the external situation is not sufficient to trigger the final action". As a consequence, this means that, according to this model, behavior C is triggered on the one hand (and usually) by its specific arousal, but on the other hand it can also be triggered by behavior B (as it were by skipping over an "external excitation"): namely then, if behavior B is blocked, for example, due to a lack of key stimuli and therefore an "excitation congestion" occurs.

Disinhibition hypothesis

In contrast to the skipping hypothesis, according to van Iersel & Bol (1958), according to the disinhibition hypothesis , the skipping movement is activated by its own energy. The disinhibition hypothesis is based on the assumption that a certain instinct can have an inhibiting effect on other instincts: If a certain instinct is being performed, it will stop all other instincts in order to avoid jumping back and forth between different behaviors. The further assumption that is decisive for the “functioning” of the disinhibition hypothesis states that certain instincts can mutually inhibit one another. If their strength is roughly the same, this means that they completely block each other: Neither one nor the other instinctive movement can then occur. Such a mutual inhibition of two drives has the consequence that a pre-existing inhibition against a third instinct is lifted. The behavior patterns associated with this third instinct can then appear due to the disinhibition and are interpreted as jumping movements.

In more general terms (after Bernhard Hassenstein ): Drive A and B inhibit each other; Behavior B also inhibits - if it occurs - the drive for behavior C; since behavior B cannot occur as long as it is blocked by A, behavior C occurs and can be classified by the observer as a skipping movement. However, in 1983 Hassenstein rejected the disinhibition hypothesis as unsuitable.

Examples

Probably the most frequently cited example of jumping movements relates to observations on roosters of almost equal strength fighting out their “pecking order” with one another : Suddenly one of the two pecks around on the floor as if he were taking in food, and the other often followed the example immediately of the rival. This situation is interpreted in the context of the instinct theory as an expression of an equally strong fight and flight motivation (actions A and B), which evokes as a jumping movement “pecking food” (action C). After such an "interlude", the fight is usually continued.

Of black-headed gulls males reported Nikolaas Tinbergen that - similar to the taps - a struggle sometimes simultaneously interrupt and the movement way of Grasabrupfens show it without but abzurupfen grass. According to Tinbergen, picking up grass is a movement that can be assigned to the functional group of nest-building; but if it occurs in connection with a fight, it is without any function ("irrelevant") and thus a skipping movement.

Another example presented by Tinbergen in a review article and cited many times since then comes from the reproductive cycle of the three- spined stickleback : When the female has laid eggs, the male often and intensely fanned his fins at the nest. The eggs in the nest are the key stimulus for this instinctive movement . If this fanning already occurs during the advertisement for a female or during the nest building - if there are no eggs yet - then Tinbergen classifies it as jumping behavior.

Terns perform cleaning movements when they are in conflict between brood care and flight or flight and attack.

Honeybees clean themselves at the feeding place when they are in conflict between staying or looking for a new feeding place.

Oystercatchers in front of a mirror perform sleep movements in the conflict between attack and flight.

Finally, the behavior of people who occasionally scratch their heads in embarrassment in the car when they don't know whether to turn right or left after the traffic light is also frequently mentioned . Many a speaker behaves in a similar way in front of a larger audience who, on the one hand, is motivated to address the audience and, on the other hand (especially if there is uncertainty), would prefer to escape the situation. As "irrelevant" actions, he performs various cleaning movements: stroking his hair, putting on and taking off glasses, cleaning glasses, tugging at the cuffs, wiping dust from clothes, organizing papers.

criticism

Both models, on the basis of which the presumed internal causes of the behaviors interpreted as jumping movements can be formally described, are further developments of the older “psychohydraulic” behavioral models to more modern models based on electrical circuits. They assume two prerequisites: “1. Every movement can be assigned to a behavior system. 2. You can tell in every situation which behavioral systems are activated and which are not. ”Another fundamental thought in these models is that animal behavior is not purely reactive, as was (and neither was) assumed by behaviorists in the 1930s mere sequence of reflexes ), but that the spontaneity of behavior is emphasized: It was assumed that there were spontaneously active nerve cells in the brain that produce excitation and thus cause an animal to show a certain behavior.

Gerard Baerends had already expressed reservations about this interpretation of observable behavior when he pointed out in 1956 in the Handbuch der Zoologie that the behaviors interpreted by ethologists as skipping movements could at least “ acquire secondary signal significance” and at the same time regretted: “However, there is a difference in the physiology of skipping movements Unfortunately, nothing else is known. ”Since no brain regions could be identified later that could be associated with the jumping behavior, Peter Sevenster took up Baerend's reference in Grzimek's animal life in 1974 and emphasized that“ we rarely (if ever) use the adaptive value or even see through the tribal history of the movement in question. "

Two decades later, the evolutionary biology reservations against the concept of skipping movements were deepened by the behavioral biologist from Bonn at the time, Hanna-Maria Zippelius, with arguments based on the theory of science ; Zippelius' criticism read: "Neither of the two hypotheses that are supposed to 'explain' the occurrence of a skipping movement offers the possibility of an empirical test," since no method is available to continuously measure the postulated changes in internal drives. Only under the condition that the strength of two drives can be measured at the same time is "a statement possible that the drives are equally strong at the point in time at which the jumping movement can be observed." At the same time, she criticized the "presumptuous claim", A behavioral scientist could decide on the basis of animal observations alone - without knowledge of the ancestral history of the behavior of the animal species in question - whether a behavior pattern in a certain situation is “irrelevant”, i.e. not expedient, a mere interruption of the “actual” action: “This is what these hypotheses are no more than imaginative considerations of individuals who, using Lorenz's concept of instinct, only pretend to 'explain' the phenomenon in question. ”In this context, Zippelius referred a. a. also to Konrad Lorenz, who had admitted in his late work that it was "downright difficult to find examples of jumping movements that do not have signal effects." But if the so-called jumping movements are consistently social signals, then they definitely have a function in the context in which they are observed: "They are then only unexpected for the observer who does not understand them, ie cannot interpret them."

For example, the example of the fighting roosters mentioned above (one of which suddenly pecks around the floor as if taking in food) can also be interpreted as a social signal that may indicate to the rival that the pecking rooster feels so superior that he can still take food even in this precarious situation.

Wolfgang Wickler , a student of Konrad Lorenz, assessed the concept of the skipping movement as outdated as early as 1990: "The action-specific energy turned out to be a modern phlogiston and the psychohydraulic model, in spite of ingenious changes, was unsuitable for adequately depicting changes in readiness and state in the animal."

See also

Individual evidence

  1. ↑ a b Peter Sevenster: The skipping movement. In: Grzimeks Tierleben , supplementary volume behavior research. Kinder Verlag, Zurich 1974, p. 225.
  2. ^ Juan D. Delius: Displacement Activities and Arousal. In: Nature. Volume 214, 1967, pp. 1259-1260, doi: 10.1038 / 2141259a0 .
  3. ↑ a b Nikolaas Tinbergen : Derived activities: their causation, biological significance, origin and emancipation during evolution. In: The Quarterly Review of Biology. Volume 27, 1952, p. 25.
  4. ↑ a b Gerard Baerends : Structure of animal behavior. In: Handbook of Zoology. Volume 8: Mammalia. 10th part, 1st half, 1956, p. 18.
  5. ^ Klaus Immelmann : Introduction to behavior research. Berlin: ³Parey 1983, p. 53.
  6. ^ I. Lindner in: Lexicon of Psychology. Volume 3, p. 2383, ISBN 978-3-451-17942-6 .
  7. ↑ Nikolaas Tinbergen: Instinct theory. Parey, Berlin 1952, p. 108.
  8. ↑ a b J. JA van Iersel, AC Angela Bol: Preening of two tern species. A study on displacement activities. In: Behavior. Volume 13, 1958, pp. 1-88, abstract .
    Peter Sevenster: A causal analysis of a displacement activity (fanning in Gasterosteus aculeatus). In: Behavior. Supplement 9, 1961, pp. 1-170.
  9. ↑ Bernhard Hassenstein : Instinct, Learning, Playing, Insight. Introduction to behavioral biology. Piper, Munich 1980.
  10. ↑ Bernhard Hassenstein: Functional diagrams as an aid for the presentation of theoretical concepts in behavioral biology. In: Zoological Yearbooks Physiology. Volume 87, 1983, pp. 181-187.
  11. ↑ a b Peter Sevenster: The skipping movement. In: Grzimeks Tierleben, supplementary volume behavior research. Kinder Verlag, Zurich 1974, p. 224.
  12. ↑ Nikolaas Tinbergen: The Skipping Movement. In: Zeitschrift für Tierpsychologie. Volume 4, 1940, pp. 1-40.
  13. ↑ Walter Flumm: On the repetition of skip-feeler cleaning movements of the honeybee when collecting differently concentrated sugar solutions. In: Insectes Sociaux. Volume 32, 1985, pp. 435-444, doi: 10.1007 / BF02224020 .
  14. ↑ psychologie.uni-wuerzburg.de (PDF; 1.0 MB) ( Memento from October 21, 2012 in the Internet Archive ), viewed on July 29, 2013: Josua Haderer: Allgemeine Psychologie I , p. 5: Elementary structures of behavior .
  15. ↑ Gerard Baerends carefully described these hypothetical, behavior-triggering nerve cells as "intra-central mechanisms that coordinate movements even without extra-central information." In: Gerard Baerends: Structure of animal behavior, p. 6.
  16. ↑ Peter Sevenster: The skipping movement. In: Grzimeks Tierleben, supplementary volume behavior research. Kinder Verlag, Zurich 1974, p. 226.
  17. ↑ a b c Hanna-Maria Zippelius : The measured theory. A critical examination of the instinct theory of Konrad Lorenz and behavioral research practice. Braunschweig: Vieweg 1992, p. 260, ISBN 978-3-528-06458-7 .
  18. ↑ Konrad Lorenz : Comparative behavior research. Basics of ethology. Springer Verlag, Vienna and New York 1978, p. 202 f.
  19. ↑ Hanna-Maria Zippelius, Die measured theory, p. 261.
  20. ↑ Wolfgang Wickler : From ethology to sociobiology. In: Jost Herbig, Rainer Hohlfeld (ed.): The second creation. Spirit and Demon in 20th Century Biology. Hanser Verlag, Munich 1990, p. 176.