Proximate and ultimate causes of behavior

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Proximate and ultimate causes of behavior are behavioral research common two, but very different approaches, behaviors to explain. The distinction goes back to the Dutch-British ethologist and Nobel Prize winner Nikolaas Tinbergen . According to Tinbergen, behavior can never be explained monocausally (by a single cause), but always multicausally (by several causes). In principle, both proximate causes (immediate reasons) and ultimate causes (evolutionary biological relationships) can be specified for each behavior.

theory

Four questions

Nikolaas Tinbergen distinguished four questions that should be considered when analyzing every life phenomenon. He called these questions "The Four Whys", in German they were later also referred to as the four basic questions of biological research . They can be formulated like this:

  1. How does behavior work on the chemical , physiological , neuroethological , psychological and social level? (Ask about the causes)
  2. How does behavior develop, how does behavior change in the course of individual life, for example due to environmental influences? (Question about ontogeny )
  3. What are the individual behaviors useful for? (Ask about the adjustment value)
  4. Which mechanisms have led to a certain behavior developing in the course of phylogenesis (tribal history)? (Question about phylogenesis)

Questions 1 and 2 concern proximate causes of the behavior, questions 3 and 4 concern ultimate causes. Three questions had already been identified by Julian Huxley , to which Tinbergen himself pointed; Tinbergen completed the scheme with the question of ontogeny. In many cases, several proximate causes can be named plausibly and often several ultimate causes.

In a similar form, Aristotle had already differentiated exactly four types of causes in natural philosophy , including the “target cause” or “end cause”, which corresponds to the question of the evolutionary advantage in the sense of Tinbergen.

Proximate causes

The proximate causes of a behavior can also be called the immediate causes or the current causes. They are often also called effective causes .

1. Proximate causes include, on the one hand, all internal (physiological, chemical, psychological, etc.) conditions that influence behavior, but also all external triggers (e.g. key stimuli ) and social conditions.

Findings on this basic level are an important prerequisite for understanding the levels above. Knowing, for example, the chemical messengers of nerve cells ( neurotransmitters ) and certain external stimulus constellations that influence behavior is not sufficient to understand the overlying levels of neuroanatomical circuit diagrams and the resulting behavior: “The whole is more than the sum of its elements Share “( Nicolai Hartmann ). In the sciences, there is often talk of an emergence relationship between micro and macro levels.

2. The proximate causes are partly dependent on the individual development of the living being ( ontogenesis ): The triggering stimuli in the environment cannot be derived from the life story, but the reactions of the living being to stimuli and environmental conditions. Tinbergen emphasizes that "many behavioral patterns are simultaneously innate and learned or partially innate and partially learned". As an example, he refers to Eibl-Eibesfeldt's observation (1955) that squirrels have to learn to crack nuts as an effective action, even though the individual components of behavior such as grasping and biting are innate. Tinbergen also praises the term “ instinct - dressage - confinement” by Konrad Lorenz (1937).

Ultimate causes

The ultimate causes of behavior can also be referred to as the evolutionary biological relationships . They are also often mentioned as basic causes .

3. One of the ultimate causes is the adaptive value of a behavior, that is, the question of the benefit for the individual.

While the proximate causes, for example, close monitoring, neurophysiological examinations or dummy experiments often empirically can be secured, make statements on the benefit of a behavior often only more or less plausible hypotheses are that are difficult to verify experimentally.

4. Phylogenesis is also one of the ultimate causes, i.e. those reasons which, in the course of tribal history, have favored the emergence of the behavior in question. They always relate to a selection advantage that the behavior resulted in for the individual or his ancestors.

According to Darwin , species change occurs in the course of tribal history as a result of variability and selection . Random mutations create new variants (mutants); within the framework of ecological limits, the selection of these mutants via the number of reproductive offspring promotes or hampers. Since many characteristics (some of the phylogenetic preconditions) remain in the course of evolution, every organism consists of characteristics of different ages; this applies equally to the construction plan and to performance of the behavior. By reconstructing the preconditions of the phylogenetic history, it is possible to interpret the being-so-and-not-different of many (behavioral) characteristics.

When it comes to the question of how behavior develops in the course of phylogenesis, the species comparison plays a central role, i.e. the comparison of animal species, but also the animal-human comparison. Through pure behavioral observation, phylogenetic relationships can mostly only be determined in relation to smaller taxonomic units; this approach is therefore only useful at the level of orders , families and genres . One example is to compare the facial expressions of apes and humans. In human ethology , similarities that are independent of culture from the comparison of cultures are considered to be indications of the possibility of innate behavior (see e.g. universals of music perception ).

Behaviors are often clearer and easier to analyze in animals whose nervous system has a simpler structure. On this basis it is then possible to investigate whether and in what way similar performance qualities exist in higher organisms and in humans - and through which performance qualities one species of animal stands out from others and humans from the animal kingdom. These aspects are discussed in the context of the theory of Konrad Lorenz et al. a. covered in his book The Back of the Mirror . It relates to the phylogenetic relationships of behavioral performance in relation to the large systematics.

Extension to cells, organs and social groups

In 1985, Nikolaas Tinbergen stated in retrospect that the exact delimitation of his four questions "has promoted the clarity of our scientific thinking about behavior - and certainly also about life processes in general". It has proven useful to examine the question of the proximate and ultimate causes not only on the levels of the individual and the group preferred by ethology and sociobiology , but also to apply it generally to all life phenomena, i.e. also to the functions of cells and the organs as well as social processes:

Causations ontogenesis Adjustment value phylogenesis
molecule
cell
organ
individual
Family + child
group
society

classification

Proximate and ultimate causes - on the one hand, direct causes as a trigger for ostensibly recognizable mechanisms, on the other hand, deeper causes ("backgrounds") - can be analyzed in many areas. These include historical events (e.g. wars), political decisions, technical incidents (e.g. a plane crash) and experiences from everyday life (e.g. the outbreak of a dispute). The four questions after Tinbergen are based on the knowledge that different types of causes work together in complex systems. The categories ontogeny , adaptive value and phylogeny in Tinbergen result from the tailoring of the general question to the field of behavioral biology; in other areas they are not applicable or only applicable in a figurative sense.

Examples

Social grooming

Question: Why do members of individual primate species show their affection through social grooming ("Lausen")?

Proximate connections

Ultimate connections

  • Adjustment value: Friendly behavior (behavior that is useful or pleasant for another member of the group) helps to create and maintain bonds. Social ties are beneficial on many occasions, e.g. B. in protection against predators , in the common hunt, in the common brood care or in the mediation of conflicts in the group.
  • Phylogenesis: brood care (a form of one-sided altruism ) and the parent-child bond of the (mammalian) ancestors of today's primates were, according to Irenäus Eibl-Eibesfeldt, phylogenetic preconditions for the evolution of individualized bonds between adults and for so-called reciprocal altruism. As part of this evolutionary development, elements of brood care behavior were adopted as socially friendly behavior in social grooming.

Smiling greetings

Question: Why do you smile when you meet good friends?

The immediate (proximate) cause is the encounter. The ultimate cause of the smile are innate movement patterns of the facial muscles, which among other things serve to signal a positive relationship.

cat and mouse

Question: Why does a mouse escape from the cat in its hole?

The immediate (proximate) cause of the escape reaction is the appearance of the cat. The ultimate cause is an inheritable behavioral program that emerged in phylogeny . Its purpose is to increase the mouse's chances of reproduction .

Infanticide in lions

Question: Why is the new alpha male in a pride of lions killing all of the young?

If a pack leader is ousted by a younger and stronger male lion , the new pack leader often kills all the young. This infanticide in lions is usually interpreted sociobiologically : the successful lion can produce offspring more quickly in this way, since those females who lose their young quickly come back into the oestrus and mate with the new male as soon as possible. John Alcock points out in his behavioral textbook that lionesses can give birth every two years. Since a male is at the head of a pack for an average of only two years, the reproductive advantage for the alpha male is obvious. This interpretation concerns the ultimate causes of behavior. Proximate causes (the immediate triggers for the infanticide) are hardly ever mentioned in this context, as they have apparently not yet been analyzed.

literature

  • Norbert Bischof : Structure and meaning . 1998, ISBN 3-456-83080-7 ( systems theory for psychologists, with an introduction to, inter alia, proximate and ultimate systems analysis , information theory , operator calculus , Z-transformation and semiotics )
  • Konrad Lorenz : Biological questions in animal psychology . In: Zeitschrift für Tierpsychologie. Volume 1, 1937, pp. 24-32.
  • Konrad Lorenz: About the formation of the concept of instinct . In: The natural sciences. Volume 25, 1937, pp. 289-300, 307-318, 324-331.
  • Konrad Lorenz: About the concept of instinctive action . In: Folia Biotheoretica. Volume 2, 1937, pp. 17-50.
  • Nikolaas Tinbergen : The Study of Instinct. Clarendon Press, Oxford 1951.
  • Nikolaas Tinbergen: On Aims and Methods of Ethology. In: Zeitschrift für Tierpsychologie. Volume 20, 1963, pp. 410-433, full text (PDF ).

Web links

  • Gerhard Medicus : Fundamentals of anthropology. An interdisciplinary science with biological roots. In: Naturwissenschaftliche Rundschau. 59th year, 2006, No. 2, pp. 65–71, full text (PDF)

Individual evidence

  1. ↑ Nikolaas Tinbergen 1951 and 1963
  2. ↑ Nikolaas Tinbergen: On Aims and Methods of Ethology. In: Zeitschrift für Tierpsychologie. Volume 20, 1963, pp. 410-433, here p. 411, full text (PDF ).
  3. ↑ Nikolaas Tinbergen: On Aims and Methods of Ethology. In: Zeitschrift für Tierpsychologie. Volume 20, 1963, p. 425, full text (PDF ), quotation: “[…] many bevore patterns can be said to be at the same time innate and learned, or partly innate and partly learned.”
  4. ↑ Nikolaas Tinbergen: Watching and wondering. In: Donald A. Dewsbury: Studying animal behavior. Autobiographies of the Founders. Chicago University Press, 1985.
  5. ↑ John Alcock : The behavior of animals from an evolutionary perspective. G. Fischer, Stuttgart, Jena and New York 1996, pp. 11-12, ISBN 978-3-437-20531-6 .