Tinbergen's four questions

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Tinbergen's four questions, named after 20th century biologist

behaviour must include ultimate (evolutionary
) explanations, in particular:

Four categories of questions and explanations

When asked about the purpose of sight in humans and animals, even elementary-school children can answer that animals have vision to help them find food and avoid danger (

behavioural ecology, comparative psychology, sociobiology, evolutionary psychology, and anthropology. Julian Huxley identified the first three questions. Niko Tinbergen gave only the fourth question, as Huxley's questions failed to distinguish between survival value and evolutionary history; Tinbergen's fourth question helped resolve this problem.[3]

Table of categories
Diachronic versus synchronic perspective
Dynamic view
Explanation of current form in terms of a historical sequence
Static view
Explanation of the current form of species
How vs. why questions Proximate view
How an individual organism's structures function
Ontogeny (development)
Developmental explanations for changes in individuals, from DNA to their current form
Mechanism (causation)
Mechanistic explanations for how an organism's structures work
Ultimate (evolutionary) view
Why a species evolved the structures (adaptations) it has
Phylogeny (evolution)
The history of the evolution of sequential changes in a species over many generations
Function (adaptation)
A species trait that solves a reproductive or survival problem in the current environment

Evolutionary (ultimate) explanations

First question: Function (adaptation)

Darwin's theory of evolution by natural selection is the only scientific explanation for why an animal's behaviour is usually well adapted for survival and reproduction in its environment. However, claiming that a particular mechanism is well suited to the present environment is different from claiming that this mechanism was selected for in the past due to its history of being adaptive.[3]

The literature conceptualizes the relationship between function and evolution in two ways. On the one hand, function and evolution are often presented as separate and distinct explanations of behaviour.

final cause.[5]

Second question: Phylogeny (evolution)

behavioural, of previous phylogenetic stages, since many traits are retained as species
evolve.

Reconstructing the phylogeny of a species often makes it possible to understand the "uniqueness" of recent characteristics: Earlier phylogenetic stages and (pre-) conditions which persist often also determine the form of more modern characteristics. For instance, the vertebrate eye (including the human eye) has a blind spot, whereas octopus eyes do not. In those two lineages, the eye was originally constructed one way or the other. Once the vertebrate eye was constructed, there were no intermediate forms that were both adaptive and would have enabled it to evolve without a blind spot.

It corresponds to Aristotle's

formal cause.[5]

Proximate explanations

Third question: Mechanism (causation)

Some prominent classes of Proximate

include:

In examining living organisms,

neurotransmitters and hormones, and the effects of such releases on behaviour, e.g. stress during birth has a tocolytic
(contraction-suppressing) effect.

However, awareness of neurotransmitters and the structure of

neurons is not by itself enough to understand higher levels of neuroanatomic structure or behaviour: "The whole is more than the sum of its parts." All levels must be considered as being equally important: cf. transdisciplinarity, Nicolai Hartmann
's "Laws about the Levels of Complexity."

It corresponds to Aristotle's

Fourth question: Ontogeny (development)

Ontogeny is the process of development of an individual organism from the zygote through the embryo to the adult form.

In the latter half of the twentieth century, social scientists debated whether human behaviour was the product of nature (genes) or nurture (environment in the developmental period, including culture).

An example of interaction (as distinct from the sum of the components) involves familiarity from childhood. In a number of species, individuals prefer to associate with familiar individuals but prefer to mate with unfamiliar ones (Alcock 2001:85–89, Incest taboo, Incest). By inference, genes affecting living together interact with the environment differently from genes affecting mating behaviour. A simple example of interaction involves plants: Some plants grow toward the light (phototropism) and some away from gravity (gravitropism).

Many forms of developmental learning have a

imprinting among geese and language acquisition
among humans. In such cases, genes determine the timing of the environmental impact.

A related concept is labeled "biased learning" (Alcock 2001:101–103) and "prepared learning" (Wilson, 1998:86–87). For instance, after eating food that subsequently made them sick, rats are predisposed to associate that food with smell, not sound (Alcock 2001:101–103). Many primate species learn to fear snakes with little experience (Wilson, 1998:86–87).[7]

See developmental biology and developmental psychology.

Explanations of Animal Behaviour: Causal Relationships; Adopted from Tinbergen (1963).

It corresponds to Aristotle's

material cause.[5]

Causal relationships

The figure shows the causal relationships among the categories of explanations. The left-hand side represents the evolutionary explanations at the species level; the right-hand side represents the proximate explanations at the individual level. In the middle are those processes' end products—genes (i.e., genome) and behaviour, both of which can be analyzed at both levels.

Evolution, which is determined by both function and phylogeny, results in the genes of a population. The genes of an individual interact with its developmental environment, resulting in mechanisms, such as a nervous system. A mechanism (which is also an end-product in its own right) interacts with the individual's immediate environment, resulting in its behaviour.

Here we return to the population level. Over many generations, the success of the species' behaviour in its ancestral environment—or more technically, the environment of evolutionary adaptedness (EEA) may result in evolution as measured by a change in its genes.

In sum, there are two processes—one at the population level and one at the individual level—which are influenced by environments in three time periods.

Examples

Vision

Four ways of explaining visual perception:

Westermarck effect

Four ways of explaining the Westermarck effect, the lack of sexual interest in one's siblings (Wilson, 1998:189–196):

  • Function: To discourage inbreeding, which decreases the number of viable offspring.
  • Phylogeny: Found in a number of mammalian species, suggesting initial evolution tens of millions of years ago.
  • Mechanism: Little is known about the neuromechanism.
  • Ontogeny: Results from familiarity with another individual early in life, especially in the first 30 months for humans. The effect is manifested in nonrelatives raised together, for instance, in kibbutzs.

Romantic love

Four ways of explaining romantic love have been used to provide a comprehensive biological definition (Bode & Kushnick, 2021):[8]

  • Function: Mate choice, courtship, sex, pair-bonding.
  • Phylogeny: Evolved by co-opting mother-infant bonding mechanisms sometime in the recent evolutionary history of humans.
  • Mechanisms: Social, psychological mate choice, genetic, neurobiological, and endocrinological mechanisms cause romantic love.
  • Ontogeny: Romantic love can first manifest in childhood, manifests with all its characteristics following puberty, but can manifest across the lifespan.

Sleep

Sleep has been described using Tinbergen's four questions as a framework (Bode & Kuula, 2021):[9]

  • Function: Energy restoration, metabolic regulation, thermoregulation, boosting immune system, detoxification, brain maturation, circuit reorganization, synaptic optimization, avoiding danger.
  • Phylogeny: Sleep exists in invertebrates, lower vertebrates, and higher vertebrates. NREM and REM sleep exist in eutheria, marsupialiformes, and also evolved in birds.
  • Mechanisms: Mechanisms regulate wakefulness, sleep onset, and sleep. Specific mechanisms involve neurotransmitters, genes, neural structures, and the circadian rhythm.
  • Ontogeny: Sleep manifests differently in babies, infants, children, adolescents, adults, and older adults. Differences include the stages of sleep, sleep duration, and sex differences.

Use of the four-question schema as "periodic table"

Konrad Lorenz, Julian Huxley and Niko Tinbergen were familiar with both conceptual categories (i.e. the central questions of biological research: 1. - 4. and the levels of inquiry: a. - g.), the tabulation was made by Gerhard Medicus.[10] The tabulated schema is used as the central organizing device in many animal behaviour, ethology, behavioural ecology and evolutionary psychology textbooks (e.g., Alcock, 2001) . One advantage of this organizational system, what might be called the "periodic table of life sciences," is that it highlights gaps in knowledge, analogous to the role played by the periodic table of elements in the early years of chemistry.

1. Mechanism 2. Ontogeny 3. Function 4. Phylogeny
a. Molecule
b. Cell
c. Organ
d. Individual
e. Family
f. Group
g. Society

This "

interdisciplinary collaboration, teaching and research (see The Four Central Questions of Biological Research Using Ethology as an ExamplePDF
).

Notes and references

  1. .
  2. .
  3. ^ a b c d Tinbergen, Niko (1963) "On Aims and Methods in Ethology," Zeitschrift für Tierpsychologie, 20: 410–433 [411].
  4. ^ Nikolaas Tinbergen, ethology, Cartwright 2000:10; Buss 2004:12)
  5. ^ a b c d Hladký, V. & Havlíček, J. (2013). Was Tinbergen an Aristotelian? Comparison of Tinbergen's Four Whys and Aristotle's Four Causes. Human Ethology Bulletin, 28(4), 3–11
  6. ^ "Phylogeny" often emphasizes the evolutionary genealogical relationships among species (Alcock 2001:492; Mayr, 2001:289) as distinct from the categories of explanations. Although the categories are more relevant in a conceptual discussion, the traditional term is retained here.
  7. ^ "Biased learning" is not necessarily limited to the developmental period.
  8. PMID 33912094
    .
  9. .

References

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Diagrams

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