Tinbergen's four questions

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Tinbergen's four questions, named after 20th century biologist Nikolaas Tinbergen, are complementary categories of explanations for animal behaviour. These are also commonly referred to as levels of analysis.[1] It suggests that an integrative understanding of behaviour must include ultimate (evolutionary) explanations, in particular:

Four categories of questions and explanations[edit]

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 (function/adaptation). Biologists have three additional explanations: sight is caused by a particular series of evolutionary steps (phylogeny), the mechanics of the eye (mechanism/causation), and even the process of an individual's development (ontogeny). This schema constitutes a basic framework of the overlapping behavioural fields of ethology, 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[edit]

First question: Function (adaptation)[edit]

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.[4] On the other hand, the common definition of adaptation is a central concept in evolution: a trait that was functional to the reproductive success of the organism and that is thus now present due to being selected for; that is, function and evolution are inseparable. However, a trait can have a current function that is adaptive without being an adaptation in this sense, if for instance the environment has changed. Imagine an environment in which having a small body suddenly conferred benefit on an organism when previously body size had had no effect on survival.[3] A small body's function in the environment would then be adaptive, but it would not become an adaptation until enough generations had passed in which small bodies were advantageous to reproduction for small bodies to be selected for. Given this, it is best to understand that presently functional traits might not all have been produced by natural selection.[3] The term "function" is preferable to "adaptation", because adaptation is often construed as implying that it was selected for due to past function. This corresponds to Aristotle's final cause.[5]

Second question: Phylogeny (evolution)[edit]

Evolution captures both the history of an organism via its phylogeny, and the history of natural selection working on function to produce adaptations.[6] There are several reasons why natural selection may fail to achieve optimal design (Mayr 2001:140–143; Buss et al. 1998). One entails random processes such as mutation and environmental events acting on small populations. Another entails the constraints resulting from early evolutionary development. Each organism harbors traits, both anatomical and 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[edit]

Third question: Mechanism (causation)[edit]

Some prominent classes of Proximate causal mechanisms include:

  • The brain: For example, Broca's area, a small section of the human brain, has a critical role in linguistic capability.
  • Hormones: Chemicals used to communicate among cells of an individual organism. Testosterone, for instance, stimulates aggressive behaviour in a number of species.
  • Pheromones: Chemicals used to communicate among members of the same species. Some species (e.g., dogs and some moths) use pheromones to attract mates.

In examining living organisms, biologists are confronted with diverse levels of complexity (e.g. chemical, physiological, psychological, social). They therefore investigate causal and functional relations within and between these levels. A biochemist might examine, for instance, the influence of social and ecological conditions on the release of certain 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 efficient cause.[5]

Fourth question: Ontogeny (development)[edit]

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 critical period, for instance, for 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[edit]

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.



Four ways of explaining visual perception:

  • Function: To find food and avoid danger.
  • Phylogeny: The vertebrate eye initially developed with a blind spot, but the lack of adaptive intermediate forms prevented the loss of the blind spot.
  • Causation: The lens of the eye focuses light on the retina.
  • Development: Neurons need the stimulation of light to wire the eye to the brain (Moore, 2001:98–99).

Westermarck effect[edit]

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[edit]

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 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"[edit]

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 "biopsychosocial" framework clarifies and classifies the associations between the various levels of the natural and social sciences, and it helps to integrate the social and natural sciences into a "tree of knowledge" (see also Nicolai Hartmann's "Laws about the Levels of Complexity"). Especially for the social sciences, this model helps to provide an integrative, foundational model for interdisciplinary collaboration, teaching and research (see The Four Central Questions of Biological Research Using Ethology as an ExamplePDF).


  1. ^ MacDougall-Shackleton, Scott A. (2011-07-27). "The levels of analysis revisited". Philosophical Transactions of the Royal Society B: Biological Sciences. 366 (1574): 2076–2085. doi:10.1098/rstb.2010.0363. PMC 3130367. PMID 21690126.
  2. ^ Daly, Martin; Wilson, Margo (1983). Sex, evolution, and behavior (2nd ed.). Boston: Willard Grant Press. ISBN 9780871507679. OCLC 9084620.
  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. ^ Bode, Adam; Kushnick, Geoff (2021). "Proximate and Ultimate Perspectives on Romantic Love". Frontiers in Psychology. 12: 573123. doi:10.3389/fpsyg.2021.573123. ISSN 1664-1078. PMC 8074860. PMID 33912094.
  9. ^ Bode, Adam; Kuula, Liisa (September 2021). "Romantic Love and Sleep Variations: Potential Proximate Mechanisms and Evolutionary Functions". Biology. 10 (9): 923. doi:10.3390/biology10090923. PMC 8468029. PMID 34571801.
  10. ^ Mapping Transdisciplinarity in Human Sciences. In: Janice W. Lee (Ed.) Focus on Gender Identity. New York, 2005, Nova Science Publishers, Inc. [1]


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