Tinbergen's four questions

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:

behavioural adaptive functions
phylogenetic history; and the proximate explanations
underlying physiological mechanisms
ontogenetic/developmental history.^[2]^{[page needed]}

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Behavior is action in response to a stimulus. My cat Cameo is now responding to both an external stimulus the sound of a bag of treats, and an internal stimulus her hunger, or at least her insatiable desire for treats. Sometimes animal behavior can seem really far out, but if you look closely enough, you can see how all behavior serves a purpose to help an animal mate, eat, avoid predators, and raise young. And since behaviors can come with advantages like these, natural selection acts on them just as it acts on physical traits ensuring the success of animals who engage in beneficial behaviors, while weeding out those that do stupid, dangerous or otherwise unhelpful stuff. The most beneficial behaviors are those that make an animal better at doing the only two things in the world that matter: eating and sex. Still, that doesn't mean all behavior is about just looking out for number one. It turns out some advantageous behavior is actually pretty selfless. More on that in a minute. But first, behavior is really just a product of a pair of factors: Morphology, or the physical structure of an animal and physiology, or the function of that morphology. Now, an animal's behavior is obviously limited by what its body is capable of doing for example, Cameo does not have opposable thumbs, so, much as she would like to get into the treat bag, by herself, she cannot. This limitation is strictly hereditary no cats can open treat bags with their thumbs because no cats have opposable thumbs. Though some cats do have thumbs. In the same way that a penguin can't fly to escape a predator; or a gazelle can't reach the same leaves as a giraffe can. Similarly, behavior is constrained by an animal's physiology. Like, Cameo's built for chasing down little critters and eating meat, not beds of lettuce. This is because her physiology, everything from her teeth to her digestive system, are geared for eating meat. If she pounced on and ate every blade of grass she came across... let's just say I would not want to be in charge of that litter box. Now the traits that make up an animal's morphology and physiology are often heritable, so we generally talk about selection acting on those traits. But as natural selection hones these traits, it's really selecting their associated behaviors. It's the USE of the trait, using wings and feathers to escape predators, or using a long neck to reach leaves, that provides the evolutionary advantage. Still, that doesn't mean all behavior is coded in an animal's genes some behaviors are learned. And even for animals that learn how to do things, natural selection has favored brain structures that are capable of learning. So one way or another, most behaviors have some genetic underpinning, and we call those behaviors adaptive. Problem is, it's not always obvious what the evolutionary advantages are for some of the nutty things that animals do. Like, why does a snapping turtle always stick out its tongue? How does a tiny Siberian hamster find its mate, miles across the unforgiving tundra? Why does a bower bird collect piles of garbage? To answer questions like those, we have to figure out what stimulus causes these behaviors, and what functions the behaviors serve. To do this, I'm going to need the help of one of the first animal behavior scientists ever, or ethologists, Niko Tinbergen. Tinbergen developed a set of four questions aimed at understanding animal behavior. The questions focus on how a behavior occurs, and why natural selection has favored this particular behavior. Determining how a behavior occurs actually involves two questions: One: what stimulus causes it? And two: what does the animal's body do in response to that stimulus? These are the causes that are closest to the specific behavior we're looking at, so they're called the proximate causes. In the case of the male Siberian hamster, the stimulus is a delicious smelling pheromone that the sexy female hamster releases when she's ready to mate. The male hamster's response, of course, is to scuttle, surprisingly quickly, over several miles if necessary to find and mate with her. So the proximate cause of this behavior was that the girl hamster signaled that she was ready to knock boots, and the male ran like crazy to get to the boot-knockin'. Asking the more complex question of why natural selection has favored this behavior requires asking two more questions: One: what about this behavior helps this animal survive and/or reproduce? And two: what is the evolutionary history of this behavior? These, as you can tell, are bigger-picture questions, and they show us the ultimate causes of the behavior. The answer to the first question, of course, is that the ability of a male hamster to detect and respond to the pheromones of an ovulating female is directly linked to his reproductive success! As for the second question, you can also see that male hamsters with superior pheromone detectors will be able to find females more successfully than other male hamsters, and thereby produce more offspring. So natural selection has honed this particular physical ability and behavior over generations of hamsters. So, who would have thought to ask these questions in the first place? And where's my chair? Niko Tinbergen was one third of a trifecta of revolutionary ethologists in the 20th century. Along with Austrians Karl von Frisch and Konrad Lorenz, he provided a foundation for studying animal behavior and applied these ideas to the study of specific behaviors and for that all three shared the Nobel Prize in 1973. You may have seen the famous photos of young graylag geese following obediently in a line behind a man. That was Lorenz, and his experiments first conducted in the 1930s introduced the world to imprinting, the formation of social bonds in infant animals, and the behavior that includes both learned and innate components. When he observed newly hatched ducklings and geese, he discovered that waterfowl in particular had no innate recognition of their mothers. In the case of graylag geese, he found the imprinting stimulus to be any nearby object moving away from the young! So when incubator-hatched goslings spent their first hours with Lorenz, not only did they follow him, but they showed no recognition of their real mother or other adults in their species! Unfortunately, Lorenz was also a member of the Nazi party from 1938 to 1943. And in response to some of his studies on degenerative features that arose in hybrid geese, Lorenz warned that it took only a small amount of "tainted blood" to have an influence on a "pure-blooded" race. Unsurprisingly, Nazi party leaders were quick to draw some insane conclusions from Lorenz's behavioral studies in the cause of what they called race hygiene. Lorenz never denied his Nazi affiliation but spent years trying to distance himself from the party and apologizing for getting caught up in that evil. Now how exactly does natural selection act on behavior out there in the world? That's where we turn to those two types of behavior that are the only things in the world that matter: eating and sex-having. Behavior associated with finding and eating food is known as foraging, which you've heard of, and natural selection can act on behaviors that allow animals to exploit food sources while using the least amount of energy possible this sweet spot is known as the optimal foraging model. And the alligator snapping turtle has optimal foraging all figured out. Rather than running around hunting down its prey, it simply sits in the water, and food comes to him. See, the alligator snapping turtle has a long, pink tongue divided into two segments, making it look like a tasty worm to a passing fish. In response to the stimulus of a passing fish, it sticks out its tongue out and wiggles it. Natural selection has, over many generations, acted not only on turtles with pinker and more wiggly tongues to catch more fish, it's also acted on those that best know how and when to wiggle those tongues to get the most food. So it's selecting both the physical trait and the behavior that best exploits it. And what could be sexier than a turtle's wiggly tongue dance? Well, how about sex? As we saw with our friend the horny Siberian hamster, some behaviors and their associated physical features are adapted to allow an animal to reproduce more, simply by being better at finding mates. But many times, animals of the same species live close together or in groups, and determining who in what group gets to mate creates some interesting behaviors and features. This is what sexual selection, is all about. Often, males of a species will find and defend a desirable habitat to raise young in, and females will choose a male based on their territory. But what about those species, and there are many of them, where the female picks a male not because of that, but because of how he dances, or even weirder, how much junk he's collected? Take the male bower bird. He builds an elaborate hut, or bower, out of twigs and bits of grass, then spends an enormous amount of time collecting stuff, sometimes piles of berries, and sometimes piles of pretty, blue, plastic crap. Ethologists believe that he's collecting the stuff to attract the female to check out his elaborate house. Once the female's been enticed to take a closer look, the male starts to sing songs and dance around, often mimicking other species, inside of his little house for her. Females will inspect a number of these bowers before choosing who to mate with. Now, doing more complex dances and having more blue objects in your bower scores bigger with females. And ethologists have shown that a higher level of problem solving, or intelligence, in males correlates to both of these activities. So yeah, it took some brawn to build that bower and collect all that junk, but chicks also dig nerds who can learn dances! So the bowerbird's brain is evolving in response to sexual selection by females. This intelligence likely also translates into other helpful behaviors like avoiding predators. So thanks to the evolution of behavior, we're really good at taking care of our nutritional and sexual needs. But what's confused scientists for a long time is why animals often look after others' needs. For instance, vampire bats in South America will literally regurgitate blood into the mouths of members of its clan who didn't get a meal that night. How do you explain animals who act altruistically like that? We actually did a whole SciShow episode on this very subject but basically, we can thank British scientist William Hamilton for coming up with an equation to explain how natural selection can simultaneously make animals fit and allow for the evolution of altruism. Hamilton found that the evolution of altruism was best understood at the level of larger communities, especially extended animal families. Basically, altruism can evolve if the benefit of a behavior is greater than its cost on an individual, because it helped the individual's relatives enough to make it worth it. Hamilton called this inclusive fitness, expanding Darwin's definition of fitness basically, how many babies somebody's making to include the offspring of relatives. So I guess the only question left is, if I forget to feed you two, who is going to regurgitate blood into the other one's mouth? Yeah, there's probably a reason that only happens with bats. Thank you for watching this episode of Crash Course Biology. Thank you to Cameo for being such a good kitty. Yeah, she finally gets her treats. There's a table of contents, of course. If you want to reinforce any of the knowledge that you gained today. If you have questions or ideas for us you can get in touch with us on Facebook or Twitter, or of course, in the comments below. We'll see you next time.

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 (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
How vs. why questions	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.^[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)

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

Third question: Mechanism (causation)

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)

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.

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:

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

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 "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 Example – PDF).

Notes and references

^ 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.
^ Daly, Martin; Wilson, Margo (1983). Sex, evolution, and behavior (2nd ed.). Boston: Willard Grant Press. ISBN 9780871507679. OCLC 9084620.
^ ^a ^b ^c ^d Tinbergen, Niko (1963) "On Aims and Methods in Ethology," Zeitschrift für Tierpsychologie, 20: 410–433 [411].
^ Nikolaas Tinbergen, ethology, Cartwright 2000:10; Buss 2004:12)
^ ^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
^ "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.
^ "Biased learning" is not necessarily limited to the developmental period.
^ 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.
^ 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.
^ Mapping Transdisciplinarity in Human Sciences. In: Janice W. Lee (Ed.) Focus on Gender Identity. New York, 2005, Nova Science Publishers, Inc. [1]

References

Alcock, John (2001) Animal Behaviour: An Evolutionary Approach, Sinauer, 7th edition. ISBN 0-87893-011-6.
Buss, David M., Martie G. Haselton, Todd K. Shackelford, et al. (1998) "Adaptations, Exaptations, and Spandrels," American Psychologist, 53:533–548. http://www.sscnet.ucla.edu/comm/haselton/webdocs/spandrels.html
Buss, David M. (2004) Evolutionary Psychology: The New Science of the Mind, Pearson Education, 2nd edition. ISBN 0-205-37071-3.
Cartwright, John (2000) Evolution and Human Behaviour, MIT Press, ISBN 0-262-53170-4.
Krebs, J.R., Davies N.B. (1993) An Introduction to Behavioural Ecology, Blackwell Publishing, ISBN 0-632-03546-3.
Lorenz, Konrad (1937) Biologische Fragestellungen in der Tierpsychologie (I.e. Biological Questions in Animal Psychology). Zeitschrift für Tierpsychologie, 1: 24–32.
Mayr, Ernst (2001) What Evolution Is, Basic Books. ISBN 0-465-04425-5.
Gerhard Medicus. "Tinbergen's four questions in behavioural Anthropology" (PDF).
Gerhard Medicus (2017) Being Human – Bridging the Gap between the Sciences of Body and Mind. Berlin: VWB 2015, ISBN 978-3-86135-584-7
Nesse, Randolph M (2013) "Tinbergen's Four Questions, Organized," Trends in Ecology and Evolution, 28:681-682.
Moore, David S. (2001) The Dependent Gene: The Fallacy of 'Nature vs. Nurture', Henry Holt. ISBN 0-8050-7280-2.
Pinker, Steven (1994) The Language Instinct: How the Mind Creates Language, Harper Perennial. ISBN 0-06-097651-9.
Tinbergen, Niko (1963) "On Aims and Methods of Ethology," Zeitschrift für Tierpsychologie, 20: 410–433.
Wilson, Edward O. (1998) Consilience: The Unity of Knowledge, Vintage Books. ISBN 0-679-76867-X.

External links

Diagrams

Derivative works

On aims and methods of cognitive ethology (pdf) by Jamieson and Bekoff.

v t e Ethology
Branches	Animal cognition Animal communication Animal consciousness Animal culture Animal sexual behaviour Animal welfare science Anthrozoology Bee learning and communication Behavioural ecology Behavioral endocrinology Behavioural genetics Cognitive ethology Comparative psychology Emotion in animals Evolutionary neuroscience Feeding Human ethology Instinct Learning Neuroethology Pain in animals Philosophical ethology Sociobiology Stereotypy Structures Hive Nest Instinct Swarm Tool use by non-humans Zoosemiotics Zoomusicology
Ethologists	Patrick Bateson Marc Bekoff Donald Broom John B. Calhoun Charles Darwin Marian Dawkins Richard Dawkins Irenäus Eibl-Eibesfeldt Dian Fossey Karl von Frisch Jane Goodall Heini Hediger Julian Huxley Konrad Lorenz Desmond Morris Thomas Sebeok William Homan Thorpe Nikolaas Tinbergen Jakob von Uexküll Wolfgang Wickler E. O. Wilson Solly Zuckerman
Societies	Association for the Study of Animal Behaviour International Society for Applied Ethology
Journals	Animal Behaviour Animal Cognition Animal Welfare Behavioral Ecology Behaviour
Category

v t e Nikolaas Tinbergen
Selected works	The Study of Instinct (book) The Herring Gull's World (book) Social Behaviour in Animals: With Special Reference to Vertebrates (book) Signals for Survival (film) The Riddle of the Rook (film)
General	Animal behaviour science Behavioural biology Behavioral ecology History of ethology Neuroethology Sociobiology Hawk/goose effect Supernormal stimulus Tinbergen's four questions
Related	Jan Tinbergen Luuk Tinbergen Konrad Lorenz Karl von Frisch Richard Dawkins Desmond Morris 1973 Nobel Prize in Physiology or Medicine

This page was last edited on 6 February 2024, at 09:24

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