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Nikolaas Tinbergen

From Wikipedia, the free encyclopedia

Niko Tinbergen
Nikolaas Tinbergen 1978.jpg
Tinbergen in 1978
Born Nikolaas Tinbergen
(1907-04-15)15 April 1907
The Hague, Netherlands
Died 21 December 1988(1988-12-21) (aged 81)
Oxford, England
Residence United Kingdom
Nationality Dutch
Alma mater Leiden University
Known for
Spouse(s) Elisabeth Rutten (1912–1990)
Children 5
Awards
Scientific career
Fields
Institutions University of Oxford
Doctoral advisor Hilbrand Boschma[2]
Doctoral students
Influenced Robert Hinde

Nikolaas 'Niko' Tinbergen FRS[1] (/ˈtɪnbɜːrɡən/; Dutch pronunciation: [ˈnikoːlaːs ˈnikoː ˈtɪnbɛrɣən]; 15 April 1907 – 21 December 1988) was a Dutch biologist and ornithologist who shared the 1973 Nobel Prize in Physiology or Medicine with Karl von Frisch and Konrad Lorenz[6][7][8][9][10] for their discoveries concerning organisation and elicitation of individual and social behaviour patterns in animals. He is regarded as one of the founders of modern ethology, the study of animal behaviour.

In 1951, he published The Study of Instinct, an influential book on animal behaviour. In the 1960s, he collaborated with filmmaker Hugh Falkus on a series of wildlife films, including The Riddle of the Rook (1972) and Signals for Survival (1969), which won the Italia prize in that year and the American blue ribbon in 1971.

YouTube Encyclopedic

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  • Animal Behavior - CrashCourse Biology #25
  • Introduction to Animal Behaviour
  • Tinbergen's Experiment - Three-Spined Stickleback
  • John Maynard Smith - The ethologists: Tinbergen and Lorenz (28/102)
  • Nikolaas Tinbergen

Transcription

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.

Contents

Education and early life

Born in The Hague, Netherlands, he was one of five children of Dirk Cornelis Tinbergen and his wife Jeannette van Eek. His brother, Jan Tinbergen, won the first Bank of Sweden Prize in Economic Sciences in Memory of Alfred Nobel in 1969.[11] They are the only siblings to each win a Nobel Prize.[12] Another brother, Luuk Tinbergen was also a noted biologist.

Tinbergen's interest in nature manifested itself when he was young. He studied biology at Leiden University and was a prisoner of war during World War II. Tinbergen's experience as a prisoner of the Nazis led to some friction with longtime intellectual collaborator Konrad Lorenz, and it was several years before the two reconciled.[13]

After the war, Tinbergen moved to England, where he taught at the University of Oxford and was a fellow first at Merton College, Oxford and later at Wolfson College, Oxford.[13] Several of his graduate students went on to become prominent biologists including Richard Dawkins,[3] Marian Dawkins,[citation needed] Desmond Morris,[5] Iain Douglas-Hamilton,[14] and Tony Sinclair.[15]

Research and career

The Study of Instinct

 Figure 1. Tinbergen's hierarchical model. Modified from The Study of Instinct (1951).
Figure 1. Tinbergen's hierarchical model. Modified from The Study of Instinct (1951).

In 1951 Tinbergen's The Study of Instinct was published. Behavioural ecologists and evolutionary biologists still recognise the contribution this book offered the field of behavioural science studies. The Study of Instinct summarises Tinbergen's ideas on innate behavioural reactions in animals and the adaptiveness and evolutionary aspects of these behaviours. By behaviour, he means the total movements made by the intact animal; innate behaviour is that which is not changed by the learning process. The major question of the book is the role of internal and external stimuli in controlling the expression of behaviour.[16]

In particular, he was interested in explaining 'spontaneous' behaviours: those that occurred in their complete form the first time they were performed and that seemed resistant to the effects of learning. He explains how behaviour can be considered a combination of these spontaneous behaviour patterns and as set series of reactions to particular stimuli. Behaviour is a reaction in that to a certain extent it is reliant on external stimuli, however it is also spontaneous since it is also dependent upon internal causal factors.[16]

His model for how certain behavioural reactions are provoked was based on work by Konrad Lorenz. Lorenz postulated that for each instinctive act there is a specific energy which builds up in a reservoir in the brain. In this model, Lorenz envisioned a reservoir with a spring valve at its base that an appropriate stimulus could act on, much like a weight on a scale pan pulling against a spring and releasing the reservoir of energy, an action which would lead an animal to express the desired behaviour.[16]

Tinbergen added complexity to this model, a model now known as Tinbergen's hierarchical model. He suggested that motivational impulses build up in nervous centres in the brain which are held in check by blocks. The blocks are removed by an innate releasing mechanism that allows the energy to flow to the next centre (each centre containing a block that needs to be removed) in a cascade until the behaviour is expressed. Tinbergen's model shows multiple levels of complexity and that related behaviours are grouped.[17]

An example is in his experiments with foraging honey bees. He showed that honey bees show curiosity for yellow and blue paper models of flowers, and suggested that these were visual stimuli causing the buildup of energy in one specific centre. However, the bees rarely landed on the model flowers unless the proper odour was also applied. In this case, the chemical stimuli of the odour allowed the next link in the chain to be released, encouraging the bee to land. The final step was for the bee to insert its mouthparts into the flower and initiate suckling. Tinbergen envisioned this as concluding the reaction set for honey bee feeding behaviour.[18]

Nobel Prize

In 1973 Tinbergen, along with Konrad Lorenz and Karl von Frisch, were awarded the Nobel Prize in Physiology or Medicine "for their discoveries concerning organization and elicitation of individual and social behaviour patterns".[19] The award recognised their studies on genetically programmed behaviour patterns, their origins, maturation and their elicitation by key stimuli. In his Nobel Lecture, Tinbergen addressed the somewhat unconventional decision of the Nobel Foundation to award the prize for Physiology or Medicine to three men who had until recently been regarded as "mere animal watchers". Tinbergen stated that their revival of the "watching and wondering" approach to studying behaviour could indeed contribute to the relief of human suffering.[20]

The studies performed by the trio on fish, insects and birds laid the foundation for further studies on the importance of specific experiences during critical periods of normal development, as well as the effects of abnormal psychosocial situations in mammals. At the time, these discoveries were stated to have caused "a breakthrough in the understanding of the mechanisms behind various symptoms of psychiatric disease, such as anguish, compulsive obsession, stereotypic behaviour and catatonic posture".[21] Tinbergen’s contribution to these studies included the testing of the hypotheses of Lorenz/von Frisch by means of "comprehensive, careful, and ingenious experiments"[22] as well as his work on supernormal stimuli. The work of Tinbergen during this time was also regarded as having possible implications for further research in child development and behaviour.[23]

He also caused some intrigue by dedicating a large part of his acceptance speech to FM Alexander, originator of the Alexander technique, a method which investigates postural reflexes and responses in human beings.[24]

Other awards and honours

In 1950 Tinbergen became member of the Royal Netherlands Academy of Arts and Sciences.[25] He was elected a Fellow of the Royal Society (FRS) in 1962.[1] He was also awarded the Godman-Salvin Medal in 1969 by the British Ornithologists' Union,[26] and in 1973 received the Swammderdam Medal[27] and Wilhelm Bölsche Medal[28] (from the Genootschap ter bervordering van Natuur-, Genees- en Heelkunde of the University of Amsterdam and the Kosmos-Gesellschaft der Naturfreunde respectively).

Approach to animal behaviour

Tinbergen described four questions he believed should be asked of any animal behaviour,[29][30][31][32] which were:

  1. Causation (mechanism): what are the stimuli that elicit the response, and how has it been modified by recent learning? How do behaviour and psyche "function" on the molecular, physiological, neuro-ethological, cognitive and social level, and what do the relations between the levels look like? (compare: Nicolai Hartmann: "The laws about the levels of complexity")
  2. Development (ontogeny): how does the behaviour change with age, and what early experiences are necessary for the behaviour to be shown? Which developmental steps (the ontogenesis follows an "inner plan") and which environmental factors play when / which role? (compare: Recapitulation theory)
  3. Function (adaptation): how does the behaviour impact on the animal's chances of survival and reproduction?
  4. Evolution (phylogeny): how does the behaviour compare with similar behaviour in related species, and how might it have arisen through the process of phylogeny? Why did structural associations (behaviour can be seen as a "time space structure") evolve in this manner and not otherwise?*

In ethology and sociobiology, causation and ontogeny are summarised as the "proximate mechanisms", while adaptation and phylogeny are the "ultimate mechanisms". They are still considered as the cornerstone of modern ethology, sociobiology and transdisciplinarity in Human Sciences.

Supernormal stimulus

A major body of Tinbergen's research focused on what he termed the supernormal stimulus. This was the concept that one could build an artificial object which was a stronger stimulus or releaser for an instinct than the object for which the instinct originally evolved. He constructed plaster eggs to see which a bird preferred to sit on, finding that they would select those that were larger, had more defined markings, or more saturated colour—and a dayglo-bright one with black polka dots would be selected over the bird's own pale, dappled eggs.

Tinbergen found that territorial male three-spined stickleback (a small freshwater fish) would attack a wooden fish model more vigorously than a real male if its underside was redder. He constructed cardboard dummy butterflies with more defined markings that male butterflies would try to mate with in preference to real females. The superstimulus, by its exaggerations, clearly delineated what characteristics were eliciting the instinctual response.

Among the modern works calling attention to Tinbergen's classic work is Deirdre Barrett's 2010 book, Supernormal Stimuli.

Autism

Tinbergen applied his observational methods to the problems of autistic children. He recommended a "holding therapy" in which parents hold their autistic children for long periods of time while attempting to establish eye contact, even when a child resists the embrace.[33] However, his interpretations of autistic behaviour, and the holding therapy that he recommended, lacked scientific support[34] and the therapy is described as controversial and potentially abusive.[35]

Publications

  • Tinbergen, N (1951). The Study of Instinct. Oxford, Clarendon Press.[ISBN missing]
  • Tinbergen, N (1953). The Herring Gull's World. London, Collins.[ISBN missing]
  • Tinbergen, N (1953). Social Behaviour in Animals: With Special Reference to Vertebrates. Methuen & Co.[ISBN missing]
  • Kruuk, H (2003). Niko's Nature: The Life of Niko Tinbergen and His Science of Animal Behaviour. Oxford, Oxford University Press. ISBN 0-19-851558-8
  • Stamp Dawkins, M; Halliday, TR; Dawkins, R (1991). The Tinbergen Legacy. London, Chapman & Hall. ISBN 0-412-39120-1
  • Burkhardt Jr., RW (2005). Patterns of Behavior : Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology. ISBN 0-226-08090-0

Personal life

Tinbergen was a member of the advisory committee to the Anti-Concorde Project[citation needed] and was also an atheist.[36]

Tinbergen married Elisabeth Rutten and they had five children. Later in life he suffered depression and feared he might, like his brother Luuk, commit suicide. He was treated by his friend, whose ideas he had greatly influenced, John Bowlby.[37] Tinbergen died on 21 December 1988, after suffering a stroke at his home in Oxford, England.[1]

References

  1. ^ a b c d Hinde, Robert A. (1990). "Nikolaas Tinbergen. 15 April 1907 – 21 December 1988". Biographical Memoirs of Fellows of the Royal Society. Royal Society. 36: 548. doi:10.1098/rsbm.1990.0043. 
  2. ^ Burkhardt, R. W., Jr. (2010). "Niko Tinbergen" (PDF). Elsevier. pp. 428–433. Retrieved 8 October 2016. 
  3. ^ a b Dawkins, Clinton Richard (1941). Selective pecking in the domestic chick. bodleian.ox.ac.uk (DPhil thesis). University of Oxford. EThOS uk.bl.ethos.710826. 
  4. ^ "Aubrey Manning". University of Edinburgh. Retrieved 8 October 2016. 
  5. ^ a b Beale, Graeme Robert (2009). Tinbergian Practice, themes and variations: the field and laboratory methods and practice of the Animal Behaviour Research Group under Nikolaas Tinbergen at Oxford University. University of Edinburgh (PhD Thesis). 
  6. ^ Tinbergen autobiography at nobelprize.org
  7. ^ The Nobel Prize in Physiology or Medicine 1973: von Frisch, Lorenz and Tinbergen
  8. ^ Tinbergen Nobel Lecture
  9. ^ Dewsbury, D. A. (2003). "The 1973 Nobel Prize for Physiology or Medicine: Recognition for behavioral science?". American Psychologist. 58 (9): 747–752. doi:10.1037/0003-066X.58.9.747. PMID 14584992. 
  10. ^ Raju, T. N. (1999). "The Nobel chronicles. 1973: Karl von Frisch (1886–1982); Konrad Lorenz (1903–89); and Nikolaas Tinbergen (1907–88)". Lancet. 354 (9184): 1130. doi:10.1016/s0140-6736(05)76931-2. PMID 10509540. 
  11. ^ Lundberg, Erik (1969). "The Sveriges Riksbank Prize in Economic Sciences in Memory of Alfred Nobel 1969". Nobelprize.org. Retrieved 27 January 2014. 
  12. ^ "Nobel Prize Facts". 
  13. ^ a b "Encyclopedia.com Nikolaas Tinbergen". Encyclopedia.com. Retrieved 16 March 2014. 
  14. ^ Daston, Lorraine; Mitman, Gregg (2005). Thinking with Animals: New Perspectives on Anthropomorphism. Columbia University Press. p. 193. ISBN 978-0-231-50377-8. 
  15. ^ Sinclair, Anthony (2012). Serengeti Story: Life and Science in the World's Greatest Wildlife Region. Oxford University Press. p. 33. ISBN 9780199645527. 
  16. ^ a b c Hinde, R. A. Ethological Models and the Concept of 'Drive'. British Journal of the Philosophy of Science, 6, 321–331 (1956)
  17. ^ Tinbergen, Nikolaas (1951). The Study of Instinct. Oxford University Press. pp. nnn–mmm. [page needed]
  18. ^ Tinbergen, Nikolaas (1951). The Study of Instinct. Oxford University Press. pp. nnn–mmm. [page needed]
  19. ^ https://www.nobelprize.org/nobel_prizes/medicine/laureates/1973/
  20. ^ Tinbergen, N. Ethology and stress diseases. Physiology Or Medicine: 1971–1980 19711980, 113 (1992)
  21. ^ Zetterström, R. The Nobel Prize for the introduction of ethology, or animal behaviour, as a new research field: possible implications for child development and behaviour: Nobel prizes of importance to Paediatrics. Acta Paediatrica 96, 1105–1108 (2007).
  22. ^ Cronholm, Börje (1973). "Award Ceremony Speech". Nobelprize.org. 
  23. ^ Zetterström, R. The Nobel Prize for the introduction of ethology, or animal behaviour, as a new research field: possible implications for child development and behaviour: Nobel prizes of importance to Paediatrics. Acta Paediatrica 96, 1105–1108 (2007)
  24. ^ Tinbergen, Nikolaas (12 December 1973). "Nikolaas Tinbergen Nobel Lecture Ethology and Stress Diseases" (PDF). NobelPrize.org. 
  25. ^ "Niko Tinbergen (1907–1988)". Royal Netherlands Academy of Arts and Sciences. Retrieved 19 July 2015. 
  26. ^ "Medals and awards " British Ornithologists' Union". bou.org.uk. Archived from the original on 31 March 2017. Retrieved 27 July 2016. 
  27. ^ Amsterdam, Universiteit van. "Swammerdam medaille – GNGH – Universiteit van Amsterdam". gngh.uva.nl. Retrieved 27 July 2016. 
  28. ^ Anderson, Ted (2013). The Life of David Lack: Father of Evolutionary Ecology. Oxford: Oxford University Press. p. 185. ISBN 0199922659 – via Google Books. 
  29. ^ Diagram on The Four Areas of Biology
  30. ^ Further Diagrams on The Four Areas of Biology (Documents No. 5, 6 and 7 in English)
  31. ^ Lorenz, K. (1937). "Biologische Fragestellung in der Tierpsychologie". Zeitschrift für Tierpsychologie. 1: 24–32. doi:10.1111/j.1439-0310.1937.tb01401.x.  (in English: Biological Questions in Animal Psychology).
  32. ^ Tinbergen, N. (1963). "On aims and methods of Ethology". Zeitschrift für Tierpsychologie. 20 (4): 410–433. doi:10.1111/j.1439-0310.1963.tb01161.x. 
  33. ^ Tinbergen N, Tinbergen EA (1986). Autistic Children: New Hope for a Cure (new ed.). Routledge. ISBN 0-04-157011-1. 
  34. ^ Bishop, D. V. M. (2008). "Forty years on: Uta Frith's contribution to research on autism and dyslexia, 1966–2006". The Quarterly Journal of Experimental Psychology. 61 (1): 16–26. doi:10.1080/17470210701508665. PMC 2409181Freely accessible. PMID 18038335. 
  35. ^ Betty Fry Williams; Randy Lee Williams (2011). Effective programs for treating autism spectrum disorder: applied behavior analysis models. Taylor & Francis. pp. 53–. ISBN 978-0-415-99931-1. Retrieved 4 February 2012. 
  36. ^ Deirdre Barrett (2010). Supernormal Stimuli: How Primal Urges Overran Their Evolutionary Purpose. W. W. Norton & Company. pp. 21–22. ISBN 978-0-393-06848-1. Tinbergen had never been a religious man. Wartime atrocities, however, had highlighted the absence of a deity for him while both sides invoked one aligned with themselves, and this turned him into a militant atheist. 
  37. ^ Van Der Horst, F. C. P. (2010). "John Bowlby's treatment of Nikolaas "Niko" Tinbergen's depressions". History of Psychology. 13 (2): 206–208. doi:10.1037/a0019381c. 

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