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{{Short description|Instinctive animal behavior}}
"'''Fixed action pattern'''" is an [[Ethology|ethological]] term describing an [[instinct]]ive behavioral sequence that is highly stereotyped and species-characteristic.<ref name=":03">{{Cite journal |last1=Páez-Rondón |first1=Oscar |last2=Aldana |first2=Elis |last3=Dickens |first3=Joseph |last4=Otálora-Luna |first4=Fernando |date=May 2018 |title=Ethological description of a fixed action pattern in a kissing bug (Triatominae): vision, gustation, proboscis extension and drinking of water and guava |journal=Journal of Ethology |volume=36 |issue=2 |pages=107–116 |doi=10.1007/s10164-018-0547-y |issn=0289-0771|doi-access=free }}</ref> Fixed action patterns are said to be produced by the innate releasing mechanism, a "hard-wired" [[neural network]], in response to a [[Fixed action pattern#Sign stimulus|sign/key stimulus]] or '''releaser'''.<ref name=":03"/><ref name=":15">{{Cite journal |last=Ronacher |first=Bernhard |date=February 2019 |title=Innate releasing mechanisms and fixed action patterns: basic ethological concepts as drivers for neuroethological studies on acoustic communication in Orthoptera |journal=Journal of Comparative Physiology A |volume=205 |issue=1 |pages=33–50 |doi=10.1007/s00359-018-01311-3 |issn=0340-7594 |pmc=6394777 |pmid=30617601}}</ref> Once released, a fixed action pattern runs to completion.<ref name=":03"/>
 
This term is often associated with [[Konrad Lorenz]], who is the founder of the concept.<ref name=":03"/> Lorenz identified six characteristics of fixed action patterns.<ref name=":22">{{Cite book |title=Studying animal behavior : autobiographies of the founders |date=1989 |publisher=University of Chicago Press |others=Dewsbury, Donald A. |isbn=0226144100 |___location=Chicago |oclc=19670401}}</ref> These characteristics state that fixed action patterns are stereotyped, complex, species-characteristic, released, triggered, and independent of experience.<ref name=":22" />
 
Fixed action patterns have been observed in many species, but most notably in fish and birds.<ref name=":03"/><ref name=":15" /> Classic studies by Konrad Lorenz and [[Nikolaas Tinbergen|Niko Tinbergen]] involve male stickleback mating behavior and greylag goose egg-retrieval behavior.<ref name=":82">{{cite journal |last1=Tinbergen, |first1=N. (1952). |title=The curiousCurious behaviorBehavior of sticklebacks.the ''Stickleback |journal=Scientific American'', |date=1952 |volume=187 |issue=6, 22–26.|pages=22–27 |jstor=24944080 }}</ref><ref name=":92">Lorenz,{{cite K.,journal Tinbergen, N|doi=10. (1970)4159/harvard. Taxis and instinct in egg-rolling response of the greylag goose9780674430389.c6 ''Studies in Animal and Human Behavior'', 1, 328–342.</ref>
 
Fixed action patterns have been shown to be evolutionarily advantageous, as they increase both fitness and speed.<ref name=":112">{{Cite web |url=https://www.jove.com/science-education/10919/fixed-action-patterns |title=Fixed Action Patterns {{!}} Protocol |website=www.jove.com |access-date=2019-10-24}}</ref> However, as a result of their predictability, they may also be used as a means of exploitation. An example of this exploitation would be brood parasitism.<ref name=":132">{{Citecite journal |last1=Peer |first1=Brian D. |last2=Robinson |first2=Scott K. |last3=Herkert |first3=James R. |date=2000-10-01 |title=Egg Rejection by Cowbird Hosts in Grasslands |journal=The Auk |date=October 2000 |volume=117 |issue=4 |pages=892–901 |doi=10.1093/auk/117.4.892 |issn=1938-4254|doi-access=free }}</ref>
 
There are four exceptions to fixed action pattern rules:<ref name=":142">{{Citecite journal |lastlast1=Schleidt |firstfirst1=Wolfgang M. |title=How "'Fixed"' is the Fixed Action Pattern? |url=https://www.academia.edu/32893025 |journal=Zeitschrift für Tierpsychologie |date=12 January 1974 |volume=36 |issue=1–51-5 |pages=184–211 |issn=0044-3573 |doi=10.1111/j.1439-0310.1974.tb02131.x |pmid=4467663 |year=2010}}</ref> reduced response threshold, [[vacuum activity]], displacement behavior, and graded response.<ref name=":142"/>
 
== Characteristics ==
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=== Male stickleback mating behavior ===
[[File:3-spined_stickleback.jpg|alt=|thumb|A three-spined stickleback like those used in Tinbergen's experiments]]
One example of fixed action patterns is the courtship and aggression behaviours of the male [[stickleback]], particularly the [[three-spined stickleback]], during mating season, described in a series of studies by Niko Tinbergen.<ref name=":82"/><ref name=":2">{{cite journal |last1=Rowland |first1=William |title=Habituation and development of response specificity to a sign stimulus: male preference for female courtship posture in stickleback |journal=Animal Behaviour |date=February 3, 2000 |volume=60 |issue=1 |pages=63–68 |doi=10.1006/anbe.2000.1462 |pmid=10924204 |s2cid=23095310 }}</ref> During the spring mating season, male sticklebacks' ventrum turns red and they establish a territory to build a nest.<ref name=":82"/> They attack other male sticklebacks that enter their territory, but court females and entice them to enter the nest to lay their eggs.<ref name=":82"/> Tinbergen used models of sticklebacks to investigate which features of male and female sticklebacks elicited attack and courtship behavior from male sticklebacks.<ref name=":82"/> Tinbergen's main findings were that male sticklebacks responded in a relatively invariant way and attacked even the most crude of models with a red belly, but in contrast, courted a model with a swollen belly.<ref name=":82"/> He also found that when presented with both a real male stickleback and a crude model, if the model's stomach was more red, the stickleback would attack the model as opposed to the other real male stickleback.<ref name=":82"/>
 
=== Greylag goose egg-retrieval behavior ===
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== Evolutionary disadvantages ==
[[File:Reed_warbler_cuckoo.jpg|thumb|Brood parasites, such as the [[European cuckoo|cuckoo]], provide a supernormal stimulus to the parenting species, in this case a [[common reed warbler]].]]
Fixed action patterns are predictable, as they are invariable, and therefore can lead to exploitation.<ref name=":122">{{Cite web |url=https://www.sciencedirect.com/topics/medicine-and-dentistry/fixed-action-pattern |title=Fixed Action Pattern – an overview {{!}} ScienceDirect Topics |website=www.sciencedirect.com |access-date=2019-10-24fact}}</ref> Some species have evolved to exploit the fixed action patterns of other species by [[mimicry]] of their sign stimuli.<ref name=":132"/> Replicating the releaser required to trigger a fixed action pattern is known as ''code-breaking''.<ref>{{Cite web|url=https://www.nature.com/scitable/knowledge/library/the-ecology-of-avian-brood-parasitism-14724491/|title=The Ecology of Avian Brood Parasitism {{!}} Learn Science at Scitable|website=www.nature.com|access-date=2019-10-24}}</ref> A well-known example of this is [[brood parasitism]], where one species will lay its eggs in the nest of another species, which will then parent its young.<ref name=":132"/> A young [[North American cowbird]], for example, provides a supernormal stimulus to its foster parent, which will cause it to forage rapidly to satisfy the larger bird's demands.<ref name=":5">Wickler, W. (1968) ''Mimicry in Plants and Animals.'' World University Library, London.</ref> A nestling will provide higher levels of stimulus with noisier, more energetic behavior, communicating its urgent need for food.<ref name=":5" /> Parents in this situation have to work harder to provide food, otherwise their own offspring are likely to die of [[starvation]].<ref name=":5" />
 
=== Brood parasitism ===
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=== Vacuum activity ===
After so long without being released, fixed action patterns are released in the absence of the sign stimulus or releaser.<ref>{{Cite web|url=https://www.psywww.com/intropsych/ch08-animals/konrad-lorenz.html#vacuum|title=The contributions of Konrad Lorenz {{!}} in Chapter 08: Animal Behavior and Cognition|website=www.psywww.com|access-date=2019-10-22}}</ref> Vacuum activity is demonstrated in courtship behavior of ring doves.<ref name=":6">{{Cite journal|last=Craig|first=Wallace|date=February 1918|journal=The Biological Bulletin|volume=34|issue=2|pages=91–107|doi=10.2307/1536346|pmid=16586767|pmc=1091358|issn=0006-3185 |jstor=1536346|title=Appetites and Aversions as Constituents of Instincts}}</ref><ref name=":7">{{Cite book|title=The Foundations of Ethology|last=Lorenz, Konrad Z.|date=1981|publisher=Springer Vienna|isbn=9783709136713|___location=Vienna|oclc=851389969}}</ref> Male blond ring doves isolated from their own species will resort to courting a pigeon, then a human's hand, and finally expressing courtship activity alone in their box, if they are left alone for a long period of time.<ref name=":6" /><ref name=":7" />
 
=== Displacement behavior ===
Fixed action patterns may be performed that are irrelevant to the stimulus present.<ref name=":10">{{cite journal |last1=Tinbergen, |first1=N., Van |last2=Iersel, |first2=J. J. A. (1947).van |title='Displacement reactionsReactions' in the threeThree-spinedSpined stickleback.Stickleback ''|journal=Behaviour'' '',|date=1947 |volume=1'', 56-63.|issue=1 {{JSTOR|pages=56–63 |jstor=4532675 }}.</ref> This can be seen in sticklebacks.<ref name=":10" /> Displacement behavior occurs when the fighting and mating urges are frustrated.<ref name=":10" /> A male stickleback chasing another male stickleback stops where their two territories meet and dives to the bottom of the tank, picking something up.<ref name=":10" /> This behavior resembles that of nest digging during mating season, but is not released by the proper sign stimulus.<ref name=":10" />
 
=== Graded response ===