Fixed action pattern: Difference between revisions

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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 |doi=10.1038/scientificamerican1252-22 |jstor=24944080 |bibcode=1952SciAm.187f..22T }}</ref><ref name=":92">Lorenz,{{cite K.,book Tinbergen, N|doi=10. (1970)4159/harvard.9780674430389.c6 |chapter=Taxis and instinctinstinctive behaviour pattern in egg-rolling response ofby the greylagGreylag goose. ''(1938) |title=Studies in Animal and Human Behavior'',Behaviour. 1,Volume 328–342.I |date=1970 |isbn=978-0-674-43038-9 |first1=Konrad |last1=Lorenz |translator-last1=Martin |translator-first1=Robert |pages=316–350 }}</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=1–5 |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 ==

There are 6six characteristics of fixed action patterns.<ref name=":22"/> Fixed action patterns are said to be stereotyped, complex, species-characteristic, released, triggered, and independent of experience.<ref name=":22"/>

*'''Stereotyped''': Fixed action patterns occur in rigid, predictable, and highly-structured sequences.<ref name=":22"/>

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==Sign stimulus==

The '''sign stimulus''', also known as a'''key stimulus''' or '''[[:simple:Releaser|releaser]]''', is the determining feature of a stimulus that produces a response. Sign stimuli are often found when observing a fixed action pattern (FAP) that is an innate behaviour with very little variance in the manner in which the actions are executed. Several examples of sign stimuli can be seen through the observation of animal behaviour in their natural environment. Experimenters have gone into these natural environments to better assess the stimuli and determine the key features of them that elicit a fixed action pattern. Scientists have also observed direct exploitation of sign stimuli in nature among different species of birds.

Fixed action patterns are released due to certain external stimuli.<ref name=":03"/> These stimuli are single or a small group of attributes of an object, not the object as a whole.<ref name=":3">{{Cite web |url=http://www.flyfishingdevon.co.uk/salmon/year1/psy128ethology_experiments/ethexpt.htm#FAP |title=Ethological studies of sign stimuli and motivation |website=www.flyfishingdevon.co.uk |access-date=2019-10-22}}</ref> These attributes may include color, shape, odor, and sound.<ref name=":03"/><ref name=":15"/>

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Exaggerated models of these attributes are called [[Supernormal stimulus|supernormal stimuli]].<ref name=":4">{{Cite book |title=The herring gull's world; a study of the social behaviour of birds |last=Tinbergen, Niko |date=1953 |publisher=Collins |isbn=0002194449 |___location=London |oclc=1029697}}</ref> A supernormal stimulus leads to an exaggerated response.<ref name=":4" /> Supernormal stimuli are more effective at releasing a response than a natural stimulus.<ref name=":4" />

An external stimulus that elicits a fixed action pattern is termed a sign stimulus if the stimuli emanates from the environment.,<ref name=":3" /> Whereas,whereas a releaser emanates from one's own species.<ref name=":3" />

===Supernormal stimuli===

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=== Male stickleback mating behavior ===

[[File:3-spined_stickleback.jpg|alt=|thumb|A 3three-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 ===

[[File:Grey_lag.jpg|thumb|A greylag goose which participates in the described egg-retrieval behavior.]]

Another example of a behavior that has been described as a fixed action pattern is the egg-retrieval behavior of the [[greylag goose]], reported in classic studies by Niko Tinbergen and Konrad Lorenz.<ref name=":92"/> Like many ground-nesting birds, if an [[Egg (biology)|egg]] becomes displaced from the nest, the greylag rolls it back to the nest with its beak.<ref name=":92"/><ref name=":0" /><ref>{{Cite web|url=http://www.pigeon.psy.tufts.edu/psych26/fap.htm|title=Fixed Action Pattern - Greylag Goose|website=pigeon.psy.tufts.edu|access-date=December 11, 2018}}</ref> The sight of the displaced egg is the sign stimulus and elicits the egg-retrieval behavior.<ref name=":92"/> First, the goose fixates its sight on the egg.<ref name=":92"/> Next, it extends its neck over the egg.<ref name=":92"/> Finally, it rolls the egg back to the nest using the underside of its beak.<ref name=":92"/> If the egg is removed from the goose during the performance of egg-rolling, the bird will continue with the behavior, pulling its head back as if an imaginary egg is still being maneuvered.<ref name=":92"/> It has been shown that the greylag will also attempt to retrieve other egg-shaped objects, such as a golf ball, door knob, or even a model egg too large to have possibly been laid by the goose itself (i.e. a supernormal stimulus).<ref>Tinbergen, N. (1951). ''The Study of Instinct''. Oxford University Press, New York.</ref>

The sight of the egg outside of the nest serves as the stimulus in this particular instance because it is only after the recognition of the egg's displacement that the fixed action pattern occurs.<ref name=":0" /><ref name=":1">{{cite web |last1=OpenStax College Biology |title=Behavioral Biology: Proximate and Ultimate Causes of Behavior |url=https://cnx.org/contents/GFy_h8cu@10.53:rZudN6XP@2/Introduction |website=cnx.org |date=21 October 2016 |access-date=20 November 2018}}</ref>

The manipulation of the sign stimulus through a series of experiments can allow scientists to understand what specific component of the stimulus is responsible for the innate behavioral sequence. If the egg were to be picked up and taken away after it is displaced from the nest, the goose still exhibits the same head moving motion even though there is no egg present.<ref name=":0" /> This was put to the test by using objects such as beer cans, and baseballs. Experimenters found that the stimulus merely had to be an object that was large enough in size, convex enough in shape, and comfortable enough for the goose to lay its neck around the edges of the object.<ref name=":0" />

These features that the stimulus has to obtain in order to trigger a resulting FAP were then given the official term of Sign Stimuli. Scientists came to the realization that there must be an innate deciphering method that the goose goes through in order to determine a suitable sign stimulus. This was defined as an [[innate releasing mechanism]] (IRM). The goose's IRM when put to the test in the natural world not being manipulated by scientific experimentation is almost always efficient in getting the desired item of an egg back into the nest.<ref name=":0" />

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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 fact|urldate=https://www.sciencedirect.com/topics/medicine-and-dentistry/fixed-action-patternNovember |title=Fixed Action Pattern – an overview {{!}} ScienceDirect Topics |website=www.sciencedirect.com |access-date=2019-10-242024}}</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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== Exceptions ==

There are 4four exceptions to fixed action pattern rules.<ref name=":142"/> These include reduced response threshold, vacuum activity, displacement behavior, and graded response.<ref name=":142"/>

=== Reduced response threshold ===

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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 ===