Editing Domestication

{{short description|Selective breeding of plants and animals to serve humans}}
{{good article}}
{{Use mdy dates|date=September 2020}}
[[File:Murgjo Sharr Mountain Dog Nedi Limani.jpg|thumb|upright=1.5|[[Dog]]s and [[sheep]] were among the first animals to be domesticated, at least 15,000 and 11,000 years ago respectively.<ref name="MacHugh Larson Orlando 2017" />]]
[[File:1962-05 1962年 海南岛乐东县 黎族民众插秧.jpg|thumb|upright=1.5|[[Rice]] was domesticated in China, some 9,000 years ago.<ref name="Fornasiero Wing Ronald 2022" />]]

'''Domestication''' is a multi-generational [[Mutualism (biology)|mutualistic]] relationship in which an animal species, such as humans or [[leafcutter ant]]s, takes over control and care of another species, such as sheep or fungi, to obtain from them a steady supply of resources, such as meat, milk, or labor. The process is gradual and geographically diffuse, based on trial and error. Domestication affected genes for behavior in animals, making them less aggressive. In plants, domestication affected genes for morphology, such as increasing seed size and stopping the [[Shattering (agriculture)|shattering]] of cereal seedheads. Such changes both make domesticated organisms easier to handle and reduce their ability to survive in the wild.

The first [[Domestication of animals|animal to be domesticated]] by humans was the [[domestication of the dog|dog]], as a [[Commensalism|commensal]], at least 15,000 years ago. Other animals, including [[goat]]s, [[sheep]], and [[cow]]s, were domesticated around 11,000 years ago. Among birds, the [[chicken]] was first domesticated in East Asia, seemingly for cockfighting, some 7,000 years ago. The horse came under domestication around 5,500 years ago in central Asia as a working animal. Among [[invertebrate]]s, the [[silkworm]] and the [[western honey bee]] were domesticated over 5,000 years ago for [[silk]] and [[honey]], respectively.

The domestication of plants began around 13,000–11,000 years ago with [[cereal]]s such as [[wheat]] and [[barley]] in the [[Middle East]], alongside crops such as [[lentil]], [[pea]], [[chickpea]], and [[flax]]. Beginning around 10,000 years ago, Indigenous peoples in the Americas began to cultivate [[peanut]]s, [[Squash (plant)|squash]], [[maize]], [[potato]]es, [[cotton]], and [[cassava]]. [[Rice]] was first domesticated in China some 9,000 years ago. In Africa, crops such as [[sorghum]] were domesticated. [[History of agriculture|Agriculture developed]] in some 13 centres around the world, domesticating different crops and animals.

Three groups of insects, namely [[ambrosia beetle]]s, [[leafcutter ant]]s, and [[fungus-growing termites]] have independently domesticated species of fungi, on which they feed. In the case of the termites, the relationship is a fully obligate symbiosis on both sides.

== Definitions ==

Domestication (not to be confused with the [[tame animal|taming]] of an individual animal<ref name="price2008">{{cite book |last1=Price |first1=Edward O. |date=2008 |title=Principles and applications of domestic animal behavior: an introductory text|publisher=Cambridge University Press |url=https://books.google.com/books?id=Ww07sIWTYAAC&q=%22taming+is%22+animal&pg=PA228 |access-date=January 21, 2016 |isbn=9781780640556}}</ref><ref name="macdonald2009">{{cite journal |doi=10.1073/pnas.0901586106 |title=From wild animals to domestic pets, an evolutionary view of domestication |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=106 |pages=9971–9978 |date=2009 |last1=Driscoll |first1=C. A. |last2=MacDonald |first2=D. W. |last3=O'Brien |first3=S. J. |issue=Supplement 1 |pmid=19528637 |pmc=2702791 |bibcode=2009PNAS..106.9971D |doi-access=free}}</ref><ref name="diamond2012">{{cite book |first=Jared |last=Diamond |author-link=Jared Diamond |editor-last=Gepts |editor-first=P. |date=2012 |title=Biodiversity in Agriculture: Domestication, Evolution, and Sustainability |publisher=[[Cambridge University Press]] |chapter=Chapter 1 |page=13}}</ref>), is from the [[Latin language|Latin]] ''{{lang|la|domesticus}}'', 'belonging to the house'.<ref>{{cite web |title=Domesticate |work=Oxford Dictionaries |publisher=[[Oxford University Press]] |date=2014 |url=http://www.oxforddictionaries.com/definition/english/domesticate |archive-url=https://web.archive.org/web/20120720025116/http://oxforddictionaries.com/definition/english/domesticate |url-status=dead |archive-date=July 20, 2012}}</ref> The term remained loosely defined until the 21st century, when the American archaeologist [[Melinda A. Zeder]] defined it as a long-term relationship in which humans take over control and care of another organism to gain a predictable supply of a resource, resulting in [[Mutualism (biology)|mutual benefits]]. She noted further that it is not synonymous with agriculture since agriculture depends on domesticated organisms but does not automatically result from domestication.<ref name="zeder2015">{{cite journal |last=Zeder |first=Melinda A. |author-link=Melinda A. Zeder |date=2015 |title=Core questions in domestication Research|journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=112 |issue=11 |pages=3191–3198 |doi=10.1073/pnas.1501711112 |pmid=25713127 |pmc=4371924 |bibcode=2015PNAS..112.3191Z |doi-access=free}}</ref>

[[File:Domestication.svg|thumb|upright=2|Diagram of the process of domestication as a process where one species actively manages another to obtain resources or services, as defined by [[Michael D. Purugganan]]<ref name="Purugganan 2022"/>]]

[[Michael D. Purugganan]] notes that domestication has been hard to define, despite the "instinctual consensus" that it means "the plants and animals found under the care of humans that provide us with benefits and which have evolved under our control."<ref name="Purugganan 2022"/> He comments that insects such as [[termite]]s, [[ambrosia beetle]]s, and [[leafcutter ant]]s have domesticated some species of [[Fungus|fungi]], and notes further that other groups such as weeds and commensals have wrongly been called domesticated.<ref name="Purugganan 2022"/> Starting from Zeder's definition, Purugganan proposes a "broad" definition: "a coevolutionary process that arises from a mutualism, in which one species (the domesticator) constructs an environment where it actively manages both the survival and reproduction of another species (the domesticate) in order to provide the former with resources and/or services."<ref name="Purugganan 2022"/> He comments that this adds [[niche construction]] to the activities of the domesticator.<ref name="Purugganan 2022">{{cite journal |last=Purugganan |first=Michael D. |author-link=Michael Purugganan |date=2022 |title=What is domestication? |journal=[[Trends in Ecology & Evolution]] |volume=37 |issue=8 |pages=663–671 |doi=10.1016/j.tree.2022.04.006 |pmid=35534288 |doi-access=free|bibcode=2022TEcoE..37..663P }}</ref>

[[Domestication syndrome]] is the suite of [[phenotype|phenotypic]] traits that arose during the initial domestication process and which distinguish crops from their [[wild ancestor]]s.<ref name="Olsen Wendel 2013" /><ref name="hammer1984">{{cite journal |last1=Hammer |first1=K. |date=1984 |title=Das Domestikationssyndrom |language=German |journal=Kulturpflanze |volume=32 |pages=11–34 |doi=10.1007/bf02098682 |s2cid=42389667}}</ref> It can also mean a set of differences now observed in domesticated mammals, not necessarily reflecting the initial domestication process. The changes include increased docility and tameness, coat coloration, reductions in tooth size, craniofacial morphology, ear and tail form (e.g., floppy ears), estrus cycles, levels of [[adrenocorticotropic hormone]] and neurotransmitters, prolongations in juvenile behavior, and reductions in brain size and of particular brain regions.<ref name="Wilkins Wrangham Fitch 2014">{{cite journal |last1=Wilkins |first1=Adam S. |last2=Wrangham |first2=Richard W. |last3=Fitch |first3=W. Tecumseh |date=July 2014 |title=The 'Domestication Syndrome' in Mammals: A Unified Explanation Based on Neural Crest Cell Behavior and Genetics |doi=10.1534/genetics.114.165423 |journal=[[Genetics (journal)|Genetics]] |volume=197 |issue=3 |pages=795–808 |pmid=25024034 |pmc=4096361 |url=https://dash.harvard.edu/bitstream/handle/1/12717449/4096361.pdf?sequence=1}}</ref>

{{anchor|History of domestication}}<!--basically, the whole of this article, see below for chapters on animals, plants, fungi, etc, it isn't a matter of a "history" subsection-->

== Cause and timing ==

{{further|Neolithic transition}}

The [[domestication of animals]] and plants was triggered by the climatic and environmental changes that occurred after the peak of the [[Last Glacial Maximum]] <!--,which was around 21,000 years ago,--> and which continue to this present day. These changes made obtaining food by [[Hunter-gatherer|hunting and gathering]] difficult.<ref name="Zalloua2017">{{cite journal |last1=Zalloua |first1=Pierre A. |last2=Matisoo-Smith |first2=Elizabeth |title=Mapping Post-Glacial expansions: The Peopling of Southwest Asia |journal=[[Scientific Reports]] |date=January 6, 2017 |volume=7 |page=40338 |doi=10.1038/srep40338| pmid=28059138 |bibcode=2017NatSR...740338P |pmc=5216412}}</ref> The first animal to be [[Domestication of the dog|domesticated was the dog]] at least 15,000 years ago.<ref name="MacHugh Larson Orlando 2017">{{cite journal |last1=MacHugh |first1=David E. |last2=Larson |first2=Greger |last3=Orlando |first3=Ludovic |title=Taming the Past: Ancient DNA and the Study of Animal Domestication |doi=10.1146/annurev-animal-022516-022747 |journal=[[Annual Review of Animal Biosciences]] |volume=5 |date=2017 |s2cid=21991146 |pmid=27813680 |pages=329–351}}</ref> The [[Younger Dryas]] 12,900 years ago was a period of intense cold and aridity that put pressure on humans to intensify their foraging strategies but did not favour agriculture. By the beginning of the [[Holocene]] 11,700 years ago, a warmer climate and increasing human populations led to small-scale animal and plant domestication and an increased supply of food.<ref name="McHugo Dover MacHugh 2019">{{Cite journal |last1=McHugo |first1=Gillian P. |last2=Dover |first2=Michael J. |last3=MacHugh |first3=David E. |date=2019-12-02 |title=Unlocking the origins and biology of domestic animals using ancient DNA and paleogenomics |journal=[[BMC Biology]] |volume=17 |issue=1 |pages=98 |doi=10.1186/s12915-019-0724-7 |pmc=6889691 |pmid=31791340 |doi-access=free }}</ref>

{| class="wikitable" style="margin:1em auto;"
|+ Timeline of some major domestication events
|-
! Event !! [[Vavilov center|Centre of origin]] !! Purpose !! Date/years ago
|-
| [[Foraging]] for wild grains || Asia || Food || > 23,000<ref name="Purugganan Fuller 2009"/>
|-
| [[Dog]] || Eurasia || [[Commensalism|Commensal]] || > 15,000<ref name="MacHugh Larson Orlando 2017"/>
|-
| [[Wheat]], [[Barley]] || Near East || Food || 13,000–11,000<ref name="Purugganan Fuller 2009"/>
|-
|| [[Flax]] || Near East || [[Textile]]s || 13,000–11,000{{sfn|Zohary|Hopf|Weiss|2012|p=139}}
|-
| [[Goat]], [[Sheep]], [[Pig]], [[Cow]] || Near East, South Asia || Food || 11,000–10,000<ref name="MacHugh Larson Orlando 2017"/>
|-
| [[Rice]] || China || Food || 9,000<ref name="Fornasiero Wing Ronald 2022">{{Cite journal |last1=Fornasiero |first1=Alice |last2=Wing |first2=Rod A. |last3=Ronald |first3=Pamela |date=2022 |title=Rice domestication |url=https://doi.org/10.1016/j.cub.2021.11.025 |journal=[[Current Biology]] |volume=32 |issue=1 |pages=R20–R24 |doi=10.1016/j.cub.2021.11.025 |pmid=35015986 |bibcode=2022CBio...32..R20F}}</ref>
|-
| [[Chicken]] || East Asia || [[Cockfighting]] || 7,000<ref name="Lawler Adler 2012"/>
|- 
| [[Horse]] || Central Asia || [[Pack animal|Draft]], [[Equestrianism|riding]] || 5,500<ref name="MacHugh Larson Orlando 2017"/>
|-
| [[Western honey bee|Honey bee]] || Ancient Egypt || [[Honey]] || > 5,000<ref name="Aizen Harder 2009"/>
|}

The appearance of the [[domestic dog]] in the archaeological record, at least 15,000 years ago, was followed by domestication of livestock and [[crop domestication|of crops]] such as [[wheat]] and [[barley]], the [[invention of agriculture]], and the transition of humans from foraging to farming in different places and times across the planet.<ref name="MacHugh Larson Orlando 2017"/><ref name="fuller2011">{{cite journal |doi=10.1080/00438243.2011.624747 |title=Cultivation and domestication had multiple origins: Arguments against the core area hypothesis for the origins of agriculture in the Near East |journal=[[World Archaeology]] |volume=43 |issue=4 |pages=628–652 |date=2011 |last1=Fuller |first1=Dorian Q. |last2=Willcox |first2=George |last3=Allaby |first3=Robin G. |s2cid=56437102}}</ref><ref name="zeder2006">{{cite book |last=Zeder |first=Melinda A. |author-link=Melinda A. Zeder |date=2006 |contribution=Archaeological approaches to documenting animal domestication |title=Documenting Domestication: New Genetic and Archaeological Paradigms |editor1-first=M. A. |editor1-last=Zeder |editor2-first=D. G. |editor2-last=Bradley |editor3-first=E. |editor3-last=Emshwiller |editor4-first=B. D. |editor4-last=Smith |pages=209–227 |location=Berkeley |publisher=[[University of California Press]]}}</ref><ref name="Galibert Ouignon Hitte 2011">{{Cite journal |last1=Galibert |first1=Francis |last2=Quignon |first2=Pascale |last3=Hitte |first3=Christophe |last4=André |first4=Catherine |date=2011-03-01 |title=Toward understanding dog evolutionary and domestication history |journal=[[Comptes Rendus Biologies]] |series=On the trail of domestications, migrations and invasions in agriculture |volume=334 |issue=3 |pages=190–196 |doi=10.1016/j.crvi.2010.12.011 |pmid=21377613|doi-access=free }}</ref> For instance, small-scale trial cultivation of cereals began some 28,000 years ago at the Ohalo II site in Israel.<ref name="Snir Nadel Groman-Yaroslavski Melamed 2015">{{cite journal |last1=Snir |first1=Ainit |last2=Nadel |first2=Dani |last3=Groman-Yaroslavski |first3=Iris |last4=Melamed |first4=Yoel |last5=Sternberg |first5=Marcelo |last6=Bar-Yosef |first6=Ofer |last7=Weiss |first7=Ehud |title=The Origin of Cultivation and Proto-Weeds, Long Before Neolithic Farming |journal=[[PLOS One]] |volume=10 |issue=7 |date=2015-07-22 |pmid=26200895 |pmc=4511808 |doi=10.1371/journal.pone.0131422 |doi-access=free |page=e0131422|bibcode=2015PLoSO..1031422S }}</ref>

In the [[Fertile Crescent]] 11,000–10,000 years ago, [[zooarchaeology]] indicates that goats, pigs, sheep, and [[taurine cattle]] were the first livestock to be domesticated. Two thousand years later, humped [[zebu]] cattle were domesticated in what is today [[Baluchistan]] in Pakistan. In [[East Asia]] 8,000 years ago, pigs were domesticated from wild boar genetically different from those found in the Fertile Crescent.<ref name="MacHugh Larson Orlando 2017"/> The [[Domestication of the cat|cat was domesticated]] in the Fertile Crescent, perhaps 10,000 years ago,<ref name="Driscoll 2009">{{Cite journal |last=Driscoll |first=Carlos |date=2009 |title=The Taming of the Cat. Genetic and Archaeological findings hint that wildcats became housecats earlier- and in different place- than previously thought |journal=[[Scientific American]] |volume=300 |issue=6 |pages=68–75 |doi=10.1038/scientificamerican0609-68 |doi-broken-date=November 1, 2024 |pmid=19485091 |pmc=5790555 |bibcode=2009SciAm.300f..68D}}</ref> from [[African wildcat]]s, possibly to control [[rodent]]s that were damaging stored food.<ref name="Driscoll Menotti-Raymond Roca 2007">{{Cite journal |last1=Driscoll |first1=Carlos A. |last2=Menotti-Raymond |first2=Marilyn |last3=Roca |first3=Alfred L. |last4=Hupe |first4=Karsten |last5=Johnson |first5=Warren E. |last6=Geffen |first6=Eli |last7=Harley |first7=Eric H. |last8=Delibes |first8=Miguel |last9=Pontier |first9=Dominique |last10=Kitchener |first10=Andrew C. |last11=Yamaguchi |first11=Nobuyuki |last12=O'Brien |first12=Stephen J. |last13=Macdonald |first13=David W. |display-authors=5 |date=2007-07-27 |title=The Near Eastern Origin of Cat Domestication |journal=[[Science (journal)|Science]] |volume=317 |issue=5837 |pages=519–523 |doi=10.1126/science.1139518 |pmc=5612713 |pmid=17600185 |bibcode=2007Sci...317..519D }}</ref>


<gallery mode="packed" heights="275" style="float:center;">
File:Centres of origin and spread of agriculture labelled.svg|[[Vavilov center|Centres of origin]] and spread of [[agriculture]] in the [[Neolithic Revolution]] as understood in 2003<ref name="DiamondandBellwood2003">{{Cite journal | doi=10.1126/science.1078208 |last1=Diamond |first1=Jared |author-link1=Jared Diamond |last2=Bellwood |first2=P. |title=Farmers and Their Languages: The First Expansions |journal=[[Science (journal)|Science]] |volume=300 |issue=5619 |pages=597–603 |date=2003 |pmid=12714734 |bibcode=2003Sci...300..597D |citeseerx=10.1.1.1013.4523 |s2cid=13350469 }}</ref>
File:Domestication Timeline.jpg|Rough timelines of domestication for 11 animal species<ref>{{cite journal |last1=McHugo |first1=Gillian P. |last2=Dover |first2=Michael J. |last3=MacHugh |first3=David E. |title=Unlocking the origins and biology of domestic animals using ancient DNA and paleogenomics |journal=BMC Biology |date=2 December 2019 |volume=17 |issue=1 |pages=98 |doi=10.1186/s12915-019-0724-7 |doi-access=free |pmid=31791340 |pmc=6889691 |issn=1741-7007}}</ref>{{efn|It includes "relevant stratigraphy and climate chronologies. For each species, the time periods of significant pre-domestication human–animal interactions are also shown." "Stratigraphy information was obtained from the International Commission on Stratigraphy website. The Quaternary temperature plot was generated from the GISP2 ice core temperature and accumulation data"}}
</gallery>

== Animals ==

=== Desirable traits ===

{{further|Domestication of vertebrates}}

[[File:Unnatural selection, 2 heads, one species.jpg|thumb |Domesticated animals tend to be smaller and less aggressive than their wild counterparts; many have other [[domestication syndrome]] traits like shorter muzzles.<ref name="Frantz Bradley Larson 2020" /> Skulls of [[grey wolf]] (left), [[Chihuahua (dog)|chihuahua dog]] (right) ]]

The domestication of vertebrate animals is the relationship between non-human vertebrates and humans who have an influence on their care and reproduction.<ref name="zeder2015" /> In his 1868 book ''[[The Variation of Animals and Plants Under Domestication]]'', [[Charles Darwin]] recognized the small number of traits that made domestic species different from their wild ancestors. He was also the first to recognize the difference between conscious [[selective breeding]] in which humans directly select for desirable traits and unconscious selection, in which traits evolve as a by-product of [[natural selection]] or from selection on other traits.<ref name="darwin1868">{{cite book |last=Darwin |first=Charles |author-link=Charles Darwin |date=1868 |title=The Variation of Animals and Plants Under Domestication |location=London |publisher=John Murray |oclc=156100686|title-link=The Variation of Animals and Plants Under Domestication}}</ref>{{sfn|Diamond|2005|p=130}}<ref name="Larson Piperno Allaby 2014">{{cite journal |doi=10.1073/pnas.1323964111 |title=Current perspectives and the future of domestication studies |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=111 |issue=17 |pages=6139–6146 |date=2014 |last1=Larson |first1=G. |last2=Piperno |first2=D. R. |last3=Allaby |first3=R. G. |last4=Purugganan |first4=M. D. |last5=Andersson |first5=L. |last6=Arroyo-Kalin |first6=M. |last7=Barton |first7=L. |last8=Climer Vigueira |first8=C. |last9=Denham |first9=T. |last10=Dobney |first10=K. |last11=Doust |first11=A. N. |last12=Gepts |first12=P. |last13=Gilbert |first13=M. T. P. |last14=Gremillion |first14=K. J. |last15=Lucas |first15=L. |last16=Lukens |first16=L. |last17=Marshall |first17=F. B. |last18=Olsen |first18=K. M. |last19=Pires |first19=J. C. |last20=Richerson |first20=P. J. |last21=Rubio De Casas |first21=R. |last22=Sanjur |first22=O. I. |last23=Thomas |first23=M. G. |last24=Fuller |first24=D. Q. |display-authors=5 |pmid=24757054 |pmc=4035915 |bibcode=2014PNAS..111.6139L |doi-access=free}}</ref>

There is a difference between domestic and wild populations; some of these differences constitute the [[Domestication syndrome#In animals|domestication syndrome]], traits presumed essential in the early stages of domestication, while others represent later improvement traits.<ref name="Olsen Wendel 2013">{{cite journal |last1=Olsen |first1=K. M. |last2=Wendel |first2=J. F. |s2cid=727983 |date=2013 |title=A bountiful harvest: genomic insights into crop domestication phenotypes |journal=[[Annual Review of Plant Biology]] |volume=64 |issue=1 |pages=47–70 |doi=10.1146/annurev-arplant-050312-120048 |pmid=23451788|bibcode=2013AnRPB..64...47O }}</ref><ref name="Doust Lukens Olsen 2014">{{cite journal |last1=Doust |first1=A. N. |last2=Lukens |first2=L. |last3=Olsen |first3=K. M. |last4=Mauro-Herrera |first4=M. |last5=Meyer |first5=A. |last6=Rogers |first6=K. |title=Beyond the single gene: How epistasis and gene-by-environment effects influence crop domestication |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=111 |issue=17 |pages=6178–6183 |date=2014 |doi=10.1073/pnas.1308940110 |pmid=24753598 |pmc=4035984 |bibcode=2014PNAS..111.6178D |doi-access=free}}</ref><ref name="larson2014" /> Domesticated mammals in particular tend to be smaller and less aggressive than their wild counterparts; other common traits are floppy ears, a smaller brain, and a shorter muzzle.<ref name="Frantz Bradley Larson 2020">{{cite journal |last1=Frantz |first1=Laurent A. F. |last2=Bradley |first2=Daniel G. |last3=Larson |first3=Greger |last4=Orlando |first4=Ludovic |title=Animal domestication in the era of ancient genomics |date=2020 |journal=[[Nature Reviews Genetics]] |volume=21 |issue=8 |pages=449–460 |doi=10.1038/s41576-020-0225-0 |pmid=32265525 |s2cid=214809393 |url=https://qmro.qmul.ac.uk/xmlui/handle/123456789/66726}}</ref> Domestication traits are generally fixed within all domesticates, and were selected during the initial episode of domestication of that animal or plant, whereas improvement traits are present only in a proportion of domesticates, though they may be fixed in individual breeds or [[Landrace|regional populations]].<ref name="Doust Lukens Olsen 2014"/><ref name="larson2014"/><ref name="meyer2013">{{cite journal |doi=10.1038/nrg3605 |pmid=24240513 |title=Evolution of crop species: Genetics of domestication and diversification |journal=[[Nature Reviews Genetics]] |volume=14 |issue=12 |pages=840–852 |date=2013 |last1=Meyer |first1=Rachel S. |last2=Purugganan |first2=Michael D. |s2cid=529535}}</ref>

Certain animal species, and certain individuals within those species, make better candidates for domestication because of their behavioral characteristics:<ref name="zeder2012">{{cite journal |last=Zeder |first=Melinda A. |author-link=Melinda A. Zeder |date=2012 |title=The domestication of animals |journal=[[Journal of Anthropological Research]] |volume=68 |issue=2 |pages=161–190 |doi=10.3998/jar.0521004.0068.201 |s2cid=85348232}}</ref><!--Fig 1--><ref name="hale1969">{{cite book |last=Hale |first=E. B. |date=1969 |chapter=Domestication and the evolution of behavior |title=The Behavior of Domestic Animals |edition=2nd |editor-first=E. S. E. |editor-last=Hafez |pages=22–42 |location=London |publisher=Bailliere, Tindall, and Cassell}}</ref><ref name="price1984">{{cite journal |last1=Price |first1=Edward O. |date=1984 |title=Behavioral aspects of animal domestication |journal=[[Quarterly Review of Biology]] |volume=59 |issue=1 |pages=1–32 |jstor=2827868 |doi=10.1086/413673 |s2cid=83908518}}</ref><ref name="price2002">{{cite book |last=Price |first=Edward O. |date=2002 |title=Animal Domestication and Behavior |location=Wallingford, UK |publisher=[[CABI Publishing]] |url=http://www.uesc.br/cursos/pos_graduacao/mestrado/animal/bibliografia2012/selene_artigo1_animal.pdf |access-date=February 29, 2016 |archive-url=https://web.archive.org/web/20170517133013/http://www.uesc.br/cursos/pos_graduacao/mestrado/animal/bibliografia2012/selene_artigo1_animal.pdf |archive-date=May 17, 2017 |url-status=dead}}</ref>

# The size and organization of their social structure<ref name="zeder2012"/>
# The availability and the degree of selectivity in their choice of mates<ref name="zeder2012"/>
# The ease and speed with which the parents bond with their young, and the maturity and mobility of the young at birth<ref name="zeder2012"/>
# The degree of flexibility in diet and habitat tolerance<ref name="zeder2012"/>
# Responses to humans and new environments, including reduced flight response and reactivity to external stimuli.<ref name="zeder2012"/>

=== Mammals ===

{{further|List of domesticated animals}}

[[File:Tibet ~ Camel Caravan (3747098653) (cropped).jpg|thumb|While dogs were [[Commensalism|commensal]]s, and sheep were kept for food, [[camel]]s were domesticated as [[pack animal|working animals]].<ref name="zeder2012" />]]

The beginnings of mammal domestication involved a protracted [[coevolution]]ary process with multiple stages along different pathways. There are three proposed<!--Zeder 2012--> major pathways that most mammal domesticates followed into domestication:<ref name="zeder2012"/><ref name="larson2014"/><ref name="marshall2014"/>

# [[Commensalism|commensals]], adapted to a human niche (e.g., [[dogs]], [[cats]], possibly [[pigs]])<ref name="zeder2012"/>
# prey animals sought for food (e.g., [[sheep]], [[goats]], [[cattle]], [[water buffalo]], [[Domestic yak|yak]], pig, [[reindeer]], [[llama]] and [[alpaca]])<ref name="zeder2012"/>
# animals targeted [[pack animal|for draft]] and [[Equestrianism|riding]] (e.g., [[horse]], [[donkey]], [[camel]]).<ref name="zeder2012"/>

Humans did not intend to domesticate mammals from either the commensal or prey pathways, or at least they did not envision a domesticated animal would result from it. In both of those cases, humans became entangled with these species as the relationship between them intensified, and humans' role in their survival and reproduction gradually led to formalized [[animal husbandry]].<ref name="larson2014"/> Although the directed pathway for draft and riding animals proceeded from capture to taming, the other two pathways are not as goal-oriented, and archaeological records suggest that they took place over much longer time frames.<ref name="larson2013">{{cite journal |last=Larson |first=Greger |date=2013 |title=A population genetics view of animal domestication |journal=[[Trends in Genetics]] |volume=29 |issue=4 |pages=197–205 |doi=10.1016/j.tig.2013.01.003 |pmid=23415592 |url=http://www.palaeobarn.com/sites/domestication.org.uk/files/downloads/98.pdf |access-date=January 2, 2016 |archive-date=June 8, 2019 |archive-url=https://web.archive.org/web/20190608065300/http://www.palaeobarn.com/sites/domestication.org.uk/files/downloads/98.pdf |url-status=dead }}</ref>

Unlike other domestic species selected primarily for production-related traits, dogs were initially selected for their behaviors.<ref name="Serpell Duffy 2014">{{cite book |last1=Serpell |first1=J. |last2=Duffy |first2=D. |contribution=Dog Breeds and Their Behavior |title=Domestic Dog Cognition and Behavior |editor-first=Alexandra |editor-last=Horowitz |location=Berlin / Heidelberg |publisher=Springer |date=2014}}</ref><ref name="Cagan Blass 2016">{{cite journal |last1=Cagan |first1=Alex |last2=Blass |first2=Torsten |title=Identification of genomic variants putatively targeted by selection during dog domestication |journal=[[BMC Evolutionary Biology]] |volume=16 |pages=10 |date=2016 |issue=1 |doi=10.1186/s12862-015-0579-7 |doi-broken-date=December 2, 2024 |pmid=26754411 |pmc=4710014 |bibcode=2016BMCEE..16...10C |doi-access=free}}</ref> The dog was domesticated long before other animals,<ref name="larson2012"/><ref name="perri2016">{{cite journal |last1=Perri |first1=Angela |title=A wolf in dog's clothing: Initial dog domestication and Pleistocene wolf variation |journal=[[Journal of Archaeological Science]] |volume=68 |pages=1–4 |date=2016 |doi=10.1016/j.jas.2016.02.003 |bibcode=2016JArSc..68....1P}}</ref> becoming established across [[Eurasia]] before the end of the [[Late Pleistocene]] era, well before [[agriculture]].<ref name="larson2012">{{cite journal |last=Larson |first=Greger |date=2012 |title=Rethinking dog domestication by integrating genetics, archeology, and biogeography |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=109 |issue=23 |pages=8878–8883 |pmid=22615366 |pmc=3384140 |doi=10.1073/pnas.1203005109 |bibcode=2012PNAS..109.8878L |doi-access=free }}</ref>

The archaeological and genetic data suggest that long-term bidirectional [[gene flow]] between wild and domestic stocks – such as in [[donkey]]s, [[horse]]s, New and Old World camelids, goats, sheep, and pigs – was common.<ref name="larson2014">{{cite journal |last1=Larson |first1=Greger |last2=Fuller |first2=Dorian Q. |date=2014 |title=The Evolution of Animal Domestication |journal=[[Annual Review of Ecology, Evolution, and Systematics]] |volume=45 |pages=115–136 |url=http://www.palaeobarn.com/sites/domestication.org.uk/files/downloads/112.pdf |doi=10.1146/annurev-ecolsys-110512-135813 |s2cid=56381833 |access-date=January 19, 2016 |archive-date=May 13, 2019 |archive-url=https://web.archive.org/web/20190513194442/http://www.palaeobarn.com/sites/domestication.org.uk/files/downloads/112.pdf |url-status=dead }}</ref><ref name="marshall2014">{{cite journal |last=Marshall |first=F. |date=2013 |title=Evaluating the roles of directed breeding and gene flow in animal domestication |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=111 |issue=17 |pages=6153–6158 |doi=10.1073/pnas.1312984110 |pmid=24753599 |pmc=4035985 |bibcode=2014PNAS..111.6153M |doi-access=free}}</ref> Human selection for domestic traits likely counteracted the homogenizing effect of gene flow from wild boars into pigs, and created [[domestication islands]] in the genome. The same process may apply to other domesticated animals.
<ref name="frantz2015">{{cite journal |last=Frantz |first=L. |s2cid=205350534 |date=2015 |title=Evidence of long-term gene flow and selection during domestication from analyses of Eurasian wild and domestic pig genomes |journal=[[Nature Genetics]] |volume=47 |issue=10 |pages=1141–1148 |doi=10.1038/ng.3394 |pmid=26323058}}</ref><ref name="pennisi2015">{{cite journal |last=Pennisi |first=E. |author-link=Elizabeth Pennisi |date=2015 |title=The taming of the pig took some wild turns |journal=[[Science (journal)|Science]] |doi=10.1126/science.aad1692}}</ref>

The 2023 [[Parasite-stress theory|parasite-mediated domestication]] hypothesis suggests that [[endoparasites]] such as [[helminths]] and [[protozoa]] could have mediated the domestication of mammals. Domestication involves taming, which has an endocrine component; and parasites can modify endocrine activity and [[microRNA]]s. Genes for resistance to parasites might be linked to those for the domestication syndrome; it is predicted that domestic animals are less resistant to parasites than their wild relatives.<ref name="Skok 2023a">{{cite journal |last=Skok |first=J. |date=2023a |title=The Parasite-Mediated Domestication Hypothesis |journal=Agricultura Scientia |doi=10.18690/agricsci.20.1.1 |volume=20 |issue=1 |pages=1–7 |doi-access=free}}</ref><ref name="Skok 2023b">{{Cite journal |last=Skok |first=J. |date=2023b |title=Addendum to "The parasite-mediated domestication hypothesis" |journal=OSF |doi=10.31219/osf.io/f92aj |doi-access=free}}</ref>

{{anchor|Insects}}

=== Birds ===

{{main|Poultry|Aviculture}}

{{multiple image
|footer=The [[chicken]] was domesticated from the red junglefowl, apparently for [[cockfighting]], some 7,000 years ago.<ref name="Lawler Adler 2012"/>
|total_width=400px
|image1=COCK FIGHT.JPG
|caption1=Cockfight in [[Tamil Nadu]], 2011
|image2=Red Junglefowl (male) - Thailand.jpg
|caption2=[[Red junglefowl]] of Southeast Asia 
}}

Domesticated birds principally mean [[poultry]], raised for meat and eggs:<ref name="AH">{{cite encyclopedia |title=Poultry |encyclopedia=The American Heritage: Dictionary of the English Language |edition=4th |publisher=Houghton Mifflin Company |date=2009 |url=http://www.thefreedictionary.com/poultry}}</ref> some [[Galliformes]] ([[chicken]], [[turkey (bird)|turkey]], [[guineafowl]]) and [[Anseriformes]] (waterfowl: [[duck]]s, [[goose|geese]], and [[swan]]s). Also widely domesticated are [[cagebirds]] such as [[songbirds]] and [[parrots]]; these are kept both for pleasure and for use in research.<ref>{{cite web |url=http://asabirds.org/history/ |title=Our History |publisher=Avicultural Society of America |access-date=3 December 2023}}</ref>
The [[domestic pigeon]] has been used both for food and as a means of communication between far-flung places through the exploitation of the pigeon's homing instinct; research suggests it was domesticated as early as 10,000 years ago.<ref name="Blechman">{{cite book |last=Blechman |first=Andrew |title=Pigeons – The fascinating saga of the world's most revered and reviled bird |publisher=University of Queensland Press |date=2007 |url=http://andrewblechman.com/pigeons/learn_more.html |isbn=9780702236419}}</ref> Chicken fossils in China have been dated to 7,400 years ago. The chicken's wild ancestor is ''[[Gallus gallus]]'', the red junglefowl of Southeast Asia. The species appears to have been kept initially for [[cockfighting]] rather than for food.<ref name="Lawler Adler 2012">{{cite journal |last1=Lawler |first1=Andrew |last2=Adler |first2=Jerry |title=How the Chicken Conquered the World |url=http://www.smithsonianmag.com/history/how-the-chicken-conquered-the-world-87583657/ |journal=[[Smithsonian (magazine)|Smithsonian Magazine]] |issue=June 2012 |date=June 2012}}</ref>

===Invertebrates ===

{{Further|Domestication of bees|Beekeeping|Sericulture}}

Two [[insect]]s, the [[silkworm]] and the [[western honey bee]], have been domesticated for over 5,000 years, often for commercial use. The silkworm is raised for the silk threads wound around its [[pupa]]l cocoon; the western honey bee, for [[honey]], and, from the 20th century, for [[pollination]] of crops.<ref name="Aizen Harder 2009">{{cite journal |last1=Aizen |first1=Marcelo A. |last2=Harder |first2=Lawrence D. |title=The Global Stock of Domesticated Honey Bees Is Growing Slower Than Agricultural Demand for Pollination |journal=[[Current Biology]] |date=2009 |volume=19 |issue=11 |pages=915–918 |doi=10.1016/j.cub.2009.03.071|pmid=19427214 |s2cid=12353259 |doi-access=free |bibcode=2009CBio...19..915A }}</ref><ref>{{cite journal |last1=Potts |first1=Simon G.|display-authors=etal |title=Global pollinator declines: Trends, impacts and drivers |journal=[[Trends in Ecology & Evolution]] |date=2010 |volume=25 |issue=6 |pages=345–353 |doi=10.1016/j.tree.2010.01.007 |url=https://www.researchgate.net/publication/41621021 |pmid=20188434 |bibcode=2010TEcoE..25..345P |citeseerx=10.1.1.693.292 }}</ref>

Several other invertebrates have been domesticated, both terrestrial and aquatic, including some such as ''[[Drosophila melanogaster]]'' fruit flies and the freshwater cnidarian ''[[Hydra (animal)|Hydra]]'' for research into genetics and physiology. Few have a long history of domestication. Most are used for food or other products such as [[shellac]] and [[cochineal]]. The [[phylum|phyla]] involved are [[Cnidaria]], [[Platyhelminthes]] (for [[biological pest control]]), [[Annelida]], [[Mollusca]], [[Arthropoda]] (marine [[crustacea]]ns as well as insects and spiders), and [[Echinodermata]]. While many marine mollusks are used for food, only a few have been domesticated, including [[Loligo|squid]], [[Sepia (genus)|cuttlefish]] and [[octopus]], all used in research on [[animal behaviour|behaviour]] and [[neurology]]. Terrestrial snails in the genera ''[[Helix (genus)|Helix]]'' are raised for food. Several parasitic or parasitoidal insects, including the fly ''[[Eucelatoria]]'', the beetle ''[[Chrysolina]]'', and the wasp ''[[Aphytis (insect)|Aphytis]]'' are raised for biological control. Conscious or unconscious artificial selection has many effects on species under domestication; variability can readily be lost by inbreeding, selection against undesired traits, or genetic drift, while in ''Drosophila'', variability in eclosion time (when adults emerge) has increased.<ref name="GonPrice1984">{{cite journal |last1=Gon III |first1=Samuel M. |last2=Price |first2=Edward O. |title=Invertebrate Domestication: Behavioral Considerations |journal=[[BioScience]] |date=October 1984 |volume=34 |issue=9 |pages=575–579 |doi=10.2307/1309600 |jstor=1309600 |url=https://www.researchgate.net/profile/Edward-Price-6/publication/240297283_Invertebrate_Domestication_Behavioral_Considerations/links/5d608a0892851c619d720388/Invertebrate-Domestication-Behavioral-Considerations.pdf <!--NOT redundant with DOI-->}}</ref>

<gallery mode=packed heights=175px>
File:Cueva arana.svg|A [[Honey hunting|honey hunter]] in a [[cave painting]] at [[Cuevas de la Araña]], Spain, c. 8,000–6,000 BC
File:Sericuturist.jpg|[[Sericulture|Sericulturalists]] preparing [[silkworm]]s for spinning of the [[silk]] 
File:02-Indian-Insect-Life - Harold Maxwell-Lefroy - Kerria-Lacca.jpg|The lac bug ''[[Kerria lacca]]'' has been kept for [[shellac]] resin.
File:Snails for Sale - Djermaa el-Fna (Central Square) - Medina (Old City) - Marrakesh - Morocco.jpg|[[Snails as food|Snails being sold as food]]
</gallery>

{{anchor|Plants}}

== Plants ==

{{further|History of agriculture|List of domesticated plants}}

Humans [[Foraging|foraged]] for wild cereals, seeds, and nuts thousands of years before they were domesticated; wild wheat and barley, for example, were gathered in the [[Levant]] at least 23,000 years ago.<ref>Weiss, E., Kislev, M.E., Simchoni, O. & Nadel, D. Small-grained wild grasses as staple food at the 23000-year-old site of Ohalo II ''Economic Botany'' 58:s125-s134.</ref><ref name="Purugganan Fuller 2009" /> [[Neolithic]] societies in West Asia first began to cultivate and then domesticate some of these plants around 13,000 to 11,000 years ago.<ref name="Purugganan Fuller 2009" /> The [[founder crops]] of the West Asian Neolithic included cereals ([[emmer]], [[einkorn wheat]], [[barley]]), [[Pulse (legume)|pulses]] ([[lentil]], [[pea]], [[chickpea]], [[Vicia ervilia|bitter vetch]]), and [[flax]].{{sfn|Zohary|Hopf|Weiss|2012|p=139}}{{sfn|Banning|2002}} Other plants were independently domesticated in 13 [[centers of origin]] (subdivided into 24 areas) of the Americas, Africa, and Asia (the Middle East, South Asia, the Far East, and New Guinea and Wallacea); in some thirteen of these regions people began to cultivate grasses and grains.{{sfn|Zohary|Hopf|Weiss|2012|p=}}<ref>{{cite book |last=Harris |first=David R. |title=The Origin and Spread of Agriculture and Pastoralism in Eurasia |location=London |publisher=University College London Press |date=1996 |pages=142–158 |isbn=9781857285376}}</ref> Rice was first cultivated in East Asia.<ref name="Normile">{{cite journal |last=Normile |first=Dennis |date=1997 |title=Yangtze seen as earliest rice site |journal=Science |volume=275 |issue=5298 |pages=309–310 |doi=10.1126/science.275.5298.309 |s2cid=140691699}}</ref><ref>"New Archaeobotanic Data for the Study of the Origins of Agriculture in China", Zhijun Zhao, Current Anthropology Vol. 52, No. S4, (October 2011), pp. S295-S306</ref> [[Sorghum]] was widely cultivated in sub-Saharan Africa,<ref>{{Cite book |last=Carney |first=Judith |title=In the Shadow of Slavery |publisher=University of California Press |date=2009 |isbn=9780520269965 |location=Berkeley and Los Angeles, California |pages=16}}</ref> while peanuts,<ref name="Dillehay Rossen Andres Williams 2007">{{cite journal |last1=Dillehay |first1=Tom D. |last2=Rossen |first2=Jack |last3=Andres |first3=Thomas C. |last4=Williams |first4=David E. |title=Preceramic Adoption of Peanut, Squash, and Cotton in Northern Peru |journal=Science |publisher=American Association for the Advancement of Science (AAAS) |volume=316 |issue=5833 |date=29 June 2007 |doi=10.1126/science.1141395 |pages=1890–1893|pmid=17600214 |bibcode=2007Sci...316.1890D |s2cid=43033764 }}</ref> squash,<ref name="Dillehay Rossen Andres Williams 2007" /><ref name="smith2006">{{cite journal |last=Smith |first=Bruce D. |date=15 August 2006 |title=Eastern North America as an Independent Center of Plant Domestication |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=103 |issue=33 |pages=12223–12228 |bibcode=2006PNAS..10312223S |doi=10.1073/pnas.0604335103 |pmc=1567861 |pmid=16894156 |doi-access=free}}</ref> cotton,<ref name="Dillehay Rossen Andres Williams 2007" /> [[maize]],<ref>{{cite journal |last1=Piperno |first1=Dolores R. |title=The Origins of Plant Cultivation and Domestication in the New World Tropics: Patterns, Process, and New Developments |journal=Current Anthropology |date=October 2011 |volume=52 |issue=S4 |pages=S453–S470 |doi=10.1086/659998 |s2cid=83061925 |quote=the Central Balsas River Valley of Mexico, maize's postulated cradle of origin ... dispersed into lower Central America by 7600 BP}}</ref> [[potato]]es,<ref name="Spooner 2005 14694–99">{{cite journal |last1=Spooner |first1=David M. |last2=McLean |first2=Karen |last3=Ramsay |first3=Gavin |last4=Waugh |first4=Robbie |last5=Bryan |first5=Glenn J. |date=29 September 2005 |title=A single domestication for potato based on multilocus amplified fragment length polymorphism genotyping |journal=[[Proceedings of the National Academy of Sciences]] |pmid=16203994 |volume=102 |issue=41 |pmc=1253605 |pages=14694–14699 |doi=10.1073/pnas.0507400102 |bibcode=2005PNAS..10214694S |doi-access=free }}</ref> and [[cassava]]<ref name="Olsen Schaal 1999">{{cite journal |last1=Olsen |first1=Kenneth M. |last2=Schaal |first2=Barbara A. |title=Evidence on the origin of cassava: Phylogeography of Manihot esculenta |journal=Proceedings of the National Academy of Sciences |volume=96 |issue=10 |date=1999-05-11 |doi=10.1073/pnas.96.10.5586 |pages=5586–5591 |pmid=10318928 |pmc=21904 |bibcode=1999PNAS...96.5586O |doi-access=free}}</ref> were domesticated in the Americas.<ref name="Dillehay Rossen Andres Williams 2007"/>

Continued domestication was gradual and geographically diffuse – happening in many small steps and spread over a wide area – on the evidence of both archaeology and genetics.<ref name="Gross Olsen 2010">{{cite journal |last1=Gross |first1=Briana L. |last2=Olsen |first2=Kenneth M. |title=Genetic perspectives on crop domestication |journal=Trends in Plant Science |volume=15 |issue=9 |date=2010 |doi=10.1016/j.tplants.2010.05.008 |pages=529–537 |pmid=20541451 |pmc=2939243 |bibcode=2010TPS....15..529G }}</ref> It was a process of intermittent trial and error and often resulted in diverging traits and characteristics.<ref name="Hughes Oliveira 2019">{{cite journal |last1=Hughes |first1=Aoife |last2=Oliveira |first2=H. R. |last3=Fradgley |first3=N. |last4=Corke |first4=F. |last5=Cockram |first5=J. |last6=Doonan |first6=J. H. |last7=Nibau |first7=C. |date=March 14, 2019 |title=μCT trait analysis reveals morphometric differences between domesticated temperate small grain cereals and their wild relatives |journal=The Plant Journal |volume=99 |issue=1 |pages=98–111 |doi=10.1111/tpj.14312 |pmc=6618119 |pmid=30868647}}</ref>

Whereas domestication of animals impacted most on the genes that controlled behavior, that of plants impacted most on the genes that controlled morphology (seed size, plant architecture, dispersal mechanisms) and physiology (timing of germination or ripening),<ref name="zeder2012"/><ref name="zeder2006"/> as in the [[Wheat#Domestication|domestication of wheat]]. Wild wheat [[Shattering (agriculture)|shatters]] and falls to the ground to reseed itself when ripe, but domesticated wheat stays on the stem for easier harvesting. This change was possible because of a random mutation in the wild populations at the beginning of wheat's [[plant cultivation|cultivation]]. Wheat with this mutation was harvested more frequently and became the seed for the next crop. Therefore, without realizing it, early farmers [[artificial selection|selected for]] this mutation. The result is domesticated wheat, which relies on farmers for its reproduction and dissemination.<ref name="Purugganan Fuller 2009">{{cite journal |last1=Purugganan |first1=Michael D. |last2=Fuller |first2=Dorian Q. |title=The nature of selection during plant domestication |journal=[[Nature (journal)|Nature]] |volume=457 |issue=7231 |date=February 1, 2009 |doi=10.1038/nature07895 |url=https://www.researchgate.net/profile/Dorian-Fuller/publication/24003450_Purugganan_MD_Fuller_DQ_The_nature_of_selection_during_plant_domestication_Nature_457_843-848/links/0912f508156a26ca22000000/Purugganan-MD-Fuller-DQ-The-nature-of-selection-during-plant-domestication-Nature-457-843-848.pdf<!--NOT redundant to DOI--> |pages=843–848 |pmid=19212403 |bibcode=2009Natur.457..843P |s2cid=205216444 }}</ref>

<gallery mode=packed heights=150px>
File:Maler der Grabkammer des Menna 012.jpg|Farmers with wheat and cattle – [[Ancient Egypt]]ian art 3,400 years ago
File:Harold f Weston - Iran11.jpg|Wild [[wheat]] ears shatter when ripe, but domesticated wheat has to be [[threshing|threshed]] and [[Winnowing|winnowed]] (as shown) to release and separate the grain. Photograph by [[Harold Weston]], Iran, 1920s
</gallery>

=== Differences from wild plants ===

{{main|Domestication syndrome#In plants}}

[[File:Usdaeinkorn1 Triticum monococcum.jpg|thumb|upright=0.8|[[Einkorn wheat]] shatters into individual [[spikelet]]s, making [[harvest]]ing difficult. Domesticated cereals do not [[Shattering (agriculture)|shatter]].<ref>Snir, Ainit and Weiss, Ehud 2014 A novel morphometric method for differentiating wild and domesticated barley through intra-rachis measurements, ''Journal of Archaeological Science'' 44: 69–75, https://doi.org/10.1016/j.jas.2014.01.014</ref><ref name="Lenser-Theissen-2013" />]]

Domesticated plants differ from their wild relatives in many ways, including

* lack of [[Shattering (agriculture)|shattering]] such as of cereal ears (ripe heads),<ref name="Purugganan Fuller 2009"/> loss of fruit [[abscission]]<ref name="Lenser-Theissen-2013"/>
* less efficient [[breeding system]] (e.g. without normal [[pollinating]] organs, making human intervention a requirement), larger seeds<!--e.g. wheat--> with lower success in the wild,<ref name="Purugganan Fuller 2009"/> or even sterility (e.g. [[Fruit#Seedless fruits|seedless fruits]]) and therefore only vegetative reproduction<ref name="Agusti citrus 2020">{{cite book |last1=Agusti |first1=Manuel |last2=Primo-Millo |first2=Eduardo |date=2020 |title=The Genus Citrus |url=https://www.sciencedirect.com/science/article/pii/B9780128121634000115 |publisher=Woodhead Publishing |pages=219–244 |isbn=978-0-12-812163-4}}</ref><ref name="Perrier banana 2009">{{cite journal |last1=Perrier |first1=Xavier |last2=Bakry |first2=Frédéric |last3=Carreel |first3=Françoise |last4=Jenny |first4=Christophe |last5=Horry |first5=Jean-Pierre |last6=Lebot |first6=Vincent |last7=Hippolyte |first7=Isabelle |display-authors=3 |title=Combining Biological Approaches to Shed Light on the Evolution of Edible Bananas |journal=Ethnobotany Research & Applications|date=2009 |volume=7 |pages=199–216 |url=http://journals.sfu.ca/era/index.php/era/article/download/362/231 |doi=10.17348/era.7.0.199-216 |hdl=10125/12515 |access-date=October 27, 2019 |archive-date=November 16, 2019 |archive-url=https://web.archive.org/web/20191116055142/http://journals.sfu.ca/era/index.php/era/article/download/362/231 |url-status=live |doi-access=free|hdl-access=free }}</ref>
* better [[palatability]] (e.g. higher sugar content, reduced bitterness), better smell, and lower toxicity<ref name="Milla Osborne Turcotte Violle 2015" /><ref name="Wu Guo Mu 2019">{{Cite journal |last1=Wu |first1=Yuye |last2=Guo |first2=Tingting |last3=Mu |first3=Qi |last4=Wang |first4=Jinyu |last5=Li |first5=Xin |last6=Wu |first6=Yun |last7=Tian |first7=Bin |last8=Wang |first8=Ming Li |last9=Bai |first9=Guihua |last10=Perumal |first10=Ramasamy |last11=Trick |first11=Harold N. |display-authors=3 |date=December 2019 |title=Allelochemicals targeted to balance competing selections in African agroecosystems |url=http://www.nature.com/articles/s41477-019-0563-0 |journal=Nature Plants |volume=5 |issue=12 |pages=1229–1236 |doi=10.1038/s41477-019-0563-0 |pmid=31792396 |bibcode=2019NatPl...5.1229W |s2cid=208539527|url-access=subscription }}</ref>
* edible part larger, e.g. cereal grains<ref name="Kantar-et-al-2016">{{cite journal |last1=Kantar |first1=Michael B. |last2=Tyl |first2=Catrin E. |last3=Dorn |first3=Kevin M. |last4=Zhang |first4=Xiaofei |last5=Jungers |first5=Jacob M. |last6=Kaser |first6=Joe M. |last7=Schendel |first7=Rachel R. |last8=Eckberg |first8=James O. |last9=Runck |first9=Bryan C. |last10=Bunzel |first10=Mirko |last11=Jordan |first11=Nick R. |last12=Stupar |first12=Robert M. |last13=Marks |first13=M. David |last14=Anderson |first14=James A. |last15=Johnson |first15=Gregg A. |last16=Sheaffer |first16=Craig C. |last17=Schoenfuss |first17=Tonya C. |last18=Ismail |first18=Baraem |last19=Heimpel |first19=George E. |last20=Wyse |first20=Donald L. |display-authors=3 |title=Perennial Grain and Oilseed Crops |journal=[[Annual Review of Plant Biology]] |publisher=[[Annual Reviews (publisher)|Annual Reviews]] |volume=67 |issue=1 |date=2016-04-29 |doi=10.1146/annurev-arplant-043015-112311 |pages=703–729 |pmid=26789233|doi-access=free |bibcode=2016AnRPB..67..703K }}</ref> or fruits<ref name="Lenser-Theissen-2013">{{cite journal |last1=Lenser |first1=Teresa |last2=Theißen |first2=Günter |title=Molecular mechanisms involved in convergent crop domestication |journal=[[Trends in Plant Science]] |publisher=[[Cell Press]] |volume=18 |issue=12 |date=2013 |doi=10.1016/j.tplants.2013.08.007 |pages=704–714 |pmid=24035234|bibcode=2013TPS....18..704L }}</ref> 
* edible part more easily separated from non-edible part<ref name="Kantar-et-al-2016" />
* increased number of fruits or grains<ref name="Lenser-Theissen-2013" />
* altered color, taste, and texture<ref name="Lenser-Theissen-2013" />
* [[daylength]] independence<ref name="Lenser-Theissen-2013" />
* [[determinate growth]]<ref name="Lenser-Theissen-2013" />
* reduced or no [[vernalization]]<ref name="Lenser-Theissen-2013" />
* less [[seed dormancy]].<ref name="Lenser-Theissen-2013" />

[[Plant defense against herbivory|Plant defenses against herbivory]], such as [[thorns, spines, and prickles]], poison, protective coverings, and sturdiness may have been reduced in domesticated plants. This would make them more likely to be eaten by herbivores unless protected by humans, but there is only weak support for most of this.<ref name="Milla Osborne Turcotte Violle 2015" /> Farmers did select for reduced bitterness and lower toxicity and for food quality, which likely increased crop palatability to herbivores as to humans.<ref name="Milla Osborne Turcotte Violle 2015">{{cite journal |last1=Milla |first1=Rubén |last2=Osborne |first2=Colin P. |last3=Turcotte |first3=Martin M. |last4=Violle |first4=Cyrille |title=Plant domestication through an ecological lens |journal=Trends in Ecology & Evolution |publisher=Elsevier BV |volume=30 |issue=8 |date=2015 |doi=10.1016/j.tree.2015.06.006 |pages=463–469|pmid=26138385 |bibcode=2015TEcoE..30..463M }}</ref> However, a survey of 29 plant domestications found that crops were as well-defended against two major insect pests ([[beet armyworm]] and [[green peach aphid]]) both chemically (e.g. with bitter substances) and morphologically (e.g. with toughness) as their wild ancestors.<ref name="Turcotte Turley Johnson 2014">{{cite journal |last1=Turcotte |first1=Martin M. |last2=Turley |first2=Nash E. |last3=Johnson |first3=Marc T. J. |title=The impact of domestication on resistance to two generalist herbivores across 29 independent domestication events |journal=New Phytologist |publisher=Wiley |volume=204 |issue=3 |date=18 July 2014 |doi=10.1111/nph.12935 |pages=671–681|pmid=25039644 |doi-access=free |bibcode=2014NewPh.204..671T }}</ref>

=== Changes to plant genome ===

[[File:Polyploid wheat origins.svg|thumb|upright=2|Domesticated [[wheat]] evolved by repeated [[Hybrid (biology)|hybridization]] and [[polyploidy]] from multiple wild ancestors, increasing the size and evolvability of the genome.<ref name="Golovnina Glushkov Blinov Mayorov 2007"/> ]]

During domestication, crop species undergo intense artificial selection that alters their genomes, establishing core traits that define them as domesticated, such as increased grain size.<ref name="Purugganan Fuller 2009"/><ref name="Gepts 2004">{{cite journal |last=Gepts |first=Paul |title=Crop Domestication as a long-term selection experiment |journal=Plant Breeding Reviews |date=2004 |volume=24 |series=2 |url=http://www.plantsciences.ucdavis.edu/gepts/LTS.pdf}}</ref> Comparison of the [[coding DNA]] of [[chromosome]] 8 in rice between fragrant and non-fragrant varieties showed that aromatic and fragrant rice, including [[basmati]] and [[Jasmine rice|jasmine]], is derived from an ancestral rice domesticate that suffered a deletion in [[exon]] 7 which altered the coding for betaine aldehyde dehydrogenase (BADH2).<ref>{{cite journal |last1=Shao |first1=G. |first2=A. |last2=Tang |first3=S. Q. |last3=Tang |first4=J. |last4=Luo |first5=G. A. |last5=Jiao |first6=J. L. |last6=Wu |first7=P. S. |last7=Hu |title=A new deletion mutation of the fragrant gene and the development of three molecular markers for fragrance in rice |journal=Plant Breeding |date=April 2011 |volume=130 |issue=2 |series=2 |doi=10.1111/j.1439-0523.2009.01764.x |pages=172–176|bibcode=2011PBree.130..172S }}</ref> Comparison of the potato genome with that of other plants located genes for resistance to potato blight caused by ''[[Phytophthora infestans]]''.<ref>{{cite journal |last=The Potato Genome Sequencing Consortium|title=Genome sequence and analysis of the tuber crop potato |journal=Nature |date=July 2011 |volume=475 |doi=10.1038/nature10158 |issue=7355 |pmid=21743474 |pages=189–195 |doi-access=free}}</ref>

In [[coconut]], genomic analysis of 10 [[Microsatellite (genetics)|microsatellite loci]] (of [[noncoding DNA]]) found two episodes of domestication based on differences between individuals in the [[Indian Ocean]] and those in the [[Pacific Ocean]].<ref name="coco">{{cite journal |last1=Gunn |first1=Bee |first2=Luc |last2=Baudouin |first3=Kenneth M. |last3=Olsen |title=Independent Origins of Cultivated Coconut (Cocos nucifera L.) in the Old World Tropics |journal=PLOS ONE |date=2011 |volume=6 |issue=6 |doi=10.1371/journal.pone.0021143 |pmid=21731660 |pmc=3120816 |page=e21143 |bibcode=2011PLoSO...621143G |doi-access=free}}</ref><ref name="zeder">{{cite journal |last1=Zeder |first1=Melinda |first2=Eve |last2=Emshwiller |first3=Bruce D. |last3=Smith |first4=Daniel G. |last4=Bradley |title=Documenting domestication: the intersection of genetics and archaeology |journal=Trends in Genetics |date=March 2006 |volume=22 |issue=3 |doi=10.1016/j.tig.2006.01.007 |url=http://www.cell.com/trends/genetics/abstract/S0168-9525(06)00026-6 |accessdate=28 November 2011 |pmid=16458995 |pages=139–55|url-access=subscription }}</ref>
The coconut experienced a [[founder effect]], where a small number of individuals with low diversity founded the modern population, permanently losing much of the genetic variation of the wild population.<ref name="coco" /> [[Population bottleneck]]s which reduced variation throughout the genome at some later date after domestication are evident in crops such as [[pearl millet]], [[cotton]], [[common bean]] and [[lima bean]].<ref name="zeder" />

In wheat, domestication involved repeated [[Hybrid (biology)|hybridization]] and [[polyploidy]]. These steps are large and essentially instantaneous changes to the genome and the [[epigenome]], enabling a rapid evolutionary response to artificial selection. Polyploidy increases the number of chromosomes, bringing new combinations of genes and alleles, which in turn [[evolvability|enable further changes]] such as by [[chromosomal crossover]].<ref name="Golovnina Glushkov Blinov Mayorov 2007">{{cite journal |last1=Golovnina |first1=K. A. |last2=Glushkov |first2=S. A. |last3=Blinov |first3=A. G. |last4=Mayorov |first4=V. I. |last5=Adkison |first5=L. R. |last6=Goncharov |first6=N. P. |title=Molecular phylogeny of the genus Triticum L |journal=Plant Systematics and Evolution |publisher=Springer |volume=264 |issue=3–4 |date=2007-02-12 |doi=10.1007/s00606-006-0478-x |pages=195–216 <!--|url=http://ressources.semencespaysannes.org/docs/triticum.pdf-->|bibcode=2007PSyEv.264..195G |s2cid=39102602 }}</ref>

=== Impact on plant microbiome ===

The [[microbiome]], the collection of [[microorganism]]s inhabiting the surface and internal tissue of plants, is affected by domestication. This includes changes in microbial species composition<ref>{{Cite journal |last1=Mutch |first1=Lesley A. |last2=Young |first2=J. Peter W. |date=2004 |title=Diversity and specificity of ''Rhizobium leguminosarum'' biovar ''viciae'' on wild and cultivated legumes |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1365-294X.2004.02259.x |journal=Molecular Ecology |volume=13 |issue=8 |pages=2435–2444 |doi=10.1111/j.1365-294X.2004.02259.x |pmid=15245415 |bibcode=2004MolEc..13.2435M |s2cid=1123490}}</ref><ref>{{Cite journal |last1=Kiers |first1=E. Toby |last2=Hutton |first2=Mark G. |last3=Denison |first3=R. Ford |date=December 22, 2007 |title=Human selection and the relaxation of legume defences against ineffective rhizobia |journal=Proceedings of the Royal Society B: Biological Sciences |volume=274 |issue=1629 |pages=3119–3126 |doi=10.1098/rspb.2007.1187 |pmc=2293947 |pmid=17939985}}</ref><ref name=":4">{{Cite journal |last1=Abdelfattah |first1=Ahmed |last2=Tack |first2=Ayco J. M. |last3=Wasserman |first3=Birgit |last4=Liu |first4=Jia |last5=Berg |first5=Gabriele |last6=Norelli |first6=John |last7=Droby |first7=Samir |last8=Wisniewski |first8=Michael |display-authors=5 |title=Evidence for host–microbiome co-evolution in apple |journal=New Phytologist |date=2021 |volume=234 |issue=6 |pages=2088–2100 |doi=10.1111/nph.17820 |pmid=34823272 |pmc=9299473 |s2cid=244661193}}</ref> and diversity.<ref>{{Cite journal |last1=Coleman-Derr |first1=Devin |last2=Desgarennes |first2=Damaris |last3=Fonseca-Garcia |first3=Citlali |last4=Gross |first4=Stephen |last5=Clingenpeel |first5=Scott |last6=Woyke |first6=Tanja |last7=North |first7=Gretchen |last8=Visel |first8=Axel |last9=Partida-Martinez |first9=Laila P. |last10=Tringe |first10=Susannah G. |display-authors=5 |date=2016 |title=Plant compartment and biogeography affect microbiome composition in cultivated and native Agave species |journal=New Phytologist |volume=209 |issue=2 |pages=798–811 |doi=10.1111/nph.13697 |pmc=5057366 |pmid=26467257|bibcode=2016NewPh.209..798C }}</ref><ref name=":4" /> Plant lineage, including [[speciation]], domestication, and [[Plant breeding|breeding]], have shaped plant [[endophyte]]s ([[phylosymbiosis]]) in similar patterns as plant genes.<ref name=":4" /><ref>{{Cite journal |last1=Bouffaud |first1=Marie-Lara |last2=Poirier |first2=Marie-Andrée |last3=Muller |first3=Daniel |last4=Moënne-Loccoz |first4=Yvan |display-authors=3 |date=2014 |title=Root microbiome relates to plant host evolution in maize and other Poaceae |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/1462-2920.12442 |journal=Environmental Microbiology |volume=16 |issue=9 |pages=2804–2814 |doi=10.1111/1462-2920.12442 |pmid=24588973 |bibcode=2014EnvMi..16.2804B|url-access=subscription }}</ref><ref>{{Cite journal |last1=Abdullaeva |first1=Yulduzkhon |last2=Ambika Manirajan |first2=Binoy |last3=Honermeier |first3=Bernd |last4=Schnell |first4=Sylvia |last5=Cardinale |first5=Massimiliano |display-authors=3 |date=July 1, 2021 |title=Domestication affects the composition, diversity, and co-occurrence of the cereal seed microbiota |journal=Journal of Advanced Research |volume=31 |pages=75–86 |pmid=34194833 |doi=10.1016/j.jare.2020.12.008 |pmc=8240117}}</ref><ref>{{Cite journal |last1=Favela |first1=Alonso |last2=O. Bohn |first2=Martin |last3=D. Kent |first3=Angela |date=August 2021 |title=Maize germplasm chronosequence shows crop breeding history impacts recruitment of the rhizosphere microbiome |journal=[[The ISME Journal]] |volume=15 |issue=8 |pages=2454–2464 |doi=10.1038/s41396-021-00923-z |pmid=33692487 |pmc=8319409|bibcode=2021ISMEJ..15.2454F }}</ref>

== Fungi ==

{{further|List of domesticated fungi and microorganisms}}
[[File:20161028-AMS-LSC-0500 (30202109333).jpg|thumb|[[Agaricus bisporus|Cultivated mushrooms]] are widely grown for food.]]

Several species of [[fungi]] have been domesticated for use directly as food, or in fermentation to produce foods and drugs. The cultivated mushroom ''[[Agaricus bisporus]]'' is widely grown for food.<ref>{{cite web |url=http://www.mushroomexpert.com/agaricus_bisporus.html |title=''Agaricus bisporus'': The Button Mushroom |publisher=MushroomExpert.com |access-date=March 25, 2017}}</ref> The yeast ''[[Saccharomyces cerevisiae]]'' have been used for thousands of years to ferment [[beer]] and [[wine]], and to leaven [[bread]].<ref name="Legras Medinoglu Cornuet 2007">{{cite journal |last1=Legras |first1=Jean-Luc |last2=Merdinoglu |first2=Didier |last3=Cornuet |first3=Jean-Marie |last4=Karst |first4=Francis |title=Bread, beer and wine: Saccharomyces cerevisiae diversity reflects human history |journal=[[Molecular Ecology (journal)|Molecular Ecology]] |volume=16 |issue=10 |pages=2091–2102 |date=2007 |doi=10.1111/j.1365-294X.2007.03266.x |pmid=17498234 |bibcode=2007MolEc..16.2091L |s2cid=13157807}}</ref> Mould fungi including ''[[Penicillium]]'' are used to mature [[cheese]]s and other dairy products, as well as to make drugs such as [[antibiotics]].<ref>{{cite web |url=https://www.acs.org/content/acs/en/pressroom/newsreleases/2008/june/pfizers-work-on-penicillin-for-world-war-ii-becomes-a-national-historic-chemical-landmark.html |date=June 12, 2008 |title=Pfizer's work on penicillin for World War II becomes a National Historic Chemical Landmark |publisher=[[American Chemical Society]]}}</ref>

== Effects ==

=== On domestic animals ===

Selection of animals for visible traits may have undesired consequences for the genetics of domestic animals.<ref>{{cite book |last=Berry |first=R. J. |chapter=The Genetical Implications of Domestication in Animals |editor1-last=Ucko |editor1-first=Peter J. |editor2-last=Dimbleby |editor2-first=G. W. |title=The Domestication and Exploitation of Plants and Animals |publisher=[[Aldine Press|Aldine]] |location=Chicago |date=1969 |pages=207–217}}</ref> A side effect of domestication has been [[zoonotic]] diseases. For example, cattle have given humanity various [[Poxviridae|viral poxes]], [[measles]], and [[tuberculosis]]; pigs and ducks have contributed [[influenza]]; and horses have brought the [[rhinovirus]]es. Many [[parasite]]s, too, have their origins in domestic animals.{{sfn|Diamond|2005|pp=198, 213}} Alongside these, the advent of domestication resulted in denser human populations, which provided ripe conditions for pathogens to reproduce, mutate, spread, and eventually find a new host in humans.<ref>{{Cite journal |last=Caldararo |first=Niccolo Leo |s2cid=87639702 |date=2012 |title=Evolutionary Aspects of Disease Avoidance: The Role of Disease in the Development of Complex Society |journal=SSRN Working Paper Series |url=http://www.ssrn.com/abstract=2001098 |doi=10.2139/ssrn.2001098|url-access=subscription }}</ref>

=== On society ===

Scholars have expressed widely differing viewpoints on domestication's effects on society. [[Anarcho-primitivism]] critiques domestication as destroying the supposed primitive state of harmony with nature in hunter-gatherer societies, and replacing it, possibly violently or by enslavement, with a social [[hierarchy]] as property and power emerged.<ref name="Boyden 1992">{{cite journal |last=Boyden |first=Stephen Vickers |title=ES&T Books |journal=Environmental Science & Technology |volume=8 |date=1992 |issue=supplement 173 |page=665 |doi=10.1021/es00028a604 |bibcode=1992EnST...26..665.}}</ref> The [[Dialectical naturalism|dialectal naturalist]] [[Murray Bookchin]] has argued that domestication of animals, in turn, meant the domestication of humanity, both parties being unavoidably altered by their relationship with each other.<ref name="Bookchin">{{cite book |last=Bookchin |first=Murray |title=The Philosophy of Social Ecology |publisher=AK Press |date=2022 |edition=3rd |isbn=9781849354400 |pages=85–87}}</ref> The sociologist [[David Nibert]] asserts that the domestication of animals involved violence against animals and damage to the environment. This, in turn, he argues, corrupted human ethics and paved the way for "conquest, extermination, displacement, repression, coerced and enslaved servitude, gender subordination and sexual exploitation, and hunger."<ref>{{cite book |last=Nibert |first=David |date=2013 |title=Animal Oppression and Human Violence: Domesecration, Capitalism, and Global Conflict |url=http://cup.columbia.edu/book/animal-oppression-and-human-violence/9780231151894 |publisher=[[Columbia University Press]] |pages=1–5 |isbn=9780231151894}}</ref>

=== On diversity ===

{{further|Sustainable agriculture}}

[[File:Wheat harvest.jpg|thumb|Industrialized agriculture on land with a simplified ecosystem]]

Domesticated ecosystems provide food, reduce predator and natural dangers, and promote commerce, but their creation has resulted in habitat alteration or loss, and multiple extinctions commencing in the Late Pleistocene.<ref name="Boivin Zeder Fuller 2016">{{cite journal |last1=Boivin |first1=Nicole L. |last2=Zeder |first2=Melinda A. |last3=Fuller |first3=Dorian Q. |last4=Crowther |first4=Alison |last5=Larson |first5=Greger |last6=Erlandson |first6=Jon M. |last7=Denham |first7=Tim |last8=Petraglia |first8=Michael D. |display-authors=3 |title=Ecological consequences of human niche construction: Examining long-term anthropogenic shaping of global species distributions |journal=Proceedings of the National Academy of Sciences |volume=113 |issue=23 |pages=6388–6396 |date=2016 |doi=10.1073/pnas.1525200113 |pmid=27274046 |pmc=4988612 |bibcode=2016PNAS..113.6388B |doi-access=free}}</ref>

Domestication reduces [[genetic diversity]] of the domesticated population, especially of alleles of genes targeted by selection.<ref name="Flint-Garcia 2013">{{Cite journal |last=Flint-Garcia |first=Sherry A. |date=2013-09-04 |title=Genetics and Consequences of Crop Domestication |journal=Journal of Agricultural and Food Chemistry |volume=61 |issue=35 |pages=8267–8276 |doi=10.1021/jf305511d |pmid=23718780 |doi-access=free |bibcode=2013JAFC...61.8267F }}</ref> One reason is a [[population bottleneck]] created by artificially selecting the most desirable individuals to breed from. Most of the domesticated strain is then born from just a few ancestors, creating a situation similar to the [[founder effect]].<ref name="Brown 2019">{{Cite journal |last=Brown |first=Terence A. |date=April 2019 |title=Is the domestication bottleneck a myth? |url=https://www.nature.com/articles/s41477-019-0404-1 |journal=Nature Plants |volume=5 |issue=4 |pages=337–338 |doi=10.1038/s41477-019-0404-1 |pmid=30962526 |bibcode=2019NatPl...5..337B |s2cid=102353100}}</ref> Domesticated populations such as of dogs, rice, sunflowers, maize, and horses have an increased [[Genetic load|mutation load]], as expected in a population bottleneck where genetic drift is enhanced by the small population size. Mutations can also be fixed in a population by a [[selective sweep]].<ref name="Allaby Ware Kistler 2019">{{Cite journal |last1=Allaby |first1=Robin G. |last2=Ware |first2=Roselyn L. |last3=Kistler |first3=Logan |date=January 2019 |title=A re-evaluation of the domestication bottleneck from archaeogenomic evidence |journal=Evolutionary Applications |volume=12 |issue=1 |pages=29–37 |doi=10.1111/eva.12680 |pmc=6304682 |pmid=30622633|bibcode=2019EvApp..12...29A }}</ref><ref name="Shepherd Lange Simon 2016">{{Cite journal |last1=Shepherd |first1=Lara D. |last2=Lange |first2=Peter J. de |last3=Cox |first3=Simon |last4=McLenachan |first4=Patricia A. |last5=Roskruge |first5=Nick R. |last6=Lockhart |first6=Peter J. |date=2016-03-24 |title=Evidence of a Strong Domestication Bottleneck in the Recently Cultivated New Zealand Endemic Root Crop, Arthropodium cirratum (Asparagaceae) |journal=PLOS ONE |volume=11 |issue=3 |pages=e0152455 |doi=10.1371/journal.pone.0152455 |pmc=4806853 |pmid=27011209 |bibcode=2016PLoSO..1152455S |doi-access=free }}</ref> Mutational load can be increased by reduced selective pressure against moderately harmful traits when reproductive fitness is controlled by human management.<ref name="Frantz Bradley Larson 2020"/> However, there is evidence against a bottleneck in crops, such as barley, maize, and sorghum, where genetic diversity slowly declined rather than showing a rapid initial fall at the point of domestication.<ref name="Allaby Ware Kistler 2019"/><ref name="Brown 2019"/> Further, the genetic diversity of these crops was regularly replenished from the natural population.<ref name="Allaby Ware Kistler 2019"/> Similar evidence exists for horses, pigs, cows, and goats.<ref name="Frantz Bradley Larson 2020"/>

== Domestication by insects ==

At least three groups of insects, namely ambrosia beetles, leafcutter ants, and fungus-growing termites, have domesticated species of [[Fungus|fungi]].<ref name="Purugganan 2022"/><ref name="Mueller Gerardo 2005"/>

=== Ambrosia beetles ===

{{further|Ambrosia beetle}}

Ambrosia beetles in the [[weevil]] subfamilies [[Scolytinae]] and [[Platypodinae]] excavate tunnels in dead or stressed trees into which they introduce fungal gardens, their sole source of nutrition. After landing on a suitable tree, an ambrosia beetle excavates a tunnel in which it releases [[Ambrosia fungi|its fungal symbiont]]. The fungus penetrates the plant's [[xylem]] tissue, extracts nutrients from it, and concentrates the nutrients on and near the surface of the beetle gallery. Ambrosia fungi are typically poor wood degraders and instead utilize less demanding nutrients.<ref>{{cite journal |last1=Kasson |first1=Matthew T. |last2=Wickert |first2=Kristen L. |last3=Stauder |first3=Cameron M. |last4=Macias |first4=Angie M. |last5=Berger |first5=Matthew C. |last6=Simmons |first6=D. Rabern |last7=Short |first7=Dylan P. G. |last8=DeVallance |first8=David B. |last9=Hulcr |first9=Jiri |title=Mutualism with aggressive wood-degrading Flavodon ambrosius (Polyporales) facilitates niche expansion and communal social structure in Ambrosiophilus ambrosia beetles |journal=Fungal Ecology |date=October 2016 |volume=23 |pages=86–96 |doi=10.1016/j.funeco.2016.07.002 |doi-access=free |bibcode=2016FunE...23...86K }}</ref> Symbiotic fungi produce and detoxify ethanol, which is an attractant for ambrosia beetles and likely prevents the growth of antagonistic pathogens and selects for other beneficial symbionts.<ref>{{cite journal |last1=Ranger |first1=Christopher M. |last2=Biedermann |first2=Peter H. W. |last3=Phuntumart |first3=Vipaporn |last4=Beligala |first4=Gayathri U. |last5=Ghosh |first5=Satyaki |last6=Palmquist |first6=Debra E. |last7=Mueller |first7=Robert |last8=Barnett |first8=Jenny |last9=Schultz |first9=Peter B. |last10=Reding |first10=Michael E. |last11=Benz |first11=J. Philipp |title=Symbiont selection via alcohol benefits fungus farming by ambrosia beetles |journal=Proceedings of the National Academy of Sciences |date=24 April 2018 |volume=115 |issue=17 |pages=4447–4452 |doi=10.1073/pnas.1716852115 |pmc=5924889 |pmid=29632193 |bibcode=2018PNAS..115.4447R |doi-access=free }}</ref> Ambrosia beetles mainly colonise wood of recently dead trees.<ref name= Hulcr2017>{{cite journal |last1=Hulcr |first1=Jiri |last2=Stelinski |first2=Lukasz L. |title=The Ambrosia Symbiosis: From Evolutionary Ecology to Practical Management |journal=Annual Review of Entomology |date=31 January 2017 |volume=62 |pages=285–303 |doi=10.1146/annurev-ento-031616-035105 |pmid=27860522 |doi-access=free }}</ref>

=== Leafcutter ants ===

{{further|Leafcutter ant}}

The leafcutter ants are any of some 47 species of leaf-chewing ants in the genera ''[[Acromyrmex]]'' and ''[[Atta (genus)|Atta]]''. The ants carry the discs of leaves that they have cut back to their nest, where they feed the leaf material to the fungi that they tend. Some of these fungi are not fully domesticated: the fungi farmed by ''[[Mycocepurus smithii]]'' constantly produce spores that are not useful to the ants, which eat fungal [[hypha]]e instead. The process of domestication by ''Atta'' ants, on the other hand, is complete; it took 30 million years.<ref name="Shik Gomez Kooij 2016">{{cite journal |last1=Shik |first1=Jonathan Z. |last2=Gomez |first2=Ernesto B. |last3=Kooij |first3=Pepijn W. |last4=Santos |first4=Juan C. |last5=Wcislo |first5=William T. |last6=Boomsma |first6=Jacobus J. |title=Nutrition mediates the expression of cultivar–farmer conflict in a fungus-growing ant |journal=Proceedings of the National Academy of Sciences |volume=113 |issue=36 |date=6 September 2016 |pmid=27551065 |pmc=5018747 |doi=10.1073/pnas.1606128113 |pages=10121–10126 |doi-access=free |bibcode=2016PNAS..11310121S }}</ref>

=== Fungus-growing termites ===

{{further|Macrotermitinae|Termitomyces}}

Some 330 fungus-growing termite species of the subfamily [[Macrotermitinae]] cultivate ''[[Termitomyces]]'' fungi to eat; domestication occurred exactly once, 25–40 [[Million years ago|mya]].<ref name="Purugganan 2022"/><ref name="Mueller Gerardo 2005">{{cite journal |last1=Mueller |first1=Ulrich G. |last2=Gerardo |first2=Nicole M. |last3=Aanen |first3=Duur K. |last4=Six |first4=Diana L. |last5=Schultz |first5=Ted R. |title=The Evolution of Agriculture in Insects |journal=Annual Review of Ecology, Evolution, and Systematics |volume=36 |issue=1 |date=1 December 2005 |doi=10.1146/annurev.ecolsys.36.102003.152626 |pages=563–595 |url=https://www.academia.edu/download/51001206/The_Evolution_of_Agriculture_in_Insects20161221-18052-s5ycda.pdf<!--not redundant to DOI-->}}</ref> The fungi, described by [[Roger Heim]] in 1942, grow on 'combs' formed from the termites' excreta, dominated by tough woody fragments.<ref name="Heim 1942">{{cite journal |last=Heim |first=Roger |author-link=Roger Heim |title=Nouvelles études descriptives sur les agarics termitophiles d'Afrique tropicale |journal=Archives du Muséum National d'Histoire Naturelle |language=French |trans-title=New Descriptive Studies on the Termitophile Mushrooms of Tropical Africa |year=1942 |volume=18 |issue=6 |pages=107–166}}</ref> The termites and the fungi are both obligate [[symbiosis|symbionts]] in the relationship.<ref name="NobreAanen2010">{{cite journal |last1=Nobre |first1=T. |last2=Aanen |first2=D. K. |journal=Communicative & Integrative Biology |title=Dispersion and colonisation by fungus-growing termites |volume=3 |issue=3 |pages=248–250 |date=1 May 2010 |url=https://doi.org/10.4161/cib.3.3.11415 |doi=10.4161/cib.3.3.11415 |pmid=20714406 |pmc=2918769 }}</ref>

<gallery class=center mode=nolines widths=180 heights=180 caption="Domestication by insects">
File:Xylosandrus crassiusculus galleryR.jpg|Gallery of the ambrosia beetle ''[[Xylosandrus crassiusculus]]'' split open, with pupae and black fungus. The fungus decomposes materials in the wood, providing food for the beetles.
File:leaf cutter ants arp.jpg|Leafcutter ants ''[[Atta cephalotes]]'' carrying discs of leaf material back to their nest to feed to their domesticated fungus
Ancistrotermes latinotus.jpg|Inside the nest of the fungus-cultivating termite ''[[Ancistrotermes]]''
File:Termitomyces heimii.jpg|''[[Termitomyces heimii]]'' growing on 'comb' inside a termite mound
File:Termitomyces reticulatus 37340.jpg|''[[Termitomyces]]'' fungi are mutually dependent on [[Macrotermitinae]] termites for their survival.
</gallery>

== See also ==

{{colbegin}}
* [[Anthrozoology]]
* [[de novo domestication|''De novo'' domestication]]
* [[Domestication theory]]
* [[Experimental evolution]]
* [[Genetic erosion]]
* [[Self-domestication]]
* [[Timeline of agriculture and food technology]]
{{colend}}{{Clear}}

== References ==
{{reflist|30em}}
{{reflist|group=lower-alpha|30em}}

== Sources ==

* {{cite encyclopedia |last=Banning |first=Edward B. |date=2002 |title=Aceramic Neolithic |encyclopedia=Encyclopedia of Prehistory, Volume 8: South and Southwest Asia |publisher=[[Kluwer Academic]]/Plenum Publishers |editor-last1=Peregrine |editor-first1=Peter N. |editor-last2=Ember |editor-first2=Melvin}}
* {{cite book |last=Diamond |first=Jared |author-link=Jared Diamond |title=Guns, Germs, and Steel: A short history of everybody for the last 13,000 years |date=2005 |orig-year=1997 |publisher=[[Chatto & Windus]] |location=London |isbn=9780099302780 |title-link=Guns, Germs, and Steel}}<!--not an academic source: use with caution-->
* {{cite magazine |last1=Hare |first1=Brian |author1-link=Brian Hare |author2-link=Vanessa Woods |last2=Woods |first2=Vanessa |title=Survival of the Friendliest: Natural selection for hypersocial traits enabled Earth's apex species to best Neandertals and other competitors |magazine=[[Scientific American]] |volume=323 |issue=2 |date=August 2020 |pages=58–63 |ref=none}}
* {{cite journal |last1=Hayden |first1=Brian |s2cid=162526285 |date=2003 |title=Were luxury foods the first domesticates? Ethnoarchaeological perspectives from Southeast Asia |journal=[[World Archaeology]] |volume=34 |issue=3 |pages=458–469 |doi=10.1080/0043824021000026459a |ref=none}}
* {{cite book |last=Marciniak |first=Arkadiusz |date=2005 |title=Placing Animals in the Neolithic: Social Zooarchaeology of Prehistoric Farming Communities |url=https://books.google.com/books?id=UnQwyhdi64oC&pg=PA1RDAN |location=London |publisher=[[UCL Press]] |isbn=9781844720927 |ref=none}}
* {{cite book |last1=Zohary |first1=Daniel |author1-link=Daniel Zohary |last2=Hopf |first2=Maria |last3=Weiss |first3=Ehud |date=2012 |title=Domestication of Plants in the Old World |edition=4 |place=Oxford |publisher=[[Oxford University Press]] |doi=10.1093/acprof:osobl/9780199549061.001.0001 |isbn=9780199549061}}

== External links ==

* [http://www.cwrdiversity.org/ Crop Wild Relative Inventory and Gap Analysis]: reliable information source on where and what to conserve ex-situ for crop gene pools of global importance
* [http://www.uh.edu/engines/epi1499.htm Discussion of animal domestication with Jared Diamond]
* [http://www.sciencemag.org/cgi/content/abstract/276/5314/932 The Initial Domestication of ''Cucurbita pepo'' in the Americas 10,000 Years Ago]
* [http://www.geochembio.com/biology/organisms/cattle/ Cattle domestication diagram] {{Webarchive |url=https://web.archive.org/web/20101219082134/http://www.geochembio.com/biology/organisms/cattle/ |date=December 19, 2010}}
* [https://www.ncbi.nlm.nih.gov/sites/entrez?Db=pubmed&Cmd=DetailsSearch&Term=domestication%5BTitle%5D+AND+%22loattrfree+full+text%22%5Bsb%5D Major topic 'domestication': free full-text articles (more than 100 plus reviews) in National Library of Medicine]

{{Prehistoric technology}}
{{Animal domestication}}

{{Authority control}}

[[Category:Domestication| ]]
[[Category:History of agriculture]]