Editing Domestication (section)

== 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}}