Domestication

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Dogs and sheep were among the first animals to be domesticated, at least 15,000 and 11,000 years ago respectively.[1]
Rice was domesticated in China, some 13,500 to 8,200 years ago.[2]

Domestication is a multi-generational mutualistic relationship between humans and other organisms, in which humans took over control and care to obtain a steady supply of resources including food. The process was gradual and geographically diffuse, based on trial and error.

The first

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

squash, maize, potatoes, cotton, and cassava. In Africa, crops such as sorghum were domesticated. Agriculture developed
in some 13 centres around the world, domesticating different crops and animals.

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 of seed-heads such as in wheat. Such changes both make domesticated organisms easier to handle, and reduce their ability to survive in the wild.

Definitions

Domestication (not to be confused with the

Latin domesticus, 'belonging to the house'.[6] 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 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.[7][8][9]

Domestication syndrome is the suite of phenotypic traits which arose during the initial domestication process, and which distinguish crops from their wild ancestors.[10][11] It can also mean a set of differences now observed in domesticated animals, 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.[12]

Cause and timing

Further information:
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 and which continue to this present day. These changes made obtaining food by hunting and gathering difficult.[13] The first animal to be domesticated was the dog at least 15,000 years ago.[1] 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.[14]

Timeline of some major domestication events
Event Centre of origin Purpose Date/years ago
Foraging for wild grains Asia Food > 23,000[15]
Dog Eurasia Commensal > 15,000[1]
Rice China Food 13,500–8,200[2]
Wheat, Barley Near East Food 13,000–11,000[15]
Flax Near East Textiles 13,000–11,000[16]
Cow
 Near East, South Asia Food 11,000–10,000[1]
Chicken East Asia
Cockfighting
 7,000[17]
Horse Central Asia Draft, riding 5,500[1]

The appearance of the

invention of agriculture, and the transition of humans from foraging to farming in different places and times across the planet.[1][18][19][20] For instance, small-scale trial cultivation of cereals began some 28,000 years ago at the Ohalo II site in Israel.[21]

In the

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.[1] The cat was domesticated in the Fertile Crescent, perhaps 10,000 years ago,[22] from European wildcats, possibly to control rodents that were damaging stored food.[23]

Neolithic revolution as understood in 2003[24]

Animals

Desirable traits

Further information: Domestication of vertebrates
chihuahua dog
(right)

The domestication of animals is the relationship between animals and humans who have influence on their care and reproduction.[7] 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.[26][27][28]

There is a difference between domestic and wild populations; some of these differences constitute the domestication syndrome, traits presumed essential in the early stages of domestication, while others represent later improvement traits.[10][29][30] Domesticated animals tend to be smaller and less aggressive than their wild counterparts; other common traits are floppy ears, a smaller brain, and a shorter muzzle.[25] 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 regional populations.[29][30][31]

Certain animal species, and certain individuals within those species, make better candidates for domestication because of their behavioral characteristics:[32][33][34][35]

  1. The size and organization of their social structure[32]
  2. The availability and the degree of selectivity in their choice of mates[32]
  3. The ease and speed with which the parents bond with their young, and the maturity and mobility of the young at birth[32]
  4. The degree of flexibility in diet and habitat tolerance[32]
  5. Responses to humans and new environments, including reduced flight response and reactivity to external stimuli.[32]

Mammals

Further information: List of domesticated animals
While dogs were commensals, and sheep were kept for food, camels, like horses and donkeys, were domesticated as working animals.[32]

The beginnings of animal domestication involved a protracted coevolutionary process with multiple stages along different pathways. There are three proposed major pathways that most animal domesticates followed into domestication:[32][30][36]

  1. pigs)[32]
  2. prey animals sought for food (e.g.,
    yak, pig, reindeer, llama and alpaca)[32]
  3. animals targeted for draft and riding (e.g., horse, donkey, camel).[32]

Humans did not intend to domesticate animals 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 led gradually to formalised animal husbandry.[30] 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.[37]

Unlike other domestic species selected primarily for production-related traits, dogs were initially selected for their behaviors.[9][38] The dog was domesticated long before other animals,[39][40] becoming established across Eurasia before the end of the Late Pleistocene era, well before agriculture.[39]

The archaeological and genetic data suggest that long-term bidirectional gene flow between wild and domestic stocks – such as in donkeys, horses, New and Old World camelids, goats, sheep, and pigs – was common. [30][36] 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. [41][42]

The 2023

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 microRNAs. 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.[43][44]

Birds

Main articles: Poultry and Aviculture
cockfighting, some 7,000 years ago.[17]

Domesticated birds principally mean

parrots; these are kept both for pleasure and for use in research.[46]
The
cockfighting rather than for food.[17]

Invertebrates

Further information:
Domestication of bees, Beekeeping, and Sericulture

Two

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 pupal cocoon; the western honey bee, for honey, and, from the 20th century, for pollination of crops.[48][49]

Several other invertebrates have been domesticated, both terrestrial and aquatic, including some such as

Helix are raised for food. Several parasitic or parasitoidal insects including the fly Eucelatoria, the beetle Chrysolina, and the wasp 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.[50]

Plants

Further information: History of agriculture and List of domesticated plants

Humans

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.[53][54] Rice was first cultivated in East Asia.[2][55] Sorghum was widely cultivated in sub-Saharan Africa,[56] while peanuts,[57] squash,[57][58] cotton, [57] maize,[59] potatoes,[60] and cassava[61] were domesticated in the Americas.[57]

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.[62] It was a process of intermittent trial and error, and often resulted in diverging traits and characteristics.[63]

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),

selected for this mutation. The result is domesticated wheat, which relies on farmers for its reproduction and dissemination.[15]

Differences from wild plants

Main article: Domestication syndrome § In plants
Einkorn wheat shatters into individual spikelets, making harvesting difficult. Domesticated cereals do not shatter.[64][65]

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

green peach aphid) both chemically (e.g. with bitter substances) and morphologically (e.g. with toughness) as their wild ancestors.[71]

Changes to plant genome

Domesticated wheat evolved by repeated hybridization and polyploidy from multiple wild ancestors, increasing the size and evolvability of the genome.[72]

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.

coding DNA of chromosome 8 in rice between fragrant and non-fragrant varieties showed that aromatic and fragrant rices, including basmati and jasmine, are derived from an ancestral rice domesticate that suffered a deletion in exon 7 which altered the coding for betaine aldehyde dehydrogenase (BADH2).[74] Comparison of the potato genome with that of other plants located genes for resistance to potato blight caused by Phytophthora infestans.[75]

In

noncoding DNA) found two episodes of domestication based on differences between individuals in the Indian Ocean and those in the Pacific Ocean.[76][77]
The coconut experienced a
common bean and lima bean.[77]

In wheat, domestication involved repeated 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 enable further changes such as by chromosomal crossover.[72]

Impact on plant microbiome

The microbiome, the collection of microorganisms inhabiting the surface and internal tissue of plants, is affected by domestication. This includes changes in microbial species composition[78][79][80] and diversity.[81][80] Plant lineage, including speciation, domestication, and breeding, have shaped plant endophytes (phylosymbiosis) in similar patterns as plant genes.[80][82][83][84]

Fungi

Further information: List of domesticated fungi and microorganisms
Cultivated mushrooms are widely grown for food.

Several species of

antibiotics.[87]

Effects

On domestic animals

Selection of animals for visible traits may have undesired consequences for the genetics of domestic animals.

parasites, too, have their origins in domestic animals.[89] 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.[90]

On society

Scholars have expressed widely differing viewpoints on domestication's effects on society.

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.[92] 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."[93]

On diversity

Further information: Sustainable agriculture
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.[94]

Domestication reduces genetic diversity of the domesticated population, especially of alleles of genes targeted by selection.[95] 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.[96] Domesticated populations such as of dogs, rice, sunflowers, maize, and horses have an increased 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.[97][98] Mutational load can be increased by reduced selective pressure against moderately harmful traitswhen reproductive fitness is controlled by human management.[25] 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.[97][96] Further, genetic diversity of these crops was regularly replenished from the natural population.[97] Similar evidence exists for horses, pigs, cows, and goats.[25]

See also

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External links
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