Domestication

Dogs and sheep were among the first animals to be domesticated.

Domestication is a sustained multi-generational relationship in which one group of organisms assumes a significant degree of influence over the reproduction and care of another group to secure a more predictable supply of resources from that second group.[1] 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 where traits evolve as a by-product of natural selection or from selection on other traits.[2][3][4] There is a genetic difference between domestic and wild populations. There is also such a difference between the domestication traits that researchers believe to have been essential at the early stages of domestication, and the improvement traits that have appeared since the split between wild and domestic populations.[5][6][7] 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.[6][7][8]

The dog was the first domesticant,[9][10][11] and was established across Eurasia before the end of the Late Pleistocene era, well before cultivation and before the domestication of other animals.[10] The archaeological and genetic data suggest that long-term bidirectional gene flow between wild and domestic stocks – including donkeys, horses, New and Old World camelids, goats, sheep, and pigs – was common.[7][12] Given its importance to humans and its value as a model of evolutionary and demographic change, domestication has attracted scientists from archaeology, palaeontology, anthropology, botany, zoology, genetics, and the environmental sciences.[13]

Etymology

Domestication (from the Latin domesticus) means 'belonging to the house'.[14]

Related terminology

Plants domesticated primarily for aesthetic enjoyment in and around the home are usually called house plants or ornamentals, while those domesticated for large-scale food production are generally called crops. A distinction can be made between those domesticated plants that have been deliberately altered or selected for special desirable characteristics, termed cultigens, and those plants that are used for human benefit, but are essentially no different from the wild populations of the species. Animals domesticated for home companionship are usually called pets, while those domesticated for food or work are called livestock or farm animals.

Definitions

Domestication

Domestication has been defined as "a sustained multi-generational, mutualistic relationship in which one organism assumes a significant degree of influence over the reproduction and care of another organism in order to secure a more predictable supply of a resource of interest, and through which the partner organism gains advantage over individuals that remain outside this relationship, thereby benefitting and often increasing the fitness of both the domesticator and the target domesticate."[1][15][16][17][18] This definition recognizes both the biological and the cultural components of the domestication process and the impacts on both humans and the domesticated animals and plants. All past definitions of domestication have included a relationship between humans with plants and animals, but their differences lay in who was considered as the lead partner in the relationship. This new definition recognizes a mutualistic relationship in which both partners gain benefits. Domestication has vastly enhanced the reproductive output of crop plants, livestock, and pets far beyond that of their wild progenitors. Domesticates have provided humans with resources that they could more predictably and securely control, move, and redistribute, which has been the advantage that had fueled a population explosion of the agro-pastoralists and their spread to all corners of the planet.[18]

This biological mutualism is not restricted to humans with domestic crops and livestock but is well-documented in nonhuman species, especially among a number of social insect domesticators and their plant and animal domesticates, for example the ant–fungus mutualism that exists between leafcutter ants and certain fungi.[1]

Domestication syndrome

Domestication syndrome is a term often used to describe the suite of phenotypic traits arising during domestication that distinguish crops from their wild ancestors.[5][19] The term is also applied to animals and includes increased docility and tameness, coat color changes, reductions in tooth size, changes in craniofacial morphology, alterations in ear and tail form (e.g., floppy ears), more frequent and nonseasonal estrus cycles, alterations in adrenocorticotropic hormone levels, changed concentrations of several neurotransmitters, prolongations in juvenile behavior, and reductions in both total brain size and of particular brain regions.[20]

Impact on humans

The domestication of animals and plants began with the wolf (Canis lupus) at least 15,000 years before present (YBP), which then led to a rapid shift in the evolution, ecology, and demography of both humans and numerous species of animals and plants.[21][7] The sudden appearance of the domestic dog (Canis lupus familiaris) in the archaeological record was followed by livestock and crop domestication, and the transition of humans from foraging to farming in different places and times across the planet.[21][22][23]

Around 10,000 YBP, a new way of life emerged for humans through the management and exploitation of plant and animal species, leading to higher-density populations in the centers of domestication,[21][24] the expansion of agricultural economies, and the development of urban communities.[21][25]

Domestication of animals

Sheep with an ear tag that is part of a national livestock identification system

The domestication of animals is the scientific theory of the mutual relationship between animals with the humans who have influence on their care and reproduction.[1] 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 where traits evolve as a by-product of natural selection or from selection on other traits.[2][3][4] There is a genetic difference between domestic and wild populations. There is also such a difference between the domestication traits that researchers believe to have been essential at the early stages of domestication, and the improvement traits that have appeared since the split between wild and domestic populations.[5][6][7] 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.[6][7][8]

Domestication should not be confused with taming. Taming is the conditioned behavioral modification of a wild-born animal when its natural avoidance of humans is reduced and it accepts the presence of humans, but domestication is the permanent genetic modification of a bred lineage that leads to an inherited predisposition toward humans.[26][27][28] Certain animal species, and certain individuals within those species, make better candidates for domestication than others because they exhibit certain behavioral characteristics: (1) the size and organization of their social structure; (2) the availability and the degree of selectivity in their choice of mates; (3) the ease and speed with which the parents bond with their young, and the maturity and mobility of the young at birth; (4) the degree of flexibility in diet and habitat tolerance; and (5) responses to humans and new environments, including flight responses and reactivity to external stimuli.[18]:Fig 1[29][30][31]

The beginnings of animal domestication involved a protracted coevolutionary process with multiple stages along different pathways.[7] It is proposed that there were three major pathways that most animal domesticates followed into domestication: (1) commensals, adapted to a human niche (e.g., dogs, cats, fowl, possibly pigs); (2) prey animals sought for food (e.g., sheep, goats, cattle, water buffalo, yak, pig, reindeer, llama and alpaca); and (3) targeted animals for draft and nonfood resources (e.g., horse, donkey, camel).[7][12][18][32][33][34][35][36][37] The dog was the first domesticant,[10][11] and was established across Eurasia before the end of the Late Pleistocene era, well before cultivation and before the domestication of other animals.[10] Humans did not intend to domesticate animals from, or at least they did not envision a domesticated animal resulting from, either the commensal or prey pathways. In both of these cases, humans became entangled with these species as the relationship between them, and the human role in their survival and reproduction, intensified.[7] Although the directed pathway proceeded from capture to taming, the other two pathways are not as goal-oriented and archaeological records suggest that they take place over much longer time frames.[13]

Unlike other domestic species which were primarily selected for production-related traits, dogs were initially selected for their behaviors.[38][39] The archaeological and genetic data suggest that long-term bidirectional gene flow between wild and domestic stocks – including donkeys, horses, New and Old World camelids, goats, sheep, and pigs – was common.[7][12] One study has concluded that 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 also apply to other domesticated animals.[40][41]

Domestication of plants

The initial domestication of animals impacted most on the genes that controlled their behavior, but the initial domestication of plants impacted most on the genes that controlled their morphology (seed size, plant architecture, dispersal mechanisms) and their physiology (timing of germination or ripening).[18][23]

Farmers with wheat and cattle - Ancient Egyptian art 1,422 BCE

The domestication of wheat provides an example. Wild wheat falls to the ground to reseed itself when ripe, but domesticated wheat stays on the stem for easier harvesting. There is evidence that this change was possible because of a random mutation that happened in the wild populations at the beginning of wheat's cultivation. Wheat with this mutation was harvested more frequently and became the seed for the next crop. Therefore, without realizing, early farmers selected for this mutation, which may otherwise have died out. The result is domesticated wheat, which relies on farmers for its own reproduction and dissemination.[42]

The earliest human attempts at plant domestication occurred in South-Western Asia. There is early evidence for conscious cultivation and trait selection of plants by pre-Neolithic groups in Syria: grains of rye with domestic traits have been recovered from Epi-Palaeolithic (c. 11,050 BCE) contexts at Abu Hureyra in Syria,[43] but this appears to be a localised phenomenon resulting from cultivation of stands of wild rye, rather than a definitive step towards domestication.[43]

By 10,000 BCE the bottle gourd (Lagenaria siceraria) plant, used as a container before the advent of ceramic technology, appears to have been domesticated. The domesticated bottle gourd reached the Americas from Asia by 8000 BCE, most likely due to the migration of peoples from Asia to America.[44]

Cereal crops were first domesticated around 9000 BCE in the Fertile Crescent in the Middle East. The first domesticated crops were generally annuals with large seeds or fruits. These included pulses such as peas and grains such as wheat. The Middle East was especially suited to these species; the dry-summer climate was conducive to the evolution of large-seeded annual plants, and the variety of elevations led to a great variety of species. As domestication took place humans began to move from a hunter-gatherer society to a settled agricultural society. This change would eventually lead, some 4000 to 5000 years later, to the first city states and eventually the rise of civilization itself.

Continued domestication was gradual, a process of trial and error that occurred intermittently. Over time perennials and small trees began to be domesticated including apples and olives. Some plants were not domesticated until recently such as the macadamia nut and the pecan.

In other parts of the world very different species were domesticated. In the Americas squash, maize, beans, and perhaps manioc (also known as cassava) formed the core of the diet. In East Asia millet, rice, and soy were the most important crops. Some areas of the world such as Southern Africa, Australia, California and southern South America never saw local species domesticated.

Domesticated plants often differ from their wild relatives in the way they spread to a more diverse environment and have a wider geographic range;[45] they may also have a different ecological preference; flower and fruit simultaneously; may lack shattering or scattering of seeds, and may have lost their dispersal mechanisms completely; have larger fruits and seeds, and so lower efficiency of dispersal; may have been converted from a perennial to annual; have lost seed dormancy and photoperiodic controls; lack normal pollinating organs; may have a different breeding system; may lack defensive adaptations such as hairs, spines and thorns, protective coverings and sturdiness; may have better palatability and chemical composition, rendering them more likely to be eaten by animals; may be more susceptible to diseases and pests; may develop seedless parthenocarpic fruits; may have undergone selection for double flowers, which may involve conversion of stamens into petals; may have become sexually sterile and therefore only reproduce vegetatively.

Negative aspects

Jared Diamond in his book Guns, Germs, and Steel discusses the universal tendency for populations that have acquired agriculture and domestic animals to first develop a large population and then to move. He recounts migrations of people armed with a suite of domestic crops overtaking, displacing or killing indigenous hunter-gatherers,[3]:p112 until after millions of years of following the hunter-gatherer lifestyle there will shortly be none remaining.[3]:p86

Selection of animals for visible "desirable" traits may make them unfit in other, unseen, ways. The consequences for the captive and domesticated animals were reduction in size, piebald color, shorter faces with smaller and fewer teeth, diminished horns, weak muscle ridges, and less genetic variability. Poor joint definition, late fusion of the limb bone epiphyses with the diaphyses, hair changes, greater fat accumulation, smaller brains, simplified behavior patterns, extended immaturity, and more pathology are a few of the defects of domestic animals. All of these changes have been documented in direct observations of the rat in the 19th century, by archaeological evidence, and confirmed by animal breeders in the 20th century.[46] A 2014, a study proposed the theory that under selection, docility in mammals and birds results partly from a slowed pace of neural crest development, that would in turn cause a reduced fear–startle response due to mild neurocristopathy that causes domestication syndrome. The theory was unable to explain curly tails nor domestication syndrome exhibited by plants.[20]

One side effect of domestication has been zoonotic diseases. For example, cattle have given humanity various viral poxes, measles, and tuberculosis; pigs and ducks have given influenza; and horses have given the rhinoviruses. Humans share over sixty diseases with dogs. Many parasites also have their origins in domestic animals.[3] 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.

Paul Shepard writes "Man substitutes controlled breeding for natural selection; animals are selected for special traits like milk production or passivity, at the expense of overall fitness and nature-wide relationships...Though domestication broadens the diversity of forms – that is, increases visible polymorphism – it undermines the crisp demarcations that separate wild species and cripples our recognition of the species as a group. Knowing only domestic animals dulls our understanding of the way in which unity and discontinuity occur as patterns in nature, and substitutes an attention to individuals and breeds. The wide variety of size, color, shape, and form of domestic horses, for example, blurs the distinction among different species of Equus that once were constant and meaningful."[47]

Some anarcho-primitivist authors describe domestication as the process by which previously nomadic human populations shifted towards a sedentary or settled existence through agriculture and animal husbandry. They claim that this kind of domestication demands a totalitarian relationship with both the land and the plants and animals being domesticated. They say that whereas, in a state of wildness, all life shares and competes for resources, domestication destroys this balance. Domesticated landscape (e.g. pastoral lands/agricultural fields and, to a lesser degree, horticulture and gardening) ends the open sharing of resources; where "this was everyone's," it is now "mine." Anarcho-primitivists state that this notion of ownership laid the foundation for social hierarchy as property and power emerged. It also involved the destruction, enslavement, or assimilation of other groups of early people who did not make such a transition.[48]

Global impact on diversity

Industrialized wheat harvest - North America today

In 2016, a study found that humans have had a major impact on global genetic diversity as well as extinction rates, including a contribution to megafaunal extinctions. Pristine landscapes no longer exist and have not existed for millennia, and humans have concentrated the planet's biomass into human-favored plants and animals. Domesticated ecosystems provide food, reduce predator and natural dangers, and promote commerce, but have also resulted in habitat loss and extinctions commencing in the Late Pleistocene. Ecologists and other researchers are advised to make better use of the archaeological and paleoecological data available for gaining an understanding the history of human impacts before proposing solutions.[49]

Genetic pollution

Main article: Genetic pollution

Genetic pollution is a controversial term for uncontrolled gene flow from domestic into wild populations.[50][51] This gene flow is undesirable according to some environmentalists and conservationists, including groups such as Greenpeace, TRAFFIC, and GeneWatch UK.

Lists

Early domestication: cow being milked in ancient Egypt.

List of domesticated animals

Domesticated animals include, or have included, a variety of mammals and birds. Some common domesticated animals includes the Cat, Cattle, Chicken, Dog, Donkey, Duck, Goat, Goose, Horse, Pig and Sheep.

For more details on this topic, see List of domesticated animals.

List of hybrid domestic animals

Hybrid domestic animals include those domestic animals that have been crossed with their wild relatives, and include the Beefalo, Bengal cat, Cama (animal), Chausie, Coydog, Dzo, Domesticated hedgehog, Sheep-goat hybrid, Hinny, Huarizo, Iron Age pig, Mule, Savannah (cat), Wolfdog, Yakalo, Zeedonk, Zorse, Zony and Zubron.

List of domesticated plants

See also

References

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Bibliography

Further reading

External links

Look up domestication or taming in Wiktionary, the free dictionary.
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