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Thursday, May 23, 2019

Genetic "pollution"

From Wikipedia, the free encyclopedia

Genetic pollution is a controversial term for uncontrolled gene flow into wild populations. It is defined as “the dispersal of contaminated altered genes from genetically engineered organisms to natural organisms, esp. by cross-pollination”, but has come to be used in some broader ways. It is related to the population genetics concept of gene flow, and genetic rescue, which is genetic material intentionally introduced to increase the fitness of a population. It is called genetic pollution when it negatively impacts on the fitness of a population, such as through outbreeding depression and the introduction of unwanted phenotypes which can lead to extinction.
 
Conservation biologists and conservationists have used the term to describe gene flow from domestic, feral, and non-native species into wild indigenous species, which they consider undesirable. They promote awareness of the effects of introduced invasive species that may "hybridize with native species, causing genetic pollution". In the fields of agriculture, agroforestry and animal husbandry, genetic pollution is used to describe gene flows between genetically engineered species and wild relatives. The use of the word “pollution” is meant to convey the idea that mixing genetic information is bad for the environment, but because the mixing of genetic information can lead to a variety of outcomes, “pollution” may not always be the most accurate descriptor.

Gene flow to wild population

Some conservation biologists and conservationists have used genetic pollution for a number of years as a term to describe gene flow from a non-native, invasive subspecies, domestic, or genetically-engineered population to a wild indigenous population.

Importance

The introduction of genetic material into the gene pool of a population by human intervention can have both positive and negative effects on populations. When genetic material is intentionally introduced to increase the fitness of a population, this is called genetic rescue. When genetic material is unintentionally introduced to a population, this is called genetic pollution and can negatively affect the fitness of a population (primarily through outbreeding depression), introduce other unwanted phenotypes, or theoretically lead to extinction.

Introduced species

An introduced species is one that is not native to a given population that is either intentionally or accidentally brought into a given ecosystem. Effects of introduction are highly variable, but if an introduced species has a major negative impact on its new environment, it can be considered an invasive species. One such example is the introduction of the Asian Longhorned beetle in North America, which was first detected in 1996 in Brooklyn, New York. It is believed that these beetles were introduced through cargo at trade ports. The beetles are highly damaging to the environment, and are estimated to cause risk to 35% of urban trees, excluding natural forests. These beetles cause severe damage to the wood of trees by larval funneling. Their presence in the ecosystem destabilizes community structure, having a negative influence on many species in the system. Introduced species are not always disruptive to an environment, however. Tomás Carlo and Jason Gleditch of Penn State University found that the number of invasive honeysuckle plants in the area correlated with the number and diversity of the birds in the Happy Valley Region of Pennsylvania, suggesting introduced honeysuckle plants and birds formed a mutually beneficial relationship. Presence of introduced honeysuckle was associated with higher diversity of the bird populations in that area, demonstrating that introduced species are not always detrimental to a given environment and it is completely context dependent.

Invasive species

Conservation biologists and conservationists have, for a number of years, used the term to describe gene flow from domestic, feral, and non-native species into wild indigenous species, which they consider undesirable. For example, TRAFFIC is the international wildlife trade monitoring network that works to limit trade in wild plants and animals so that it is not a threat to conservationist goals. They promote awareness of the effects of introduced invasive species that may "hybridize with native species, causing genetic pollution". Furthermore, the Joint Nature Conservation Committee, the statutory adviser to the UK government, has stated that invasive species "will alter the genetic pool (a process called genetic pollution), which is an irreversible change."

Invasive species can invade both large and small native populations and have a profound effect. Upon invasion, invasive species interbreed with native species to form sterile or more evolutionarily fit hybrids that can outcompete the native populations. Invasive species can cause extinctions of small populations on islands that are particularly vulnerable due to their smaller amounts of genetic diversity. In these populations, local adaptations can be disrupted by the introduction of new genes that may not be as suitable for the small island environments. For example, the Cercocarpus traskiae of the Catalina Island off the coast of California has faced near extinction with only a single population remaining due to the hybridization of its offspring with Cercocarpus betuloides.

Domestic populations

Increased contact between wild and domesticated populations of organisms can lead to reproductive interactions that are detrimental to the wild population's ability to survive. A wild population is one that lives in natural areas and is not regularly looked after by humans. This contrast with domesticated populations that live in human controlled areas and are regularly, and historically, in contact with humans. Genes from domesticated populations are added to wild populations as a result of reproduction. In many crop populations this can be the result of pollen traveling from farmed crops to neighboring wild plants of the same species. For farmed animals, this reproduction may happen as the result of escaped or released animals.

Aquaculture

Aquaculture is the practice of farming aquatic animals or plants for the purpose of consumption. This practice is becoming increasingly common for the production of salmon. This is specifically termed aquaculture of salmonoids. One of the dangers of this practice is the possibility of domesticated salmon breaking free from their containment. The occurrence of escaping incidents is becoming increasingly common as aquaculture gains popularity. Farming structures may be ineffective at holding the vast number of fast growing animals they house. Natural disasters, high tides, and other environmental occurrences can also trigger aquatic animal escapes. The reason these escapes are considered dangers is the impact they pose for the wild population they reproduce with after escaping. In many instances the wild population experiences a decreased likelihood of survival after reproducing with domesticated populations of salmon.

The Washington Department of Fish and Wildlife cites that "commonly expressed concerns surrounding escaped Atlantic salmon include competition with native salmon, predation, disease transfer, hybridization, and colonization" A report done by that organization in 1999 did not find that escaped salmon posed a significant risk to wild populations.

Crops

Crops refer to groups of plants grown for consumption. Despite domestication over many years, these plants are not so far removed from their wild relatives that they could reproduce if brought together. Many crops are still grown in the areas they originated and gene flow between crops and wild relatives impacts the evolution of wild populations. Farmers can avoid reproduction between the different populations by timing their planting of crops so that crops are not flowering when wild relatives would be. Domesticated crops have been changed through artificial selection and genetic engineering. The genetic make up of many crops is different than that of its wild relatives, but the closer they grow to one another the more likely they are to share genes through pollen. Gene flow persists between crops and wild counterparts.

Genetically engineered organisms

Genetically engineered organisms are genetically modified in a laboratory, and therefore distinct from those that were bred through artificial selection. In the fields of agriculture, agroforestry and animal husbandry, genetic pollution is being used to describe gene flows between GE species and wild relatives. An early use of the term "genetic pollution" in this later sense appears in a wide-ranging review of the potential ecological effects of genetic engineering in The Ecologist magazine in July 1989. It was also popularized by environmentalist Jeremy Rifkin in his 1998 book The Biotech Century. While intentional crossbreeding between two genetically distinct varieties is described as hybridization with the subsequent introgression of genes, Rifkin, who had played a leading role in the ethical debate for over a decade before, used genetic pollution to describe what he considered to be problems that might occur due the unintentional process of (modernly) genetically modified organisms (GMOs) dispersing their genes into the natural environment by breeding with wild plants or animals.

Concerns about negative consequences from gene flow between genetically engineered organisms and wild populations are valid. Most corn and soybean crops grown in the midwestern USA are genetically modified. There are corn and soybean varieties that are resistant to herbicides like glyphosate and corn that produces neonicotinoid pesticide within all of its tissues. These genetic modifications are meant to increase yields of crops but there is little evidence that yields actually increase. While scientists are concerned genetically engineered organisms can have negative effects on surrounding plant and animal communities, the risk of gene flow between genetically engineered organisms and wild populations is yet another concern. Many farmed crops may be weed resistant and reproduce with wild relatives. More research is necessary to understand how much gene flow between genetically engineered crops and wild populations occurs, and the impacts of genetic mixing.

Mutated organisms

Mutations within organisms can be executed through the process of exposing the organism to chemicals or radiation in order to generate mutations. This has been done in plants in order to create mutants that have a desired trait. These mutants can then be bred with other mutants or individuals that are not mutated in order to maintain the mutant trait. However, similar to the risks associated with introducing individuals to a certain environment, the variation created by mutated individuals could have a negative impact on native populations as well.

Preventative measures

Since 2005 there has existed a GM Contamination Register, launched for GeneWatch UK and Greenpeace International that records all incidents of intentional or accidental release of organisms genetically modified using modern techniques.

Genetic use restriction technologies (GURTs) were developed for the purpose of property protection, but could be beneficial in preventing the dispersal of transgenes. GeneSafe technologies introduced a method that became known as “Terminator.” This method is based on seeds that produce sterile plants. This would prevent movement of transgenes into wild populations as hybridization would not be possible. However, this technology has never been deployed as it disproportionately negatively affects farmers in developing countries, who save seeds to use each year (whereas in developed countries, farmers generally buy seeds from seed production companies).

Physical containment has also been utilized to prevent the escape of transgenes. Physical containment includes barriers such as filters in labs, screens in greenhouses, and isolation distances in the field. Isolation distances have not always been successful, such as transgene escape from an isolated field into the wild in herbicide-resistant bentgrass Agrostis stolonifera.

Another suggested method that applies specifically to protection traits (e.g. pathogen resistance) is mitigation. Mitigation involves linking the positive trait (beneficial to fitness) to a trait that is negative (harmful to fitness) to wild but not domesticated individuals. In this case, if the protection trait was introduced to a weed, the negative trait would also be introduced in order to decrease overall fitness of the weed and decrease possibility of the individual’s reproduction and thus propagation of the transgene.

Risks

Not all genetically engineered organisms cause genetic pollution. Genetic engineering has a variety of uses and is specifically defined as a direct manipulation of the genome of an organism. Genetic pollution can occur in response to the introduction of a species that is not native to a particular environment, and genetically engineered organisms are examples of individuals that could cause genetic pollution following introduction. Due to these risks, studies have been done in order to assess the risks of genetic pollution associated with organisms that have been genetically engineered:
  1. Genetic In a 10-year study of four different crops, none of the genetically engineered plants were found to be more invasive or more persistent than their conventional counterparts. An often cited claimed example of genetic pollution is the reputed discovery of transgenes from GE maize in landraces of maize in Oaxaca, Mexico. The report from Quist and Chapela, has since been discredited on methodological grounds. The scientific journal that originally published the study concluded that "the evidence available is not sufficient to justify the publication of the original paper."  More recent attempts to replicate the original studies have concluded that genetically modified corn is absent from southern Mexico in 2003 and 2004.
  2. A 2009 study verified the original findings of the controversial 2001 study, by finding transgenes in about 1% of 2000 samples of wild maize in Oaxaca, Mexico, despite Nature retracting the 2001 study and a second study failing to back up the findings of the initial study. The study found that the transgenes are common in some fields, but non-existent in others, hence explaining why a previous study failed to find them. Furthermore, not every laboratory method managed to find the transgenes.
  3. A 2004 study performed near an Oregon field trial for a genetically modified variety of creeping bentgrass (Agrostis stolonifera) revealed that the transgene and its associate trait (resistance to the glyphosate herbicide) could be transmitted by wind pollination to resident plants of different Agrostis species, up to 14 km from the test field. In 2007, the Scotts Company, producer of the genetically modified bentgrass, agreed to pay a civil penalty of $500,000 to the United States Department of Agriculture (USDA). The USDA alleged that Scotts "failed to conduct a 2003 Oregon field trial in a manner which ensured that neither glyphosate-tolerant creeping bentgrass nor its offspring would persist in the environment".
Not only are there risks in terms of genetic engineering, but there are risks that emerge from species hybridization In Czechoslovakia, ibex were introduced from Turkey and Sinai to help promote the ibex population there, which caused hybrids that produced offspring too early, which caused the overall population to disappear completely. The genes of each population of the ibex in Turkey and Sinai were locally adapted to their environments so when placed in a new environmental context did not flourish. Additionally, the environmental toll that may arise from the introduction of a new species may be so disruptive that the ecosystem is no longer able to sustain certain populations.

Controversy

Environmentalist perspectives

The use of the word “pollution” in the term genetic pollution has a deliberate negative connotation and is meant to convey the idea that mixing genetic information is bad for the environment. However, because the mixing of genetic information can lead to a variety of outcomes, “pollution” may not be the most accurate descriptor. Gene flow is undesirable according to some environmentalists and conservationists, including groups such as Greenpeace, TRAFFIC, and GeneWatch UK:
"Invasive species have been a major cause of extinction throughout the world in the past few hundred years. Some of them prey on native wildlife, compete with it for resources, or spread disease, while others may hybridize with native species, causing "genetic pollution". In these ways, invasive species are as big a threat to the balance of nature as the direct overexploitation by humans of some species."
It can also be considered undesirable if it leads to a loss of fitness in the wild populations. The term can be associated with the gene flow from a mutation bred, synthetic organism or genetically engineered organism to a non GE organism, by those who consider such gene flow detrimental. These environmentalist groups stand in complete opposition to the development and production of genetically engineered organisms. 

Seeds of Destruction: Hidden Agenda of Genetic Manipulation is a book by American writer, Frederick William Engdahl. In his book, Engdahl explains how the (oil-rich) Rockefeller family is planing to control world farming by the Green Revolution plan. Rockefeller family is trying to finish other family farms in the United States and also is trying do same as this plan to other countries and finally take the whole world food security on their hands. Engdahl believes that green revolution is happening by the new world empire. He also sees the Rockefeller family hand behind the great Genetically modified organism (GMO) project to control world population.

Governmental definition

From a governmental perspective, genetic pollution is defined as follows by the Food and Agriculture Organization of the United Nations:
"Uncontrolled spread of genetic information (frequently referring to transgenes) into the genomes of organisms in which such genes are not present in nature."

Scientific perspectives

Use of the term 'genetic pollution' and similar phrases such as genetic deterioration, genetic swamping, genetic takeover, and genetic aggression, are being debated by scientists as many do not find it scientifically appropriate. Rhymer and Simberloff argue that these types of terms:
"...imply either that hybrids are less fit than the parentals, which need not be the case, or that there is an inherent value in "pure" gene pools."
They recommend that gene flow from invasive species be termed genetic mixing since:
"Mixing" need not be value-laden, and we use it here to denote mixing of gene pools whether or not associated with a decline in fitness.
Environmentalists such as Patrick Moore, an ex-member and cofounder of Greenpeace, questions if the term genetic pollution is more political than scientific. The term is considered to arouse emotional feelings towards the subject matter. In an interview he comments:
"If you take a term used quite frequently these days, the term "genetic pollution," otherwise referred to as genetic contamination, it is a propaganda term, not a technical or scientific term. Pollution and contamination are both value judgments. By using the word "genetic" it gives the public the impression that they are talking about something scientific or technical--as if there were such a thing as genes that amount to pollution.
Thus, using the term “genetic pollution” is inherently political. A scientific approach to discussing gene flow between introduced and native species would be to use terms like genetic mixing or gene flow. Such mixing can definitely have negative consequences on the fitness of native populations, so it is important not to minimize the risk. However, because genetic mixing can also lead to fitness recovery in cases that could be described as “genetic rescue”, it is important to distinguish that just mixing genes from introduced into native populations can lead to variable outcomes for the fitness of native populations.

Aurochs

From Wikipedia, the free encyclopedia

Aurochs
Temporal range: From early Pleistocene to
1627 (wild form) resp. Present (domestic form)
Aurochse.jpg
Mounted skeleton of a bull found in Vig, Denmark, National Museum of Denmark

Extinct  (1627) (IUCN 3.1)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Bovidae
Subfamily: Bovinae
Genus: Bos
Species:
B. primigenius
Binomial name
Bos primigenius
(Bojanus, 1827)
Subspecies
Wild:
domestic:
Bos primigenius map.jpg
Distribution of the three subspecies

The aurochs (/ˈɔːrɒks/ or /ˈrɒks/; pl. aurochs, or rarely aurochsen, aurochses), also known as urus or ure (Bos primigenius), is an extinct species of large wild cattle that inhabited Europe, Asia, and North Africa. It is the ancestor of domestic cattle; it has also been suggested as an ancestor genetically to the modern European bison, which have been crossbred with steppe bison. The species survived in Europe until 1627, when the last recorded aurochs died in the Jaktorów Forest, Poland.

During the Neolithic Revolution, which occurred during the early Holocene, at least two aurochs domestication events occurred: one related to the Indian subspecies, leading to zebu cattle, and the other one related to the Eurasian subspecies, leading to taurine cattle. Other species of wild bovines were also domesticated, namely the wild water buffalo, gaur, wild yak and banteng. In modern cattle, numerous breeds share characteristics of the aurochs, such as a dark colour in the bulls with a light eel stripe along the back (the cows being lighter), or a typical aurochs-like horn shape.

Taxonomy

Illustration from Sigismund von Herberstein's book published in 1556 captioned : "I am 'urus', tur in Polish, aurox in German (dunces call me bison) lit. (the) ignorant (ones) had given me the name (of) Bison"; Latin original: Urus sum, polonis Tur, germanis Aurox: ignari Bisontis nomen dederant
 
The aurochs was variously classified as Bos primigenius, Bos taurus, or, in old sources, Bos urus. However, in 2003, the International Commission on Zoological Nomenclature "conserved the usage of 17 specific names based on wild species, which are predated by or contemporary with those based on domestic forms", confirming Bos primigenius for the aurochs. Taxonomists who consider domesticated cattle a subspecies of the wild aurochs should use B. primigenius taurus; those who consider domesticated cattle to be a separate species may use the name B. taurus, which the Commission has kept available for that purpose.

Etymology

The words aurochs, urus, and wisent have all been used synonymously in English. But the extinct aurochs/urus is a completely separate species from the still-extant wisent, also known as European bison. The two were often confused, and some 16th-century illustrations of aurochs and wisents have hybrid features. The word urus (/ˈjʊərəs/; plural uri) is a Latin word, but was borrowed into Latin from Germanic (cf. Old English/Old High German ūr, Old Norse úr). In German, OHG ūr was compounded with ohso "ox", giving ūrohso, which became early modern Aurochs. The modern form is Auerochse.

The word aurochs was borrowed from early modern German, replacing archaic urochs, also from an earlier form of German. The word is invariable in number in English, though sometimes a back-formed singular auroch and/or innovated plural aurochses occur. The use in English of the plural form aurochsen is nonstandard, but mentioned in The Cambridge Encyclopedia of the English Language. It is directly parallel to the German plural Ochsen (singular Ochse) and recreates by analogy the same distinction as English ox (singular) and oxen (plural).

Evolution

The Prejlerup-aurochs, a bull at the Zoological Museum in Copenhagen from 7400 BC
 
Life restoration of an aurochs bull found in Braunschweig, Germany
 
Speculative life restoration of the Indian aurochs (B. p. namadicus)
 
During the Pliocene, the colder climate caused an extension of open grassland, which led to the evolution of large grazers, such as wild bovines. Bos acutifrons is an extinct species of cattle that has been suggested as an ancestor for the aurochs.

The oldest aurochs remains have been dated to about 2 million years ago, in India. The Indian subspecies was the first to appear. During the Pleistocene, the species migrated west into the Middle East (western Asia), as well as to the east. They reached Europe about 270,000 years ago. The South Asian domestic cattle, or zebu, descended from Indian aurochs at the edge of the Thar Desert; the zebu is resistant to drought. Domestic yak, gayal, and Bali cattle do not descend from aurochs.

The first complete mitochondrial genome (16,338 base pairs) DNA sequence analysis of Bos primigenius from an archaeologically verified and exceptionally well preserved aurochs bone sample was published in 2010, followed by the publication in 2015 of the complete genome sequence of Bos primigenius using DNA isolated from a 6,750-year-old British aurochs bone. Further studies using the Bos primigenius whole genome sequence have identified candidate microRNA-regulated domestication genes.

Three wild subspecies of aurochs are recognised. Only the Eurasian subspecies survived until recent times.
  • The Eurasian aurochs (B. p. primigenius) once ranged across the steppes and taigas of Europe, Siberia and Central Asia, and East Asia. It is noted as part of the Pleistocene megafauna, and declined in numbers along with other megafauna species by the end of Pleistocene. The Eurasian aurochs were domesticated into modern taurine cattle breeds around the sixth millennium BC in the Middle East, and possibly also at about the same time in the Far East. Aurochs were still widespread in Europe during the time of the Roman Empire, when they were widely popular as a battle beast in Roman arenas. Excessive hunting began and continued until the species was nearly extinct. By the 13th century, aurochs existed only in small numbers in Eastern Europe, and the hunting of aurochs became a privilege of nobles, and later royal households. The aurochs were not saved from extinction, and the last recorded live aurochs, a female, died in 1627 in the Jaktorów Forest, Poland, from natural causes. Aurochs were found to have lived on the island of Sicily, having migrated via a land bridge from Italy. After the disappearance of the land bridge, Sicilian aurochs evolved to be 20% smaller than their mainland relatives due to insular dwarfism. Fossilized specimens were found in Japan, possibly herded with steppe bison.
  • The Indian aurochs (B. p. namadicus) once inhabited India. It was the first subspecies of the aurochs to appear, at 2 million years ago, and from about 9000 years ago, it was domesticated as the zebu. Fossil remains indicate wild Indian aurochs besides domesticated zebu cattle were in Gujarat and the Ganges area until about 4–5000 years ago. Remains from wild aurochs 4400 years old are clearly identified from Karnataka in South India.
  • The North African aurochs (B. p. africanus) once lived in the woodland and shrubland of North Africa. It descended from aurochs populations migrating from the Middle East. The North African aurochs was morphologically very similar to the Eurasian subspecies, so this taxon may exist only in a biogeographic sense. However, evidence indicates it was genetically distinct from the Eurasian subspecies. Depictions show that North African aurochs may have had a light saddle marking on its back. This subspecies may have been extinct before the Middle Ages.

Description

The appearance of the aurochs has been reconstructed from skeletal material, historical descriptions, and contemporaneous depictions, such as cave paintings, engravings, or Sigismund von Herberstein’s illustration. The work by Charles Hamilton Smith is a copy of a painting owned by a merchant in Augsburg, which may date to the 16th century. Scholars have proposed that Smith's illustration was based on a cattle/aurochs hybrid, or an aurochs-like breed. The aurochs was depicted in prehistoric cave paintings and described in Julius Caesar's The Gallic War, Book 6, Ch. 28.

Size

Holocene aurochs bull skull in Berlin
 
The aurochs were one of the largest herbivores in postglacial Europe, comparable to the wisent (European bison). The size of an aurochs appears to have varied by region; in Europe, northern populations were bigger on average than those from the south. For example, during the Holocene, aurochs from Denmark and Germany had an average height at the shoulders of 155–180 cm (61–71 in) in bulls and 135–155 cm (53–61 in) in cows, while aurochs populations in Hungary had bulls reaching 155–160 cm (61–63 in). The body mass of aurochs appears to have shown some variability. Some individuals were comparable in weight to the wisent and the banteng, reaching around 700 kg (1,540 lb), whereas those from the late-middle Pleistocene are estimated to have weighed up to 1,500 kg (3,310 lb), as much as the largest gaur (the largest extant bovid). The sexual dimorphism between bulls and cows was strongly expressed, with the cows being significantly shorter than bulls on average. 

Restoration of the aurochs based on a bull skeleton from Lund and a cow skeleton from Cambridge, with chart of characteristic external features of the aurochs

Horns

Because of the massive horns, the frontal bones of aurochs were elongated and broad. The horns of the aurochs were characteristic in size, curvature, and orientation. They were curved in three directions: upwards and outwards at the base, then swinging forwards and inwards, then inwards and upwards. Aurochs horns could reach 80 cm (31 in) in length and between 10 and 20 cm (3.9 and 7.9 in) in diameter. The horns of bulls were larger, with the curvature more strongly expressed than in cows. The horns grew from the skull at a 60° angle to the muzzle, facing forwards.

Body shape

The proportions and body shape of the aurochs were strikingly different from many modern cattle breeds. For example, the legs were considerably longer and more slender, resulting in a shoulder height that nearly equalled the trunk length. The skull, carrying the large horns, was substantially larger and more elongated than in most cattle breeds. As in other wild bovines, the body shape of the aurochs was athletic, and especially in bulls, showed a strongly expressed neck and shoulder musculature. Therefore, the fore hand was larger than the rear, similar to the wisent, but unlike many domesticated cattle. Even in carrying cows, the udder was small and hardly visible from the side; this feature is equal to that of other wild bovines.

Aurochs in a cave painting in Lascaux, France

Coat colour

The coat colour of the aurochs can be reconstructed by using historical and contemporary depictions. In his letter to Conrad Gesner (1602), Anton Schneeberger describes the aurochs, a description that agrees with cave paintings in Lascaux and Chauvet. Calves were born a chestnut colour. Young bulls changed their coat colour at a few months old to a very deep brown or black, with a white eel stripe running down the spine. Cows retained the reddish-brown colour. Both sexes had a light-coloured muzzle. Some North African engravings show aurochs with a light-coloured "saddle" on the back, but otherwise no evidence of variation in coat colour is seen throughout its range. A passage from Mucante (1596), describing the “wild ox” as gray, but is ambiguous and may refer to the wisent. Egyptian grave paintings show cattle with a reddish-brown coat colour in both sexes, with a light saddle, but the horn shape of these suggest that they may depict domesticated cattle. Remains of aurochs hair were not known until the early 1980s.

Colour of forelocks

Some primitive cattle breeds display similar coat colours to the aurochs, including the black colour in bulls with a light eel stripe, a pale mouth, and similar sexual dimorphism in colour. A feature often attributed to the aurochs is blond forehead hairs. Historical descriptions tell that the aurochs had long and curly forehead hair, but none mentions a certain colour for it. Cis van Vuure (2005) says that, although the colour is present in a variety of primitive cattle breeds, it is probably a discolouration that appeared after domestication. The gene responsible for this feature has not yet been identified. Zebu breeds show lightly coloured inner sides of the legs and belly, caused by the so-called zebu-tipping gene. It has not been tested if this gene is present in remains of the wild form of the zebu, the Indian aurochs.

Behaviour and ecology

A painting by Heinrich Harder showing an aurochs fighting off a wolf pack
 
Like many bovids, aurochs formed herds for at least a part of the year. These probably did not number much more than 30. If aurochs had social behaviour similar to their descendants, social status was gained through displays and fights, in which cows engaged as well as bulls. Indeed, aurochs bulls were reported to often have had severe fights. As in other wild cattle, ungulates that form unisexual herds, considerable sexual dimorphism was expressed. Ungulates that form herds containing animals of both sexes, such as horses, have more weakly developed sexual dimorphism.

During the mating season, which probably took place during the late summer or early autumn, the bulls had severe fights, and evidence from the forest of Jaktorów shows these could lead to death. In autumn, aurochs fed up for the winter and got fatter and shinier than during the rest of the year, according to Schneeberger. Calves were born in spring. According to Schneeberger, the calf stayed at the cow's side until it was strong enough to join and keep up with the herd on the feeding grounds.

Calves were vulnerable to wolves and, to an extent, bears, while healthy adult aurochs probably did not have to fear these predators. In prehistoric Europe, North Africa, and Asia, big cats, such as lions and tigers, and hyenas were additional predators that probably preyed on aurochs.

Historical descriptions, like Caesar's Commentarii de Bello Gallico or Schneeberger, tell that aurochs were swift and fast, and could be very aggressive. According to Schneeberger, aurochs were not concerned when a man approached, but when teased or hunted, an aurochs could get very aggressive and dangerous, and throw the teasing person into the air, as he described in a 1602 letter to Gesner.

Habitat and distribution

Floodplain forests like this one in Germany were the aurochs' last refuge during its final centuries of existence.
 
No consensus exists concerning the habitat of the aurochs. While some authors think that the habitat selection of the aurochs was comparable to the African forest buffalo, others describe the species as inhabiting open grassland and helping maintain open areas by grazing, together with other large herbivores. With its hypsodont jaw, the aurochs was probably a grazer and had a food selection very similar to domesticated cattle. It was not a browser like many deer species, nor a semi-intermediary feeder like the wisent. Comparisons of the isotope levels of Mesolithic aurochs and domestic cattle bones showed that aurochs probably inhabited wetter areas than domestic cattle. Schneeberger describes that during winter, the aurochs ate twigs and acorns in addition to grasses. 

After the beginning of the Common Era, the habitat of aurochs became more fragmented because of the steadily growing human population. During the last centuries of its existence, the aurochs was limited to remote regions, such as floodplain forests or marshes, with no competing domestic herbivores and less hunting pressure.

Relationship with humans

Domestication

The Vig-aurochs, one of two very well-preserved aurochs skeletons found in Denmark. The circles indicate where the animal was wounded by arrows.
 
Genetic analyses show that the Texas Longhorn breed of cattle originated from an Iberian hybrid of domesticated wild aurochs from the Middle East and domesticated Indian aurochs.
 
The aurochs, which ranged throughout much of Eurasia and Northern Africa during the late Pleistocene and early Holocene, is the wild ancestor of modern cattle. Archaeological evidence shows that domestication occurred independently in the Near East and the Indian subcontinent between 10,000 and 8,000 years ago, giving rise to the two major domestic taxa observed today: humpless Bos taurus (taurine) and humped Bos indicus (zebu), respectively. This is confirmed by genetic analyses of matrilineal mitochondrial DNA sequences, which reveal a marked differentiation between modern B. taurus and B. indicus haplotypes, demonstrating their derivation from two geographically and genetically divergent wild populations. A third domestication event possibly occurred from another form of the aurochs in Africa. The sanga cattle, a zebu-like cattle breed with no back hump, is commonly believed to originate from crosses between humped zebus with taurine cattle breeds. However, some archaeological evidence indicates these cattle were domesticated independently in Africa and that bloodlines of taurine and zebu cattle were introduced only within the last few hundreds years.

Domestication of the aurochs began in the southern Caucasus and northern Mesopotamia from about the sixth millennium BC. Genetic evidence suggests that aurochs were independently domesticated in India and possibly also in northern Africa. Domesticated cattle and aurochs are so different in size that they have been regarded as separate species; however, large ancient cattle and aurochs have more similar morphological characteristics, with significant differences only in the horns and some parts of the cranium.

A mitochondrial DNA study suggests that all domesticated taurine cattle originated from about 80 wild female aurochs in the Near East.

Charles Hamilton Smith's copy of a painting possibly dating to the 16th century
 
Comparison of aurochs bones with those of modern cattle has provided many insights about the aurochs. Remains of the beast, from specimens believed to have weighed more than a ton, have been found in Mesolithic sites around Goldcliff, Wales. Though aurochs became extinct in Britain during the Bronze Age, analysis of bones from aurochs that lived about the same time as domesticated cattle traditionally suggested no genetic contribution to modern breeds. More recent work has pointed to substantial aurochs contributions to indigenous British cattle breeds, with the most material found in Kerry cattle.

Indian zebu, although domesticated eight to ten thousand years ago, are related to aurochs that diverged from the Near Eastern ones some 200,000 years ago. African cattle are thought to have descended from aurochs more closely related to the Near Eastern ones. The Near East and African aurochs groups are thought to have split some 25,000 years ago, probably 15,000 years before domestication. The "Turano-Mongolian" type of cattle now found in northern China, Mongolia, Korea, and Japan may represent a fourth domestication event (and a third event among B. taurus–type aurochs). This group may have diverged from the Near East group some 35,000 years ago. Whether these separate genetic populations would have equated to separate subspecies is unclear.

The maximum range of the aurochs was from Europe (excluding Ireland and northern Scandinavia), to northern Africa, the Middle East, India, and Central Asia. Until at least 3,000 years ago, the aurochs was also found in eastern China, where it is recorded at the Dingjiabao Reservoir in Yangyuan County. Most remains in China are known from the area east of 105°E, but the species has also been reported from the eastern margin of the Tibetan plateau, close to the Heihe River.[39] In Japan, excavations in various locations, such as in Iwate and Tochigi prefectures, have found aurochs which may have herded with steppe bison.

Extinction

The ornamented horn of the last aurochs bull that belonged to King Sigismund III of Poland
 
The inscription reads: "The Aurochs – Bos primigenius bojanus, the ancestor of domestic cattle, lived in this forest Jaktorów until the year 1627."
 
By the time of Herodotus (fifth century BC), aurochs had disappeared from southern Greece, but remained common in the area north and east of the Echedorus River close to modern Thessaloniki. The last reports of the species in the southern tip of the Balkans date to the first century BC, when Varro reported that fierce wild oxen lived in Dardania (southern Serbia) and Thrace. By the 13th century AD, the aurochs' range was restricted to Poland, Lithuania, Moldavia, Transylvania, and East Prussia. The right to hunt large animals on any land was restricted first to nobles, and then gradually, to only the royal households. As the population of aurochs declined, hunting ceased, and the royal court used gamekeepers to provide open fields for grazing for the aurochs. The gamekeepers were exempted from local taxes in exchange for their service. Poaching aurochs was punishable by death.
According to a Polish royal survey in 1564, the gamekeepers knew of 38 animals. The last recorded live aurochs, a female, died in 1627 in the Jaktorów Forest, Poland, from natural causes. The causes of extinction were unrestricted hunting, a narrowing of habitat due to the development of farming, and diseases transmitted by domesticated cattle.

Breeding of aurochs-like cattle

While all the wild subspecies are extinct, B. primigenius lives on in domesticated cattle, and attempts are being made to breed similar types suitable for filling the extinct subspecies' role in the wild.

The idea of breeding back the aurochs was first proposed in the 19th century by Feliks Paweł Jarocki. In the 1920s, a first attempt was undertaken by the Heck brothers in Germany with the aim of breeding an effigy (a look-alike) of the aurochs. Starting in the 1990s, grazing and rewilding projects brought new impetus to the idea and new breeding-back efforts came underway, this time with the aim of recreating an animal not only with the looks, but also with the behaviour and the ecological impact of the aurochs, to be able to fill the ecological role of the aurochs. 

Heck cattle: the first attempt to breed a look-alike from modern cattle from the 1920s

Heck cattle

In the early 1920s, two German zoo directors (in Berlin and Munich), the brothers Heinz and Lutz Heck, began a selective breeding program to breed back the aurochs into existence from the descendant domesticated cattle. Their plan was based on the concept that a species is not extinct as long as all its genes are still present in a living population. The result is the breed called Heck cattle. It resembles what is known about the appearance of the aurochs in colour, and in some cases, also horn shape.

Taurus Project

The Arbeitsgemeinschaft Biologischer Umweltschutz, a conservation group in Germany, started to crossbreed Heck cattle with southern-European primitive breeds in 1996, with the goal of increasing the aurochs-likeness of certain Heck cattle herds. These crossbreeds are called Taurus cattle. It is intended to bring in aurochs-like features that are supposedly missing in Heck cattle using Sayaguesa Cattle and Chianina, and to a lesser extent Spanish Fighting Cattle (Lidia). The same breeding program is being carried out in Latvia, in Lille Vildmose National Park in Denmark, and in the Hungarian Hortobágy National Park. The program in Hungary also includes Hungarian Grey cattle and Watusi.

Tauros Programme

First-generation cross bull from Tauros Programme

The Dutch-based Tauros Programme, (initially TaurOs Project) is trying to DNA-sequence breeds of primitive cattle to find gene sequences that match those found in "ancient DNA" from aurochs samples. The modern cattle would be selectively bred to try to produce the aurochs-type genes in a single animal. Starting around 2007, Tauros Programme selected a number of primitive breeds mainly from Iberia and Italy, such as Sayaguesa cattle, Maremmana primitivo, Pajuna cattle, Limia cattle, Maronesa cattle, Tudanca cattle, and others, which already bear considerable resemblance to the aurochs in certain features. Tauros Programme started collaborations with Rewilding Europe and European Wildlife, two European organizations for ecological restoration and rewilding, and now has breeding herds not only in the Netherlands but also in Portugal, Croatia, Romania, and the Czech Republic. Numerous crossbred calves of the first, second, and third offspring generations have already been born. An ecologist working on the Tauros programme has estimated it will take 7 generations for the project to achieve its aims, possibly by 2025.

Uruz Project

A further back-breeding effort, the Uruz project, was started in 2013 by the True Nature Foundation, an organization for ecological restoration and rewilding. It differs from the other projects in that it is planning to make use of genome editing. Its preliminary plans called for the use of Sayaguesa, Maremmana primitive, or Hungarian Grey (Steppe) cattle, and Texas Longhorn with wildtype colour or Barrosã cattle. The finalised plans now call for setting up two breeding lines, Sayaguesa × Maremmana primitiva/Hungarian Steppe cattle and Watusi × Chianina, and later crossing these lines. Two Watusi × Chianina breeding herds have been set up in Boxmeer and Breda in the Netherlands, another herd using Barrosã is being set up in northern Portugal.

Auerrind Project

The newest of the back-breeding efforts, the Auerrindprojekt, was started in 2015 as a conjoint effort of the Experimentalarchäologisches Freilichtlabor Lauresham (Lauresham Experimental-Archaeological Open-air Laboratory, run by Lorsch Abbey), the Förderkreis Große Pflanzenfresser im Kreis Bergstraße e.V. (Promoting Association Megaherbivores in Bergstraße District) and the Landschaftspflegebetrieb Hohmeyer (Landscape Preservation Company Hohmeyer). In accordance with the breeding aims, the Auerrindprojekt has already set up two breeding herds of Watusi × Chianina and one breeding herd of Sayaguesa x Podolian cattle; a second breeding herd of Sayaguesa × Podolian cattle will be started in 2017.

Other projects

Scientists of the Polish Foundation for Recreating the Aurochs (PFOT) in Poland hope to use DNA from bones in museums to recreate the aurochs. They plan to return this animal to the forests of Poland. The project has gained the support of the Polish Ministry of the Environment. They plan research on ancient preserved DNA. Other research projects have extracted "ancient" DNA over the past 20 years and their results have been published in such periodicals as Nature and PNAS. Polish scientists Ryszard Słomski and Jacek A. Modliński believe that modern genetics and biotechnology make it possible to recreate an animal almost identical to the aurochs. They say this research will lead to examining the causes of the extinction of the aurochs, and help prevent a similar occurrence with domesticated cattle.

Breeding back the aurochs

Highland cattle – their genome has been secondarily introgressed by European aurochs.
 
Approaches that aim to breed an aurochs-like phenotype do not equate to an aurochs-like genotype. In 2015, researchers mapped the draft genome of a British aurochs dated to 6,750 years before present. Researchers compared the genome to the genomes of 73 modern cattle populations and found that traditional or landrace cattle breeds of Scottish, Irish, Welsh, and English origin – such as Highland, Dexter, Kerry, Welsh Black, and White Park – carry the ancestry of the sequenced aurochs, but the other populations did not. Another study concluded that because of this genomic introgression of the aurochs into these breeds, if this reflects "the bigger picture across the aurochs/cattle range, perhaps several subpopulations of aurochs are not extinct at all." The study proposed that it will be possible to consider breeding back cattle "that are genetically akin to specific original aurochs populations, through selective cross-breeding of local cattle breeds bearing local aurochs-genome ancestry."

Cultural significance

Cro-Magnon graffito of Bos primigenius in Grotta del Romito, Papasidero, Italy
 
Replica of Chauvet cave art depicting aurochs, woolly rhino, and wild horses
 
The aurochs was an important game animal appearing in both Paleolithic European and Mesopotamian cave paintings, such as those found at Lascaux and Livernon in France. Aurochs existed into the Iron Age in Anatolia and the Near East, where it was worshiped as a sacred animal, the Lunar Bull, associated with the Great Goddess and later with Mithras. In 2012, an archaeological mission of the British Museum, led by Lebanese archaeologist Claude Doumet Serhal, discovered at the site of the old American school in Sidon, Lebanon, the remains of wild animal bones, including those of an aurochs, dating from the late-fourth to early-third millennium. A 1999 archaeological dig in Peterborough, England, uncovered the skull of an aurochs. The front part of the skull had been removed, but the horns remained attached. The supposition is that the killing of the aurochs in this instance was a sacrificial act. 

The violent cup of Vaphio showing aurochs hunting, Greece, (15th century BC)
 
Mural from Çatalhöyük excavated by James Mellaart showing neolithic hunters attacking an aurochs
 
An aurochs bull used in heraldry: Coat of arms of Mecklenburg region, Germany
 
Also during antiquity, the aurochs was regarded as an animal of cultural value. Aurochs are depicted on the Ishtar Gate. In the Peloponnese there is a 15th-century BC depiction on the so-called violent cup of Vaphio, of hunters trying to capture with nets three wild bulls being probably aurochs, in a possibly Cretan date palm stand. The one of the bulls throws one hunter on the ground while attacking the second with its horns. The cup despite the older perception of being Minoan seems to be Mycenaean. Greeks and Paeonians were hunting aurochs (wild oxen/bulls) and used their huge horns as trophies, cups for wine, and offers to the gods and heroes. For example, as mentioned by Samus, Philippus of Thessalonica and Antipater when Philip V of Macedon killed an aurochs on the foothills of mountain Orvilos, he offered the horns which were 105 cm long and the skin to a temple of Hercules. Aurochs horns were often used by Romans as hunting horns. Aurochs were among those wild animals caught for fights (venationes) in arenas. Julius Caesar described aurochs in Gaul:
... those animals which are called uri. These are a little below the elephant in size, and of the appearance, colour, and shape of a bull. Their strength and speed are extraordinary; they spare neither man nor wild beast which they have espied. These the Germans take with much pains in pits and kill them. The young men harden themselves with this exercise, and practice themselves in this sort of hunting, and those who have slain the greatest number of them, having produced the horns in public, to serve as evidence, receive great praise. But not even when taken very young can they be rendered familiar to men and tamed. The size, shape, and appearance of their horns differ much from the horns of our oxen. These they anxiously seek after, and bind at the tips with silver, and use as cups at their most sumptuous entertainments.
— Julius Caesar. Gallic War Commentaries, Chapter 6.28
They survived in the wild in Europe till late in the Roman Empire and were occasionally captured and exhibited live in shows in the colosseum.

The Hebrew Bible contains numerous references to the untameable strength of re'em, translated as "bullock" or "wild-ox" in Jewish translations and translated rather poorly in the King James Version as "unicorn", but recognised from the last century by Hebrew scholars as the aurochs.

When the aurochs became rarer, hunting it became a privilege of the nobility and a sign of a high social status. The "Nibelungenlied" describes Siegfried killing aurochs: "Darnach schlug er schiere einen Wisent und einen Elch, starker Ure viere und einen grimmen Schelch", meaning "After that, he defeated one wisent and one elk, four aurochs, and one Schelch" - the background of the "Schelch" is dubious. Aurochs horns were commonly used as drinking horns by the nobility, which led to the fact that many aurochs horn sheaths are preserved today (albeit often discoloured). The drinking horn at Corpus Christi College, Cambridge, given to the college on its foundation in 1352, probably by the college's founders, the Guilds of Corpus Christi and the Blessed Virgin Mary, is thought to come from an aurochs. A painting by Willem Kalf depicts an aurochs horn. The horns of the last aurochs bulls, which died in 1620, were ornamented with gold and are located at the Livrustkammaren in Stockholm today. 

Schneeberger writes that aurochs were hunted with arrows, nets, and hunting dogs. With immobilised aurochs, a ritual was practised that might be regarded as cruel nowadays: the curly hair on the forehead was cut from the skull of the living animal. Belts were made out of this hair and were believed to increase the fertility of women. When the aurochs was slaughtered, a cross-like bone (os cardis) was extracted from the heart. This bone, which is also present in domesticated cattle, contributed to the mystique of the animal and magical powers have been attributed to it.

A 16th-century illustration by Teodoro Ghisi, believed to show an aurochs, although the horns and muzzle differ from those of an aurochs.
 
In eastern Europe, where it survived until nearly 400 years ago, the aurochs has left traces in fixed expressions. In Russia, a drunken person behaving badly was described as "behaving like an aurochs", whereas in Poland, big, strong people were characterized as being "a bloke like an aurochs".

In Central Europe, the aurochs features in toponyms and heraldic coats of arms. For example, the names Ursenbach and Aurach am Hongar are derived from the aurochs. An aurochs head, the traditional arms of the German region Mecklenburg, figures in the coat of arms of Mecklenburg-Vorpommern. The aurochs (Romanian bour, from Latin būbalus) was also the symbol of Moldavia; nowadays, they can be found in the coat of arms of both Romania and Moldova. An aurochs head is featured on an 1858 series of Moldavian stamps, the so-called Bull's Heads (cap de bour in Romanian), renowned for their rarity and price among collectors. In Romania there are still villages named Boureni, after the Romanian word for the auroch. The horn of the aurochs is a charge of the coat of arms of Tauragė, Lithuania, (the name of Tauragė is a compound of taũras "auroch" and ragas "horn"). It is also present in the emblem of Kaunas, Lithuania, and was part of the emblem of Bukovina during its time as an Austro-Hungarian Kronland. The Swiss Canton of Uri is named after the aurochs; its yellow flag shows a black aurochs head. East Slavic surnames Turenin, Turishchev, Turov, and Turovsky originate from the Slavic name of the species tur. In Slovakia, toponyms such as Turany, Turíčky, Turie, Turie Pole, Turík, Turová (villages), Turiec (river and region), Turská dolina (valley) and others are used. Turopolje, a large lowland floodplain south of the Sava River in Croatia, got its name from the once-abundant aurochs (Croatian: tur). The ancient name of the Estonian town of Rakvere, Tarwanpe or Tarvanpea, probably derives from "Aurochs head" (Tarvan pea) in ancient Estonian

In 2002, a 3.5-m-high and 7.1-m-long statue of an aurochs was erected in Rakvere, Estonia, for the town's 700th birthday. The sculpture, by artist Tauno Kangro, has become a symbol of the town.

Aurochs are frequently mentioned in the A Song of Ice and Fire series of fantasy novels by George R. R. Martin; in particular, roasted aurochs are sometimes served at banquets.

In the 2012 movie Beasts of the Southern Wild, the six-year-old main character imagines aurochs, though the fantasy creatures are portrayed by "costumed" Vietnamese Pot-bellied piglets.

Inequality (mathematics)

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