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Wheat is a group of wild and domesticated grasses of the genus Triticum (Template:IPAc-en).<ref>Template:Cite Merriam-Webster</ref> They are cultivated for their cereal grains, which are staple foods around the world. Well-known wheat species and hybrids include the most widely grown common wheat (T. aestivum), spelt, durum, emmer, einkorn, and Khorasan or Kamut. The archaeological record suggests that wheat was first cultivated in the regions of the Fertile Crescent around 9600 BC.

Wheat is grown on a larger area of land than any other food crop (Template:Convert in 2021). World trade in wheat is greater than that of all other crops combined. In 2021, world wheat production was Template:Convert, making it the second most-produced cereal after maize (known as corn in North America and Australia; wheat is often called corn in countries including Britain).<ref name="Mencken-1984">Template:Cite book</ref> Since 1960, world production of wheat and other grain crops has tripled and is expected to grow further through the middle of the 21st century. Global demand for wheat is increasing because of the usefulness of gluten to the food industry.

Wheat is an important source of carbohydrates. Globally, it is the leading source of vegetable proteins in human food, having a protein content of about 13%, which is relatively high compared to other major cereals but relatively low in protein quality (supplying essential amino acids). When eaten as the whole grain, wheat is a source of multiple nutrients and dietary fibre. In a small part of the general population, gluten – which comprises most of the protein in wheat – can trigger coeliac disease, noncoeliac gluten sensitivity, gluten ataxia, and dermatitis herpetiformis.

DescriptionEdit

File:Triticum aestivum - Köhler–s Medizinal-Pflanzen-274.jpg
A: Plant; B ripe ear of corn; 1 spikelet before flowering; 2 the same, flowering and spread, enlarged; 3 flowers with glumes; 4 stamens 5 pollen; 6 and 7 ovaries with juice scales; 8 and 9 parts of the scar; 10 fruit husks; 11, 12, 13 seeds, natural size and enlarged; 14 the same cut up, enlarged.

Wheat is a stout grass of medium to tall height. Its stem is jointed and usually hollow, forming a straw. There can be many stems on one plant. It has long narrow leaves, their bases sheathing the stem, one above each joint. At the top of the stem is the flower head, containing some 20 to 100 flowers. Each flower contains both male and female parts.<ref name="Brittanica"/> The flowers are wind-pollinated, with over 99% of pollination events being self-pollinations and the rest cross-pollinations.<ref>Template:Cite journal</ref> The flower is housed in a pair of small leaflike glumes. The two (male) stamens and (female) stigmas protrude outside the glumes. The flowers are grouped into spikelets, each with between two and six flowers. Each fertilised carpel develops into a wheat grain or berry; botanically a caryopsis fruit, it is often called a seed. The grains ripen to a golden yellow; a head of grain is called an ear.<ref name="Brittanica">Template:Cite encyclopedia</ref>

Leaves emerge from the shoot apical meristem in a telescoping fashion until the transition to reproduction i.e. flowering.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The last leaf produced by a wheat plant is known as the flag leaf. It is denser and has a higher photosynthetic rate than other leaves, to supply carbohydrate to the developing ear. In temperate countries the flag leaf, along with the second and third highest leaf on the plant, supply the majority of carbohydrate in the grain and their condition is paramount to yield formation.<ref name="Pajević-1999">Template:Cite journal</ref><ref name="Araus-1986">Template:Cite book</ref> Wheat is unusual among plants in having more stomata on the upper (adaxial) side of the leaf, than on the under (abaxial) side.<ref>Template:Cite journal</ref> It has been theorised that this might be an effect of it having been domesticated and cultivated longer than any other plant.<ref>Template:Cite journal</ref> Winter wheat generally produces up to 15 leaves per shoot and spring wheat up to 9<ref name="AHDB">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> and winter crops may have up to 35 tillers (shoots) per plant (depending on cultivar).<ref name="AHDB"/>

Wheat roots are among the deepest of arable crops, extending as far down as Template:Convert.<ref>Template:Cite book</ref> While the roots of a wheat plant are growing, the plant also accumulates an energy store in its stem, in the form of fructans,<ref>Template:Cite journal</ref> which helps the plant to yield under drought and disease pressure,<ref>Template:Cite journal</ref> but it has been observed that there is a trade-off between root growth and stem non-structural carbohydrate reserves. Root growth is likely to be prioritised in drought-adapted crops, while stem non-structural carbohydrate is prioritised in varieties developed for countries where disease is a bigger issue.<ref>Template:Cite journal</ref>

Depending on variety, wheat may be awned or not awned. Producing awns incurs a cost in grain number,<ref>Template:Cite journal</ref> but wheat awns photosynthesise more efficiently than their leaves with regards to water usage,<ref>Template:Cite journal</ref> so awns are much more frequent in varieties of wheat grown in hot drought-prone countries than those generally seen in temperate countries. For this reason, awned varieties could become more widely grown due to climate change. In Europe, however, a decline in climate resilience of wheat has been observed.<ref>Template:Cite journal</ref>

HistoryEdit

File:Wheatareal.PNG
Origin and 21st century production areas of wheat

DomesticationEdit

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Hunter-gatherers in West Asia harvested wild wheats for thousands of years before they were domesticated,<ref>Template:Cite journal</ref> perhaps as early as 21,000 BC,<ref>Template:Cite journal</ref> but they formed a minor component of their diets.<ref name="Arranz-Otaegui-2018">Template:Cite journal</ref> In this phase of pre-domestication cultivation, early cultivars were spread around the region and slowly developed the traits that came to characterise their domesticated forms.<ref>Template:Cite journal</ref>

Repeated harvesting and sowing of the grains of wild grasses led to the creation of domestic strains, as mutant forms ('sports') of wheat were more amenable to cultivation. In domesticated wheat, grains are larger, and the seeds (inside the spikelets) remain attached to the ear by a toughened rachis during harvesting.<ref>Template:Cite journal</ref> In wild strains, a more fragile rachis allows the ear to shatter easily, dispersing the spikelets.<ref>Template:Cite journal</ref> Selection for larger grains and non-shattering heads by farmers might not have been deliberately intended, but simply have occurred because these traits made gathering the seeds easier; nevertheless such 'incidental' selection was an important part of crop domestication. As the traits that improve wheat as a food source involve the loss of the plant's natural seed dispersal mechanisms, highly domesticated strains of wheat cannot survive in the wild.<ref name="Purugganan-2009">Template:Cite journal</ref>

Wild einkorn wheat (T. monococcum subsp. boeoticum) grows across Southwest Asia in open parkland and steppe environments.<ref name="Zohary-2012">Template:Cite book</ref> It comprises three distinct races, only one of which, native to Southeast Anatolia, was domesticated.<ref name="Ozkan-2002">Template:Cite journal</ref> The main feature that distinguishes domestic einkorn from wild is that its ears do not shatter without pressure, making it dependent on humans for dispersal and reproduction.<ref name="Zohary-2012"/> It also tends to have wider grains.<ref name="Zohary-2012"/> Wild einkorn was collected at sites such as Tell Abu Hureyra (Template:Circa) and Mureybet (Template:Circa), but the earliest archaeological evidence for the domestic form comes after Template:Circa in southern Turkey, at Çayönü, Cafer Höyük, and possibly Nevalı Çori.<ref name="Zohary-2012"/> Genetic evidence indicates that it was domesticated in multiple places independently.<ref name="Ozkan-2002"/>

Wild emmer wheat (T. turgidum subsp. dicoccoides) is less widespread than einkorn, favouring the rocky basaltic and limestone soils found in the hilly flanks of the Fertile Crescent.<ref name="Zohary-2012"/> It is more diverse, with domesticated varieties falling into two major groups: hulled or non-shattering, in which threshing separates the whole spikelet; and free-threshing, where the individual grains are separated. Both varieties probably existed in prehistory, but over time free-threshing cultivars became more common.<ref name="Zohary-2012"/> Wild emmer was first cultivated in the southern Levant, as early as 9600 BC.<ref>Template:Cite journal</ref><ref name="Colledge-2007">Template:Cite book</ref> Genetic studies have found that, like einkorn, it was domesticated in southeastern Anatolia, but only once.<ref name="Ozkan-2002"/><ref>Template:Cite journal</ref> The earliest secure archaeological evidence for domestic emmer comes from Çayönü, Template:Circa, where distinctive scars on the spikelets indicated that they came from a hulled domestic variety.<ref name="Zohary-2012"/> Slightly earlier finds have been reported from Tell Aswad in Syria, Template:Circa, but these were identified using a less reliable method based on grain size.<ref name="Zohary-2012"/>

Early farmingEdit

File:NHM - Jungsteinzeit Sichel 2.jpg
Sickles with stone microblades were used to harvest wheat in the Neolithic period, Template:Circa

Einkorn and emmer are considered two of the founder crops cultivated by the first farming societies in Neolithic West Asia.<ref name="Zohary-2012" /> These communities also cultivated naked wheats (T. aestivum and T. durum) and a now-extinct domesticated form of Zanduri wheat (T. timopheevii),<ref>Template:Cite journal</ref> as well as a wide variety of other cereal and non-cereal crops.<ref name="Arranz-Otaegui-2023">Template:Cite journal</ref> Wheat was relatively uncommon for the first thousand years of the Neolithic (when barley predominated), but became a staple after around 8500 BC.<ref name="Arranz-Otaegui-2023"/> Early wheat cultivation did not demand much labour. Initially, farmers took advantage of wheat's ability to establish itself in annual grasslands by enclosing fields against grazing animals and re-sowing stands after they had been harvested, without the need to systematically remove vegetation or till the soil.<ref>Template:Cite journal</ref> They may also have exploited natural wetlands and floodplains to practice décrue farming, sowing seeds in the soil left behind by receding floodwater.<ref>Template:Cite journal</ref><ref>Template:Cite book</ref><ref>Template:Cite book</ref> It was harvested with stone-bladed sickles.<ref>Template:Cite journal</ref> The ease of storing wheat and other cereals led farming households to become gradually more reliant on it over time, especially after they developed individual storage facilities that were large enough to hold more than a year's supply.<ref name="Weide-2021">Template:Cite journal</ref>

Wheat grain was stored after threshing, with the chaff removed.<ref name="Weide-2021"/> It was then processed into flour using ground stone mortars.<ref>Template:Cite journal</ref> Bread made from ground einkorn and the tubers of a form of club rush (Bolboschoenus glaucus) was made as early as 12,400 BC.<ref>Template:Cite journal</ref> At Çatalhöyük (Template:Circa), both wholegrain wheat and flour was used to prepare bread, porridge and gruel.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> Apart from food, wheat may also have been important to Neolithic societies as a source of straw, which could be used for fuel, wicker-making, or wattle and daub construction.<ref>Template:Cite book</ref>

SpreadEdit

Domestic wheat was quickly spread to regions where its wild ancestors did not grow naturally. Emmer was introduced to Cyprus as early as 8600 BC and einkorn Template:Circa;<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> emmer reached Greece by 6500 BC, Egypt shortly after 6000 BC, and Germany and Spain by 5000 BC.<ref>Template:Cite book</ref> "The early Egyptians were developers of bread and the use of the oven and developed baking into one of the first large-scale food production industries."<ref>Template:Cite encyclopedia</ref> By 4000 BC, wheat had reached the British Isles and Scandinavia.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name="Smith-2015">Template:Cite journal</ref><ref name="Brace-2019">Template:Cite journal</ref> Wheat was also cultivated in India around 3500 BC.<ref>Template:Cite journal</ref> Wheat likely appeared in China's lower Yellow River around 2600 BC.<ref>Template:Cite journal</ref>

The oldest evidence for hexaploid wheat has been confirmed through DNA analysis of wheat seeds, dating to around 6400–6200 BC, recovered from Çatalhöyük.<ref name="Bilgic-2016">Template:Cite journal</ref> Template:As of the earliest known wheat with sufficient gluten for yeasted breads was found in a granary at Assiros in Macedonia dated to 1350 BC.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> From the Middle East, wheat continued to spread across Europe and to the Americas in the Columbian exchange. In the British Isles, wheat straw (thatch) was used for roofing in the Bronze Age, and remained in common use until the late 19th century.<ref>Template:Cite book</ref><ref>Template:Cite book</ref> White wheat bread was historically a high status food, but during the nineteenth century it became in Britain an item of mass consumption, displacing oats, barley and rye from diets in the North of the country. It became "a sign of a high degree of culture".<ref>Template:Cite book</ref> After 1860, the enormous expansion of wheat production in the United States flooded the world market, lowering prices by 40%, and (along with the expansion of potato growing) made a major contribution to the nutritional welfare of the poor.<ref>Template:Cite book</ref>

EvolutionEdit

PhylogenyEdit

File:Polyploid wheat origins.svg
Wheat origins by repeated hybridization and polyploidy.<ref name="Golovnina-2007"/> Not all species are shown.

Some wheat species are diploid, with two sets of chromosomes, but many are stable polyploids, with four sets of chromosomes (tetraploid) or six (hexaploid).<ref name="Golovnina-2007"/> Einkorn wheat (Triticum monococcum) is diploid (AA, two complements of seven chromosomes, 2n=14).<ref name="Belderok-2000"/> Most tetraploid wheats (e.g. emmer and durum wheat) are derived from wild emmer, T. dicoccoides. Wild emmer is itself the result of a hybridization between two diploid wild grasses, T. urartu and a wild goatgrass such as Ae. speltoides.<ref>Template:Cite journal</ref> The hybridization that formed wild emmer (AABB, four complements of seven chromosomes in two groups, 4n=28) occurred in the wild, long before domestication, and was driven by natural selection. Hexaploid wheats evolved in farmers' fields as wild emmer hybridized with another goatgrass, Ae. squarrosa or Ae. tauschii, to make the hexaploid wheats including bread wheat.<ref name="Golovnina-2007"/><ref name="Dvorak-2012">Template:Cite journal</ref>

A 2007 molecular phylogeny of the wheats gives the following not fully-resolved cladogram of major cultivated species; the large amount of hybridisation makes resolution difficult. Markings like "6N" indicate the degree of polyploidy of each species:<ref name="Golovnina-2007">Template:Cite journal</ref>

Template:Clade

TaxonomyEdit

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During 10,000 years of cultivation, numerous forms of wheat, many of them hybrids, have developed under a combination of artificial and natural selection. This complexity and diversity of status has led to much confusion in the naming of wheats.<ref>Template:Cite journal</ref><ref>Template:Citation</ref>

Major speciesEdit

Hexaploid species (6N)

  • Common wheat or bread wheat (T. aestivum) – The most widely cultivated species in the world.<ref name="Yang-2022">Template:Cite journal</ref>
  • Spelt (T. spelta) – Another species largely replaced by bread wheat, but in the 21st century grown, often organically, for artisanal bread and pasta.<ref>Template:Cite news</ref>

Tetraploid species (4N)

  • Durum (T. durum) – A wheat widely used today, and the second most widely cultivated wheat.<ref name="Yang-2022" />
  • Emmer (T. turgidum subsp. dicoccum and T. t. conv. durum) – A species cultivated in ancient times, derived from wild emmer, T. dicoccoides, but no longer in widespread use.<ref name="USDA_ARS">Template:GRIN</ref>
  • Khorasan or Kamut (T. turgidum ssp. turanicum, also called T. turanicum) is an ancient grain type; Khorasan is a historical region in modern-day Afghanistan and the northeast of Iran. The grain is twice the size of modern wheat and has a rich nutty flavor.<ref name="Khlestkina-2006">Template:Cite journal</ref>

Diploid species (2N)

  • Einkorn (T. monococcum). Domesticated from wild einkorn, T. boeoticum, at the same time as emmer wheat.<ref>Template:Cite book</ref>

Hulled versus free-threshing speciesEdit

File:Naked and hulled wheat.jpg
Hulled wheat and einkorn. Note how the einkorn ear breaks down into intact spikelets.

The wild species of wheat, along with the domesticated varieties einkorn,<ref name="Potts-1996">Template:Cite book</ref> emmer<ref>Template:Cite book</ref> and spelt,<ref>Template:Cite book</ref> have hulls. This more primitive morphology (in evolutionary terms) consists of toughened glumes that tightly enclose the grains, and (in domesticated wheats) a semi-brittle rachis that breaks easily on threshing. The result is that when threshed, the wheat ear breaks up into spikelets. To obtain the grain, further processing, such as milling or pounding, is needed to remove the hulls or husks. Hulled wheats are often stored as spikelets because the toughened glumes give good protection against pests of stored grain.<ref name="Potts-1996"/> In free-threshing (or naked) forms, such as durum wheat and common wheat, the glumes are fragile and the rachis tough. On threshing, the chaff breaks up, releasing the grains.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

As a foodEdit

Grain classesEdit

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Classification of wheat greatly varies by the producing country.Template:Sfn

Argentina's grain classes were formerly related to the production region or port of shipment: Rosafe (grown in Santa Fe province, shipped through Rosario), Bahia Blanca (grown in Buenos Aires and La Pampa provinces and shipped through Bahia Blanca), Buenos Aires (shipped through the port of Buenos Aires). While mostly similar to the US Hard Red Spring wheat, the classification caused inconsistencies, so Argentina introduced three new classes of wheat, with all names using a prefix Trigo Dura Argentina (TDA) and a number.Template:Sfn

The grain classification in Australia is within the purview of its National Pool Classification Panel. Australia chose to measure the protein content at 11% moisture basis.Template:Sfn

The decisions on the wheat classification in Canada are coordinated by the Variety Registration Office of the Canadian Food Inspection Agency. Like in the US system, the eight classes in Western Canada and six classes in Eastern Canada are based on colour, season, and hardness. Canada has a unique requirement that the varieties of wheat grains should allow for purely visual identification.Template:Sfn

The wheat grain classes used in the United States are named by colour, season, and hardness:<ref name="Bridgwater-1966">Template:Cite encyclopedia</ref><ref name="NYTimes-1981">Template:Cite news</ref><ref name="USDA">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Food value and usesEdit

File:USDA wheat.jpg
Wheat is used in a wide variety of foods.

Template:Nutritionalvalue

Wheat is a staple cereal worldwide.<ref name="Mauseth-2014">Template:Cite book</ref><ref name="Belderok-2000">Template:Cite book</ref> Raw wheat berries can be ground into flour or, using hard durum wheat only, can be ground into semolina; germinated and dried creating malt; crushed or cut into cracked wheat; parboiled (or steamed), dried, and de-branned into groats, then crushed into bulgur.<ref>Template:Cite book</ref> If the raw wheat is broken into parts at the mill, as is usually done, the outer husk or bran can be used in several ways. Wheat is a major ingredient in baked foods, such as bread, rolls, crackers, biscuits, pancakes, pasta, pies, pastries, pizza, cakes, cookies, and muffins; in fried foods, such as doughnuts; in breakfast cereals, gravy, porridge, and muesli; in semolina; and in drinks such as beer, vodka, and boza (a fermented beverage).<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In manufacturing wheat products, gluten is valuable to impart viscoelastic functional qualities in dough,<ref name="Shewry-2002">Template:Cite journal</ref> enabling the preparation of diverse processed foods such as breads, noodles, and pasta that facilitate wheat consumption.<ref name="EUFIC-2009" /><ref name="Shewry-2015" />

NutritionEdit

Raw red winter wheat is 13% water, 71% carbohydrates including 12% dietary fiber, 13% protein, and 2% fat (table). Some 75–80% of the protein content is as gluten.<ref name="Shewry-2002"/> In a reference amount of Template:Convert, wheat provides Template:Convert of food energy and is a rich source (20% or more of the Daily Value, DV) of multiple dietary minerals, such as manganese, phosphorus, magnesium, zinc, and iron (table). The B vitamins, niacin (36% DV), thiamine (33% DV), and vitamin B6 (23% DV), are present in significant amounts (table).

Wheat is a significant source of vegetable proteins in human food, having a relatively high protein content compared to other major cereals.<ref name="European Community-2016">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> However, wheat proteins have a low quality for human nutrition, according to the DIAAS protein quality evaluation method.<ref name="FAO-2013">Template:Cite book</ref><ref name="Wolfe-2015">Template:Cite journal</ref> Though they contain adequate amounts of the other essential amino acids, at least for adults, wheat proteins are deficient in the essential amino acid lysine.<ref name="Shewry-2015" /><ref name="Shewry-2022">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Because the proteins present in the wheat endosperm (gluten proteins) are particularly poor in lysine, white flours are more deficient in lysine compared with whole grains.<ref name="Shewry-2015"/> Significant efforts in plant breeding are made to develop lysine-rich wheat varieties, without success, Template:As of.<ref name="Vasal">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Supplementation with proteins from other food sources (mainly legumes) is commonly used to compensate for this deficiency,<ref name="FAO">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> since the limitation of a single essential amino acid causes the others to break down and become excreted, which is especially important during growth.<ref name="Shewry-2015"/>

Health advisoriesEdit

Consumed worldwide by billions of people, wheat is a significant food for human nutrition, particularly in the least developed countries where wheat products are primary foods.<ref name="Shewry-2015">Template:Cite journal</ref><ref name="Shewry-2009">Template:Cite journal</ref> When eaten as the whole grain, wheat supplies multiple nutrients and dietary fiber recommended for children and adults.<ref name="EUFIC-2009">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name="Shewry-2015"/><ref name="USDA-2014">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name="USDA-2016">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In genetically susceptible people, wheat gluten can trigger coeliac disease.<ref name="Shewry-2002"/><ref name="WGOGG-2016"/> Coeliac disease affects about 1% of the general population in developed countries.<ref name="WGOGG-2016">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name="NIDDK-2016">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The only known effective treatment is a strict lifelong gluten-free diet.<ref name="WGOGG-2016"/> While coeliac disease is caused by a reaction to wheat proteins, it is not the same as a wheat allergy.<ref name="WGOGG-2016" /><ref name="NIDDK-2016"/> Other diseases triggered by eating wheat are non-coeliac gluten sensitivity<ref name="NIDDK-2016" /><ref name="Ludvigsson-2012">Template:Cite journal</ref> (estimated to affect 0.5% to 13% of the general population<ref name="Molina-Infante-2015">Template:Cite journal</ref>), gluten ataxia, and dermatitis herpetiformis.<ref name="Ludvigsson-2012" /> Certain short-chain carbohydrates present in wheat, known as FODMAPs (mainly fructose polymers), may be the cause of non-coeliac gluten sensitivity. Template:As of, reviews have concluded that FODMAPs only explain certain gastrointestinal symptoms, such as bloating, but not the extra-digestive symptoms that people with non-coeliac gluten sensitivity may develop.<ref name="Volta-2019">Template:Cite journal</ref><ref name="Verbeke-2018">Template:Cite journal</ref><ref name="Fasano-2015">Template:Cite journal</ref> Other wheat proteins, amylase-trypsin inhibitors, have been identified as the possible activator of the innate immune system in coeliac disease and non-coeliac gluten sensitivity.<ref name="Verbeke-2018" /><ref name="Fasano-2015" /> These proteins are part of the plant's natural defense against insects and may cause intestinal inflammation in humans.<ref name="Verbeke-2018" /><ref name="Barone-2014">Template:Cite journal</ref>

Production and consumptionEdit

GlobalEdit

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Wheat production, 2023
Country Millions of tonnes
Template:CHN 136.6
{{#invoke:flag India}} 110.6
{{#invoke:flag }} 91.5
Template:USA 49.3
Template:AUS 41.2
{{#invoke:flag France}} 35.9
Template:CAN 31.9
World 799
Source: UN Food and Agriculture Organization<ref>{{#invoke:citation/CS1|citation CitationClass=web

}}</ref>

In 2023, world wheat production was 799 million tonnes, led by China, India, and Russia which collectively provided 42.4% of the world total.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Template:As of, the largest exporters were Russia (32 million tonnes), United States (27), Canada (23) and France (20), while the largest importers were Indonesia (11 million tonnes), Egypt (10.4) and Turkey (10.0).<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In 2021, wheat was grown on Template:Convert worldwide, more than any other food crop.<ref name="FAOStat-2023">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> World trade in wheat is greater than for all other crops combined.<ref name="Curtis-2002">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Global demand for wheat is increasing due to the unique viscoelastic and adhesive properties of gluten proteins, which facilitate the production of processed foods, whose consumption is increasing as a result of the worldwide industrialization process and westernization of diets.<ref name="Shewry-2015" /><ref name="Day-2006">Template:Cite journal</ref>

19th centuryEdit

File:Wheat prices in England, OWID.svg
CitationClass=web }}</ref>

Wheat became a central agriculture endeavor in the worldwide British Empire in the 19th century, and remains of great importance in Australia, Canada and India.<ref>Template:Cite book</ref> In Australia, with vast lands and a limited work force, expanded production depended on technological advances, especially regarding irrigation and machinery. By the 1840s there were 900 growers in South Australia. They used "Ridley's Stripper", a reaper-harvester perfected by John Ridley in 1843,<ref>Template:Cite book</ref> to remove the heads of grain. In Canada, modern farm implements made large scale wheat farming possible from the late 1840s. By 1879, Saskatchewan was the center, followed by Alberta, Manitoba and Ontario, as the spread of railway lines allowed easy exports to Britain. By 1910, wheat made up 22% of Canada's exports, rising to 25% in 1930 despite the sharp decline in prices during the worldwide Great Depression.<ref>Template:Cite journal</ref> Efforts to expand wheat production in South Africa, Kenya and India were stymied by low yields and disease. However, by 2000 India had become the second largest producer of wheat in the world.<ref>Template:Cite journal</ref> In the 19th century the American wheat frontier moved rapidly westward. By the 1880s 70% of American exports went to British ports. The first successful grain elevator was built in Buffalo in 1842.<ref>Template:Cite book</ref> The cost of transport fell rapidly. In 1869 it cost 37 cents to transport a bushel of wheat from Chicago to Liverpool. In 1905 it was 10 cents.<ref>Template:Cite book</ref>

Late 20th century yieldsEdit

In the 20th century, global wheat output expanded by about 5-fold, but until about 1955 most of this reflected increases in wheat crop area, with lesser (about 20%) increases in crop yields per unit area. After 1955 however, there was a ten-fold increase in the rate of wheat yield improvement per year, and this became the major factor allowing global wheat production to increase. Thus technological innovation and scientific crop management with synthetic nitrogen fertilizer, irrigation and wheat breeding were the main drivers of wheat output growth in the second half of the century. There were some significant decreases in wheat crop area, for instance in North America.<ref>Template:Cite book</ref> Better seed storage and germination ability (and hence a smaller requirement to retain harvested crop for next year's seed) is another 20th-century technological innovation. In medieval England, farmers saved one-quarter of their wheat harvest as seed for the next crop, leaving only three-quarters for food and feed consumption. By 1999, the global average seed use of wheat was about 6% of output.<ref>Template:Cite book</ref> In the 21st century, rising temperatures associated with global warming are reducing wheat yield in several locations.<ref>Template:Cite journal</ref>

AgronomyEdit

Growing wheat Edit

Wheat is an annual crop. It can be planted in autumn and harvested in early summer as winter wheat in climates that are not too severe, or planted in spring and harvested in autumn as spring wheat. It is normally planted after tilling the soil by ploughing and then harrowing to kill weeds and create an even surface. The seeds are then scattered on the surface, or drilled into the soil in rows. Winter wheat lies dormant during a winter freeze. It needs to develop to a height of 10 to 15 cm before the cold intervenes, so as to be able to survive the winter; it requires a period with the temperature at or near freezing, its dormancy then being broken by the thaw or rise in temperature. Spring wheat does not undergo dormancy. Wheat requires a deep soil, preferably a loam with organic matter, and available minerals including soil nitrogen, phosphorus, and potassium. An acid and peaty soil is not suitable. Wheat needs some 30 to 38 cm of rain in the growing season to form a good crop of grain.<ref name="EOS-2023"/>

The farmer may intervene while the crop is growing to add fertilizer, water by irrigation, or pesticides such as herbicides to kill broad-leaved weeds or insecticides to kill insect pests. The farmer may assess soil minerals, soil water, weed growth, or the arrival of pests to decide timely and cost-effective corrective actions, and crop ripeness and water content to select the right moment to harvest. Harvesting involves reaping, cutting the stems to gather the crop; and threshing, breaking the ears to release the grain; both steps are carried out by a combine harvester. The grain is then dried so that it can be stored safe from mould fungi.<ref name="EOS-2023">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Crop developmentEdit

File:Wheat developmental stages.tif
Wheat developmental stages on the BBCH and Zadok's scales

Wheat normally needs between 110 and 130 days between sowing and harvest, depending upon climate, seed type, and soil conditions. Optimal crop management requires that the farmer have a detailed understanding of each stage of development in the growing plants. In particular, spring fertilizers, herbicides, fungicides, and growth regulators are typically applied only at specific stages of plant development. For example, it is currently recommended that the second application of nitrogen is best done when the ear (not visible at this stage) is about 1 cm in size (Z31 on Zadoks scale). Knowledge of stages is also important to identify periods of higher risk from the climate. Farmers benefit from knowing when the 'flag leaf' (last leaf) appears, as this leaf represents about 75% of photosynthesis reactions during the grain filling period, and so should be preserved from disease or insect attacks to ensure a good yield. Several systems exist to identify crop stages, with the Feekes and Zadoks scales being the most widely used. Each scale is a standard system which describes successive stages reached by the crop during the agricultural season.<ref>Template:Cite book</ref> For example, the stage of pollen formation from the mother cell, and the stages between anthesis and maturity, are susceptible to high temperatures, and this adverse effect is made worse by water stress.<ref>Template:Cite journal</ref>

Farming techniquesEdit

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Technological advances in soil preparation and seed placement at planting time, use of crop rotation and fertilizers to improve plant growth, and advances in harvesting methods have all combined to promote wheat as a viable crop. When the use of seed drills replaced broadcasting sowing of seed in the 18th century, another great increase in productivity occurred. Yields of pure wheat per unit area increased as methods of crop rotation were applied to land that had long been in cultivation, and the use of fertilizers became widespread.<ref>Template:Cite book</ref>

Improved agricultural husbandry has more recently included pervasive automation, starting with the use of threshing machines,<ref>Template:Cite journal</ref> and progressing to large and costly machines like the combine harvester which greatly increased productivity.<ref>Template:Cite book</ref> At the same time, better varieties such as Norin 10 wheat, developed in Japan in the 1930s,<ref>Template:Cite journal</ref> or the dwarf wheat developed by Norman Borlaug in the Green Revolution, greatly increased yields.<ref name="Shindler-2016">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>Template:Cite journal</ref>

In addition to gaps in farming system technology and knowledge, some large wheat grain-producing countries have significant losses after harvest at the farm and because of poor roads, inadequate storage technologies, inefficient supply chains and farmers' inability to bring the produce into retail markets dominated by small shopkeepers. Some 10% of total wheat production is lost at farm level, another 10% is lost because of poor storage and road networks, and additional amounts are lost at the retail level.<ref>Template:Cite journal</ref>

In the Punjab region of the Indian subcontinent, as well as North China, irrigation has been a major contributor to increased grain output. More widely over the last 40 years, a massive increase in fertilizer use together with the increased availability of semi-dwarf varieties in developing countries, has greatly increased yields per hectare.<ref name="Godfray-2010">Template:Cite journal</ref> In developing countries, use of (mainly nitrogenous) fertilizer increased 25-fold in this period. However, farming systems rely on much more than fertilizer and breeding to improve productivity. A good illustration of this is Australian wheat growing in the southern winter cropping zone, where, despite low rainfall (300 mm), wheat cropping is successful even with relatively little use of nitrogenous fertilizer. This is achieved by crop rotation with leguminous pastures. The inclusion of a canola crop in the rotations has boosted wheat yields by a further 25%.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In these low rainfall areas, better use of available soil-water (and better control of soil erosion) is achieved by retaining the stubble after harvesting and by minimizing tillage.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Pests and diseasesEdit

Pests and diseases consume 21.47% of the world's wheat crop annually.<ref name="Savary-2019">Template:Cite journal</ref>

DiseasesEdit

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There are many wheat diseases, mainly caused by fungi, bacteria, and viruses.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Plant breeding to develop new disease-resistant varieties, and sound crop management practices are important for preventing disease. Fungicides, used to prevent the significant crop losses from fungal disease, can be a significant variable cost in wheat production. Estimates of the amount of wheat production lost owing to plant diseases vary between 10 and 25% in Missouri.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> A wide range of organisms infect wheat, of which the most important are viruses and fungi.<ref>Template:Cite encyclopedia</ref>

The main wheat-disease categories are:

A historically significant disease of cereals including wheat, though commoner in rye is ergot; it is unusual among plant diseases in also causing sickness in humans who ate grain contaminated with the fungus involved, Claviceps purpurea.<ref name="Harveson-2017">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Animal pestsEdit

Among insect pests of wheat is the wheat stem sawfly, a chronic pest in the Northern Great Plains of the United States and in the Canadian Prairies.<ref>Template:Cite journal</ref> Wheat is the food plant of the larvae of some Lepidoptera (butterfly and moth) species including the flame, rustic shoulder-knot, setaceous Hebrew character and turnip moth. Early in the season, many species of birds and rodents feed upon wheat crops. These animals can cause significant damage to a crop by digging up and eating newly planted seeds or young plants. They can also damage the crop late in the season by eating the grain from the mature spike. Recent post-harvest losses in cereals amount to billions of dollars per year in the United States alone, and damage to wheat by various borers, beetles and weevils is no exception.<ref>Biological Control of Stored-Product Pests. Biological Control News Volume II, Number 10 October 1995 Template:Webarchive

  • Post-harvest Operations Compendium, FAO.</ref> Rodents can also cause major losses during storage, and in major grain growing regions, field mice numbers can sometimes build up explosively to plague proportions because of the ready availability of food.<ref>CSIRO Rodent Management Research Focus: Mice plagues Template:Webarchive</ref> To reduce the amount of wheat lost to post-harvest pests, Agricultural Research Service scientists have developed an "insect-o-graph", which can detect insects in wheat that are not visible to the naked eye. The device uses electrical signals to detect the insects as the wheat is being milled. The new technology is so precise that it can detect 5–10 infested seeds out of 30,000 good ones.<ref>{{#invoke:citation/CS1|citation

|CitationClass=web }}</ref>

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Breeding objectivesEdit

In traditional agricultural systems, wheat populations consist of landraces, informal farmer-maintained populations that often maintain high levels of morphological diversity. Although landraces of wheat are no longer extensively grown in Europe and North America, they continue to be important elsewhere. The origins of formal wheat breeding lie in the nineteenth century, when single line varieties were created through selection of seed from a single plant noted to have desired properties. Modern wheat breeding developed in the first years of the twentieth century and was closely linked to the development of Mendelian genetics. The standard method of breeding inbred wheat cultivars is by crossing two lines using hand emasculation, then selfing or inbreeding the progeny. Selections are identified (shown to have the genes responsible for the varietal differences) ten or more generations before release as a variety or cultivar.<ref name="Bajaj-1990" />

Major breeding objectives include high grain yield, good quality, disease- and insect resistance and tolerance to abiotic stresses, including mineral, moisture and heat tolerance.<ref name="MVGS_IAEA"/><ref name="Sarkar-2021">Template:Cite journal</ref> Wheat has been the subject of mutation breeding, with the use of gamma-, x-rays, ultraviolet light (collectively, radiation breeding), and sometimes harsh chemicals. The varieties of wheat created through these methods are in the hundreds (going as far back as 1960), more of them being created in higher populated countries such as China.<ref name="MVGS_IAEA">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Bread wheat with high grain iron and zinc content has been developed through gamma radiation breeding,<ref>Template:Cite journal</ref> and through conventional selection breeding.<ref name="MacNeil-2021">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> International wheat breeding is led by the International Maize and Wheat Improvement Center in Mexico. ICARDA is another major public sector international wheat breeder, but it was forced to relocate from Syria to Lebanon in the Syrian Civil War.<ref name="ICARDA">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Pathogens and wheat are in a constant process of coevolution.<ref name="Fabre-2022" /> Spore-producing wheat rusts are substantially adapted towards successful spore propagation, which is essentially to say its R0.<ref name="Fabre-2022"/> These pathogens tend towards high-R0 evolutionary attractors.<ref name="Fabre-2022">Template:Cite journal</ref>

For higher yieldsEdit

File:Long-term wheat yields in Europe, OWID.svg
Breeding has increased yields over time

The presence of certain versions of wheat genes has been important for crop yields. Genes for the 'dwarfing' trait, first used by Japanese wheat breeders to produce Norin 10 short-stalked wheat, have had a huge effect on wheat yields worldwide, and were major factors in the success of the Green Revolution in Mexico and Asia, an initiative led by Norman Borlaug.<ref>Template:Cite journal</ref> Dwarfing genes enable the carbon that is fixed in the plant during photosynthesis to be diverted towards seed production, and they also help prevent the problem of lodging.<ref>Template:Cite journal</ref> "Lodging" occurs when an ear stalk falls over in the wind and rots on the ground, and heavy nitrogenous fertilization of wheat makes the grass grow taller and become more susceptible to this problem.<ref>Template:Cite book</ref> By 1997, 81% of the developing world's wheat area was planted to semi-dwarf wheats, giving both increased yields and better response to nitrogenous fertilizer.<ref>Template:Cite journal</ref>

T. turgidum subsp. polonicum, known for its longer glumes and grains, has been bred into main wheat lines for its grain size effect, and likely has contributed these traits to Triticum petropavlovskyi and the Portuguese landrace group Arrancada.<ref name="Adamski-2021">Template:Cite journal</ref> As with many plants, MADS-box influences flower development, and more specifically, as with other agricultural Poaceae, influences yield. Despite that importance, Template:As of little research has been done into MADS-box and other such spikelet and flower genetics in wheat specifically.<ref name="Adamski-2021" />

The world record wheat yield is about Template:Convert, reached in New Zealand in 2017.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> A project in the UK, led by Rothamsted Research has aimed to raise wheat yields in the country to Template:Convert by 2020, but in 2018 the UK record stood at Template:Convert, and the average yield was just Template:Convert.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

For disease resistanceEdit

File:Stem rust on differential lines wheat.jpg
Different strains have been infected with the stem rust fungus. The strains bred to be resistant have their leaves unaffected or relatively unaffected by the fungus.

Wild grasses in the genus Triticum and related genera, and grasses such as rye have been a source of many disease-resistance traits for cultivated wheat breeding since the 1930s.<ref>Template:Cite journal</ref> Some resistance genes have been identified against Pyrenophora tritici-repentis, especially races 1 and 5, those most problematic in Kazakhstan.<ref name="Dahm-2021">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Wild relative, Aegilops tauschii is the source of several genes effective against TTKSK/Ug99 - Sr33, Sr45, Sr46, and SrTA1662 - of which Sr33 and SrTA1662 are the work of Olson et al., 2013, and Sr45 and Sr46 are also briefly reviewed therein.<ref name="Bohra-2021">Template:Cite journal</ref>

Template:Visible anchor (FHB, Fusarium ear blight) is also an important breeding target. Marker-assisted breeding panels involving kompetitive allele specific PCR can be used. Singh et al. 2019 identify a KASP genetic marker for a pore-forming toxin-like gene providing FHB resistance.<ref name="Kaur-2020">Template:Cite journal</ref>

In 2003 the first resistance genes against fungal diseases in wheat were isolated.<ref name="Feuillet-2003">Template:Cite journal</ref><ref name="Yahiaoui-2004">Template:Cite journal</ref> In 2021, novel resistance genes were identified in wheat against powdery mildew and wheat leaf rust.<ref name="Sánchez-Martín-2021">Template:Cite journal</ref><ref name="Kolodziej-2021">Template:Cite journal</ref> Modified resistance genes have been tested in transgenic wheat and barley plants.<ref name="Koller-2023">Template:Cite journal</ref>

To create hybrid vigorEdit

Because wheat self-pollinates, creating hybrid seed to provide the possible benefits of heterosis, hybrid vigor (as in the familiar F1 hybrids of maize), is extremely labor-intensive; the high cost of hybrid wheat seed relative to its moderate benefits have kept farmers from adopting them widely<ref>Mike Abram for Farmers' Weekly. 17 May 2011. Hybrid wheat to make a return</ref><ref>Bill Spiegel for agriculture.com 11 March 2013 Hybrid wheat's comeback</ref> despite nearly 90 years of effort.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name="Bajaj-1990">Bajaj, Y.P.S. (1990) Wheat. Springer Science+Business Media. pp. 161–163. Template:ISBN.</ref> Commercial hybrid wheat seed has been produced using chemical hybridizing agents, plant growth regulators that selectively interfere with pollen development, or naturally occurring cytoplasmic male sterility systems. Hybrid wheat has been a limited commercial success in Europe (particularly France), the United States and South Africa.<ref>Basra, Amarjit S. (1999) Heterosis and Hybrid Seed Production in Agronomic Crops. Haworth Press. pp. 81–82. Template:ISBN.</ref>

Synthetic hexaploids made by crossing the wild goatgrass wheat ancestor Aegilops tauschii,<ref>Template:Cite journal</ref> and other Aegilops,<ref name="Kishii-2019">Template:Cite journal</ref> and various durum wheats are now being deployed, and these increase the genetic diversity of cultivated wheats.<ref>(12 May 2013) Cambridge-based scientists develop 'superwheat' BBC News UK, Retrieved 25 May 2013</ref><ref>Synthetic hexaploids Template:Webarchive</ref><ref>(2013) Synthetic hexaploid wheat Template:Webarchive UK National Institute of Agricultural Botany, Retrieved 25 May 2013</ref>

For gluten contentEdit

Modern bread wheat varieties have been cross-bred to contain greater amounts of gluten,<ref>Template:Cite journal</ref> which affords significant advantages for improving the quality of breads and pastas from a functional point of view.<ref name="Delcour-2012">Template:Cite journalTemplate:Open access</ref> However, a 2020 study that grew and analyzed 60 wheat cultivars from between 1891 and 2010 found no changes in albumin/globulin and gluten contents over time. "Overall, the harvest year had a more significant effect on protein composition than the cultivar. At the protein level, we found no evidence to support an increased immunostimulatory potential of modern winter wheat."<ref name="Pronin-2020">Template:Cite journal</ref>

For water efficiencyEdit

Stomata (or leaf pores) are involved in both uptake of carbon dioxide gas from the atmosphere and water vapor losses from the leaf due to water transpiration. Basic physiological investigation of these gas exchange processes has yielded carbon isotope based method used for breeding wheat varieties with improved water-use efficiency. These varieties can improve crop productivity in rain-fed dry-land wheat farms.<ref>Template:Cite journal</ref>

For insect resistanceEdit

The complex genome of wheat has made its improvement difficult. Comparison of hexaploid wheat genomes using a range of chromosome pseudomolecule and molecular scaffold assemblies in 2020 has enabled the resistance potential of its genes to be assessed. Findings include the identification of "a detailed multi-genome-derived nucleotide-binding leucine-rich repeat protein repertoire" which contributes to disease resistance, while the gene Sm1 provides a degree of insect resistance,<ref name="Walkowiak-2020">Template:Cite journal</ref> for instance against the orange wheat blossom midge.<ref name="Kassa-2016">Template:Cite journal</ref>

GenomicsEdit

Decoding the genomeEdit

In 2010, 95% of the genome of Chinese Spring line 42 wheat was decoded.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> This genome was released in a basic format for scientists and plant breeders to use but was not fully annotated.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In 2012, an essentially complete gene set of bread wheat was published.<ref name="Hall-2012">Template:Cite journal</ref> Random shotgun libraries of total DNA and cDNA from the T. aestivum cv. Chinese Spring (CS42) were sequenced to generate 85 Gb of sequence (220 million reads) and identified between 94,000 and 96,000 genes.<ref name="Hall-2012"/> In 2018, a more complete Chinese Spring genome was released by a different team.<ref name="UniSaskatchewan-2018">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In 2020, 15 genome sequences from various locations and varieties around the world were reported, with examples of their own use of the sequences to localize particular insect and disease resistance factors.<ref name="UniSaskatchewan-2020">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Template:Visible anchor is controlled by R genes which are highly race-specific.<ref name="Kumar-2020"/>

Genetic engineeringEdit

For decades, the primary genetic modification technique has been non-homologous end joining (NHEJ). However, since its introduction, the [[CRISPR/Cas9|Template:Visible anchor/Template:Visible anchor]] tool has been extensively adopted, for example:

Template:As of these examples illustrate the rapid deployment and results that CRISPR/Cas9 has shown in wheat disease resistance improvement.<ref name="Li-2021" />

In artEdit

The Dutch artist Vincent van Gogh created the series Wheat Fields between 1885 and 1890, consisting of dozens of paintings made mostly in different parts of rural France. They depict wheat crops, sometimes with farm workers, in varied seasons and styles, sometimes green, sometimes at harvest. Wheatfield with Crows was one of his last paintings, and is considered to be among his greatest works.<ref>Template:Cite book</ref><ref>Template:Cite book</ref>

In 1967, the American artist Thomas Hart Benton made his oil on wood painting Wheat, showing a row of uncut wheat plants, occupying almost the whole height of the painting, between rows of freshly-cut stubble. The painting is held by the Smithsonian American Art Museum.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

In 1982, the American conceptual artist Agnes Denes grew a two-acre field of wheat at Battery Park, Manhattan. The ephemeral artwork has been described as an act of protest. The harvested wheat was divided and sent to 28 world cities for an exhibition entitled "The International Art Show for the End of World Hunger".<ref>Template:Cite news</ref>

See alsoEdit

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ReferencesEdit

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SourcesEdit

Further readingEdit

  • The World Wheat Book : A History of Wheat Breeding

External linksEdit

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