This grain is grown on more land area than any other commercial food.[citation needed] World trade in wheat is greater than for all other crops combined.[5] Globally, wheat is the leading source of vegetable protein in human food, having a higher protein content than other major cereals, maize (corn) or rice.[6] In terms of total production tonnages used for food, it is currently second to rice as the main human food crop and ahead of maize, after allowing for maize's more extensive use in animal feeds.
Wheat was a key factor enabling the emergence of city-based societies at the start of civilization because it was one of the first crops that could be easily cultivated on a large scale, and had the additional advantage of yielding a harvest that provides long-term storage of food. Wheat contributed to the emergence of city-states in the Asian Fertile Crescent, including the Babylonian and Assyrian empires. Wheat grain is a staple food used to make flour for leavened, flat and steamed breads, biscuits, cookies, cakes, breakfast cereal, pasta, noodles, couscous[7] and for fermentation to make beer,[8] other alcoholic beverages,[9] and biofuel.[10]
There are six wheat classifications: 1) hard red winter, 2) hard red spring, 3) soft red winter, 4) durum (hard), 5) hard white, and 6) soft white wheat.[citation needed] The hard wheats have the most amount of gluten and are used for making bread, rolls and all-purpose flour. The soft wheats are used for making flat bread, cakes, pastries, crackers, muffins, and biscuits. A high percentage of wheat production in the EU is used as animal feed, often surplus to human requirements or low-quality wheat.[11]
Wheat is planted to a limited extent as a forage crop for livestock, although the straw cannot be used as feed.[12] Its straw can be used as a construction material for roofing thatch.[13][14] The whole grain can be milled to leave just the endosperm for white flour. The by-products of this are bran and germ. The whole grain is a concentrated source of vitamins, minerals, and protein, while the refined grain is mostly starch.
Wheat is one of the first cereals known to have been domesticated, and wheat's ability to self-pollinate greatly facilitated the selection of many distinct domesticated varieties. The archaeological record suggests that this first occurred in the regions known as the Fertile Crescent. Recent findings estimate the first domestication of wheat down to a small region of southeastern Turkey,[15] and domesticated Einkorn wheat at Wadi el Jilat in Jordan—has been dated to 7,500-7,300 BCE.[16]
Origin
Cultivation and repeated harvesting and sowing of the grains of wild grasses led to the creation of domestic strains, as mutant forms ('sports') of wheat were preferentially chosen by farmers. In domesticated wheat, grains are larger, and the seeds (inside the spikelets) remain attached to the ear by a toughened rachis during harvesting. In wild strains, a more fragile rachis allows the ear to easily shatter and disperse the spikelets.[17] Selection for these traits 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 also involve the loss of the plant's natural seed dispersal mechanisms, highly domesticated strains of wheat cannot survive in the wild.Cultivation of wheat began to spread beyond the Fertile Crescent after about 8000 BCE. Jared Diamond traces the spread of cultivated emmer wheat starting in the Fertile Crescent sometime before 8800 BCE. Archaeological analysis of wild emmer indicates that it was first cultivated in the southern Levant with finds dating back as far as 9600 BCE.[18][19] Genetic analysis of wild einkorn wheat suggests that it was first grown in the Karacadag Mountains in southeastern Turkey. Dated archeological remains of einkorn wheat in settlement sites near this region, including those at Abu Hureyra in Syria, suggest the domestication of einkorn near the Karacadag Mountain Range.[20] With the anomalous exception of two grains from Iraq ed-Dubb, the earliest carbon-14 date for einkorn wheat remains at Abu Hureyra is 7800 to 7500 years BCE.[21]
Remains of harvested emmer from several sites near the Karacadag Range have been dated to between 8600 (at Cayonu) and 8400 BCE (Abu Hureyra), that is, in the Neolithic period. With the exception of Iraq ed-Dubb, the earliest carbon-14 dated remains of domesticated emmer wheat were found in the earliest levels of Tell Aswad, in the Damascus basin, near Mount Hermon in Syria. These remains were dated by Willem van Zeist and his assistant Johanna Bakker-Heeres to 8800 BCE. They also concluded that the settlers of Tell Aswad did not develop this form of emmer themselves, but brought the domesticated grains with them from an as yet unidentified location elsewhere.[22]
The cultivation of emmer reached Greece, Cyprus and India by 6500 BCE, Egypt shortly after 6000 BCE, and Germany and Spain by 5000 BCE.[23] "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." [24] By 3000 BCE, wheat had reached England and Scandinavia. A millennium later it reached China. The first identifiable bread wheat (Triticum aestivum) with sufficient gluten for yeasted breads has been identified using DNA analysis in samples from a granary dating to approximately 1350 BCE at Assiros in Greek Macedonia.[25]
From Asia, wheat continued to spread throughout Europe. In England, wheat straw (thatch) was used for roofing in the Bronze Age, and was in common use until the late 19th century.[26]
Farming techniques
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. Agricultural cultivation using horse collar leveraged plows (at about 3000 BCE) was one of the first innovations that increased productivity. Much later, 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 long cultivated land, and the use of fertilizers became widespread. Improved agricultural husbandry has more recently included threshing machines and reaping machines (the 'combine harvester'), tractor-drawn cultivators and planters, and better varieties (see Green Revolution and Norin 10 wheat). Great expansion of wheat production occurred as new arable land was farmed in the Americas and Australia in the 19th and 20th centuries.
Genetics
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| Green wheat a month before harvest |
- Einkorn wheat (T. monococcum) is diploid (AA, two complements of seven chromosomes, 2n=14).[1]
- 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 Aegilops searsii or Ae. speltoides. The unknown grass has never been identified among now surviving wild grasses, but the closest living relative is Aegilops speltoides.[citation needed] The hybridization that formed wild emmer (AABB) occurred in the wild, long before domestication,[27] and was driven by natural selection.
- Hexaploid wheats evolved in farmers' fields. Either domesticated emmer or durum wheat hybridized with yet another wild diploid grass (Aegilops tauschii) to make the hexaploid wheats, spelt wheat and bread wheat.[27] These have three sets of paired chromosomes, three times as many as in diploid wheat.
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.[28]
Heterosis, or hybrid vigor (as in the familiar F1 hybrids of maize), occurs in common (hexaploid) wheat, but it is difficult to produce seed of hybrid cultivars on a commercial scale (as is done with maize) because wheat flowers are perfect and normally self-pollinate. Commercial hybrid wheat seed has been produced using chemical hybridizing agents; these chemicals 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 USA and South Africa.[29] F1 hybrid wheat cultivars should not be confused with the standard method of breeding inbred wheat cultivars by crossing two lines using hand emasculation, then selfing or inbreeding the progeny many (ten or more) generations before release selections are identified to be released as a variety or cultivar.
Synthetic hexaploids made by crossing the wild goatgrass wheat ancestor Aegilops tauschii and various durum wheats are now being deployed, and these increase the genetic diversity of cultivated wheats.[30][31][32]
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 valuable carbon isotope based methods that are used for breeding wheat varieties with improved water-use efficiency. These varieties can improve crop productivity in rain-fed dry-land wheat farms.[33]
In 2010, a team of UK scientists funded by BBSRC announced they had decoded the wheat genome for the first time (95% of the genome of a variety of wheat known as Chinese Spring line 42).[34] This genome was released in a basic format for scientists and plant breeders to use but was not a fully annotated sequence which was reported in some of the media.[35]
On 29 November 2012, an essentially complete gene set of bread wheat has been published.[36] Random shotgun libraries of total DNA and cDNA from the T. aestivum cv. Chinese Spring (CS42) were sequenced in Roche 454 pyrosequencer using GS FLX Titanium and GS FLX+ platforms to generate 85 Gb of sequence (220 million reads), equivalent to 5X genome coverage and identified between 94,000 and 96,000 genes.[36]
This sequence data provides direct access to about 96,000 genes, relying on orthologous gene sets from other cereals. and represents an essential step towards a systematic understanding of biology and engineering the cereal crop for valuable traits. Its implications in cereal genetics and breeding includes the examination of genome variation, association mapping using natural populations, performing wide crosses and alien introgression, studying the expression and nucleotide polymorphism in transcriptomes, analyzing population genetics and evolutionary biology, and studying the epigenetic modifications. Moreover, the availability of large-scale genetic markers generated through NGS technology will facilitate trait mapping and make marker-assisted breeding much feasible.[37]
Moreover, the data not only facilitate in deciphering the complex phenomena such as heterosis and epigenetics, it may also enable breeders to predict which fragment of a chromosome is derived from which parent in the progeny line, thereby recognizing crossover events occurring in every progeny line and inserting markers on genetic and physical maps without ambiguity. In due course, this will assist in introducing specific chromosomal segments from one cultivar to another. Besides, the researchers had identified diverse classes of genes participating in energy production, metabolism and growth that were probably linked with crop yield, which can now be utilized for the development of transgenic wheat. Thus whole genome sequence of wheat and the availability of thousands of SNPs will inevitably permit the breeders to stride towards identifying novel traits, providing biological knowledge and empowering biodiversity-based breeding.[37]
Plant breeding
In traditional agricultural systems wheat populations often consist of landraces, informal farmer-maintained populations that often maintain high levels of morphological diversity. Although landraces of wheat are no longer 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.[38]The major breeding objectives include high grain yield, good quality, disease and insect resistance and tolerance to abiotic stresses, including mineral, moisture and heat tolerance. The major diseases in temperate environments include the following, arranged in a rough order of their significance from cooler to warmer climates: eyespot, Stagonospora nodorum blotch (also known as glume blotch), yellow or stripe rust, powdery mildew, Septoria tritici blotch (sometimes known as leaf blotch), brown or leaf rust, Fusarium head blight, tan spot and stem rust. In tropical areas, spot blotch (also known as Helminthosporium leaf blight) is also important.
Wheat has also been the subject of mutation breeding, with the use of gamma, x-rays, ultraviolet light, and sometimes harsh chemicals. The varieties of wheat created through this methods are in the hundreds (varieties being as far back as 1960), more of them being created in higher populated countries such as China.[39]
Hybrid wheat
Because wheat self-pollinates, creating hybrid varieties is extremely labor-intensive; the high cost of hybrid wheat seed relative to its moderate benefits have kept farmers from adopting them widely[40][41] despite nearly 90 years of effort.[42] F1 hybrid wheat cultivars should not be confused with wheat cultivars deriving from standard plant breeding. Heterosis or hybrid vigor (as in the familiar F1 hybrids of maize) occurs in common (hexaploid) wheat, but it is difficult to produce seed of hybrid cultivars on a commercial scale as is done with maize because wheat flowers are perfect in the botanical sense, meaning they have both male and female parts, and normally self-pollinate.[38] 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.[43]Hulled versus free-threshing wheat
The four wild species of wheat, along with the domesticated varieties einkorn,[44] emmer[45] and spelt,[46] 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. In contrast, 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. Hulled wheats are often stored as spikelets because the toughened glumes give good protection against pests of stored grain.[44]Naming
There are many botanical classification systems used for wheat species, discussed in a separate article on Wheat taxonomy. The name of a wheat species from one information source may not be the name of a wheat species in another.Within a species, wheat cultivars are further classified by wheat breeders and farmers in terms of:
- Growing season, such as winter wheat vs. spring wheat.[14]
- Protein content. Bread wheat protein content ranges from 10% in some soft wheats with high starch contents, to 15% in hard wheats.
- The quality of the wheat protein gluten. This protein can determine the suitability of a wheat to a particular dish. A strong and elastic gluten present in bread wheats enables dough to trap carbon dioxide during leavening, but elastic gluten interferes with the rolling of pasta into thin sheets. The gluten protein in durum wheats used for pasta is strong but not elastic.
- Grain color (red, white or amber). Many wheat varieties are reddish-brown due to phenolic compounds present in the bran layer which are transformed to pigments by browning enzymes. White wheats have a lower content of phenolics and browning enzymes, and are generally less astringent in taste than red wheats. The yellowish color of durum wheat and semolina flour made from it is due to a carotenoid pigment called lutein, which can be oxidized to a colorless form by enzymes present in the grain.
Major cultivated species of wheat[citation needed]
Hexaploid Species- Common wheat or Bread wheat (T. aestivum) – A hexaploid species that is the most widely cultivated in the world.
- Spelt (T. spelta) – Another hexaploid species cultivated in limited quantities. Spelt is sometimes considered a subspecies of the closely related species common wheat (T. aestivum), in which case its botanical name is considered to be Triticum aestivum subsp. spelta.
- Durum (T. durum) – The only tetraploid form of wheat widely used today, and the second most widely cultivated wheat.
- Emmer (T. dicoccon) – A tetraploid species, cultivated in ancient times but no longer in widespread use.
- Khorasan (Triticum turgidum ssp. turanicum also called Triticum turanicum) is a tetraploid wheat species.[2] It is an ancient grain type; Khorasan refers to a historical region in modern-day Afghanistan and the northeast of Iran. This grain is twice the size of modern-day wheat and is known for its rich nutty flavor.[3]
- Einkorn (T. monococcum) – A diploid species with wild and cultivated variants. Domesticated at the same time as emmer wheat, but never reached the same importance.
- Durum – Very hard, translucent, light-colored grain used to make semolina flour for pasta & bulghur; high in protein, specifically, gluten protein.
- Hard Red Spring – Hard, brownish, high-protein wheat used for bread and hard baked goods. Bread Flour and high-gluten flours are commonly made from hard red spring wheat. It is primarily traded at the Minneapolis Grain Exchange.
- Hard Red Winter – Hard, brownish, mellow high-protein wheat used for bread, hard baked goods and as an adjunct in other flours to increase protein in pastry flour for pie crusts. Some brands of unbleached all-purpose flours are commonly made from hard red winter wheat alone. It is primarily traded on the Kansas City Board of Trade. One variety is known as "turkey red wheat", and was brought to Kansas by Mennonite immigrants from Russia.[47]
- Soft Red Winter – Soft, low-protein wheat used for cakes, pie crusts, biscuits, and muffins. Cake flour, pastry flour, and some self-rising flours with baking powder and salt added, for example, are made from soft red winter wheat. It is primarily traded on the Chicago Board of Trade.
- Hard White – Hard, light-colored, opaque, chalky, medium-protein wheat planted in dry, temperate areas. Used for bread and brewing.
- Soft White – Soft, light-colored, very low protein wheat grown in temperate moist areas. Used for pie crusts and pastry. Pastry flour, for example, is sometimes made from soft white winter wheat.
As a food
Raw wheat 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, crushed and de-branned into bulgur also known as groats. If the raw wheat is broken into parts at the mill, as is usually done, the outer husk or bran can be used several ways. Wheat is a major ingredient in such foods as bread, porridge, crackers, biscuits, Muesli, pancakes, pies, pastries, cakes, cookies, muffins, rolls, doughnuts, gravy, boza (a fermented beverage), and breakfast cereals (e.g., Wheatena, Cream of Wheat, Shredded Wheat, and Wheaties).
Commercial use
Harvested wheat grain that enters trade is classified according to grain properties for the purposes of the commodity markets. Wheat buyers use these to decide which wheat to buy, as each class has special uses, and producers use them to decide which classes of wheat will be most profitable to cultivate.Wheat is widely cultivated as a cash crop because it produces a good yield per unit area, grows well in a temperate climate even with a moderately short growing season, and yields a versatile, high-quality flour that is widely used in baking. Most breads are made with wheat flour, including many breads named for the other grains they contain, for
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