Triticeae

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Triticeae
Hordeum jubatum - close-up (aka).jpg
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Clade: Commelinids
Order: Poales
Family: Poaceae
Clade: BOP clade
Subfamily: Pooideae
Supertribe: Triticodae
Tribe: Triticeae
L.
Genera

See text.

Triticeae is a botanical tribe within the subfamily Pooideae of grasses that includes genera with many domesticated species. Major crop genera found in this tribe include wheat (see wheat taxonomy), barley, and rye; crops in other genera include some for human consumption, and others used for animal feed or rangeland protection. Among the world's cultivated species, this tribe has some of the most complex genetic histories. An example is bread wheat, which contains the genomes of three species with only one being a wheat Triticum species. Seed storage proteins in the Triticeae are implicated in various food allergies and intolerances.

Contents

Genera of Triticeae

Genera recognized in Triticeae according to Robert Soreng et al.: [1]

Cultivated or edible species

4 different commercial forms of Triticeae cultivars. Clockwise from top: common wheat flour, European spelt, barley corns, rolled rye Gluten Sources.png
4 different commercial forms of Triticeae cultivars. Clockwise from top: common wheat flour, European spelt, barley corns, rolled rye

Aegilops

Amblyopyrum

Elymus

Various species are cultivated for pastoral purposes or to protect fallow land from opportunistic or invasive species

Hordeum

Many barley cultivars

Leymus

Secale

Ryes

Triticum

(Wheat)

Genetics

Genomes of some Triticeae genera and species
Genera & Species1st2nd3rd
Triticum boeoticum AA
Triticum monococcum AMAM
Triticum urartu AUAU
Aegilops speltoides var. speltoidesBB
Aegilops caudataCC
Aegilops tauschii DD
Lophopyrum elongatumEE
Hordeum vulgare HH
Thinopyrum bessarabicumJJ
Aegilops comosaMM
Aegilops uniaristataNN
Henrardia persicaOO
Agropyrum cristatumPP
Secale cereale RR
Aegilops bicornisSS
Amblyopyrum muticumTT
Aegilops umbellulataUU
DasypyrumVV
PsathyrostachysNsNs
PseudoroegneriaStSt
Triticum zhukovskyiAAAMAMGG
Triticum turgidumAABB
Triticum aestivum AABBDD
Triticum timopheevii AAGG
Aegilops cylindrica CCDD
Stenostachys sp.HHWW
Elmyus canadensisHHStSt
Elmyus aboliniiYYStSt
Thinopyrum Vjd =(V/J/D)JJStStVjdVjd
Leymus tricoidesNsNsXmXm

Triticeae and its sister tribe Bromeae (bromes or cheat grasses) when joined form a sister clade with Poeae and Aveneae (Oats). Inter-generic gene flow characterized these taxa from the early stages. For example, Poeae and Aveneae share a mtDNA genetic marker with barley and 10 other members of Triticeae, whereas all 19 genera of Triticeae bear a wheat marker along with Bromeae. [4]

Genera within Triticeae contain diploid, allotetraploid and/or allohexaploid genomes, the capacity to form allopolyploid genomes varies within the tribe. In this tribe, the majority of diploid species tested are closely related to Aegilops, the more distal members (earliest branch points) include Hordeum (Barley), Eremian, Psathyrostachys. The broad distribution of cultivars within the Tribe and the properties of the proteins have implication in the treatment of certain digestive diseases and autoimmune disorders.[ citation needed ]

Evolution of the tribe

One of the earliest branches in Triticeae, to Pseudoroegeneria, produces the genome StSt and another Hordeum then genome = HH. Allotetraploid combinations of Pseudoroegeneria and Hordeum and are seen in Elmyus (HHStSt), [5] but also shows introgression from Australian and Agropyron wheatgrasses. [6] Elymus contains mostly Pseudoroegeneria mtDNA. [7]

Many genera and species of Triticeae are allopolyploids, having more chromosomes than seen in typical diploids. Typical allopolyploids are tetraploid or hexaploid, XXYY or XXYYZZ. The creation of polyploid species results from natural random events tolerated by polyploid-capable plants. Natural allopolyploid plants may have selective advantages and some may permit the recombination of distantly related genetic material. Poulard wheat is an example of a stable allotetraploid wheat.[ citation needed ]

The Secale (domesticated rye) may be a very early branch from the goat grass clad (or goat grasses are a branch of early rye grasses), as branch these are almost contemporary with the branching between monoploid wheat and Aegilops tauschii. Studies in Anatolia now suggest Rye ( Secale ) was cultivated, but not domesticated, prior to the holocene and to evidence for the cultivation of wheat. As climate changed the favorability of Secale declined. At that time other strains of barley and wheat may have been cultivated, but humans did little to change them.[ citation needed ]

Goat grasses and the evolution of bread wheat

BreadWheatEvolution.svg

Tetraploidization in wild emmer wheat

Aegilops appears to be basal to several taxa such as Triticum, Amblyopyrum, and Crithopsis. Certain species such as Aegilops speltoides could potentially represent core variants of the taxa. The generic placement may be more a matter of nomenclature. Genera Aegilops and Triticum are very closely related; as the adjacent image illustrates, the Aegilops species occupy most of the basal branch points in bread wheat evolution indicating that genus Triticum evolved from Aegilops after an estimated 4 million years ago. [8] The divergence of the genomes is followed by allotetraploidization of a speltoid goatgrass x basal wheat species Triticum boeoticum with strains in the middle eastern region giving rise to cultivated emmer wheat. [9]

Hexaploidization of tetraploid wheat

Hybridization of tetraploid wheat with Ae. tauschii produced a hulled wheat similar to spelt, suggesting T. spelta is basal. The tauschii species can be subdivided into subspecies tauschii (eastern Turkey to China or Pakistan) and strangulata (Caucasus to S. Caspian, N. Iran). The D genome of bread wheat is closer to A.t. strangulata than A.t. tauschii. It is suggested that Ae. tauschii underwent rapid selective evolution prior to combining with tetraploid wheat.[ citation needed ]

Wild Triticeae use by humans

Intense use of wild Triticeae can be seen in the Levant as early as 23,000 years ago. [10] This site, Ohala II (Israel), also shows that Triticeae grains were processed and cooked. [11] Many cultivars appear to have been domesticated in the region of the upper Fertile Crescent, Levant and central Anatolia. [12] [13] More recent evidence suggests that cultivation of wheat from emmer's wheat required a longer period with wild seeding maintaining a presence in archaeological finds. [14]

Pastoral grasses

Triticeae has a pastoral component that some contend goes back to the Neolithic period and is referred to as the Garden Hunting Hypothesis. In this hypothesis grains could be planted or shared for the purpose of attracting game animals so that they could be hunted close to settlements.[ citation needed ]

Today, rye and other Triticeae cultivars are used to graze animals, particularly cattle. Rye grasses in the New World have been used selectively as fodder, but also to protect grasslands without the introduction of invasive Old World species.[ citation needed ]

Triticeae and health

Glutens (storage proteins) in the Triticeae tribe have been linked to gluten-sensitive diseases. While it was once believed that oats carried similar potentials, recent studies indicate that most oat sensitivity is the result of contamination.[ citation needed ] Triticeae glutens studies are important in determining the links between gluten and gastrointestinal, allergic, and autoimmune diseases. [15] Some of the recently discovered biochemical and immunochemical properties of these proteins suggest they evolved for protection against dedicated or continuous consumption by mammalian seed-eaters. [16] [17] One recent publication even raises doubts about wheat's safety for anyone to eat. [18] Overlapping properties with regard to food preparation[ clarification needed ] have made these proteins much more useful as cereal cultivars, and a balanced perspective suggests a variable tolerance to Triticeae glutens reflects early childhood environment and genetic predisposition. [19] [20] [21] [22]

Related Research Articles

<span class="mw-page-title-main">Wheat</span> Genus of grass cultivated for grain

Wheat is a grass widely cultivated for its seed, a cereal grain that is a staple food around the world. The many species of wheat together make up the genus Triticum ; the most widely grown is common wheat. The archaeological record suggests that wheat was first cultivated in the regions of the Fertile Crescent around 9600 BC. Botanically, the wheat kernel is a caryopsis, a type of fruit.

<span class="mw-page-title-main">Oat</span> Cool weather staple grain, animal feed

The oat, sometimes called the common oat, is a species of cereal grain grown for its seed, which is known by the same name. Oats appear to have been domesticated as a secondary crop, as their seeds resembled those of other cereals closely enough for them to be included by early cultivators. Oats tolerate cold winters less well than cereals such as wheat, barley, and rye, but need less summer heat and more rain, making them important in areas such as Northwest Europe that have cool wet summers. They can tolerate low-nutrient and acid soils. Oats grow thickly and vigorously, allowing them to outcompete many weeds, and compared to other cereals are relatively free from diseases.

<span class="mw-page-title-main">Rye</span> Species of grain

Rye is a grass grown extensively as a grain, a cover crop and a forage crop. It is grown principally in an area from Eastern and Northern Europe into Russia. It is much more tolerant of cold weather and poor soil than other cereals, making it useful in those regions; its vigorous growth suppresses weeds and provides abundant forage for animals early in the year. It is a member of the wheat tribe (Triticeae) which includes the cereals wheat and barley. Rye grain is used for bread, beer, rye whiskey, and animal fodder. In Scandinavia, rye was a staple food in the Middle Ages, and rye crispbread remains a popular food in the region. Europe produces around half of the world's rye; relatively little is traded between countries. A wheat-rye hybrid, triticale, combines the qualities of the two parent crops and is produced in large quantities worldwide. In European folklore, the Roggenwolf is a carnivorous corn demon or Feldgeist.

<span class="mw-page-title-main">Einkorn</span> Primitive wheat

Einkorn wheat is either a wild species of wheat (Triticum) or its domesticated form. The wild form is T. boeoticum, and the domesticated form is T. monococcum. Einkorn is a diploid species of hulled wheat, with tough glumes (husks) that tightly enclose the grains. The cultivated form is similar to the wild, except that the ear stays intact when ripe and the seeds are larger. The domestic form is known as petit épeautre in French, Einkorn in German, "einkorn" or "littlespelt" in English, piccolo farro in Italian and escanda menor in Spanish. The name refers to the fact that each spikelet contains only one grain.

<span class="mw-page-title-main">Triticale</span> Hybrid wheat/rye crop

Triticale is a hybrid of wheat (Triticum) and rye (Secale) first bred in laboratories during the late 19th century in Scotland and Germany. Commercially available triticale is almost always a second-generation hybrid, i.e., a cross between two kinds of primary (first-cross) triticales. As a rule, triticale combines the yield potential and grain quality of wheat with the disease and environmental tolerance of rye. Only recently has it been developed into a commercially viable crop. Depending on the cultivar, triticale can more or less resemble either of its parents. It is grown mostly for forage or fodder, although some triticale-based foods can be purchased at health food stores and can be found in some breakfast cereals.

<i>Secale</i> Genus of grasses

Secale is a genus of the grass tribe Triticeae, which is related to barley (Hordeum) and wheat (Triticum). The genus includes cultivated species such as rye as well as weedy and wild rye species. The best-known species of the genus is the cultivated rye, S. cereale, which is grown as a grain and forage crop. Wild and weedy rye species help provide a huge gene pool that can be used for improvement of the cultivated rye.

<span class="mw-page-title-main">Emmer</span> Type of wheat

Emmer or hulled wheat is an awned type of wheat. It is a tetraploid. The domesticated types are Triticum turgidum subsp. dicoccum and T. t. conv. durum. The wild plant is called T. t. subsp. dicoccoides. The principal difference between the wild and the domestic forms is that the ripened seed head of the wild plant shatters and scatters the seed onto the ground, while in the domesticated emmer, the seed head remains intact, thus making it easier for people to harvest the grain.

<span class="mw-page-title-main">Durum wheat</span> Species of wheat used for food

Durum wheat, also called pasta wheat or macaroni wheat, is a tetraploid species of wheat. It is the second most cultivated species of wheat after common wheat, although it represents only 5% to 8% of global wheat production. It was developed by artificial selection of the domesticated emmer wheat strains formerly grown in Central Europe and the Near East around 7000 BC, which developed a naked, free-threshing form. Like emmer, durum wheat is awned. It is the predominant wheat that grows in the Middle East.

<span class="mw-page-title-main">Spelt</span> Species of wheat

Spelt, also known as dinkel wheat or hulled wheat, is a species of wheat. It is a relict crop, eaten in Central Europe and northern Spain. It is high in protein and may be considered a health food.

<span class="mw-page-title-main">Khorasan wheat</span> Species of grass

Khorasan wheat or Oriental wheat is a tetraploid wheat species. The grain is twice the size of modern-day wheat, and has a rich, nutty flavor.

<span class="mw-page-title-main">Gliadin</span> Protein in wheat & other cereals

Gliadin is a class of proteins present in wheat and several other cereals within the grass genus Triticum. Gliadins, which are a component of gluten, are essential for giving bread the ability to rise properly during baking. Gliadins and glutenins are the two main components of the gluten fraction of the wheat seed. This gluten is found in products such as wheat flour. Gluten is split about evenly between the gliadins and glutenins, although there are variations found in different sources.

<i>Aegilops</i> Genus of grasses

Aegilops is a genus of Eurasian and North American plants in the grass family, Poaceae. They are known generally as goatgrasses. Some species are known as invasive weeds in parts of North America.

<span class="mw-page-title-main">Common wheat</span> Species of plant

Common wheat, also known as bread wheat, is a cultivated wheat species. About 95% of wheat produced worldwide is common wheat; it is the most widely grown of all crops and the cereal with the highest monetary yield.

<span class="mw-page-title-main">Taxonomy of wheat</span> Classification of wheat

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 diversity has led to much confusion in the naming of wheats. Genetic and morphological characteristics of wheat influence its classification; many common and botanical names of wheat are in current use.

<span class="mw-page-title-main">Wheat allergy</span> Medical condition

Wheat allergy is an allergy to wheat that typically presents itself as a food allergy, but can also be a contact allergy resulting from occupational exposure. Wheat allergy may be immunoglobulin E mediated or not and may involve mast cell response. Wheat allergy is rare. Prevalence in adults was estimated to be 0.21% in a 2012 study in Japan.

<i>Aegilops tauschii</i> Species of grass

Aegilops tauschii, the Tausch's goatgrass or rough-spike hard grass, is an annual grass species. It is native to Crimea, the Caucasus region, western and Central Asia, Afghanistan, Pakistan, the western Himalaya, and parts of China, and has been introduced to other locales, including California.

<span class="mw-page-title-main">Triticeae glutens</span> Seed storage protein in mature wheat seeds

Gluten is the seed storage protein in mature wheat seeds. It is the sticky substance in bread wheat which allows dough to rise and retain its shape during baking. The same, or very similar, proteins are also found in related grasses within the tribe Triticeae. Seed glutens of some non-Triticeae plants have similar properties, but none can perform on a par with those of the Triticeae taxa, particularly the Triticum species. What distinguishes bread wheat from these other grass seeds is the quantity of these proteins and the level of subcomponents, with bread wheat having the highest protein content and a complex mixture of proteins derived from three grass species.

Anti-gliadin antibodies are produced in response to gliadin, a prolamin found in wheat. In bread wheat it is encoded by three different alleles, AA, BB, and DD. These alleles can produce slightly different gliadins, which can cause the body to produce different antibodies. Some of these antibodies can detect proteins in specific grass taxa such as Triticeae, while others react sporadically with certain species in those taxa, or over many taxonomically defined grass tribes.

<span class="mw-page-title-main">Barley</span> Cereal grain

Barley, a member of the grass family, is a major cereal grain grown in temperate climates globally. It was one of the first cultivated grains; it was domesticated in the Fertile Crescent around 9000 BC, giving it nonshattering spikelets and making it much easier to harvest. Its use then spread throughout Eurasia by 2000 BC. Barley prefers relatively low temperatures and well-drained soil to grow. It is relatively tolerant of drought and soil salinity but is less winter-hardy than wheat or rye.

<i>Triticum carthlicum</i> Species of grass

Triticum carthlicum Nevski, 1934, the Persian wheat, is a wheat with a tetraploid genome.

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