Triticeae | |
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Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Monocots |
Clade: | Commelinids |
Order: | Poales |
Family: | Poaceae |
Clade: | BOP clade |
Subfamily: | Pooideae |
Supertribe: | Triticodae |
Tribe: | Triticeae L. |
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.
Genera of Triticeae
Genera recognized in Triticeae according to Robert Soreng et al.:
- Aegilops
- Agropyron
- Amblyopyrum
- Anthosachne
- Australopyrum
- Connorochloa
- Crithopsis
- Dasypyrum
- Douglasdeweya
- Elymus (syn. Campeiostachys, Elytrigia, Hystrix, Roegneria, Sitanion)
- Eremopyrum
- Festucopsis
- Henrardia
- Heteranthelium
- Hordelymus
- Hordeum (syn. Critesion)
- Kengyilia
- Leymus (syn. Aneurolepidium, Eremium, Macrohystrix, Microhystrix)
- Pascopyrum
- Peridictyon
- Psathyrostachys
- Pseudoroegneria
- Secale
- Stenostachys
- Taeniatherum
- Thinopyrum
- Triticum
Cultivated or edible species
Aegilops
- Various species (rarely identifiable to species in archaeological material) occur in pre-agrarian archaeobotanical remains from Near Eastern sites. Their edible grains were doubtless harvested as wild food resources.
- speltoides - ancient food grain, putative source of B genome in bread wheat and G genome in T. timopheevii
- tauschii - Source of D genome in wheat
Amblyopyrum
- muticum - Source of T genome.
Elymus
Various species are cultivated for pastoral purposes or to protect fallow
land from opportunistic or invasive species
Hordeum
Many barley cultivars
- vulgare - common barley (6 subspecies, ~100 cultivars)
- bulbosum - edible seeds
- murinum (mouse barley) - cooked as piƱole, bread-flour capable, medicinal: diuretic.
Leymus
- arenarius (Lyme grass) - bread-flour capable, possible food additive
- racemosus (Volga Wild Rye) - drought tolerant cereal, used in Russia
- condensatus (Giant Wild Rye) - Edible seeds, harvesting problematic small seeds
- triticoides (Squaw grass) - used in North America, seed hairs must be singed
Secale
Ryes
- cereale (Cereal Rye) - Livestock feed and sour dough bread - 6 subspecies.
- cornutum-ergot (Ergot of Spurred Rye) - herbal medicine at very low doses, deadly poisonous as food.
- strictum - actively cultivated
- sylvestre - (Tibetan Rye) - actively cultivated in Tibet and China highlands.
- vavilovii (Armenian Wild Rye) - edible seeds, thickener.
Triticum
Wheat
- aestivum (bread wheat) - (AABBDD Genome)
- compactum (club wheat)
- macha (hulled)
- spelta (hulled, spelt)
- sphaerococcum (shot wheat)
- monococcum (Einkorn wheat) (A Genome)
- timopheevii (Sanduri wheat)
- turgidum (poulard wheat) (AABB Genome)
- carthlicum (Persian black wheat)
- dicoccoides (wild emmer wheat)
- dicoccum (cultivated emmer wheat) - used to make Farro
- durum (durum wheat)
- paleocolchicum
- polonicum (Polish wheat)
- turanicum
Genetics
Genera & Species | 1st | 2nd | 3rd |
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Triticum boeoticum | AA |
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Triticum monococcum | AMAM |
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Triticum urartu | AUAU |
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Aegilops speltoides var. speltoides | BB |
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Aegilops caudata | CC |
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Aegilops tauschii | DD |
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Lophopyrum elongatum | EE |
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Hordeum vulgare | HH |
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Thinopyrum bessarabicum | JJ |
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Aegilops comosa | MM |
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Aegilops uniaristata | NN |
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Henrardia persica | OO |
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Agropyrum cristatum | PP |
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Secale cereale | RR |
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Aegilops bicornis | SS |
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Amblyopyrum muticum | TT |
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Aegilops umbellulata | UU |
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Dasypyrum | VV |
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Psathyrostachys | NsNs |
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Pseudoroegneria | StSt |
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Triticum zhukovskyi | AA | AMAM | GG | |
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Triticum turgidum | AA | BB |
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Triticum aestivum | AA | BB | DD | |
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Triticum timopheevii | AA | GG |
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Aegilops cylindrica | CC | DD |
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Stenostachys sp. | HH | WW |
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Elmyus canadensis | HH | StSt |
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Elmyus abolinii | YY | StSt |
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Thinopyrum Vjd =(V/J/D) | JJ | StSt | VjdVjd | |
| Leymus tricoides | NsNs | XmXm |
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 genetic marker with barley and 10 other members of Triticeae, whereas all 19 genera of Triticeae bear a wheat marker along with Bromeae. 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.
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), but also shows introgression from Australian and Agropyron wheatgrasses. Elymus contains mostly Pseudoroegeneria mtDNA.
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.
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 favorablitiy of Secale declined. At that time other strains of barley and wheat may have been cultivated, but humans did little to change them.
Goat grasses and the evolution of bread wheat
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. 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.
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.
Wild Triticeae use by humans
Intense use of wild Triticeae can be seen in the Levant as early as 23,000 years ago. This site, Ohala II (Israel), also shows that Triticeae grains were processed and cooked. Many cultivars appear to have been domesticated in the region of the upper Fertile Crescent, Levant and central Anatolia. 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.
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.
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.
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. Triticeae glutens studies are important in determining the links between gluten and gastrointestinal, allergic, and autoimmune diseases.
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. One recent publication even raises doubts about wheat's safety for anyone to eat. Overlapping properties with regard to food preparation
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.