词条 | Taxonomy of wheat | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
释义 |
During 10,000 years of cultivation, numerous forms of wheat, many of them hybrids, have developed under a combination of artificial and natural selection.[1][2] This diversity has led to much confusion in the naming of wheats. This article explains how genetic and morphological characteristics of wheat influence its classification, and gives the most common botanical names of wheat in current use (see Table of wheat species). Information on the cultivation and uses of wheat is at the main wheat page. Aegilops and TriticumThe genus Triticum includes the wild and domesticated species usually thought of as wheat. In the 1950s growing awareness of the genetic similarity of the wild goatgrasses (Aegilops) led botanists such as Bowden to amalgamate Aegilops and Triticum as one genus, Triticum.[3] This approach is still followed by some (mainly geneticists), but has not been widely adopted by taxonomists.[4] Aegilops is morphologically highly distinct from Triticum, with rounded rather than keeled glumes.[5] Aegilops is important in wheat evolution because of its role in two important hybridisation events. Wild emmer (T. dicoccoides and T. araraticum) resulted from the hybridisation of a wild wheat, T. urartu, and an as yet unidentified goatgrass, probably closely related to Ae. speltoides.[6] Hexaploid wheats (e.g. T. aestivum and T. spelta) are the result of a hybridisation between a domesticated tetraploid wheat, probably T. dicoccum or T. durum, and another goatgrass, Ae. tauschii (also known as Ae. squarrosa).[7]Early taxonomyBotanists of the classical period, such as Columella, and in sixteenth and seventeenth century herbals, divided wheats into two groups, Triticum corresponding to free-threshing wheats, and Zea corresponding to hulled ('spelt') wheats.[4] Carl Linnaeus recognised five species, all domesticated:[4]
Later classifications added to the number of species described, but continued to give species status to relatively minor variants, such as winter vs. spring forms. The wild wheats were not described until the mid-19th century because of the poor state of botanical exploration in the Near East, where they grow.[4] The development of a modern classification depended on the discovery, in the 1920s, that wheat was divided into 3 ploidy levels.[8] Important characters in wheatPloidy levelAs with many grasses, polyploidy is common in wheat.[9] There are two wild diploid (non-polyploid) wheats, T. boeoticum and T. urartu. T. boeoticum is the wild ancestor of domesticated einkorn, T. monococcum.[10] Cells of the diploid wheats each contain 2 complements of 7 chromosomes, one from the mother and one from the father (2n=2x=14, where 2n is the number of chromosomes in each somatic cell, and x is the basic chromosome number). The polyploid wheats are tetraploid (4 sets of chromosomes, 2n=4x=28), or hexaploid (6 sets of chromosomes, 2n=6x=42). The tetraploid wild wheats are wild emmer, T. dicoccoides, and T. araraticum. Wild emmer is the ancestor of all the domesticated tetraploid wheats, with one exception: T. araraticum is the wild ancestor of T. timopheevi.[11] There are no wild hexaploid wheats, although feral forms of common wheat are sometimes found. Hexaploid wheats developed under domestication. Genetic analysis has shown that the original hexaploid wheats were the result of a cross between a tetraploid domesticated wheat, such as T. dicoccum or T. durum, and a wild goatgrass, Ae. tauschii.[7] Polyploidy is important to wheat classification for three reasons:
GenomeObservation of chromosome behaviour during meiosis, and the results of hybridisation experiments, have shown that grass genomes (complete complements of genetic matter) can be grouped into distinctive types. Each type has been given a name, e.g. B or D. Grasses sharing the same genome will be more-or-less interfertile, and might be treated by botanists as one species. Identification of genome types is obviously a valuable tool in investigating hybridisation. For example, if two diploid plants hybridise to form a new polyploid form (an allopolyploid), the two original genomes will be present in the new form. Many thousands of years after the original hybridisation event, identification of the component genomes will allow identification of the original parent species.[14] In Triticum, five genomes, all originally found in diploid species, have been identified:
The genetic approach to wheat taxonomy (see below) takes the genome composition as defining each species.[15] As there are five known combinations in Triticum this translates into five super species:
DomesticationThere are four wild species, all growing in rocky habitats in the fertile crescent of the Near East.[16] All the other species are domesticated. Although relatively few genes control domestication, and wild and domesticated forms are interfertile, wild and domesticated wheats occupy entirely separate habitats. Traditional classification gives more weight to domesticated status. Hulled vs. Free-threshingAll wild wheats are hulled: they have tough glumes (husks) that tightly enclose the grains. Each package of glumes, lemma and palaea, and grain(s) is known as a spikelet. At maturity the rachis (central stalk of the cereal ear) disarticulates, allowing the spikelets to disperse.[17] The first domesticated wheats, einkorn and emmer, were hulled like their wild ancestors, but with rachises that (while not entirely tough) did not disarticulate at maturity. During the Pre-Pottery Neolithic B period, at about 8000 BC, free-threshing forms of wheat evolved, with light glumes and fully tough rachis. Hulled or free-threshing status is important in traditional classification because the different forms are usually grown separately, and have very different post-harvesting processing. Hulled wheats need substantial extra pounding or milling to remove the tough glumes. For more information, see Wheat: Hulled vs. free-threshing wheat MorphologyIn addition to hulled/free-threshing status, other morphological criteria, e.g. spike laxness or glume wingedness, are important in defining wheat forms. Some of these are covered in the individual species accounts linked from this page, but Floras must be consulted for full descriptions and identification keys. Traditional vs. genetic classificationsAlthough the range of recognised types of wheat has been reasonably stable since the 1930s, there are now sharply differing views as to whether these should be recognised at species level (traditional approach) or at subspecific level (genetic approach). The first advocate of the genetic approach was Bowden, in a 1959 classification (now historic rather than current).[18] He, and subsequent proponents (usually geneticists), argued that forms that were interfertile should be treated as one species (the biological species concept). Thus emmer and hard wheat should both be treated as subspecies (or at other infraspecific ranks) of a single tetraploid species defined by the genome BAu. Van Slageren's 1994 classification is probably the most widely used genetic-based classification at present.[19] Users of traditional classifications give more weight to the separate habitats of the traditional species, which means that species that could hybridise do not, and to morphological characters. There are also pragmatic arguments for this type of classification: it means that most species can be described in Latin binomials, e.g. Triticum aestivum, rather than the trinomials necessary in the genetic system, e.g. Triticum aestivum subsp. aestivum. Both approaches are widely used. Infraspecific classificationIn the nineteenth century, elaborate schemes of classification were developed in which wheat ears were classified to botanical variety on the basis of morphological criteria such as glume hairiness and colour or grain colour. These variety names are now largely abandoned, but are still sometimes used for distinctive types of wheat such as miracle wheat, a form of T. turgidum with branched ears, known as T. turgidum L. var. mirabile Körn. The term cultivar (abbreviated as cv.) is often confused with species or domesticate. In fact, it has a precise meaning in botany: it is the term for a distinct population of a crop, usually commercial and resulting from deliberate plant-breeding. Cultivar names are always capitalised, often placed between apostrophes, and not italicised. An example of a cultivar name is T. aestivum cv. 'Pioneer 2163'. A cultivar is often referred to by farmers as a variety, but this is best avoided in print, because of the risk of confusion with botanical varieties. The term landrace is applied to informal, farmer-maintained populations of crop plants. NamingBotanical names for wheat are generally expected to follow an existing classification, such as those listed as current at the Wheat Classification Tables Site [https://web.archive.org/web/20000310130245/http://www.ksu.edu/wgrc/Taxonomy/taxintro.html]. The classifications given in the following table are among those suitable for use. If a genetic classification is favoured, the GRIN classification is comprehensive, based on van Slageren's work but with some extra taxa recognised. If the traditional classification is favoured, Dorofeev's work is a comprehensive scheme that meshes well with other less complete treatments.Wikipedia's wheat pages generally follow a version of the Dorofeev scheme – see the taxobox on the Wheat page. A general rule is that different taxonomic schemes should not be mixed in one context. In a given article, book or web page, only one scheme should be used at a time. Otherwise, it will be unclear to others how the botanical name is being used. Table of wheat species
Note: Blank common name indicates that no common name is in use in the English language. Explanatory notes on selected names
Artificial species and mutantsRussian botanists have given botanical names to hybrids developed during genetical experiments. As these only occur in the laboratory environment, it is questionable whether botanical names (rather than lab. numbers) are justified. Botanical names have also been given to rare mutant forms. Examples include:
See also
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External links{{Commonscat|Wheat|Wheat}} Taxonomy
Genetics
Morphology
2 : Wheat|Plant taxonomies |
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