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''Avena barbata'' is a species of
wild oat ''Avena'' is a genus of Eurasian and African plants in the grass family. Collectively known as the oats, they include some species which have been cultivated for thousands of years as a food source for humans and livestock. They are widespread ...
known by the common name slender wild oat. It has edible seeds. It is a diploidized autotetraploid grass (2n=4x=28). Its diploid ancestors are ''A. hirtula'' Lag. and ''A. wiestii'' Steud (2n=2x=14), which are considered Mediterranean and desert
ecotypes In evolutionary ecology, an ecotype,Greek: ''οίκος'' = home and ''τύπος'' = type, coined by Göte Turesson in 1922 sometimes called ecospecies, describes a genetically distinct geographic variety, population, or race within a specie ...
, respectively, comprising a single species. ''A westie'' and ''A. hirtula'' are widespread in the Mediterranean Basin, growing in mixed stands with ''A. barbata'', though they are difficult to tell apart. This is a winter annual grass with thin tillers (stems) growing up to 60 to 80 centimeters in maximum height, but known to sometimes grow taller. The bristly spikelets are 2 to 3 centimeters long, not counting the bent
awn AWN may stand for: * Awn Access to Justice Network in Gaza Strip, Legal Aid Network operate in Gaza Strip, Palestine * Animation World Network, an online organization for animators * Avant Window Navigator, a dock-like bar that tracks open wind ...
which is up to 4 centimeters in length. ''Avena barbata'' largely reproduces by selfing in natural populations, with very low rates of outcrossing. ''A. barbata'' is native to central Asia (as far east as
Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries and dependencies by population, fifth-most populous country, with a population of almost 24 ...
) and the
Mediterranean Basin In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and w ...
. As an introduced species it also occurs in other Mediterranean-like habitats of
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 List of islands of New Zealand, smaller islands. It is the ...
, Australia,
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring count ...
,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, t ...
,
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the eas ...
,
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, and
Uruguay Uruguay (; ), officially the Oriental Republic of Uruguay ( es, República Oriental del Uruguay), is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast; while bordering ...
. In Europe it has been reported in
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,
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,
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,
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,
Bulgaria Bulgaria (; bg, България, Bǎlgariya), officially the Republic of Bulgaria,, ) is a country in Southeast Europe. It is situated on the eastern flank of the Balkans, and is bordered by Romania to the north, Serbia and North Mac ...
, and
Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
. In North America it is an
introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived the ...
and
noxious weed A noxious weed, harmful weed or injurious weed is a weed that has been designated by an agricultural or other governing authority as a plant that is injurious to agricultural or horticultural crops, natural habitats or ecosystems, or humans or liv ...
, where it is especially widespread in
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the ...
. In California it has displaced native species of grass. It is also found in
Oregon Oregon () is a state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington, while the Snake River delineates much of its eastern boundary with Idah ...
,
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,
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,
Massachusetts Massachusetts (Massachusett language, Massachusett: ''Muhsachuweesut assachusett writing systems, məhswatʃəwiːsət'' English: , ), officially the Commonwealth of Massachusetts, is the most populous U.S. state, state in the New England ...
,
Nevada Nevada ( ; ) is a state in the Western region of the United States. It is bordered by Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. Nevada is the 7th-most extensive, ...
,
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a U.S. state, state in the Southwestern United States. It is the list of U.S. states and territories by area, 6th largest and the list of U.S. states and territories by population, 14 ...
, and
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe, New Mexico, Santa Fe , LargestCity = Albuquerque, New Mexico, Albuquerque , LargestMetro = Albuquerque metropolitan area, Tiguex , Offi ...
. Genetic evidence indicates that ''A. barbata'' in Argentina and California originated from Spain, during the
Spanish colonization of the Americas Spain began colonization of the Americas, colonizing the Americas under the Crown of Castile and was spearheaded by the Spanish . The Americas were invaded and incorporated into the Spanish Empire, with the exception of Colonial Brazil, Braz ...
.


Genetic studies of Californian populations

Californian populations of ''Avena barbata'' represent one of the most extensively studied examples of putative "
ecotype In evolutionary ecology, an ecotype,Greek: ''οίκος'' = home and ''τύπος'' = type, coined by Göte Turesson in 1922 sometimes called ecospecies, describes a genetically distinct geographic variety, population, or race within a specie ...
s" in the plant literature. Its
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and pop ...
and
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
have been extensively examined since 1967, primarily in the laboratories of R.W. Allard and Subodh Jain and their many students in the 1960s, 1970s, 1980s, and 1990s at U.C. Davis, and more recently by Robert Latta at the University of Dalhousie University in Nova Scotia. The general pattern that emerged from these earlier studies was that throughout the Central Valley of California, consisting of semiarid grasslands and oak savannahs, and extending south to San Diego, populations of this species were dominated by a
monomorphic Monomorphic or Monomorphism may refer to: *Monomorphism, an injective homomorphism in mathematics * Monomorphic QRS complex, a wave pattern seen on an electrocardiogram * Monomorphic, a linguistic term meaning "consisting of only one morpheme" *Mo ...
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological prop ...
possessing dark/black seeds with hairy
lemma Lemma may refer to: Language and linguistics * Lemma (morphology), the canonical, dictionary or citation form of a word * Lemma (psycholinguistics), a mental abstraction of a word about to be uttered Science and mathematics * Lemma (botany), a ...
s, as well as smooth
leaf sheath A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
s; these morphological characters were correlated with a specific
isozyme In biochemistry, isozymes (also known as isoenzymes or more generally as multiple forms of enzymes) are enzymes that differ in amino acid sequence but catalyze the same chemical reaction. Isozymes usually have different kinetic parameters (e.g. dif ...
pattern as well as a specific ribosomal DNA genotype. This "ecotype" is called the "
xeric Deserts and xeric shrublands are a biome defined by the World Wide Fund for Nature. Deserts and xeric (ancient Greek xērós, “dry") shrublands form the largest terrestrial biome, covering 19% of Earth's land surface area. Ecoregions in this h ...
" type. Populations outside the Central Valley, along the coastal strip, the intermontane regions of the coast ranges, and the higher foothills of the Sierra Nevada mountains, were either monomorphic for white seeds with generally smooth lemmas and hairy leaf sheaths or were polymorphic with varying mixtures of the seed and leaf sheath characters. These populations were also either monomorphic or polymorphic for isozyme patterns and
ribosomal DNA Ribosomal DNA (rDNA) is a DNA sequence that codes for ribosomal RNA. These sequences regulate transcription initiation and amplification, and contain both transcribed and non-transcribed spacer segments. In the human genome there are 5 chrom ...
genotypes other than the xeric type; they are called the "
mesic Mesic may refer to: * Mesic, North Carolina, a town in the United States * Mesic habitat, a type of habitat See also *Mesić (disambiguation) *Mešić Mešić is a Bosnian surname, a patronymic derived from the masculine given name '' Meša'', it ...
" type. The mesic type has never apparently been observed south of approximately the same latitude as Monterey, either in coastal ranges, the Central Valley, or the foothills of the Sierras. When the morphological traits as well as the
allozyme Alloenzymes (or also called allozymes) are variant forms of an enzyme which differ structurally but not functionally from other allozymes coded for by different alleles at the same locus. These are opposed to isozymes, which are enzymes that perfo ...
and
ribosomal DNA Ribosomal DNA (rDNA) is a DNA sequence that codes for ribosomal RNA. These sequences regulate transcription initiation and amplification, and contain both transcribed and non-transcribed spacer segments. In the human genome there are 5 chrom ...
genotypes were considered together, it is argued that there are six ecotypes in the otherwise "mesic" classification. Whole-plant studies also showed that the xeric and mesic types differed from each other for many characters such as the flag leaf, primary stem height, number of tillers, weight and number of seeds, dry weight, and flowering time, with the music ecotype being generally larger and more fecund, overall, than the xeric type; further, the flag leaves of the xeric type were consistently smaller than the mesic type under many conditions. It was further shown that xeric populations that were monomorphic for the seed and leaf sheath characters and allozymes had a less genetic variation for
quantitative genetics Quantitative genetics deals with phenotypes that vary continuously (such as height or mass)—as opposed to discretely identifiable phenotypes and gene-products (such as eye-colour, or the presence of a particular biochemical). Both branches ...
characters than mesic populations; however, quantitative genetic variation existed in all xeric or mesic populations that were studied. Consequently, with all genetic characters studied, xeric populations of the xeric ecotype were more similar to each other than they were to the music ecotype, and the evidence indicated that the various ecotypes represented significant
linkage disequilibrium In population genetics, linkage disequilibrium (LD) is the non-random association of alleles at different loci in a given population. Loci are said to be in linkage disequilibrium when the frequency of association of their different alleles is h ...
and ''coadapted genetic complexes''. For field identification purposes, the leaf sheath pubescence in the seedling stage and
lemma Lemma may refer to: Language and linguistics * Lemma (morphology), the canonical, dictionary or citation form of a word * Lemma (psycholinguistics), a mental abstraction of a word about to be uttered Science and mathematics * Lemma (botany), a ...
color at seed maturity as well as the flag leaf dimensions would reliably separate the xeric from the mesic ecotypes throughout California. Early on it was speculated that the genetic patterns observed in ''A. barbata'' were highly
correlated In statistics, correlation or dependence is any statistical relationship, whether causal or not, between two random variables or bivariate data. Although in the broadest sense, "correlation" may indicate any type of association, in statisti ...
with rainfall and temperature. The general pattern at both a macro and micro geographical scale was that the monomorphic "xeric" type occurred in those regions with between 250mm and 500mm of rainfall, while the polymorphic and monomorphic "mesic" populations occurred in those areas of California with greater than 500mm. Regardless of the correlations found with the mesic and xeric genotypes with rainfall in California, greenhouse experiments have not shown that the xeric type has greater reproductive capacity or other physiologic superiority to the mesic under artificially induced wet or dry conditions. In fact Latta argues that the mesic type is superior to the xeric, and may be supplanting the xeric in those areas where the xeric has been dominant, at least in Northern California.


Genetic studies of Mediterranean populations

''Avena barbata'' has been studied in Spain, Israel, and Morocco by students and colleagues of R.W. Allard at U.C. Davis, Pèrez de la Vega and Pedro Garcia of the University of Leon, and E. Nevo in Israel (). The general pattern that has emerged is that there is more genetic variability in the Mediterranean populations than there are in Californian populations. Further, the multi-
locus Locus (plural loci) is Latin for "place". It may refer to: Entertainment * Locus (comics), a Marvel Comics mutant villainess, a member of the Mutant Liberation Front * ''Locus'' (magazine), science fiction and fantasy magazine ** ''Locus Award ...
genotypes The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
found in California are unique to California. The Mediterranean populations have their own unique sets of multi-locus genotypes. There is a unique 14-locus
allozyme Alloenzymes (or also called allozymes) are variant forms of an enzyme which differ structurally but not functionally from other allozymes coded for by different alleles at the same locus. These are opposed to isozymes, which are enzymes that perfo ...
genotype specific to the colder regions of Spain. Neither the "xeric" nor monomorphic "mesic" genotypes described in California are found in Spain.


Genetic studies of Argentinian populations

Both Californian and Argentinian populations represent a subset of the genetic variability found in Spain, on a locus by locus comparison. However, unlike Spain, Argentina has one widespread 14 locus allozyme genotype called the "Pampeano" type, which is not found in Spain; it differs from the Californian "xeric" type at three of the 14 loci examined. The "xeric" Californian type is found in 6% of the plants examined in Argentina; the "xeric" type also occurs in Chile. The genetic evidence indicates that ''Avena barbata'' came to both Argentina and California from southwest Spain.


Gallery

File:Avena barbata (3823399576).jpg, Avena barbata spikelet


References


External links


Jepson Manual TreatmentGrass Manual TreatmentPhoto gallery
{{Taxonbar, from=Q4828032
barbata Barbata (Bergamasque: ) is a '' comune'' (municipality) in the Province of Bergamo in the Italian region of Lombardy, located about east of Milan and about southeast of Bergamo. Barbata borders the following municipalities: Antegnate, Camis ...
Flora of Malta