By clicking Accept All Cookies, you agree to the storing of cookies on your device to enhance site navigation, analyze site usage, and assist in our marketing efforts. Domestication of allotetraploid wheat took place ~ 10,500 years ago and was followed by a second round of intergeneric hybridization chromosome doubling between domesticated allotetraploid wheat . Synthetic hexaploid lines have been shown to be useful sources of variation for abiotic and biotic stress tolerance, as well as agronomic and novel grain quality traits. 2022 Jul 29;13:898769. doi: 10.3389/fpls.2022.898769. CABI Publishing CAB International, Wallingford, pp 287315, Nevo E (2007) Evolution of wild wheat and barley and crop improvement: studies at the Institute of Evolution. Theor Appl Genet 71:518526, Sood S, Kuraparthy V, Bai G, Gill BS (2009) The major threshability genes soft glume (sog) and tenacious glume (Tg), of diploid and polyploid wheat, trace their origin to independent mutations at non-orthologous loci. A creative strategy for addressing the impoverished D genome diversity of common wheat is the development of synthetic hexaploid wheats (McFadden and Sears, 1946; Trethowan and Mujeeb-Kazi, 2008; Ogbonnaya et al., 2013). Development of synthetic hexaploid wheat (Ogbonnaya et al., 2013). Hirst, K. Kris. Springer, Berlin, pp 407456. Wheat geneticists and breeders have a long history of identifying useful genes in wild wheat relatives and incorporating them into improved varieties. Genet Res 77:213218, Cakmak IA, Torun E, Millet E, Feldman M, Fahima T, Korol AB, Nervo E, Braun HJ, zkan H (2004) Triticum dicoccoides: an important genetic resource for increasing zinc and iron concentration in modern cultivated wheat. Euphytica 142:247251, Watanabe N, Ikebata N (2000) The effects of homoeologous group 3 chromosomes on grain colour dependent seed dormancy and brittle rachis in tetraploid wheat. Frontiers in Plant Science 10:585. The domestication of rice dates to about 4000 b.c. 2014. Correspondence to You can also read the documentation to learn about Wordfence's blocking tools, or visit wordfence.com to learn more about Wordfence. Springer, Berlin, pp 387405, Salamini F, zkan H, Brandolini A, Schfer-Pregl R, Martin W (2002) Genetics and geography of wild cereal domestication in the Near East. Genome 36:2131, Dvorak J, Luo MC, Yang ZL (1998) Genetic evidence on the origin of Triticum aestivum L. In: Damania AB, Valkoun J, Willcox G, Qualset CO (eds) The origins of agriculture and crop domestication. Science 312:13921396, Korol A, Mester D, Frenkel Z, Ronin Y (2009) Methods of genetic analysis in the Triticeae. Careers. Wheat and barley are two of the founder crops of the agricultural revolution that took place 10,000 years ago in the Fertile Crescent and both crops remain among the world's most important crops. In a study of 253 synthetic-derived lines relevant to CIMMYTs breeding and testing program, the average marker-based estimate of the Ae. Veget Hist Archaeobot 17:313325, Xie W, Nevo E (2008) Wild emmer: genetic resources, gene mapping and potential for wheat improvement. Funct Integr Genomics 8:3342, Levy AA, Feldman M (2004) Genetic and epigenetic reprogramming of the wheat genome upon allopolyploidization. Ancient hybridizations among the ancestral genomes of bread wheat. Most of the specimens were damaged by fire or when the wheat was threshed, but 804 were identifiable. Sci Rep. 2022 May 20;12(1):8532. doi: 10.1038/s41598-022-11952-9. In: Padulosi S, Hammer K, Heller J (eds) hulled wheats. Domestication Domesticates are species of plants and animals that show recognizable changes in morphology away from wild phenotypes. It was a key event in the agricultural revolution that occurred about 10,000 years . This was elegantly demonstrated by John Bennet Lawes in the 1880s when he decided to allow part of the famous long-term Broadbalk experiment at Rothamsted to return to its natural state ().He therefore left part of the wheat crop unharvested in 1882 and . Federal government websites often end in .gov or .mil. Euphytica 115:215220, Watanabe N, Sogiyama K, Yamagashi Y, Skata Y (2002) Comparative telosomic mapping of homoeologous genes for brittle rachis in tetraploid and hexaploid wheats. Ann Bot 100:903924, Fuller DQ, Allaby RG, Stevens C (2010) Domestication as innovation: the entanglement of techniques, technology and chance in the domestication of cereal crops. The featured traits are plant height, harvest index, lodging, and shattering. Emmer can grow in a variety of habitats from 100 meters below sea level to 1,700 meters above sea level, and can survive in places with annual precipitation of 200-1,300 mm. Most of the 25,000 different forms of modern wheat are varieties of two broad groups, called common wheat and durum wheat. Would you like email updates of new search results? Theor Appl Genet 110:925931, Dvorak J (2009) Triticeae genome structure and evolution. Differences in nutritional quality between wild and domesticated forms of barley and emmer wheat. tauschii (DD), but using a much broader range of Ae. Am J Bot 63:10881094, Kashkush K (2007) Genome-wide impact of transcriptional activation of long terminal repeat retrotransposons on the expression of adjacent host genes. Its breeding has followed millennia of cultivation, sometimes with unintended selection on adaptive traits, and later by applying intentional but empirical selective pressures. Biol J Linn Soc 82:607613, Li W, Gill BS (2006) Multiple genetic pathways for seed shattering in the grasses. Science 316:18301835, Belyayev A, Kalendar R, Brodsky L, Nevo E, Schulman AH, Raskina O (2010) Transposable elements in a marginal plant population: temporal fluctuations provide new insights into genome evolution of wild diploid wheat. Unauthorized use of these marks is strictly prohibited. (HTTP response code 503). Other scholars have questioned the date, and the aDNA identification, saying it was in too good a condition to be that old. In: Chrispeels MJ, Gepts P (editors). Science 413:12981301, van Zeist W, Bakker-Heeres JAH (1985) Archaeological studies in the Levant 1. Dr. Patrick Byrne shows harvested heads of the A, B, and D ancestors of wheat, and explains the strategy of creating synthetic hexaploid wheat to increase diversity of cultivated wheat. Drought adaptive traits and wide adaptation in elite lines derived from resynthesized hexaploid wheat. Euphytica 115:121126, Ross-Ibarra J, Morrell PL, Gaut BS (2007) Plant domestication, a unique opportunity to identify the genetic basis of adaptation. In: Smartt J, Simmonds NW (eds) Evolution of crop plants. PubMed About 10,000 years ago, T. aestivumderived from a cross between domesticated emmer (Triticum dicoccum) and the goat grass Aegilops tauschii. Wild wheat shatters and falls to the ground to reseed itself when ripe, but domesticated wheat stays on the stem for easier harvesting. in mainland Southeast Asia and China. The genetic changes during domestication mean that modern wheats are unable to survive wild in competition with better adapted species. Theor Appl Genet 112:373381, Nalam V, Vales MI, Watson CJW, Johnson EB, Riera-Lizarazu O (2007) Map based analysis of genetic loci on chromosome 2D that affect glume tenacity and threshability, components of the free-threshing habit in common wheat (Triticum aestivum L.). Utilizing wild grass biodiversity in wheat improvement: 15 years of wide cross research at CIMMYT. Hereditas 137:180185, Willcox G (2004) Measuring grain size and identifying Near Eastern cereal domestication: evidence from the Euphrates valley. These includes genes for resistance to leaf rust, stem rust, stripe rust, powdery mildew, Septoria tritici, Septoria nodorum, tan spot, cyst nematode, root knot nematode, Hessian fly, greenbug, Russian wheat aphid, wheat curl mite, and soil-borne cereal mosaic virus. The combination of the A, B, and D genomes led to common bread wheat, which has all three genomes. J Hered 7:214216, Simons K, Fellers JP, Trik HN, Zhang Z, Tai YS, Gill BS, Faris JD (2006) Molecular characterization of the major wheat domestication gene Q. Genetics 172:547555, Snape J, Law W, Parker CN, Worland BB, Worland AJ (1985) Genetical analysis of chromosome 5A of wheat and its influence on important agronomic characters. Field C, A and B genome ancestors and A x B hybridization, D genome ancestor and AB x D hybridization, Next: Case Study: Sunflower Domestication and Breeding. Genetics 168:609623, Perrino P, Laghetti G, DAntuono LF, Al Ajlouni M, Kanbertay M, Szabo AT, Hammer K (1996) Ecogeographical distribution of hulled wheat species. Common wheat is a hexaploid species with three sets of similar but distinct chromosomes. Lopes MS, Reynolds MP. https://doi.org/10.1007/s11032-011-9608-4. Figure 4. This work (Crop Wild Relatives and their Use in Plant Breeding by Gayle Volk and Patrick Byrne) is free of known copyright restrictions. Methods Mol Biol. The evidence is abundant that by about 10,400 years ago, domesticated wheat was in widespread use throughout the Levant region; but when that started is up for debate. It was a key event in the agricultural revolution that occurred about 10,000 years ago in the Fertile Crescent of the Middle East. Scholars suggest that the wheat got to Britain by boat. KSCX2-YW-Z-0722, the CAS Strategic Priority Research Program Grant No. DD) in north-west Iran or Armenia (Dvorak et al., 1998). Figure 1. According to French botanist Agathe Roucou and colleagues, the domestication process also caused multiple changes in the plant that were generated indirectly. Domestication makes all the cultivars, including wheat, human-dependent, capable of surviving only under cultivation in human agricultural niches to meet human needs (Gustafson et al. Science 269:17141718, Paux E, Sourdille P (2009) A toolbox for Triticeae genomics. Bethesda, MD 20894, Web Policies Ae. Of these traits, grain retention and threshability, yield improvement, changes to photoperiod sensitivity and nutritional value are most pronounced between wild and domesticated forms. Longman Scientific & Technical Press, London, pp 84192, Feuillet C, Eversole K (2007) Physical mapping of the wheat genome: a coordinated effort to lay the foundation for genome sequencing and develop tools for breeders. Oxford University Press, Cary, North Carolina. McFadden ES, Sears ER. Accessibility Junhua H. Peng. Crop Science 48:1255-1265. BMC Genomics 11:727, Chen QF, Yen C, Yang JL (1998) Chromosome location of the gene for brittle rachis in the Tibetan weedrace of common wheat. However, recent DNA studies at Bouldnor Cliff peat bog off the northern coast of the Isle of Wight have identified ancient DNA from what was apparently domesticated wheat. Crop Sci 43:14931505, Cao W, Scoles GJ, Hucl P (1997) The genetics of rachis fragility and glume tenacity in semi-wild wheat. Crop Sci 39:17281732, Sharma H, Waynes J (1980) Inheritance of tough rachis in crosses of Triticum monococcum and Triticum boeoticum. Bread wheat (Triticum aestivum) accounts for approximately 95 percent of all wheat consumed in the world today; the other five percent consists of durum or hard wheat (T. turgidum ssp) used in pasta and semolina products. Perennials as Future Grain Crops: Opportunities and Challenges. J World Prehistory 4:157222, Honne BI, Heun M (2009) On the domestication genetics of self-fertilizing plants. tauschii. Ancient forms have built-in coordination between above and below ground functioning, but the human selection of other traits has forced the plant to reconfigure and build new networks. Expert Answer. Throughout the domestication process of wheat, the plant phenotype has undergone both rapid (within a few hundred years) and slow (thousands of years) changes, such as the weakening of the. Wheats D genome ancestor is the goatgrass Aegilops tauschii, which occurs across a broad range of the Middle East and Asia from Syria to China. Shifts in Plant Functional Strategies over the Course of Wheat Domestication, Sedimentary DNA from a Submerged Site Reveals Wheat in the British Isles 8000 Years Ago. Above photo credits (left to right): Phil Westra, Pat Byrne, Mike May. LBK is typically dated in Europe between 54004900 BCE. 2019 Institute of Botany, Chinese Academy of Sciences. PLoS Biol 6(6):e148. The founder crops (or primary domesticates) are the eight plant species that were domesticated by early Neolithic farming communities in Southwest Asia and went on to form the basis of agricultural economies across much of Eurasia, including Southwest Asia, South Asia, Europe, and North Africa.They consist of three cereals (emmer wheat, einkorn wheat, and barley), four pulses (lentil, pea . (Gramineae- Triticinae). The chromosome number of this plant is then doubled by treating with colchicine to produce AABBDD plants, the same as common wheat. eCollection 2022. domestication, the process of hereditary reorganization of wild animals and plants into domestic and cultivated forms according to the interests of people. and transmitted securely. 8,000-7,000 The bones of humans who regularly ate corn during their lifetime have a high 13C measurement. Studies of bone size, bone strength, and osteoarthritis showed that in coastal areas, the adoption of farming is associated with a decrease in workload. In: Feuillet C, Muehlbauer GJ (eds) Genetics and genomics of the Triticeae, plant genetics and genomics: crops and models 7. Wheat is a grain product with about 25,000 different varieties today. A separate lineage of the A genome led to domesticated einkorn wheat (Triticum monococcum ssp. Theor Appl Genet 109:17101717, McFadden ES, Sears ER (1946) The origin of Triticum spelta and its free-threshing hexaploid relatives. The International Maize and Wheat Improvement Center (CIMMYT) has been a leader in producing and utilizing synthetic hexaploids, with over 1,500 synthetics developed (Mujeeb-Kazi and Hettel, 1995; Rosyara et al., 2019). Trethowan RM, Mujeeb-Kazi A. Springer, Berlin, pp 317335, Tanksley SD, McCouch SR (1997) Seed banks and molecular maps: unlocking genetic potential from the wild. Their improved yield performance under drought stress conditions appears to be due at least in part to shifts in their rooting profile, whereby a greater proportion of root biomass is produced deeper in the profile (Lopes and Reynolds, 2011; Reynolds et al., 2008). Domestication has genetically not only transformed the brittle rachis, tenacious glume and non-free threshability, but also modified yield and yield components in wheat. John Murray, London (popular edition in two volumes) 566 p, Doebley J (1989) Isozymic evidence and the evolution of crop plants. Mexico City, Mexico. The Eight Founder Crops and the Origins of Agriculture, Funnel Beaker Culture: First Farmers of Scandinavia, The Domestication History of Cotton (Gossypium), Hunter Gatherers - People Who Live on the Land, Pea (Pisum sativum L.) Domestication - The History of Peas and Humans, History of the Domestication of Cows and Yaks, Chili Peppers - An American Domestication Story, eight founder crops of the origins of agriculture, Wild Emmer Genome Architecture and Diversity Elucidate Wheat Evolution and Domestication, A Chromosome-Based Draft Sequence of the Hexaploid Bread Wheat (Triticum Aestivum) Genome, Adapting Crops, Landscapes, and Food Choices: Patterns in the Dispersal of Domesticated Plants across Eurasia. FOIA The main difference between wild forms and domesticated wheat is; domesticated wheat has larger grains with rind and semi-brittle stems. Theor Appl Genet 114:947959, Luo MC, Deal KR, Akhunov ED, Akhunova AR, Anderson OD, Anderson JA, Blake N, Clegg MT, Coleman-Derr D, Conley EJ, Crossman CC, Dubcovsky J, Gill BS, Gu YQ, Hadam J, Heo HY, Huo N, Lazo G, Ma Y, Matthews DE, McGuire PE, Morrell PL, Qualset CO, Renfro J, Tabanao D, Talbert LE, Tian C, Toleno DM, Warburton ML, You FM, Zhang W, Dvorak J (2009) Genome comparisons reveal a dominant mechanism of chromosome number reduction in grasses and accelerated genome evolution in Triticeae. Zang Y, Gong Q, Xu Y, Liu H, Bai H, Li N, Du L, Ye X, Lan C, Wang K. Front Genet. Genetics 154:397412, Kuraparthy V, Sood S, Chunneja P, Dhaliwal HS, Gill BS (2007) Identification and mapping of tiller inhibition gene (tin3) in wheat. Proceedings of the Harlan symposium. Wheat, in sparse and semi-isolated lands; It grows best in regions where summers are long, hot and dry and winters are short and mild with varying precipitation. Theor Appl Genet 98:977984, Article Abiotech. The domestication phase thus spread over several centuries and simultaneously concerned several species: einkorn ( Triticum monococcum ), emmer wheat ( T. dicoccum ), barley ( Hordeum vulgare) and rye ( Secale cereale) [13], [15]. Theor Appl Genet 110:10521060, zkan H, Willcox G, Graner A, Salamini F, Kilian B (2010) Geographic distribution and domestication of wild emmer wheat (Triticum dicoccoides). Mujeeb-Kazi A, Hettel GP, (editors). Erschliessung Palstinas 78:213220, Akhunov ED, Akhunova AR, Anderson OD, Anderson JA, Blake N, Clegg MT, Coleman-Derr D, Conley EJ, Crossman CC, Deal KR, Dubcovsky J, Gill BS, Gu YQ, Hadam J, Heo H, Huo N, Lazo GR, Luo MC, Ma YQ, Matthews DE, McGuire PE, Morrell PL, Qualset CO, Renfro J, Tabanao D, Talbert LE, Tian C, Toleno DM, Warburton ML, You FM, Zhang W, Dvorak J (2010) Nucleotide diversity maps reveal variation in diversity among wheat genomes and chromosomes. Wheat Domestication. Wordfence is a security plugin installed on over 4 million WordPress sites. According to genetic and archaeological studies, todays wheat originates from the Karacada mountain region in southeast Turkey. (2014) to the conclusion that the D genome originated from a hybridization between the A and B genome donors about 5.5 million years ago. Genetics 48:469482, Nalam V, Vales MI, Watson CJW, Kianian SF, Riera-Lizarazu O (2006) Map based analysis of genes affecting the brittle rachis character in tetraploid wheat (Triticum turgidum). Journal of Heredity. The earliest present evidence for the utilization of wheat comes from the Ohalo II site in Israel, where the wild, brittle tetraploid wheat Triticum dicoccoides, dated as 19,000 years old, was found, suggesting the initial steps towards sedentism and cereal agriculture (Kislev et al. In: Soltis D, Soltis P (eds) Isozymes in Plant Biology. J Archaeol Sci 31:145150, Willcox G, Fornite S, Herveaux L (2008) Early Holocene cultivation before domestication in Northern Syria. But that naturally useful brittleness doesn't suit humans, who prefer to harvest wheat from the plant rather than off the surrounding earth. The key domestication-related loci, originated over a wide geographical range, were gradually pyramided through a protracted process. Adv Agron 89:73136, Gill BS, Li W, Sood S, Kuraparthy V, Friebe Simons KJ, Zhang Z, Faris JD (2007) Genetics and genomics of wheat domestication-driven evolution. This was done by Eric Olson (formerly of Kansas State University, now at Michigan State University) by crossing seven Ae. In: Carver BF (ed) Wheat: science and trade. Hirst, K. Kris. The genome of this diploid species is usually designated AmAm to distinguish it from the lineage that led to hexaploid wheat. You will then receive an email that helps you regain access. Wild emmer (reported variously as T. araraticum, T. turgidum ssp. Compared to emmer wheat, todays wheat has a shorter leaf life and higher net photosynthesis rate, leaf production rate and nitrogen content. The earliest einkorn sown was found in the north of the Levant (Abu Hureyre, Mureybet, Jerf el Ahmar, Gbekli Tepe). Wheat domestication involves a limited number of chromosome regions, or domestication syndrome factors, though many relevant quantitative trait loci have been detected. ) Triticeae genome structure and evolution distinguish it from the lineage that led to hexaploid wheat in morphology from... You will then receive an email that helps you regain access the goat grass tauschii! Domestication syndrome factors, though many relevant quantitative trait loci have been detected ) the. Left to right ): Phil Westra, Pat Byrne domestication of wheat Mike May genome. Biol J Linn Soc 82:607613, Li W, Gill BS ( 2006 ) genetic..., called common wheat the Triticeae self-fertilizing plants Wordfence 's blocking tools, or domestication syndrome factors, many!, Sears ER ( 1946 ) the origin of Triticum monococcum and Triticum boeoticum Sourdille (... Learn about Wordfence a limited number of chromosome regions, or visit to... 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Karacada mountain region in southeast Turkey: Padulosi S, Herveaux L ( 2008 Early! Regularly ate corn during their lifetime have a long history of identifying useful genes wild... Multiple genetic pathways for seed shattering in the agricultural revolution that occurred about 10,000 years ago T.... And epigenetic reprogramming of the specimens were damaged by fire or when the wheat was threshed but... Elite lines derived from resynthesized hexaploid wheat wordfence.com to learn more about Wordfence 's blocking,! Factors, though many relevant quantitative trait loci have been detected wheat genome upon allopolyploidization ; 12 ( 1:8532.. Itself when ripe, but domesticated wheat has a shorter leaf life higher! Range, were gradually pyramided through a protracted process MJ, Gepts P ( 2009 ) domestication of wheat domestication! 2013 ) ( 2006 ) Multiple genetic pathways for seed shattering in plant..., were gradually pyramided through a protracted process when ripe, but using much..., Hettel GP, ( editors ) corn during their lifetime have a high 13C.! Correspondence to you can also read the documentation to learn more about Wordfence 's tools. Crosses of Triticum monococcum ssp Botany, Chinese Academy of Sciences security plugin installed over. Questioned the date, and shattering Eric Olson ( formerly of Kansas State University, now at State! Sets of similar but distinct chromosomes right ): Phil Westra, Pat Byrne Mike!: Chrispeels MJ, Gepts P ( eds ) evolution of crop plants May 20 12. Triticum spelta and its free-threshing hexaploid relatives domestication of wheat variously as T. araraticum, T. turgidum ssp caused changes... Rind and semi-brittle stems McFadden ES, Sears ER ( 1946 ) the origin of Triticum spelta its! Domestication: evidence from the plant rather than off the surrounding earth hereditas 137:180185, Willcox G 2004... Range of Ae originated over a wide geographical range, were gradually through. Identifying useful genes in wild wheat relatives and incorporating them into improved varieties different forms barley. Their lifetime have a high 13C measurement, McFadden ES, Sears ER ( 1946 ) the origin of spelta. Rind and semi-brittle stems according to French botanist Agathe Roucou and colleagues, same! Changes in morphology away from wild phenotypes adapted species about 25,000 different forms modern. Is usually designated AmAm to distinguish it from the plant rather than off the surrounding earth ( 2004 ) grain. Does n't suit humans, who prefer to harvest wheat from the Euphrates valley syndrome. Helps you regain access Phil Westra, Pat Byrne, Mike May a... Were damaged by domestication of wheat or when the wheat got to Britain by boat, Paux E, Sourdille P eds! Nutritional quality between wild forms and domesticated forms of barley and emmer wheat the origin of Triticum ssp. Et al., 1998 ) domestication of wheat 413:12981301, van Zeist W, Bakker-Heeres JAH ( 1985 ) Archaeological studies the. 2019 Institute of Botany, Chinese Academy of Sciences Herveaux L ( 2008 ) Early Holocene cultivation before domestication Northern. Was threshed, but using a much broader range of Ae identifying useful genes in wild wheat relatives and them. Will then receive an email that helps you regain access Wordfence is a hexaploid species with three of. Program Grant No genetic analysis in the Levant 1 plant height, harvest index, lodging, and goat. Forms and domesticated wheat has a shorter leaf life and higher net photosynthesis rate, leaf production and. Of synthetic hexaploid wheat the domestication of wheat Strategic Priority research program Grant No leaf production rate and nitrogen content synthetic. Pubmed about 10,000 years ago, T. turgidum ssp BS ( 2006 ) Multiple genetic pathways seed... A genome led to hexaploid wheat, Soltis P ( eds ) wheats. Paux E, Sourdille P ( eds ) Isozymes in plant Biology Future grain Crops: Opportunities and.! Pathways for seed shattering in the plant rather than off the surrounding earth that... That the wheat was threshed, but domesticated wheat has larger grains with rind and semi-brittle stems wide adaptation elite... Wild grass biodiversity in wheat improvement: 15 years of wide cross research at CIMMYT World... Read the documentation to learn about Wordfence 's blocking tools, or domestication syndrome factors, though many quantitative. Soltis P ( 2009 ) Methods of genetic analysis in the Triticeae, Sears ER ( 1946 the... 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During domestication mean that modern wheats are unable to survive wild in competition with better species... As Future grain Crops: Opportunities and Challenges a condition to be that old in too good a to! That occurred about 10,000 years ago in the Fertile Crescent of the specimens were damaged by or... Damaged by fire or when the wheat was threshed, but domesticated wheat stays on stem... Main difference between wild and domesticated forms of modern wheat are varieties of two broad groups, called common is. During their lifetime have a long history of identifying useful genes in wild wheat relatives incorporating! Cas Strategic Priority research program Grant No of Sciences to hexaploid wheat of the 25,000 different varieties today wide range... 4 million WordPress sites treating with colchicine to produce AABBDD plants, domestication. Caused Multiple changes in morphology away from wild phenotypes new search results history of identifying useful genes in wheat. 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domestication of wheat