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The ‘Wheat Puzzle’ and Kartvelians route to the Caucasus Tengiz Beridze Genetic Resources and Crop Evolution An International Journal ISSN 0925-9864 Genet Resour Crop Evol DOI 10.1007/s10722-019-00759-9 1 23 Your article is protected by copyright and all rights are held exclusively by Springer Nature B.V.. This e-offprint is for personal use only and shall not be self-archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Genet Resour Crop Evol https://doi.org/10.1007/s10722-019-00759-9 (0123456789().,-volV)( 0123456789().,-volV) RESEARCH ARTICLE The ‘Wheat Puzzle’ and Kartvelians route to the Caucasus Tengiz Beridze Received: 4 October 2018 / Accepted: 21 February 2019 Ó Springer Nature B.V. 2019 Abstract Wheat (Triticum L.) originated in the Introduction Fertile Crescent approximately 10 kya BP and has since spread worldwide. The ‘Wheat Puzzle’ was South Caucasus (notably Georgia) and their earlier termed the observation that wild predecessors of five residents played an important role in wheat formation. Georgian endemic wheat subspecies are found in 17 domesticated species and subspecies of Triticum Fertile crescent quite far from the South Caucasus. are known (Table 1). Diploid-1; Tetraploid-9, from Kartvelian peoples are the ethno-linguistic group of which Georgian endemic 3; Hexaploid-7, from which speakers of Kartvelian languages (Georgian, Megrel, Georgian endemic 2. (https://en.wikipedia.org/wiki/ Laz, Svan). Kartvelian language is one of the seven Taxonomy_of_wheat). language families of Eurasiatic superfamily, that may Georgian endemic wheat species include one have arisen from a common ancestor over 15 kya BP Triticum species and four subspecies (Menabde (Pagel et al. in Proc Natl Acad Sci USA 1948; Menabde 1961; Hammer et al. 2011): 110:8471–8476, 2013). One of the possibility to 1. Triticum turgidum subsp. palaeocolchicum (Men- explain ‘Wheat Puzzle’ is that speakers of Pro- abde) A´ .Lo¨ve tokartvelian language could be separated from Pro- 2. Triticum turgidum subsp. carthlicum (Nevski) A´ . toeurasiatic language speakers after migration from Lo¨ve Africa to the Arabian peninsula and later moved to the 3. Triticum timopheevii subsp. zhukovskyi (Menabde northern part of Mesopotamia where wheat was & Ericzjan) L. B. Cai domesticated. Kartvelian speakers could migrate fur- 4. Triticum zhukovskyi Menabde & Ericzjan ther to South Caucasus together with domesticated 5. Triticum aestivum subsp. macha (Dekapr. & wheat subspecies. Menabde) McKey Keywords Wheat Á Domestication Á Kartvelians Á Some of these species bear an evolutionarily close Caucasus Á Migration affinity to wild wheat species or have retained some of their features (Menabde 1961). All these cultivated species and subspecies have been found in Western Georgia, except subsp. carthlicum which was pre- dominantly distributed in Eastern Georgia. All these & T. Beridze ( ) species and subspecies grew in the territory of Georgia Institute of Molecular Genetics, Agricultural University of Georgia, Tbilisi, Georgia until the middle of the last century. e-mail: [email protected] 123 Author's personal copy Genet Resour Crop Evol Table 1 Domesticated Botanical name (http://www.theplantlist.org/tpl1.1/search?q=Triticum) Genome wheat species and subspecies Diploid (29) 1 Triticum monococcum L. Am Tetraploid (49) 2 Triticum turgidum subsp. dicoccon (Schrank) Thell. BAu 3 Triticum ispahanicum Heslot BAu 4 Triticum turgidum subsp. palaeocolchicum (Menabde) A´ .Lo¨ve BAu 5 Triticum turgidum subsp. durum (Desf.) Husn. BAu 6 Triticum turgidum subsp. turgidum BAu 7 Triticum turgidum subsp. polonicum (L.) Thell. BAu 8 Triticum turgidum subsp. turanicum (Jakubz.) A´ .Lo¨ve & D. Lo¨ve BAu 9 Triticum turgidum subsp. carthlicum (Nevski) A´ .Lo¨ve BAu 10 Triticum timopheevii subsp. zhukovskyi (Menabde & Ericzjan) L. B. Cai GAm Hexaploid (69) 11 Triticum aestivum subsp. spelta (L.) Thell. BAuD 12 Triticum aestivum subsp. macha (Dekapr. & Menabde) McKey BAuD 13 Triticum aestivum subsp. vavilovii (Jakubz.) Sears BAuD 14 Triticum aestivum subsp. aestivum BAuD 15 Triticum aestivum subsp. compactum (Host) Mackey BAuD 16 Triticum sphaerococcum Percival BAuD 17 Triticum zhukovskyi Menabde & Ericzjan GAmAu On the basis of genetic and morphological evi- Earlier we termed as the ‘Zanduri Puzzle’, the dence, Georgian wheat (T. turgidum subsp. palaeo- observation that the wild T. timopheevii subsp. arme- colchicum) is assumed to be a segregant from a hybrid niacum (A´ .Lo¨ve) van Slageren was not found in cross between wild emmer wheat and T. aestivum Georgia, though cultivated T. timopheevii is detected subsp. aestivum (Dvorak and Luo 2001), whereas only here (Gogniashvili et al. 2015). This statement is subsp. carthlicum may be a segregant from a hybrid true also for both domesticated tetraploid wheats cross between domesticated emmer wheat and T. subsp. paleocolchicum and subsp. carthlicum. The aestivum (Kuckuck 1979). wild emmer was not found in Georgia (and in South The third subspecies of wheat detected in Western Caucasus), though cultivated tetraploid wheats were Georgia is hexaploid domesticated, hulled spelt wheat only detected there (Gogniashvili et al. 2018). T. aestivum subsp. macha (Dekaprelevich and Men- Mori et al. (2009) analyzed the molecular variation abde 1932). This subspecies is endemic to Georgia and at 23 microsatellite loci in the chloroplast genome of is cultivated along with tetraploid West Georgian Timopheevi wheats. None of the T. araraticum wheat T. turgidum subsp. palaeocolchicum. plastotypes collected in South Caucasus was closely Another line of Triticum species is the Timopheevi related to the T. timopheevii plastotype. However, the group, which contains the G genome. T. timopheevii plastotypes found in northern Syria and southern wheat was discovered in Western Georgia, where it Turkey showed closer relationships with T. timophee- was termed Zanduri wheat. In the past, the Zanduri vii. These results suggested that the domestication of population was a set of diploid-T. monococcum var. T. timopheevii wheat might have occurred in regions hornemannii (2n = 14) (Gvatsa Zanduri), tetraploid T. including southern Turkey and northern Syria (Mori timopheevii (2n = 28) (Chelta Zanduri) and hexaploid et al. 2009). T. zhukovskyi (2n = 42). Hexaploid wheat T. zhukovs- kyi is the result of hybridization of T. timopheevii subsp. zhukovskyi with T. monococcum L. var. horne- mannii (Menabde and Eritsian 1960). 123 Author's personal copy Genet Resour Crop Evol Wheat history data and on their own findings. The authors suggest that modern domestic tetraploid wheats derived from Wheat (Triticum L.) originated in the Fertile Crescent wild emmer lines from southeast Turkey (Ozkan et al. approximately 10 kya ago and has since spread 2011; Allaby et al. 2017). worldwide. According to Tsunewaki et al. the wheat Northeast expansion of domesticated emmer culti- genus Triticum contains six species (Wang et al. vation resulted in sympatry with Aegilops tauschii 1997): Coss. (genome DD). Approximately 7 kya ago, the hexaploid bread wheat T. aestivum L. (BBAuAuDD) 1. Diploid species: Triticum monococcum L. and arose in the South Caucasus region by allopoly- Triticum urartu Thumanjan ex Gandilyan. ploidization of the cultivated Emmer wheat Triticum 2. Tetraploid species: T. turgidum L. and T. dicoccum Schrank with the Caucasian Ae. tauschii timopheevii. subsp. strangulata (Eig) Tzvelev (Dvorak et al. 1998; 3. Hexaploid species: T. aestivum L. and T. Dubcovsky and Dvorak 2007). zhukovskyi. All Triticum species are native to the ‘Fertile Crescent’ of the Near East, which encompasses the Kartvelian ethnolinguistic history eastern Mediterranean, southeastern Turkey, northern Iraq and western Iran, and its neighboring regions of One of the central questions of wheat domestication is the South Caucasus, and northern Iran (Matsuoka which people (s) participated in wheat domestication. 2011). Who lived in Fertile Crescent 10 kya ago in the area of There are four wild species, which grow in the wheat domestication? Fertile Crescent of the Near East: The areas of wheat domestication: 1. Triticum monococcum subsp. aegilopoides Asch. T. monococcum-the Karacadag Mountain of south- & Graebn. eastern Turkey (Heun et al. 1997). 2. Triticum urartu Thumanjan ex Gandilyan. T. turgidum-southeastern Turkey (Ozkan et al. 3. Triticum turgidum subsp. dicoccoides (Ko¨rn.) 2011). Thell. T. timopheevii-northern Syria and southern Turkey 4. Triticum timopheevii subsp. armeniacum (A´ . (Mori et al. 2009). Lo¨ve) van Slageren. One of the possibility to explain ‘Wheat Puzzle’ is The diploid wheat T. monococcum L. (einkorn) is that Kartvelians lived (with other peoples) in the area among the first crops domesticated by humans in the of Fertile Crescent (possibly participated in wheat Fertile Crescent. The site of domestication of diploid domestication) and brought some wheat species and wheat (einkorn) was identified (Heun et al. 1997). subspecies further north to South
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    Proceedings of the First International Workshop on Hulled Wheats 21-22 July 1995 Castelvecchio Pascoli, Tuscany, Italy S. Padulosi, K. Hammer and J. Heller, editors ii HULLED WHEATS The International Plant Genetic Resources Institute (IPGRI) is an autonomous international scientific organization operating under the aegis of the Consultative Group on International Agricultural Research (CGIAR). The international status of IPGRI is conferred under an Establishment Agreement which, by December 1995, had been signed by the Governments of Australia, Belgium, Benin, Bolivia, Burkina Faso, Cameroon, China, Chile, Congo, Costa Rica, Côte d’Ivoire, Cyprus, Czech Republic, Denmark, Ecuador, Egypt, Greece, Guinea, Hungary, India, Iran, Israel, Italy, Jordan, Kenya, Mauritania, Morocco, Pakistan, Panama, Peru, Poland, Portugal, Romania, Russia, Senegal, Slovak Republic, Sudan, Switzerland, Syria, Tunisia, Turkey, Ukraine and Uganda. IPGRI's mandate is to advance the conservation and use of plant genetic resources for the benefit of present and future generations. IPGRI works in partnership with other organizations, undertaking research, training and the provision of scientific and technical advice and information, and has a particularly strong programme link with the Food and Agriculture Organization of the United Nations. Financial support for the agreed research agenda of IPGRI is provided by the Governments of Australia, Austria, Belgium, Canada, China, Denmark, France, Germany, India, Italy, Japan, the Republic of Korea, Mexico, the Netherlands, Norway, Spain, Sweden, Switzerland, the UK and the USA, and by the Asian Development Bank, IDRC, UNDP and the World Bank. The Institute of Plant Genetics and Crop Plant Research (IPK) is operated as an independent foundation under public law. The foundation statute assigns to IPK the task of conducting basic research in the area of plant genetics and research on cultivated plants.