Theoretical and Applied Genetics

, Volume 114, Issue 6, pp 947–959 | Cite as

The structure of wild and domesticated emmer wheat populations, gene flow between them, and the site of emmer domestication

  • M.-C. Luo
  • Z.-L. Yang
  • F. M. You
  • T. Kawahara
  • J. G. Waines
  • J. DvorakEmail author
Original Paper


The domestication of emmer wheat (Triticum turgidum spp. dicoccoides, genomes BBAA) was one of the key events during the emergence of agriculture in southwestern Asia, and was a prerequisite for the evolution of durum and common wheat. Single- and multilocus genotypes based on restriction fragment length polymorphism at 131 loci were analyzed to describe the structure of populations of wild and domesticated emmer and to generate a picture of emmer domestication and its subsequent diffusion across Asia, Europe and Africa. Wild emmer consists of two populations, southern and northern, each further subdivided. Domesticated emmer mirrors the geographic subdivision of wild emmer into the northern and southern populations and also shows an additional structure in both regions. Gene flow between wild and domesticated emmer occurred across the entire area of wild emmer distribution. Emmer was likely domesticated in the Diyarbakir region in southeastern Turkey, which was followed by subsequent hybridization and introgression from wild to domesticated emmer in southern Levant. A less likely scenario is that emmer was domesticated independently in the Diyarbakir region and southern Levant, and the Levantine genepool was absorbed into the genepool of domesticated emmer diffusing from southeastern Turkey. Durum wheat is closely related to domesticated emmer in the eastern Mediterranean and likely originated there.


Amplify Fragment Length Polymorphism Durum Wheat Emmer Wheat Wild Emmer Emmer Accession 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



We thank O.D. Anderson, M.D. Gale, A. Graner, G.E. Hart, A. Kleinhofs, M.E. Sorrells, and M.K. Walker-Simmons for sharing clones with us, H.E. Bockelman, S. Jana, B.L. Johnson, C.O. Qualset, R. Papa, and J. Valkoun for supplying the seeds of plants used in this study. We also thank E.D. Akhunov for valuable assistance with statistical analyses and P. Morrell for valuable discussions. Financial support from USDA/ACSREES/NRICGP by grant 99-35301-7905 to J. Dvorak is acknowledged.

Supplementary material

122_2006_474_MOESM1_ESM.rtf (52 kb)
S1. Plant accessions used in the study and their original locations (RTF 52.1 kb)


  1. Allaby RG, Brown TA (2003) AFLP data and the origins of domesticated crops. Genome 46:448–453PubMedCrossRefGoogle Scholar
  2. Ammerman AJ, Cavalli-Sforza LL (1984) The neolithic transition and the genetics of populations in Europe. Princeton University Press, PrincetonGoogle Scholar
  3. Badr A, Muller K, Schafer-Pregl R, El Rabey H, Effgen S, Ibrahim HH, Pozzi C, Rohde W, Salamini F (2000) On the origin and domestication history of barley (Hordeum vulgare). Mol Biol Evol 17:499–510PubMedGoogle Scholar
  4. Blumler MA (1998) Introgression of durum in to wild emmer and the agricultural origin question. In: Damania AB, Valkoun, J, Willcox G, Qualset CO (eds) The origins of agriculture and crop domestication, ICARDA, Aleppo, Syria, ICARDA, IPGRI, FAO and UC/GRCP, pp 252–268Google Scholar
  5. de Moulins D (1993) Les restes de plantes carbonisees de Cafer Hoyuk. Cahiers de l’Euphrate 7:191–234Google Scholar
  6. Dubcovsky J, Luo MC, Zhong GY, Bransteitter R, Desai A, Kilian A, Kleinhofs A, Dvorak J (1996) Genetic map of diploid wheat, Triticum monococcum L., and its comparison with maps of Hordeum vulgare L. Genetics 143:983–999PubMedGoogle Scholar
  7. Dvorak J (1980) Homoeology between Agropyron elongatum chromosomes and Triticum aestivum chromosomes. Can J Genet Cytol 22:237–259Google Scholar
  8. Dvorak J, McGuire PE, Cassidy B (1988) Apparent sources of the A genomes of wheats inferred from the polymorphism in abundance and restriction fragment length of repeated nucleotide sequences. Genome 30:680–689Google Scholar
  9. Dvorak J, Luo M-C, Yang Z-L (1998) Restriction fragment length polymorphism and divergence in the genomic regions of high and low recombination in self-fertilizing and cross-fertilizing Aegilops species. Genetics 148:423–434PubMedGoogle Scholar
  10. Dvorak J, Yang Z-L, You FM, Luo MC (2004) Deletion polymorphism in wheat chromosome regions with contrasting recombination rates. Genetics 168:1665–1675PubMedCrossRefGoogle Scholar
  11. Dvorak J, Akhunov ED, Akhunov AR, Deal KR, Luo MC (2006) Molecular characterization of a diagnostic DNA marker for domesticated tetraploid wheat provides evidence for gene flow from wild tetraploid wheat to hexaploid wheat. Mol Biol Evol 23:1386–1396PubMedCrossRefGoogle Scholar
  12. Falush D, Pritchard J, Stephens M (2003) Inference of population structure using multilocus genotype data: linked loci and correlated allele frequencies. Genetics 164:1567–1587PubMedGoogle Scholar
  13. Heun M, Schafer-Pregel R, Klawan D, Castagna R, Accerbi M, Borghi B, Salamini F (1997) Site of einkorn wheat domestication identified by DNA fingerprinting. Science 278:1312–1314CrossRefGoogle Scholar
  14. Hopf M (1983) The plants found at Jericho. In: Kenyon KM, Holland TA (eds) Excavations at Jericho V. British School of Archaeology in Jerusalem, London, pp 580–621Google Scholar
  15. Hourani GF (1963) Arab seafaring in the Indian Ocean in ancient and early medieval times. Princeton Oriental Studies, vol 13. Princeton University Press, PrincetonGoogle Scholar
  16. Lewis PO, Zaykin D (1997) Genetic data analysis: Computer program for the analysis of allelic data. Version 1.0. A free program distributed by the authors over the Internet from the GDA Home Page at
  17. Luo MC, Deal KR, Young ZL, Dvorak J (2005) Comparative genetic maps reveal extreme crossover localization in the Aegilops speltoides chromosomes. Theor Appl Genet 111:1098–1106PubMedCrossRefGoogle Scholar
  18. Mori N, Ishi T, Ishido T, Hirosawa S, Watatani H, Kawahara T, Nesbitt M, Belay G, Takumi S, Ogihara Y, Nakamura C (2003) Origins of domesticated emmer and common wheat inferred from chloroplast DNA fingerprinting. In: Pogna NE, Romano M, Pogna EA, Galterio G (eds) Proceedings of the 10th international wheat genet symposium, Paestum, Italy. Instituto Sperimentale per la Cerealicoltura, Rome, Italy, pp 25–28Google Scholar
  19. Nei M (1978) Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89:583–590PubMedGoogle Scholar
  20. Nesbitt M, Samuel D (1996) From staple crop to extinction? The archaeology and history of hulled wheats. In: Padulosi S, Hammer K, Heller J (eds) Hulled wheats. Promoting the conservation and use of underutilized and neglected crops 4 Proc 1st Internatl Workshop on hulled wheats, Castelvecchio Pacoli, Tuscany, Italy. International Plant Genetic Resources Institute, Rome, ItalyGoogle Scholar
  21. Ozkan H, Brandolini A, Schafer-Pregl R, Salamini F (2002) AFLP analysis of a collection of tetraploid wheat indicated the origin of emmer and hard wheat domestication in southeastern Turkey. Mol Biol Evol 19:1797–1801PubMedGoogle Scholar
  22. Ozkan H, Brandolini A, Pozzi C, Effgen S, Wunder J, Salamini F (2005) A reconsideration of the domestication geography of tetraploid wheat. Theor Appl Genet 110:1052–1060PubMedCrossRefGoogle Scholar
  23. Page RDM (1996) TREEVIEW: an application to display phylogenetic trees on personal computers. Comp Appl Biosci 12:357–358PubMedGoogle Scholar
  24. Pritchard JK, Wen W (2004) Documentation for structure software,
  25. Pritchard J, Stephens M, Donnelly P (2000) Inference of population structure using multilocus genotype data. Genetics 155:945–959PubMedGoogle Scholar
  26. Rosenberg NA, Pritchard JK, Weber JL, Cann HM, Kidd KK, Zhivotovsky LA, Feldman MW (2002) Genetic structure of human populations. Science 298:2381–2385PubMedCrossRefGoogle Scholar
  27. Ryan WBF, Pitman WC (2000) Noah’s flood. Simon and Schuster, New YorkGoogle Scholar
  28. Sears ER (1966) Nullisomic-tetrasomic combinations in hexaploid wheat. In: Riley R, Lewis KR (eds) Chromosome manipulations and plant genetics. Oliver & Boyd, Edinburgh, pp 29–44Google Scholar
  29. Thuillet AC, Bru D, David J, Roumet P, Santomi S, Sourdille P, Bataillon T (2002) Direct estimation of mutation rate for 10 microsatellite loci in durum wheat, Triticum turgidum (L.) Thell. ssp durum Desf. Mol Biol Evol 19:122–125PubMedGoogle Scholar
  30. van Zeist W, Bakker-Heeres JAH (1975) Archaeobotanical studies in the Levant. 1. Neolithic sites in the Damascus basin: Aswad, Ghoraife, Ramad. Palaeohistoria 24:165–256Google Scholar
  31. van Zeist W, de Roller GJ (1991/1992) The plant husbandry of aceramic Cayonu, SE Turkey. Palaeohistoria 33/34:65–96Google Scholar
  32. Weir BB (1996) Genetic data analysis. Sinauer Assoc, SunderlandGoogle Scholar
  33. Willcox G (1998) Archaeobotanical evidence for the beginnings of agriculture in Southwest Asia. In: Damania AB, Valkoun, J, Willcox G, Qualset CO (eds) The origins of agriculture and crop domestication, ICARDA, Aleppo, Syria, ICARDA, IPGRI, FAO and UC/GRCPGoogle Scholar
  34. Willcox J (1991) Cafer Hoyuk (Turquie): Les Charbons de bois neolothiques. Cahiers de l’Euphrate 5–6:139–150Google Scholar

Copyright information

© Springer-Verlag 2007

Authors and Affiliations

  • M.-C. Luo
    • 1
  • Z.-L. Yang
    • 1
    • 2
  • F. M. You
    • 1
  • T. Kawahara
    • 3
  • J. G. Waines
    • 4
  • J. Dvorak
    • 1
    Email author
  1. 1.Department of Plant SciencesUniversity of CaliforniaDavisUSA
  2. 2.Seminis Vegetable SeedsWoodlandUSA
  3. 3.Plant Germplasm Institute, Graduate School of AgricultureKyoto UniversityKyotoJapan
  4. 4.Department of Botany & Plant SciencesUniversity of CaliforniaRiversideUSA

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