Theoretical and Applied Genetics

, Volume 126, Issue 11, pp 2777–2789 | Cite as

Cbf14 copy number variation in the A, B, and D genomes of diploid and polyploid wheat

  • Taniya Dhillon
  • Eric J. StockingerEmail author
Original Paper


Freezing tolerance and winter hardiness are complex traits. In the Triticeae, two loci on the group 5 chromosome homoeologs are repeatedly identified as having major effects on these traits. Recently, we found that segments of the genomic region at one of these loci, Frost resistance-2 (Fr-2) is copy number variable in barley. Freezing-tolerant winter-hardy genotypes have greater tandem copy numbers of the genomic region encompassing the C-repeat binding factor genes Cbf2A and Cbf4B at Fr-H2 than the less freezing-tolerant nonwinter-hardy genotypes. Here we report that in wheat the Cbf14 gene at Fr-2 is copy number variable. Using DNA blot hybridizations, we estimated copy numbers of Cbf14 across the different genomes of diploid and polyploid wheat. Copy numbers of Cbf14 are lower in the B genome than in the A and D genomes across all ploidy levels. Among hexaploid red wheats, winter genotypes harbor greater Cbf14 copy numbers than spring genotypes. Cbf14 copy numbers also vary across the red winter wheats such that hard wheats harbor greater copy numbers than soft wheats. Analysis of hexaploid wheat chromosome 5 substitution lines indicates that Cbf14 copy numbers in the introgressions are stable in the different backgrounds. Taken together our data suggest that higher copy number states existed in the diploid wild ancestors prior to the polyploidization events and that the loss of Cbf14 copies occurred in the cultivated germplasm.


Hexaploid Wheat Freezing Tolerance Substitution Line Tetraploid Wheat Diploid Wheat 
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 Drs. Harold E. Bockelman, Bikram S. Gill, David F. Garvin, P. Stephen Baenziger, and Carl A. Griffey for providing wheat accessions and genetic stocks. We thank Alexandra Shaffner for DNA extractions. We also thank the anonymous reviewers for helpful suggestions and Laura J. Chapin for help in making changes to some of the figures during the revision. This work was supported by grants from SEEDS: OARDC Research Enhancement Competitive Grants Program (2009127) and the Ohio Plant Biotechnology Consortium (2010-011). Salaries and research support in the Stockinger laboratory provided by state and federal funds appropriated to The Ohio State University, Ohio Agricultural Research and Development Center.

Supplementary material

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Supplementary material (DOCX 29113 kb)


  1. Akhunov ED, Akhunova AR, Dvorak J (2005) BAC libraries of Triticum urartu, Aegilops speltoides and Ae. tauschii, the diploid ancestors of polyploid wheat. Theor Appl Genet 111:1617–1622PubMedCrossRefGoogle Scholar
  2. Ausubel FM, Brent R, Kingston RE, Moore DD, Seidmen JG, Smith JA, Struhl K (1993) Current protocols in molecular biology. Greene Publishing Associates/Wiley, New YorkGoogle Scholar
  3. Båga M, Chodaparambil SV, Limin AE, Pecar M, Fowler DB, Chibbar RN (2007) Identification of quantitative trait loci and associated candidate genes for low-temperature tolerance in cold-hardy winter wheat. Funct Integr Genomics 7:53–68PubMedCrossRefGoogle Scholar
  4. Braun HJ, Sãulescu NN (2002) Breeding winter and facultative wheat. In: Curtis BC, Rajaram S, Gómez Macpherson H (eds) Bread wheat: improvement and production. Food and Agriculture Organization of the United Nations, Rome, pp 217–226Google Scholar
  5. Cahalan C, Law CN (1979) The genetical control of cold resistance and vernalisation requirement in wheat. Heredity 42:125–132CrossRefGoogle Scholar
  6. Carver BF, Krenzer EG, Hunger RM, Martin TJ, Klatt AR, Porter DR, Verchot J, Rayas-Duarte P, Guenzi AC, Martin BC, Bai G (2003) Registration of ‘Intrada’ wheat. Crop Sci 43:1135–1136CrossRefGoogle Scholar
  7. Chantret N, Salse J, Sabot F, Rahman S, Bellec A, Laubin B, Dubois I, Dossat C, Sourdille P, Joudrier P, Gautier MF, Cattolico L, Beckert M, Aubourg S, Weissenbach J, Caboche M, Bernard M, Leroy P, Chalhoub B (2005) Molecular basis of evolutionary events that shaped the hardness locus in diploid and polyploid wheat species (Triticum and Aegilops). Plant Cell 17:1033–1045PubMedCrossRefGoogle Scholar
  8. Charmet G (2011) Wheat domestication: lessons for the future. CR Biol 334:212–220CrossRefGoogle Scholar
  9. Clark JA (1931) Registration of improved wheat varieties, VI. J Am Soc Agron 23:1010–1012CrossRefGoogle Scholar
  10. Cook DE, Lee TG, Guo X, Melito S, Wang K, Bayless AM, Wang J, Hughes TJ, Willis DK, Clemente TE, Diers BW, Jiang J, Hudson ME, Bent AF (2012) Copy number variation of multiple genes at Rhg1 mediates nematode resistance in soybean. Science 338:1206–1209PubMedCrossRefGoogle Scholar
  11. Cox TS (1991) The contribution of introduced germplasm to the development of U.S. wheat cultivars. In: Shands HL, Wiesner LE (eds) Use of plant introductions in cultivar development: proceedings of a symposium. Crop Science Society of America, Inc., Madison, pp 25–47Google Scholar
  12. Curtis BC (2002) Wheat in the world. In: Curtis BC, Rajaram S, Gómez Macpherson H (eds) Bread wheat: improvement and production. Food and Agriculture Organization of the United Nations, Rome, pp 1–17Google Scholar
  13. DeBolt S (2010) Copy number variation shapes genome diversity in Arabidopsis over immediate family generational scales. Genome Biol Evol 2:441–453PubMedCrossRefGoogle Scholar
  14. Dhillon T, Pearce S, Stockinger E, Distelfeld A, Li C, Knox AK, Vashegyi I, Vagujfalvi A, Galiba G, Dubcovsky J (2010) Regulation of freezing tolerance and flowering in temperate cereals: the VRN-1 connection. Plant Physiol 153:1846–1858PubMedCrossRefGoogle Scholar
  15. Díaz A, Zikhali M, Turner AS, Isaac P, Laurie DA (2012) Copy number variation affecting the Photoperiod-B1 and Vernalization-A1 genes is associated with altered flowering time in wheat (Triticum aestivum). PLoS ONE 7:e33234PubMedCrossRefGoogle Scholar
  16. Distelfeld A, Li C, Dubcovsky J (2009) Regulation of flowering in temperate cereals. Curr Opin Plant Biol 12:178–184PubMedCrossRefGoogle Scholar
  17. Flood RG, Halloran GM (1986) Genetics and physiology of vernalization response in wheat. Adv Agron 39:87–125CrossRefGoogle Scholar
  18. Fowler DB, Gusta LV (1979) Selection for winterhardiness in wheat. 1. Identification of genotypic variability. Crop Sci 19:769–772CrossRefGoogle Scholar
  19. Fowler DB, Limin AE (2004) Interactions among factors regulating phenological development and acclimation rate determine low-temperature tolerance in wheat. Ann Bot (Lond) 94:717–724CrossRefGoogle Scholar
  20. Fowler DB, Dvorak J, Gusta LV (1977) Comparative cold hardiness of several Triticum species and Secale cereale L. Crop Sci 17:941–943CrossRefGoogle Scholar
  21. Francia E, Rizza F, Cattivelli L, Stanca AM, Galiba G, Toth B, Hayes PM, Skinner JS, Pecchioni N (2004) Two loci on chromosome 5H determine low-temperature tolerance in a ‘Nure’ (winter) × ‘Tremois’ (spring) barley map. Theor Appl Genet 108:670–680PubMedCrossRefGoogle Scholar
  22. Francia E, Barabaschi D, Tondelli A, Laido G, Rizza F, Stanca AM, Busconi M, Fogher C, Stockinger EJ, Pecchioni N (2007) Fine mapping of a HvCBF gene cluster at the frost resistance locus Fr-H2 in barley. Theor Appl Genet 115:1083–1091PubMedCrossRefGoogle Scholar
  23. Galiba G, Quarrie SA, Sutka J, Morounov A, Snape JW (1995) RFLP mapping of the vernalization (Vrn1) and frost resistance (Fr1) genes on chromosome 5A of wheat. Theor Appl Genet 90:1174–1179CrossRefGoogle Scholar
  24. Galiba G, Stockinger EJ, Francia E, Milc J, Kocsy G, Pecchioni N (2013) Freezing tolerance in the Triticeae. In: Varshney RK, Tuberosa R (eds) Translational genomics for crop breeding. Wiley, USA, pp 99–124 Google Scholar
  25. Gautier MF, Cosson P, Guirao A, Alary R, Joudrier P (2000) Puroindoline genes are highly conserved in diploid ancestor wheats and related species but absent in tetraploid Triticum species. Plant Sci 153:81–91CrossRefGoogle Scholar
  26. Giles RJ, Brown TA (2006) GluDy allele variations in Aegilops tauschii and Triticum aestivum: implications for the origins of hexaploid wheats. Theor Appl Genet 112:1563–1572PubMedCrossRefGoogle Scholar
  27. Gill BS, Appels R, Botha-Oberholster AM, Buell CR, Bennetzen JL, Chalhoub B, Chumley F, Dvorak J, Iwanaga M, Keller B, Li W, McCombie WR, Ogihara Y, Quetier F, Sasaki T (2004) A workshop report on wheat genome sequencing: international Genome Research on Wheat Consortium. Genetics 168:1087–1096PubMedCrossRefGoogle Scholar
  28. Gilmour SJ, Zarka DG, Stockinger EJ, Salazar MP, Houghton JM, Thomashow MF (1998) Low temperature regulation of the Arabidopsis CBF family of AP2 transcriptional activators as an early step in cold-induced COR gene expression. Plant J 16:433–442PubMedCrossRefGoogle Scholar
  29. Greenup A, Peacock WJ, Dennis ES, Trevaskis B (2009) The molecular biology of seasonal flowering-responses in Arabidopsis and the cereals. Ann Bot 103:1165–1172PubMedCrossRefGoogle Scholar
  30. Gusta LV, Oconnor BJ, MacHutcheon MG (1997) The selection of superior winter-hardy genotypes using a prolonged freeze test. Can J Plant Sci 77:15–21CrossRefGoogle Scholar
  31. Gusta LV, O’Connor BJ, Gao YP, Jana S (2001) A re-evaluation of controlled freeze-tests and controlled environment hardening conditions to estimate the winter survival potential of hardy winter wheats. Can J Plant Sci 81:241–246CrossRefGoogle Scholar
  32. Haley SD, Quick JS, Martin TJ, Johnson JJ, Peairs FB, Stromberger JA, Clayshulte SR, Clifford BL, Rudolph JB (2003) Registration of ‘Avalanche’ wheat. Crop Sci 43:432 Google Scholar
  33. Hall MD, Rohrer-Perkins W, Griffey CA, Liu SY, Thomason WE, Abaye AO, Bullard-Schilling A, Gundrum PG, Fanelli JK, Chen J, Brooks WS, Seago JE, Will BC, Hokanson EG, Behl HD, Pitman RM, Kenner JC, Vaughn ME, Corbin RA, Dunaway DW, Lewis TR, Starner DE, Gulick SA, Beahm BR, Whitt DL, Lafferty JB (2011) Registration of ‘Snowglenn’ winter durum wheat. J Plant Reg 5:81–86CrossRefGoogle Scholar
  34. Hayes PM, Blake T, Chen THH, Tragoonrung S, Chen F, Pan A, Liu B (1993) Quantitative trait loci on barley (Hordeum vulgare L.) chromosome 7 associated with components of winterhardiness. Genome 36:66–71PubMedCrossRefGoogle Scholar
  35. Ibrahim AMH, Haley SD, Baenziger PS, Jin Y, Langham MAC, Rickertsen J, Kalsbeck S, Little R, Ingemansen J, Chung OK, Seabourn BW, Bai GH, Chen MS, McVey DV (2008) Registration of ‘Alice’ Wheat. J Plant Reg 2:110–114CrossRefGoogle Scholar
  36. Knox AK, Li C, Vagujfalvi A, Galiba G, Stockinger EJ, Dubcovsky J (2008) Identification of candidate CBF genes for the frost tolerance locus Fr-A m 2 in Triticum monococcum. Plant Mol Biol 67:257–270PubMedCrossRefGoogle Scholar
  37. Knox AK, Dhillon T, Cheng H, Tondelli A, Pecchioni N, Stockinger EJ (2010) CBF gene copy number variation at Frost Resistance-2 is associated with levels of freezing tolerance in temperate-climate cereals. Theor Appl Genet 121:21–35PubMedCrossRefGoogle Scholar
  38. Korbel JO, Kim PM, Chen X, Urban AE, Weissman S, Snyder M, Gerstein MB (2008) The current excitement about copy-number variation: how it relates to gene duplications and protein families. Curr Opin Struct Biol 18:366–374PubMedCrossRefGoogle Scholar
  39. Limin AE, Fowler DB (1981) Cold hardiness of some relatives of hexaploid wheat. Can J Bot 59:572–573CrossRefGoogle Scholar
  40. Limin AE, Fowler DB (1982) The expression of cold hardiness in Triticum species amphiploids. Can J Genet Cytol 24:51–56Google Scholar
  41. Liu Q, Kasuga M, Sakuma Y, Abe H, Miura S, Yamaguchi-Shinozaki K, Shinozaki K (1998) Two transcription factors, DREB1 and DREB2, with an EREBP/AP2 DNA binding domain separate two cellular signal transduction pathways in drought- and low-temperature-responsive gene expression, respectively, in Arabidopsis. Plant Cell 10:1391–1406PubMedGoogle Scholar
  42. Luo MC, Yang ZL, You FM, Kawahara T, Waines JG, Dvorak J (2007) The structure of wild and domesticated emmer wheat populations, gene flow between them, and the site of emmer domestication. Theor Appl Genet 114:947–959PubMedCrossRefGoogle Scholar
  43. MacDonald MD (1987) Registration of two winter wheat disomic whole chromosome substitution germplasm lines. Crop Sci 27:1097CrossRefGoogle Scholar
  44. Martin TJ, Sears RG, Seifers DL, Harvey TL, Witt MD, Schlegel AJ, McCluskey PJ, Hatchett JH (2001) Registration of ‘Trego’ wheat. Crop Sci 41:929–930CrossRefGoogle Scholar
  45. Matsuoka Y (2011) Evolution of polyploid triticum wheats under cultivation: the role of domestication, natural hybridization and allopolyploid speciation in their diversification. Plant Cell Physiol 52:750–764PubMedCrossRefGoogle Scholar
  46. Matus IA, Hayes PM (2002) Genetic diversity in three groups of barley germplasm assessed by simple sequence repeats. Genome 45:1095–1106PubMedCrossRefGoogle Scholar
  47. McHale LK, Haun WJ, Xu WW, Bhaskar PB, Anderson JE, Hyten DL, Gerhardt DJ, Jeddeloh JA, Stupar RM (2012) Structural variants in the soybean genome localize to clusters of biotic stress-response genes. Plant Physiol 159:1295–1308PubMedCrossRefGoogle Scholar
  48. Miller AK, Galiba G, Dubcovsky J (2006) A cluster of 11 CBF transcription factors is located at the frost tolerance locus Fr-A m2 in Triticum monococcum. Mol Genet Genomics 275:193–203PubMedCrossRefGoogle Scholar
  49. Morris R, Schmidt JA, Mattern PJ, Johnson VA (1966) Chromosomal location of genes for flour quality in the wheat variety ‘Cheyenne’ using substitution lines. Crop Sci 6:119–122CrossRefGoogle Scholar
  50. Ozkan H, Willcox G, Graner A, Salamini F, Kilian B (2011) Geographic distribution and domestication of wild emmer wheat (Triticum dicoccoides). Genet Resour Crop Evol 58:11–53CrossRefGoogle Scholar
  51. Pike PR, MacRitchie F (2004) Protein composition and quality of some new hard white winter wheats. Crop Sci 44:173–176CrossRefGoogle Scholar
  52. Quisenberry KS, Reitz LP (1974) Turkey wheat: the cornerstone of an Empire. Agr His 48:98–110Google Scholar
  53. Ratnayaka I, Baga M, Fowler DB, Chibbar RN (2005) Construction and characterization of a BAC library of a cold-tolerant hexaploid wheat cultivar. Crop Sci 45:1571–1577CrossRefGoogle Scholar
  54. Roberts DWA (1986) Chromosomes in ‘Cadet’ and ‘Rescue’ wheats carrying loci for cold hardiness and vernalization response. Can J Genet Cytol 28:991–997Google Scholar
  55. Roberts DWA (1990) Identification of loci on chromosome 5A of wheat involved in control of cold hardiness, vernalization, leaf length, rosette growth habit, and height of hardened plants. Genome 33:247–259Google Scholar
  56. Roberts DWA, MacDonald MD (1988) Role of chromosome 5A in wheat in control of some traits associated with cold hardiness of winter wheat. Can J Bot 66:658–662CrossRefGoogle Scholar
  57. Sandve SR, Fjellheim S (2010) Did gene family expansions during the Eocene-Oligocene boundary climate cooling play a role in Pooideae adaptation to cool climates? Mol Ecol 19:2075–2088PubMedCrossRefGoogle Scholar
  58. Schilling AS, Abaye AO, Griffey CA, Brann DE, Alley MM, Pridgen TH (2003) Adaptation and performance of winter durum wheat in Virginia. Agron J 95:642–651CrossRefGoogle Scholar
  59. Sears ER (1966) Nullisomic-tetrasomic combinations in hexaploid wheat. In: Riley R, Lewis KR (eds) Chromosome manipulation and plant genetics; the contributions to a symposium held during the Tenth International Botanical Congress, Edinburgh, 1964. Oliver & Boyd, Edinburgh, pp 29–45Google Scholar
  60. Skinner JS, von Zitzewitz J, Szucs P, Marquez-Cedillo L, Filichkin T, Amundsen K, Stockinger EJ, Thomashow MF, Chen TH, Hayes PM (2005) Structural, functional, and phylogenetic characterization of a large CBF gene family in barley. Plant Mol Biol 59:533–551PubMedCrossRefGoogle Scholar
  61. Skinner JS, Szucs P, von Zitzewitz J, Marquez-Cedillo L, Filichkin T, Stockinger EJ, Thomashow MF, Chen TH, Hayes PM (2006) Mapping of barley homologs to genes that regulate low temperature tolerance in Arabidopsis. Theor Appl Genet 112:832–842PubMedCrossRefGoogle Scholar
  62. Snape JW, Semikhodskii A, Fish L, Sarma RN, Quarrie SA, Galiba G, Sutka J (1997) Mapping frost tolerance loci in wheat and comparative mapping with other cereals. Acta Agric Hung 45:265–270Google Scholar
  63. Stankiewicz P, Lupski JR (2010) Structural variation in the human genome and its role in disease. Annu Rev Med 61:437–455PubMedCrossRefGoogle Scholar
  64. Stockinger EJ, Mulinix CA, Long CM, Brettin TS, Iezzoni AF (1996) A linkage map of sweet cherry based on RAPD analysis of a microspore-derived callus culture population. J Hered 87:214–218PubMedCrossRefGoogle Scholar
  65. Stockinger EJ, Gilmour SJ, Thomashow MF (1997) Arabidopsis thaliana CBF1 encodes an AP2 domain-containing transcriptional activator that binds to the C-repeat/DRE, a cis-acting DNA regulatory element that stimulates transcription in response to low temperature and water deficit. Proc Natl Acad Sci USA 94:1035–1040PubMedCrossRefGoogle Scholar
  66. Stockinger EJ, Cheng H, Skinner JS (2006) Structural organization of barley CBF genes coincident with QTLs for cold hardiness. In: Chen THH, Uemura M, Fujikawa S (eds) Cold hardiness in plants: molecular genetics, cell biology and physiology. CABI Publishing Oxon, UK, pp 53–63Google Scholar
  67. Stockinger EJ, Skinner JS, Gardner KG, Francia E, Pecchioni N (2007) Expression levels of barley Cbf genes at the Frost resistance-H2 locus are dependent upon alleles at Fr-H1 and Fr-H2. Plant J 51:308–321PubMedCrossRefGoogle Scholar
  68. Sutka J, Snape JW (1989) Location of a gene for frost resistance on chromosome 5A of wheat. Euphytica 42:41–44CrossRefGoogle Scholar
  69. Thomashow MF (1999) Plant cold acclimation: freezing tolerance genes and regulatory mechanisms. Annu Rev Plant Physiol Plant Mol Biol 50:571–599PubMedCrossRefGoogle Scholar
  70. Thomashow MF (2010) Molecular basis of plant cold acclimation: insights gained from studying the CBF cold response pathway. Plant Physiol 154:571–577PubMedCrossRefGoogle Scholar
  71. Tóth B, Galiba G, Feher E, Sutka J, Snape JW (2003) Mapping genes affecting flowering time and frost resistance on chromosome 5B of wheat. Theor Appl Genet 107:509–514PubMedCrossRefGoogle Scholar
  72. Trevaskis B (2010) The central role of the VERNALIZATION1 gene in the vernalization response of cereals. Funct Plant Biol 37:479–487CrossRefGoogle Scholar
  73. Vágújfalvi A, Galiba G, Cattivelli L, Dubcovsky J (2003) The cold-regulated transcriptional activator Cbf3 is linked to the frost-tolerance locus Fr-A2 on wheat chromosome 5A. Mol Genet Genomics 269:60–67PubMedGoogle Scholar
  74. Vágújfalvi A, Aprile A, Miller A, Dubcovsky J, Delugu G, Galiba G, Cattivelli L (2005) The expression of several Cbf genes at the Fr-A2 locus is linked to frost resistance in wheat. Mol Genet Genomics 274:506–514PubMedCrossRefGoogle Scholar
  75. Veisz O, Sutka J (1989) The relationships of hardening period and the expression of frost resistance in chromosome subsitution lines of wheat. Euphytica 43:41–45CrossRefGoogle Scholar
  76. von Bothmer R, Sato K, Komatsuda T, Yasuda S, Fischbeck G (2003) The domestication of cultivated barley. In: von Bothmer R, van Hintum T, Knüpffer H, Sato K (eds) Diversity in barley (Hordeum vulgare), 1st edn. Elsevier, New York, pp 9–27CrossRefGoogle Scholar
  77. Wilen RW, Fu P, Robertson AJ, Gusta LV (1996) A comparison of the cold hardiness potential of spring cereals and vernalized and non-vernalized winter cereals. In: Li PH, Chen THH (eds) Fifth international plant cold hardiness seminar. Plenum Press, Corvallis, pp 191–201Google Scholar
  78. Xue GP (2003) The DNA-binding activity of an AP2 transcriptional activator HvCBF2 involved in regulation of low-temperature responsive genes in barley is modulated by temperature. Plant J 33:373–383PubMedCrossRefGoogle Scholar
  79. Yan L, Loukoianov A, Tranquilli G, Helguera M, Fahima T, Dubcovsky J (2003) Positional cloning of the wheat vernalization gene VRN1. Proc Natl Acad Sci USA 100:6263–6268PubMedCrossRefGoogle Scholar
  80. Yan L, Loukoianov A, Blechl A, Tranquilli G, Ramakrishna W, SanMiguel P, Bennetzen JL, Echenique V, Dubcovsky J (2004) The wheat VRN2 gene is a flowering repressor down-regulated by vernalization. Science 303:1640–1644PubMedCrossRefGoogle Scholar
  81. Zemetra RS, Morris R (1988) Effects of an intercultivaral chromosome substitution on winterhardiness and vernalization in wheat. Genetics 119:453–456PubMedGoogle Scholar
  82. Zemetra RS, Morris R, Schmidt JW (1986) Gene locations for heading date using reciprocal chromosome substitutions in winter wheat. Crop Sci 26:531–533CrossRefGoogle Scholar
  83. Zeven AC (1986) Landrace groups of bread wheat (Triticum aestivum L. em. Thell.). Acta Hortic 45:365–376 Google Scholar
  84. Zeven AC, van Hintum TJL (1992) Classification of landraces and improved cultivars of hexaploid wheats (Triticum aestivum, T. compactum and T. spelta) grown in the USA and described in 1922. Euphytica 59:33–47Google Scholar
  85. Zohary D (1969) The progenitors of wheat and barley in relation to domestication and agriculture dispersal in the Old World. In: Ucko PJ, Dimbleby GW (eds) The domestication and exploitation of plants and animals. Aldine Publishing Company, Chicago, pp 47–66Google Scholar
  86. Zohary D, Hopf M (2000) Domestication of plants in the old world: the origin and spread of cultivated plants in West Asia, Europe, and the Nile Valley, 3rd edn. Oxford University Press, OxfordGoogle Scholar

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© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  1. 1.Department of Horticulture and Crop ScienceThe Ohio State University/Ohio Agricultural Research and Development Center (OARDC)WoosterUSA

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