Register      Login
Crop and Pasture Science Crop and Pasture Science Society
Plant sciences, sustainable farming systems and food quality
RESEARCH ARTICLE

Triticum (Aegilops) tauschii in the natural and artificial synthesis of hexaploid wheat

G. M. Halloran A , F. C. Ogbonnaya B C E and E. S. Lagudah D
+ Author Affiliations
- Author Affiliations

A 4 James Service Place, South Melbourne, Vic. 3205, Australia.

B Primary Industries Research Victoria (PIRVic), Department of Primary Industries, Private Bag 260, Horsham, Vic. 3401, Australia.

C Molecular Plant Breeding Cooperative Research Centre, Bundoora, Vic. 3083, Australia.

D CSIRO Division of Plant Industry, Canberra, ACT 2600, Australia.

E Corresponding author. Current address: International Centre for Agricultural Research in the Dry Areas (ICARDA), PO Box 5466, Aleppo, Syria. Email: F.Ogbonnaya@cgiar.org

Australian Journal of Agricultural Research 59(5) 475-490 https://doi.org/10.1071/AR07352
Submitted: 25 August 2007  Accepted: 20 March 2008   Published: 12 May 2008

Abstract

An account is given of the possible time(s) and place(s) of the origin of hexaploid wheat from natural hybridisation between Triticum tauschii (Ae. tauschii) and both wild and cultivated forms of tetraploid wheat. A recapitulation is presented of the likely genotypic and phenotypic status of the newly arisen natural hexaploid and the likely path of hybridisation from whence it arose. Recent substantial contributions of T. tauschii to wheat improvement indicate the likelihood that introgession en masse from T. tauschii has not occurred throughout its natural and agricultural associations with wheat. This has been substantiated in comparative studies revealing higher levels of genetic variation in T. tauschii compared with the D genome of hexaploid wheat.

A case is made for a widening of the concept of the gene pool of T. tauschii for wheat improvement and the notion of a secondary gene pool is proposed to include variation in T. tauschii as it occurs in several polyploid forms of ‘grass Triticum’.

The likely differentiation of growth habit forms, conditioned by vernalisation (i.e. vrn) genes, in hexaploid wheat synthesis, including the interaction of these genes in hexaploid wheat, is discussed. It is speculated that growth habit differentiation was of significance to the hexaploid’s yield contribution and survival in tetraploid-hexaploid mixtures (likely to be a common constitution of wheat crops of early agriculture), and in the Neolithic spread of agriculture to the higher latitude, and colder environments of NW Europe and central Asia.

The significance of the contribution of T. tauschii to the unique milling and bread-making properties of hexaploid wheat is discussed in the light of Roman discernment of its closer fulfilment of the requirements of leavened bread-making compared with tetraploid wheat. The significance of the contribution of T. tauschii to the evolution of wheat appears to have been much delayed (by ~6500 years) in that hexaploid wheat did not receive singular attention and cultivation until during the Roman era, from whence it gradually rose in popularity to eventually achieve its current pre-eminent status.

Continuing systematic evaluation of genetic variation in both the primary and secondary gene pools of T. tauschii for wheat improvement, using both conventional and genetic analysis and contemporary genomic tools, is advocated. The latter approach is particularly important for quantitative traits in the light of wide divergence in plant phenotype of their representatives from that of hexaploid wheat.

Additional keywords: genetic diversity, synthetic hexaploid wheat, Aegilops tauschii, durum, Triticum aestivum.


References


Anderson JA , Waldron BL , Moreno-Sevilla B , Stack RW , Frohberg RC (1998) Detection of Fusarium head blight resistance QTL in wheat using AFLPs and RFLPs. In ‘Proceedings of the 9th International Wheat Genetics Symposium’. Vol. 1. (Ed. AE Slinkard) (University of Saskatchewan: Saskatoon, Canada)

Appels R, Lagudah ES (1990) Manipulation of chromosomal segments from wild wheat for the improvement of bread wheat. Australian Journal of Plant Physiology 17, 253–266.
Crossref | GoogleScholarGoogle Scholar | open url image1

Assefa S, Fehrmann H (2000) Resistance to wheat leaf rust in Aegilops tauschii Coss. and inheritance of resistance in hexaploid wheat. Genetic Resources and Crop Evolution 47, 135–140.
Crossref | GoogleScholarGoogle Scholar | open url image1

Assefa S, Fehrmann H (2004) Evaluation of Aegilops tauschii Coss. for resistance to wheat stem rust and inheritance of resistance genes in hexaploid wheat. Genetic Resources and Crop Evolution 51, 663–669.
Crossref | GoogleScholarGoogle Scholar | open url image1

Azizinya S, Ghanadha MR, Zali AA, Samadi BY, Ahmadi A (2005) An evaluation of quantitative traits related to drought resistance in synthetic wheat genotypes in stress and non-stress conditions. Iranian Journal of Agricultural Sciences 36, 281–293 [Persian]. open url image1

Bai D, Knott DR (1992) Suppression of resistance to leaf rust (Puccinia recondita f. sp. tritici Rob. Ex. Desm.) and stem rust (Puccinia graminis f.sp. tritici Eriks and Henn.). Genome 35, 276–282. open url image1

Bálint A, Röder MS, Hell R, Galiba G, Börner A (2007) Mapping of QTLs affecting copper tolerance and the Cu, Fe, Mn and Zn concentrations in the shoots of wheat seedlings. Biologia Plantarum 51, 129–134.
Crossref | GoogleScholarGoogle Scholar | open url image1

Brown AHD, Zohary D, Nevo E (1978) Outcrossing rates and heterozygosity in natural populations of Hordeum spontaneum koch. in Israel. Heredity 41, 49–62.
Crossref | GoogleScholarGoogle Scholar | open url image1

Cox TS, Hatchett JH, Gill BS, Raupp WJ, Sears RG (1990) Agronomic performance of hexaploid wheat lines derived from direct crosses between wheat and Aegilops squarrosa. Plant Breeding 105, 271–277.
Crossref |
open url image1

Cox TS, Bequette RK, Bowden RL, Sears RG (1997) Grain yield and breadmaking quality of wheat lines with the leaf rust resistance gene Lr41. Crop Science 37, 154–161. open url image1

Cox TS, Raupp WJ, Gill BS (1994) Leaf rust-resistance genes Lr41, Lr42, and Lr43 transferred from Triticum tauschii to common wheat. Crop Science 34, 339–343. open url image1

Cox TS, Raupp WJ, Wilson DL, Gill BS, Leath S, Bockus WW, Browder LE (1992) Resistance to foliar diseases in a collection of Triticum tauschii germplasm. Plant Disease 76, 1061–1064. open url image1

Damania AB , Tahir M (1993) Heat and cold tolerance in wild relatives and primitive forms of wheat. In ‘Evaluation and utilisation of biodiversity in wild wheat relatives and primitive forms for wheat improvement’. (Ed. AB Damania) pp. 217–224. (John Wiley and Sons: Chichester, UK)

del Blanco IA, Rajaram S, Kronstad WE (2001) Agronomic potential of synthetic hexaploid wheat-derived populations. Crop Science 41, 676–679. open url image1

Dreccer FM, Borgognone GM, Ogbonnaya FC, Trethowan RM, Winter B (2007) CIMMYT-selected derived synthetic bread wheats for rainfed environments: yield evaluation in Mexico and Australia. Field Crops Research 100, 218–228.
Crossref | GoogleScholarGoogle Scholar | open url image1

Dreccer MF , Ogbonnaya FC , Borgognone MG (2004) Sodium exclusion in primary synthetic wheats. In ‘Proceedings of the 54th Australian Cereal Chemistry Conference and 11th Wheat Breeders Assembly’. Canberra, ACT. (Eds CK Black, JF Panozzo, GJ Rebetzke) pp. 118–121. (The Royal Australian Chemical Institute, Inc.)

Dvõrák J, Luo MC, Yang ZL, Zhang AB (1998) The structure of the Aegilops tauschii genepool and the evolution of hexaploid wheat. Theoretical and Applied Genetics 97, 657–670.
Crossref | GoogleScholarGoogle Scholar | open url image1

Eastwood RF (1995) Genetics of resistance to Heterodera avenae in Triticum tauschii and its transfer to bread wheat. PhD thesis, The University of Melbourne, Australia.

Enjalbert J , Goldringer I , David J , Brabant P (1977) The relevance of outcrossing for the dynamic management of genetic resources of predominantly selfing Triticum aestivum L. (bread wheat). In ‘Colloque BRG/USTL methodologies digestion et de conservation des resources, genetiques’. Lille, France, 8–10 October 1997. Genetic, Selection, Evolution 1998 (Supplement S197-S211).

Faris JD, Anderson JA, Francl LJ, Jordahl JG (1997) RFLP mapping of resistance to chlorosis induction by Pyrenophora tritici-repentis in wheat. Theoretical and Applied Genetics 94, 98–103.
Crossref | GoogleScholarGoogle Scholar | open url image1

Feldman M, Lui B, Segal G, Abbo S, Levy AA, Vega JM (1977) Rapid elimination of low copy DNA sequences in polyploid wheat. A possible mechanism of differentiation of homoeologous chromosomes. Genetics 147, 137–139. open url image1

Flood RG, Halloran GM (1986) Genetics and physiology of vernalisation response in wheat. Advances in Agronomy 39, 87–125. open url image1

Friesen TL, Faris JD (2004) Molecular mapping of resistance to Pyrenophora tritici-repentis race 5 and sensitivity to Ptr ToxB in wheat. Theoretical and Applied Genetics 109, 464–471.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Fritz AK, Cox TX, Gill BS, Sears RG (1995) Molecular marker-facilitated analysis of introgression in winter wheat × Triticum tauschii populations. Crop Science 35, 1691–1695. open url image1

Gatford KT (2004) Seed dormancy mechanisms in diploid wheat (Triticum tauschii (Cos.) Schmalh.). PhD thesis, The University of Melbourne, Australia.

Gatford KT, Eastwood RF, Halloran GM (2002a) Germination inhibitors in bracts surrounding the grain of Triticum tauschii. Functional Plant Biology 29, 881–890.
Crossref | GoogleScholarGoogle Scholar | open url image1

Gatford KT, Hearden P, Ogbonnaya F, Eastwood RF, Halloran GM (2002b) Novel resistance to pre-harvest sprouting in Australian wheat from the wild relative Triticum tauschii. Euphytica 126, 67–76.
Crossref | GoogleScholarGoogle Scholar | open url image1

Gianibelli MC, Gupta RB, Lafiandra D, Margiofta B, MacRitchie F (2001) Polymorphism of high molecular weight glutenin subunits in Triticum tauschii; charaterisation by chromatography and electrophoretic methods. Journal of Cereal Science 33, 39–52.
Crossref | GoogleScholarGoogle Scholar | open url image1

Gill BS, Bernd F, Raupp JW, Wilson DL, Cox ST, Sears RG, Brown-Guedira GL, Fritz AK (2006) Wheat genetic resource centre: the first 25 years. Advances in Agronomy 89, 73–136. open url image1

Gill BS, Raupp WJ (1987) Direct genetic transfers from Aegilops squarrosa L. to hexaploid wheat. Crop Science 27, 445–450. open url image1

Gororo NN (1999) Growth, development and yield of wheat lines bearing Triticum tauschii germplasm. PhD thesis, The University of Melbourne, Australia.

Gororo NN, Eagles HA, Eastwood RF, Nicolas ME, Flood RG (2002) Use of Triticum tauschii to improve yield of wheat in low-yielding environments. Euphytica 123, 241–254.
Crossref |
open url image1

Guadagnuolo R, Savova-Biarichi D, Felber F (2001) Gene flow from wheat (Triticum aestivum L.) to jointed goat-grass (Aegilops cylindrica Host.) as revealed by RAPD and microsatellite markers. Theoretical and Applied Genetics 103, 1–8.
Crossref | GoogleScholarGoogle Scholar | open url image1

Halloran GM (1977) Developmental basis of maturity differences in spring wheat. Agronomy Journal 69, 889–902. open url image1

Halloran GM (1981) Genetic, physiological and adaptability studies in wheat and certain other agricultural species. Vol. 1, D. Agric. Sc. thesis, The University of Melbourne, Australia.

Halloran GM (1996) Genetic and physiological studies in crop and pasture species. Vol. 2, DSc thesis, The University of Melbourne, Australia.

Hartel KD, Berzonsky WA, Kianian SF, Ali S (2004) Expression of a Triticum turgidum var. dicoccoides source of Fusarium head blight resistance transferred to synthetic hexaploid wheat. Plant Breeding 123, 516–519.
Crossref | GoogleScholarGoogle Scholar | open url image1

He P, Friebe BR, Gill BS, Zhou JM (2003) Allopolyploidy alters gene expression in the highly stable hexaploid wheat. Plant Molecular Biology 52, 401–414.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Huang XQ, Coster H, Ganal MW, Röder MS (2003) Advanced backcross QTL analysis for the identification of quantitative trait loci alleles from wild relatives of wheat (Triticum aestivum L.). Theoretical and Applied Genetics 106, 1379–1389.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Huang XQ, Kempf H, Ganal MW, Röder MS (2004) Advanced backcross QTL analysis in progenies derived from a cross between a German elite winter wheat variety and a synthetic wheat (Triticum aestivum L.). Theoretical and Applied Genetics 109, 933–943.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Hucl P (1996) Outcrossing rates for 10 Canadian spring wheat cultivars. Canadian Journal of Plant Science 76, 423–427. open url image1

Imtiaz M, Ogbonnaya FC, Oman J, van Ginkel M (2008) Characterization of QTL controlling genetic variation for pre-harvest sprouting in synthetic backcross derived wheat lines. Genetics 178, 1725–1736.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Jaaska V (1980) Electrophoretic survey of seedling esterases in wheat in relation to their phylogeny. Theoretical and Applied Genetics 56, 273–284.
Crossref | GoogleScholarGoogle Scholar | open url image1

Jasny N (1942) Competition among grasses in classical antiquity. The American Historical Review 47, 747–764.
Crossref | GoogleScholarGoogle Scholar | open url image1

Kashkush K, Feldman M, Levy AA (2002) Gene loss, silencing and activation in a newly synthetised wheat allotetraploid. Genetics 160, 1651–1659.
PubMed |
open url image1

Kihara H (1944) Discovery of the DD analyser, one of the ancestors of Triticum vulgare. Agriculture and Horticulture 19, 889–890. open url image1

Kihara H, Yamashita K, Tanaka M, Tabushi J (1957) Some aspects of the new amphidiploids synthesized from the hybrids, Emmer wheats × Aegilops squarrosa var. strangulata. Wheat Information Service 6, 14–15. open url image1

Kimber G , Tsunewaki K (1988) Genome symbols and plasma types in the wheat group. In ‘Proceedings of the 7th International Wheat Genetics Symposium’. pp. 1209–1210. (Eds TE Miller, RMD Koebner) (Bath Press: Bath, UK)

Knaggs P, Ambrose MJ, Reader SM, Miller TE (2000) Morphological characterisation and evaluation of the subdivision of Aegilops tauschii Coss. Wheat Information Service 91, 15–19. open url image1

Kunert A, Naz AA, Dedeck O, Pillen K, Léon J (2007) AB-QTL analysis in winter wheat: I. Synthetic hexaploid wheat (T. turgidum ssp. dicoccoides × T. tauschii) as a source of favourable alleles for milling and baking quality traits. Theoretical and Applied Genetics 115, 683–695.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Kushnir U, Halloran GM (1982a) Evidence for Aegilops sharonensis Eig as the donor of the B genome of wheat. Genetics 99, 495–512. open url image1

Kushnir U, Halloran GM (1982b) Quantitative studies of the amphidiploid (Aegilops sharonensis × Triticum monococcum) and the origin of the B genome of wheat. Wheat Information Service 54, 12–16. open url image1

Lagudah ES , Appels R , McNeil D , Schachtman DP (1993) Exploiting the diploid D genome chromatin for wheat improvement. In ‘Gene Conservation and Exploitation’. (Eds JP Gustafson, R Appels, P Raven) pp. 87–107. (Plenum Press: New York)

Lagudah ES, Halloran GM (1989) Phylogenetic relationships of Triticum tauschii, the D genome donor to hexaploid wheat. 3. Variation in and the genetics of seed esterases (Est-5). Theoretical and Applied Genetics 77, 851–856.
Crossref | GoogleScholarGoogle Scholar | open url image1

Landjeva S, Korzun V, Börner A (2007) Molecular markers: actual and potential contributions to wheat genome characterization and breeding. Euphytica 156, 271–296.
Crossref | GoogleScholarGoogle Scholar | open url image1

Law CN , Young CF , Brown JWS , Snape JW , Worland AJ (1978) The study of grain protein control in wheat using whole chromosome substitution lines. In ‘Seed protein improvement by nuclear techniques’. pp. 483–502. (Atomic Energy Agency: Vienna)

Lelley T , Stachel M , Grausberger H , Vollman H (1999) Views about the origin of wheat used for bread. In ‘Bericht űber die Arbeitstagung (Jubilaumstagung) der Vereinigung Osterreichircher Pflanzenzüchter’. (Gumpenstein)

Lelley T, Stachel M, Grausberger H, Vollman J (2000) Analysis of relationships between Aegilops tauschii and the D genome of wheat utilising microsatellites. Genome 43, 661–668.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Liu B, Vega JM, Segal G, Abbo S, Rodova M, Feldman M (1998) Rapid genomic changes in newly synthetised amphiploids of Triticum and Aegilops. I. Changes in low copy non-coding DNA sequences. Genome 41, 272–277.
Crossref | GoogleScholarGoogle Scholar | open url image1

Liu S, Zhou R, Dong Y, Li P, Jia J (2006) Development, utilization of introgression lines using synthetic wheat as donor. Theoretical and Applied Genetics 112, 1360–1373.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Loughman R, Lagudah ES, Trottet M, Wilson RE, Mathews A (2001) Septoria nodorum blotch resistance in Aegilops tauschii and its expression in synthetic amphiploids. (Special issue: wheat breeding in the new century: applying molecular genetic analyses of key quality and agronomic traits). Australian Journal of Agricultural Research 52, 1393–1402.
Crossref | GoogleScholarGoogle Scholar | open url image1

Lubbers EL, Gill KS, Cox TS, Gill BS (1991) Variation of molecular markers among geographically diverse accessions of Triticum tauschii. Genome 34, 354–361. open url image1

Mabille F, Grit J, Abecarsis J (2001) Mechanical properties of wheat seed coats. Cereal Chemistry 78, 231–235.
Crossref | GoogleScholarGoogle Scholar | open url image1

Mackie AM, Lagudah ES, Lafiandra D (1996) Molecular and biochemical characterisation of HMW glutenin subunits from T. tauschii and the D genome of hexaploid wheat. Journal of Cereal Science 23, 213–225.
Crossref | GoogleScholarGoogle Scholar | open url image1

Marino CL, Nelson JC, Lu YH, Sorrels ME, Leroy P, Lopes CR, Hart GE (1996) RFLP-based linkage maps of the homeologous group 6 chromosomes of hexaploid wheat (Triticum aestivum L. em Thell.). Genome 39, 359–366.
Crossref |
open url image1

Matsuoka Y, Nasuda S (2004) Durum wheat as a candidate for the unknown female progenitor of bread wheat: an empirical study with a highly fertile F1 hybrid with Aegilops tauschii Coss. Theoretical and Applied Genetics 109, 1710–1717.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Mohabbati F (2005) Effects of salinity on synthetic wheat genotypes. Czech Journal of Genetics and Plant Breeding 41, 268–272. open url image1

Morgan JM (1983) Osmoregulation as a selection criterion for drought tolerance in wheat. Australian Journal of Agricultural Research 34, 607–614.
Crossref | GoogleScholarGoogle Scholar | open url image1

Morgan JM (1999) Pollen grain expression of a gene controlling differences in osmoregulation in wheat leaves: a simple breeding method. Australian Journal of Agricultural Research 50, 953–962.
Crossref | GoogleScholarGoogle Scholar | open url image1

Mujeeb-Kazi A (1993) Interspecific and intergeneric hybridization in the Triticeae for wheat improvement. In ‘Biodiversity and wheat improvement’. (Ed. A Damania) pp. 95–102. (John Wiley & Sons: Chichester, UK)

Murphy NEA, Loughman R, Wilson RE, Lagudah ES, Appels R, Jones MGK (2001) A single gene controls resistance to Septoria nodorum blotch in the Aegilops tauschii accession AUS21712. (Special issue: Wheat breeding in the new century: applying molecular genetic analyses of key quality and agronomic traits.). Australian Journal of Agricultural Research 52, 1403–1407.
Crossref | GoogleScholarGoogle Scholar | open url image1

Nakai Y (1973) Isozyme variation in Aegilops and Triticum. 2. Esterase and acid phosphate isozymes studies by gel isoelectrofocusing method. Seiken Jiho 24, 45–73. open url image1

Narasimhamoorthy B, Gill BS, Fritz AK, Nelson JC, Brown-Guedira GL (2006) Advanced backcross QTL analysis of a hard winter wheat-synthetic wheat population. Theoretical and Applied Genetics 112, 787–796.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Nelson JC, Autrique JE, Fuentes-Dávila G, Sorrells ME (1998) Chromosomal location of genes for resistance to Karnal bunt in wheat. Crop Science 38, 231–236. open url image1

Nelson JC, Sorrells ME, Van Deynze AE, Lu YH, Atkinson M, Bernard M, Leroy P, Faris JD, Anderson JA (1995a) Molecular mapping of wheat: major genes and rearrangements in homoeologous groups 4, 5 and 7. Genetics 141, 721–731. open url image1

Nelson JC, Van Deynze AE, Autrique E, Sorrells ME, Lu YH, Merlino M, Atkinson M, Leroy P (1995b) Molecular mapping of wheat homoeologous group 2. Genome 38, 516–524. open url image1

Nelson JC, Van Deynze AE, Autrique E, Sorrells ME, Lu YH, Negre S, Bernard M, Leroy P (1995c) Molecular mapping of wheat homeologous group 3. Genome 38, 525–533. open url image1

Nishikawa K, Furuta Y, Wada T (1980) Genetic studies on α-amylase isozymes in wheat III. Intraspeciic variation in Aegilops squarrosa and birthplace of hexaploid wheat. Japanese Journal of Genetics 55, 325–336.
Crossref | GoogleScholarGoogle Scholar | open url image1

Ogbonnaya FC , Halloran GM , Lagudah ES (2005) D genome of wheat- 60 years on from Kihara, Sears and McFadden. In ‘Frontiers of Wheat Bioscience’. Wheat Information Service No. 100. (Ed. K Tsunewaki) pp. 205–220. (Kihara Memorial Yokohama Foundation for the Advancement of Life Sciences: Yokohama, Japan)

Ogbonnaya FC , Imtiaz M , Hearnden P , Wilson J , Eastwood RF , Gatford KT , van Ginkel M (2006) Identification of a novel gene for seed dormancy in wheat. In ‘Breeding for Success: Diversity in Action. Proceedings of the 13th Australasian Plant Breeding Conference’. 18–21 April 2006, Christchurch, New Zealand. (Ed. CF Mercer) pp. 634–639. (CD-ROM)

Ogbonnaya FC, Ye G, Trethowan R, Dreccer F, Shepperd J, van Ginkel M (2007) Yield of synthetic backcross-derived lines in rainfed environments of Australia. Euphytica 157, 321–336.
Crossref | GoogleScholarGoogle Scholar | open url image1

Ogihara Y, Tsunewaki K (1988) Diversity and evolution of chloroplast DNA in Triticum and Aegilops as revealed by restriction fragment analysis. Theoretical and Applied Genetics 76, 321–332.
Crossref | GoogleScholarGoogle Scholar | open url image1

Okamoto M (1957) Asynaptic effect of chromosome V. Wheat Information Service 5, 6. open url image1

Ozkan H, Levy AA, Feldman M (2001) Allopolyploidy induced rapid genome evolution in the wheat (Aegilops-Triticum) group. The Plant Cell 13, 1735–1747.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Payne PI, Holt LM, Jackson EA, Law CN (1984) Wheat storage proteins: their genetics and their potential for manipulation by plant breeders. Philosophical Transactions of the Royal Society of London - A 304, 359–371.
Crossref | GoogleScholarGoogle Scholar | open url image1

Pestova E, Korzun V, Goncharov NP, Hammer K, Ganal MW, Röder MS (2000) Microsatellite analysis of Aegilops tauschii germplasm. Theoretical and Applied Genetics 101, 100–106.
Crossref | GoogleScholarGoogle Scholar | open url image1

Pestsova EG, Börner A, Röder MS (2001) Development of a set of Triticum aestivum–Aegilops tauschii introgression lines. Hereditas 135, 139–143.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Pestsova EG, Börner A, Röder MS (2006) Development and QTL assessment of Triticum aestivum–Aegilops tauschii introgression lines. Theoretical and Applied Genetics 112, 634–647.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Pomeranz Y , Williams PL (1990) Wheat hardness: its genetic, structural and biochemical background measurement and significance. Chapter 8. Advances in Cereal Science and Technology 10, 471–548.

Pritchard DJ, Hollington PA, Davies WP, Gorham J, de Leon JLD, Mujeeb-Kazi A (2002) K+/Na+ discrimination in synthetic hexaploid wheat lines: Transfer of the trait for K+/Na+ discrimination from Aegilops tauschii into a Triticum turgidum background. Cereal Research Communications 30, 261–267. open url image1

Renfrew JM (1969) The archaeological evidence for the domestication of plants: methods and problems. In ‘The domestication and exploitation of plants and animals’. (Eds PJ Ucko, GW Dimbleby) pp. 149–172. (Aldine: Chicago, IL)

Rickman G (1980) ‘The corn supply of ancient Rome.’ (Oxford University Press: New York)

Riley R, Chapman V (1958) Genetic control of the cytologically diploid behaviour of hexaploid wheat. Nature 182, 713–715.
Crossref | GoogleScholarGoogle Scholar | open url image1

Riley R, Kimber G, Chapman V (1961) Origin of the genetic control of diploid-like behaviour of polyploid wheat. The Journal of Heredity 52, 22–25. open url image1

Röder MS, Korzun V, Wendehake K, Plaschke J, Tixier MH, Leroy P, Ganal MW (1998) A microsatellite map of wheat. Genetics 149, 2007–2023.
PubMed |
open url image1

Schachtman DP, Lagudah ES, Munns R (1992) The expression of salt tolerance from Triticum tauschii in hexaploid wheat. Theoretical and Applied Genetics 84, 714–719.
Crossref | GoogleScholarGoogle Scholar | open url image1

Schachtman DP, Munns R (1992) Sodium accumulation in leaves of Triticum species that differ in salt tolerance. Australian Journal of Plant Physiology 19, 331–340.
Crossref | GoogleScholarGoogle Scholar | open url image1

Shaked H, Kashkush K, Ozkan H, Feldman M, Levy AA (2001) Sequence elimination and cytosine methylation are rapid and reproducible responses of the genome to wide hybridization and allopolyploidy in wheat. The Plant Cell 13, 1749–1759.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Simón MR, Ayala FM, Cordo CA, Röder MS, Börner A (2004) Molecular mapping of quantitative trait loci determining resistance to Septoria tritici blotch caused by Mycosphaerella graminicola in wheat. Euphytica 138, 41–48.
Crossref | GoogleScholarGoogle Scholar | open url image1

Somers DJ, Isaac P, Edwards K (2004) A high density microsatellite consensus map for bread wheat (Triticum aestivum L.). Theoretical and Applied Genetics 109, 1105–1114.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Song QJ, Shi JR, Singh S, Fickus EW, Costa JM, Lewis J, Bill BS, Ward R, Cregan PB (2005) Development and mapping of microsatellite (SSR) markers in wheat. Theoretical and Applied Genetics 110, 550–560.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Sourdille P, Perretant MR, Charmet G, Leroy P, Gautier MF, Joudrier P, Nelson JC, Sorrells ME, Bernard M (1996) Linkage between RFLP markers and genes affecting kernel hardness in wheat. Theoretical and Applied Genetics 93, 580–586.
Crossref | GoogleScholarGoogle Scholar | open url image1

Tanksley SD, Nelson JC (1996) Advanced backcross QTL analysis: a method for the simultaneous discovery and transfer of valuable QTLs from unadapted germplasm into elite breeding lines. Theoretical and Applied Genetics 92, 191–203.
Crossref | GoogleScholarGoogle Scholar | open url image1

Ter Steege MW, den Ouden FM, Lambers H, Stam P, Peeters AJM (2005) Genetic and physiological architecture of early vigor in Aegilops tauschii, the D-genome donor of hexaploid Wheat. A quantitative trait loci analysis. Plant Physiology 139, 1078–1094.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Tsuji S, Tsunewaki K (1976) Genetic diversity of the cytoplasm in Triticum and Aegilops. III. On the origin of the cytoplasm of two hexaploid Aegilops species. Japanese Journal of Genetics 51, 149–159.
Crossref | GoogleScholarGoogle Scholar | open url image1

Tsunewaki K, Ogihara Y (1983) The molecular basis of genetic diversity among cytoplasms of Triticum and Aegilops species. II. On the origin of polyploidy wheat cytoplasms as suggested by chloroplast DNA restriction fragment patterns. Genetics 104, 155–171.
PubMed |
open url image1

Turner NC (1997) Further progress in crop water relations. Advances in Agronomy 58, 293–338. open url image1

Van Deynze AE, Dubcovsky J, Gill KS, Nelson JC, Sorrells ME, Dvorák J, Gill BS, Lagudah ES, McCouch SR, Appels R (1995) Molecular-genetic maps for group 1 chromosomes of Triticeae species and their relation to chromosomes in rice and oat. Genome 38, 45–59. open url image1

van Ginkel M, Ogbonnaya F (2007) Novel genetic diversity from synthetic wheats in breeding cultivars for changing production conditions. Field Crops Research 104, 86–94.
Crossref | GoogleScholarGoogle Scholar | open url image1

Vedel F, Quetier F, Cauderon Y, Dosba F, Doussinault G (1981) Studies on maternal inheritance in polyploidy wheats with cytoplasmic DNA as genetic markers. Theoretical and Applied Genetics 59, 239–245. open url image1

Waines JG (1994) High temperature stress in wild wheats and spring wheats. Australian Journal of Plant Physiology 21, 705–715.
Crossref | GoogleScholarGoogle Scholar | open url image1

Wardlaw IF, Moncur (1995) The response of wheat to high temperature following anthesis I. The rate and duration of kernel filling. Australian Journal of Plant Physiology 22, 391–397.
Crossref | GoogleScholarGoogle Scholar | open url image1

Weng Y, Li W, Devkota RN, Rudd JC (2005) Microsatellite markers associated with two Aegilops tauschii-derived greenbug resistance loci in wheat. Theoretical and Applied Genetics 110, 462–469.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Wernstedt FL (1972) ‘World Climatic Data.’ (Climatic Data Press: Lemont, PA)

Xiao J, Li J, Yuan L, Tanksley SD (1996) Identification of QTLs affecting traits of agronomic importance in a recombinant inbred population derived from a subspecific rice cross. Theoretical and Applied Genetics 92, 230–244.
Crossref |
open url image1

Xu X, Monneveux P, Damania AB, Zahavieva M (1993) Evaluation of salt tolerance in genetic resoures of Triticum and Aegilops species. Plant Genetic Resources Newsletter 96, 11–16. open url image1

Yang W, Yu Y, Zhang Y, Hu X, Wang Y, Zhou Y, Lu B (2003) Inheritance and expression of stripe rust resistance in common wheat (Triticum aestivum) transferred from Aegilops tauschii and its utilization. Hereditas 139, 49–55.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Zhang HB, Dvórák J (1992) The genome origin and evolution of Triticum crassum and Triticum syriacum determined from variation in repeated nucleotide sequences. Genome 35, 806–814. open url image1

Zhao YH, Kimber G (1984) New hybrids with D-genome wheat relatives. Genetics 106, 509–515.
PubMed |
open url image1

Zohary D (1989) Domestication of the Southweat Asian Neolithic crop assemblage of cereals, pulses and flax: evidence from the living plants. In ’Foraging and farming – the evolution of plant exploitation’. (Eds DR Harris, GC Hillman) pp. 358–373. (Unwin Hyman: London)