Plant Cell Reports

, Volume 33, Issue 4, pp 595–603 | Cite as

The place of asymmetric somatic hybridization in wheat breeding

  • Shuwei Liu
  • Guangmin XiaEmail author


Since its first development some 40 years ago, the application of the somatic hybridization technique has generated a body of hybrid plant material involving a wide combination of parental species. Until the late 1990s, the technique was ineffective in wheat, as regeneration from protoplasts was proving difficult to achieve. Since this time, however, a successful somatic hybridization protocol for wheat has been established and used to generate a substantial number of both symmetric and asymmetric somatic hybrids and derived materials, especially involving the parental combination bread wheat and tall wheatgrass (Thinopyrum ponticum). This review describes the current state of the art for somatic hybridization in wheat and focuses on its potential application for wheat improvement.


Asymmetric somatic hybridization Wheat germplasm Genomic variation Alien introgression Protoplast fusion 



This work was supported by funds from the Natural Science Foundation of China (No. 31271707; 31000568) and the Doctoral Foundation of Shandong Province (BS2010NY013).

Conflict of interest

The authors declare that they have no conflict of interest.


  1. Austin S, Pohlman JD, Brown CR, Mojahedi H, Santo GS, Douches DS, Helgeson JP (1988) Interspecific somatic hybridization between Solanum tuberosum L. and S. bulbocastanum dun. as a means of traits of transferring nematode resistance. Am Potato J 70:485–495CrossRefGoogle Scholar
  2. Bauer-Weston B, Keller W, Webb J, Gleddie S (1993) Production and characterization of asymmetric somatic hybrids between Arabidopsis thaliana and Brassica napus. Theor Appl Genet 86:150–158PubMedGoogle Scholar
  3. Begum F, Paul S, Bag N, Sikdar SR, Sen SK (1995) Somatic hybrids between Brassica juncea (L.) Czern. and Diplotaxis harra (Forsk.) Boiss and the generation of backcross progenies. Theor Appl Genet 91:1167–1172PubMedGoogle Scholar
  4. Bijoya B, Aniruddha PS, Hari SG (1999) Transfer of wild abortive cytoplasmic male sterility through protoplast fusion in rice. Mol Breed 5:319–327CrossRefGoogle Scholar
  5. Cai YF, Xiang FN, Zhi DY, Liu H, Xia GM (2007) Genotyping of somatic hybrids between Festuca arundinacea Schreb. and Triticum aestivum L. Plant Cell Rep 26:1809–1819PubMedCrossRefGoogle Scholar
  6. Cardi T, Bastia T, Monti L, Earle ED (1999) Organelle DNA and male fertility variation in Solanum ssp. and interspecific somatic hybrids. Theor Appl Genet 99:819–828CrossRefGoogle Scholar
  7. Carlson PS, Smith HH, Dearing R (1972) Parasexual interspecific plant hybridization. Proc Natl Acad Sci USA 69:2292–2294PubMedCentralPubMedCrossRefGoogle Scholar
  8. Chen WP, Wu QS, Liu DJ, Jones MGK (1992) Callus formation from somatic hybridization of wheat (Triticum aestivum L.) and Ryegrass (Lolium perenne L.) by electrofusion. Acta Bot Sin 34:284–290Google Scholar
  9. Chen SY, Xia GM, Quan TY, Xiang FN (2004a) Studies on the salt-tolerance of F3–F6 hybrid lines originated from somatic hybridization between common wheat and Thinopyrum ponticum. Plant Sci 167:773–779CrossRefGoogle Scholar
  10. Chen SY, Liu SW, Xu CH, Chen YZ, Xia GM (2004b) Heredity of chloroplast and nuclear genomes of asymmetric somatic hybrid lines between wheat and couch grass. Acta Bot Sin 46:110–115Google Scholar
  11. Chen FG, Xu CH, Chen MZ, Wang YH, Xia GM (2008a) A new α-gliadin gene family for wheat breeding: somatic introgression line II-12 derived from Triticum aestivum and Agropyron elongatum. Mol Breed 22:675–685CrossRefGoogle Scholar
  12. Chen FG, Zhao F, Xu CH, Xia GM (2008b) Molecular characterization of LMW-GS genes from a somatic hybrid introgression line II-12 between Triticum aestivum and Agropyron elongatum in relation to quick evolution. J Genet Genomics 35:743–749PubMedCrossRefGoogle Scholar
  13. Chen FG, Zhao F, Liu SW, Xia GM (2009) The γ-gliadin gene content of a derivative from a somatic hybrid between bread wheat and tall wheatgrass. Mol Breed 24:117–126CrossRefGoogle Scholar
  14. Cheng AX, Xia GM (2004) Somatic hybridization between common wheat and Italian ryegrass. Plant Sci 166:1219–1226CrossRefGoogle Scholar
  15. Cheng AX, Xia GM, Chen HM (2004a) DNA transfer from wild millet to common wheat by asymmetric somatic hybridization. Acta Bot Sin 46:1114–1121Google Scholar
  16. Cheng AX, Xia GM, Zhi DY, Chen HM (2004b) Intermediate fertile Triticum aestivum (+) Agropyron elongatum somatic hybrids are generated by low doses of UV irradiation. Cell Res 14:86–91PubMedCrossRefGoogle Scholar
  17. Comai L (2000) Genetic and epigenetic interactions in allopolyploid plants. Plant Mol Biol 43:387–399PubMedCrossRefGoogle Scholar
  18. Cui HF, Yu ZY, Deng JY, Gao X, Sun S, Xia GM (2009) Introgression of bread wheat chromatin into tall wheatgrass via somatic hybridization. Planta 229:323–330PubMedCrossRefGoogle Scholar
  19. Deng JY, Cui HF, Zhi DY, Zhou CE, Xia GM (2007) Analysis of remote asymmetric somatic hybrids between common wheat and Arabidopsis thaliana. Plant Cell Rep 26:1233–1241PubMedCrossRefGoogle Scholar
  20. Derks FHM, Wijbrandi J, Koornneef M (1991) Organelle analysis of symmetric and asymmetric hybrids between Lycopersicon peruvianum and Lycopersicon esculentum. Theor Appl Genet 81:199–204PubMedCrossRefGoogle Scholar
  21. Dong YC, Zheng DS (2000) Genetic resource of Chinese wheat. China Agricultural Publishing Company, BeijingGoogle Scholar
  22. Dong W, Wang MC, Xu F, Quan TY, Peng KQ, Xiao LT, Xia GM (2013) Wheat oxophytodienoate reductase gene TaOPR1 confers salinity tolerance via enhancement of abscisic acid signaling and reactive oxygen species scavenging. Plant Physiol 161:1217–1228PubMedCentralPubMedCrossRefGoogle Scholar
  23. Dudits D, Fejér O, Hadlaczky G, Koncz C, Lázár GB, Horváth G (1980) Intergeneric gene transfer mediated by plant protoplast fusion. Mol Gen Genet 179:283–288CrossRefGoogle Scholar
  24. Feldman M, Levy AA (2005) Allopolyploidy—a shaping force in the evolution of wheat genomes. Cytogenet Genome Res 109:250–258PubMedCrossRefGoogle Scholar
  25. Feldman M, Lupton FGH, Miller TE (1995) Wheat Triticum spp. (Gramineae-Triticinae). In: Smartt J, Simmonds NW (eds) Evolution of crop plants. Longman Scientific and Technical, Harlow, pp 184–192Google Scholar
  26. Feng DS, Zhao SY, Chen FG, Xia GM (2004) Two quality-associated HMW glutenin subunits in a somatic hybrid line between Triticum aestivum and Agropyron elongatum. Theor Appl Genet 110:136–144PubMedCentralPubMedCrossRefGoogle Scholar
  27. Gao X, Liu SW, Sun Q, Xia GM (2010) High frequency of HMW-GS sequence variation through somatic hybridization between Agropyron elongatum and common wheat. Planta 231:245–250PubMedCrossRefGoogle Scholar
  28. Glimelius K, Fahleson J, Landgren M, Sjodin C, Sundberg E (1991) Gene transfer via somatic hybridization in plants. Trends Biotechnol 9:24–30CrossRefGoogle Scholar
  29. Guo WW, Cheng YJ, Deng XX (2000a) Regeneration and molecular characterization of intergeneric somatic hybrids between Citrus reticulate and Poncirus trifoliate. Plant Cell Rep 20:829–834Google Scholar
  30. Guo WW, Zhou CH, Yi HL, Deng XX (2000b) Intergeneric somatic hybrid plants between citrus and Poncirus trifoliate and evaluation of their root rot resistance. Acta Bot Sin 42:668–672Google Scholar
  31. Guo WW, Wu RC, Cheng YJ, Deng XX (2007) Production and molecular characterization of Citrus intergeneric somatic hybrids between red tangerine and citrange. Plant Breed 126:72–76CrossRefGoogle Scholar
  32. Han FP, Fedak G, Ouellet T, Liu B (2003) Rapid genomic changes in interspecific and intergeneric hybrids and allopolyploids of Triticeae. Genome 46:716–723PubMedCrossRefGoogle Scholar
  33. Handley LW, Nickels RL, Cameron MW, Moore PP, Sink KC (1986) Somatic hybrid plants between Lycopersicon esculentum and Solanum lycopersicoides. Theor Appl Genet 71:691–697PubMedCrossRefGoogle Scholar
  34. Hansen LN, Earle ED (1995) Transfer of resistance to Xanthomonas campestris by campestris into Brassica oleracea by protoplast fusion. Theor Appl Genet 91:1293–1300PubMedCrossRefGoogle Scholar
  35. He P, Friebe BR, Gill BS, Zhou JM (2003) Allopolyploidy alters gene expression in the highly stable hexaploid wheat. Plant Mol Biol 52:401–414PubMedCrossRefGoogle Scholar
  36. He YN, Li W, Lv J, Jia YB, Wang MC, Xia GM (2012) Ectopic expression of a wheat MYB transcription factor gene, TaMYB73, improves salinity stress tolerance in Arabidopsis thaliana. J Exp Bot 63:1511–1522PubMedCrossRefGoogle Scholar
  37. Jauhar PP (2006) Modern biotechnology as an integral supplement to conventional plant breeding: the prospects and challenges. Crop Sci 46:1841–1859CrossRefGoogle Scholar
  38. Jauhar PP, Chibbar RN (1999) Chromosome mediated and direct gene transfers in wheat. Genome 42:570–583CrossRefGoogle Scholar
  39. Kalendar R, Grob T, Regina M, Suoniemi A, Schulman A (1999) IRAP and REMAP: two new retrotransposon-based DNA fingerprinting techniques. Theor Appl Genet 98:704–711CrossRefGoogle Scholar
  40. Kisaka H, Kisaka M, Kanno A, Kameya T (1998) Intergeneric somatic hybridization of rice (Oryza sativa L.) and barley (Hordeum vulgare L.) by protoplast fusion. Plant Cell Rep 17:362–367CrossRefGoogle Scholar
  41. Leino M, Teixeira R, Landgren M, Glimelius K (2003) Brassica napus lines with rearranged Arabidopsis mitochondria display CMS and a range of developmental aberrations. Theor Appl Genet 106:1156–1163PubMedGoogle Scholar
  42. Li YG, Stoutjestijk PA, Larkin PJ (1999) Somatic hybridization for plant improvement. In: Soh W-Y, Bhojwani SS (eds) Morphogenesis in plant tissue cultures. Kluwer Academic Publishers, Dordrecht, pp 363–418CrossRefGoogle Scholar
  43. Li CL, Xia GM, Xiang FN, Zhou CE, Cheng AX (2004) Regeneration of asymmetric somatic hybrid plants from the fusion of two types of wheat with Russian wild rye. Plant Cell Rep 23:461–467PubMedCrossRefGoogle Scholar
  44. Li AM, Wei CX, Jiang JJ, Zhang YT, Snowdon RJ, Wang YP (2009) Phenotypic variation in the progenies of somatic hybrids between Brassica napus and Sinapis alba. Euphytica 170:289–296CrossRefGoogle Scholar
  45. Li CL, Lv J, Zhao X, Ai XH, Zhu XL, Wang MC, Zhao SY, Xia GM (2010a) TaCHP: a wheat zinc finger protein gene down-regulated by abscisic acid and salinity stress plays a positive role in stress tolerance. Plant Physiol 154:211–221PubMedCentralPubMedCrossRefGoogle Scholar
  46. Li S, Xu CH, Yang YN, Xia GM (2010b) Functional analysis of TaDi19A, a salt-responsive gene in wheat. Plant Cell Environ 33:117–129PubMedCrossRefGoogle Scholar
  47. Liu JH, Deng XX (2000) Preliminary analysis of cytoplasmic genome of diploid somatic hybrid derived from fusion between rough lemon and Hamlin sweet orange. Acta Bot Sin 42:102–104Google Scholar
  48. Liu B, Liu DJ (1995) Transfer of a partial nuclear genome of Avena nuda L. into Triticum aestivum L. by ‘donor-recipient’ protoplast fusion. Acta Biol Exp Sin 28:95–102Google Scholar
  49. Liu B, Liu ZL, Li XW (1999) Production of a highly asymmetric somatic hybrid between rice and Zizania latifolia (Griseb): evidence for inter-genomic exchange. Theor Appl Genet 98:1099–1103CrossRefGoogle Scholar
  50. Liu H, Shi L, Zhao JS, Xia GM (2006) Genetic characteristic of HMW-GS in somatic hybrid wheat lines—potential application to wheat breeding. J Agric Food Chem 54:5007–5013CrossRefGoogle Scholar
  51. Liu SW, Zhao SY, Chen FG, Xia GM (2007) Generation of novel high quality HMW-GS genes in two introgression lines of Triticum aestivum/Agropyron elongatum. BMC Evol Biol 7:76PubMedCentralPubMedCrossRefGoogle Scholar
  52. Liu H, Liu SW, Xia GM (2009) Generation of high frequency of novel alleles of the high molecular weight glutenin in somatic hybridization between bread wheat and tall wheatgrass. Theor Appl Genet 118:1193–1198PubMedCrossRefGoogle Scholar
  53. Liu C, Li S, Wang MC, Xia GM (2012) A transcriptomic analysis reveals the nature of salinity tolerance of a wheat introgression line. Plant Mol Biol 78:159–169PubMedCrossRefGoogle Scholar
  54. Louzada E, Grosser J, Gmitter FJ (1993) Intergeneric somatic hybridization of sexually incompatible Citrus sinensis and Atalantia ceylanica. Plant Cell Rep 12:687–690PubMedCrossRefGoogle Scholar
  55. Ma XF, Fang P, Gustafson JP (2004) Polyploidization-induced genome variation in triticale. Genome 47:839–848PubMedCrossRefGoogle Scholar
  56. Madlung A, Masuelli RW, Watson B, Reynolds SH, Davison J, Comai L (2002) Remodeling of DNA methylation and phenotypic and transcriptional changes in synthetic Arabidopsis allotetraploids. Plant Physiol 129:733–746PubMedCentralPubMedCrossRefGoogle Scholar
  57. Melchers G, Labib G (1974) Somatic hybridisation of plants by fusion of protoplasts. Mol Gen Genet 135:277–294CrossRefGoogle Scholar
  58. Melchers G, Sacristan MD, Holder AA (1978) Somatic hybrid plants of potato and tomato regenerated from fused protoplasts. Carlsberg Res Commun 43:203–218CrossRefGoogle Scholar
  59. Mohapatra T, Kiriti PB, Dinesh Kumar V, Prakash S, Chopra VL (1998) Random chloroplast segregation and mitochondrial genome recombination in somatic hybrid plants of Diplotaxis catholica + Brassica juncea. Plant Cell Rep 17:814–818CrossRefGoogle Scholar
  60. Ohgawara T, Kobayashi S, Ohgawara E, Uchimaya H, Ishii S (1985) Somatic hybrid plants obtained by protoplast fusion between Citrus sinensis and Poncirus trifoliata. Theor Appl Genet 71:1–4PubMedCrossRefGoogle Scholar
  61. Ozias-Akins P, Ferl RJ, Vasil IK (1986) Somatic hybridization in the gramineae: Pennisetum americanum (L.) K. Schum. (Pearl millet) + Panicum maximum Jacq. (Guinea grass). Mol Gen Genet 203:365–370CrossRefGoogle Scholar
  62. Ozkan H, Levy AA, Feldman M (2001) Alloploidy-induced rapid genome evolution in the wheat (Aegilops-Triticum) group. Plant Cell 13:1735–1747PubMedCentralPubMedCrossRefGoogle Scholar
  63. Park YG, Kim JH, Son SH (1992) Induction of somatic hybrid by protoplast fusion between Populus koreana × P. nigra var. italica and P. euramericana cv. Guardi. J Korean Soc 81:273–279Google Scholar
  64. Parokonny AS, Kenton AY, Gleba YY, Bennett MD (1992) Genome reorganization in Nicotiana asymmetric somatic hybrids analysed by in situ hybridization. Plant J 2:863–874PubMedGoogle Scholar
  65. Peng ZY, Wang MC, Li F, Lv HJ, Li CL, Xia GM (2009) A proteomic study of the response to salinity and drought stress in an introgression strain of bread wheat. Mol Cell Proteomics 8:2676–2686PubMedCentralPubMedCrossRefGoogle Scholar
  66. Qin Z, Lv HJ, Zhu XL, Meng C, Quan TY, Wang MC, Xia GM (2013) Ectopic expression of a wheat WRKY transcription factor gene TaWRKY71-1 results in hyponastic leaves in Arabidopsis thaliana. PLoS One 8:e63033PubMedCentralPubMedCrossRefGoogle Scholar
  67. Riley R, Chapman V (1958) Genetic control of the cytologically diploid behavior of hexaploid wheat. Nature 182:713–715CrossRefGoogle Scholar
  68. Salmon A, Ainouche ML, Wendel JF (2005) Genetic and epigenetic consequences of recent hybridization and polyploidy in Spartina (Poaceae). Mol Ecol 14:1163–1175PubMedCrossRefGoogle Scholar
  69. Sears ER (1993) Use of radiation to transfer alien chromosome segments to wheat. Crop Sci 33:897–901CrossRefGoogle Scholar
  70. 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. Plant Cell 13:1749–1759PubMedCentralPubMedCrossRefGoogle Scholar
  71. Shan L, Zhao SY, Quan TY, Xia GM (2004) Isolation and analysis of cDNA fragments responded to salt-stress from a new somatic hybrid line between Triticum aestivum and Agropyron elongatum by differential display. Hi-Tech Lett 14:29–33 (In Chinese with an English abstract)Google Scholar
  72. Shan L, Zhao SY, Xia GM (2005) Cloning of the full-length cDNA of the wheat involved in salt stress: root hair defective 3 gene (RHD3). J Integr Plant Biol 47:881–891CrossRefGoogle Scholar
  73. Shan L, Li C, Chen F, Zhao SY, Xia GM (2008) A Bowman–Birk type protease inhibitor is involved in the tolerance to salt stress in wheat. Plant Cell Environ 31:1128–1137PubMedCrossRefGoogle Scholar
  74. Shepard JF, Bidney D, Barsby T, Kemble A (1983) Genetic transfer in plants through interspecific protoplast fusion. Science 219:683–688PubMedCrossRefGoogle Scholar
  75. Sjodin C, Glimelius K (1989) Brassica naponigra, a somatic hybrid resistant to Phoma lingam. Theor Appl Genet 77:651–656PubMedGoogle Scholar
  76. Song K, Lu P, Tang K, Osborn TC (1995) Rapid genome change in synthetic polyploids of Brassica and its implications for polyploid evolution. Proc Natl Acad Sci USA 92:7719–7723PubMedCentralPubMedCrossRefGoogle Scholar
  77. Spangenberg G, Valles MP, Wang ZY, Montavon P, Nagel J, Potrykus I (1994) Asymmetric somatic hybridization between tall fescue (Festuca arundinaceae Schreb) and irradiated Italian ryegrass (Lolium multiforum Lam) protoplast. Theor Appl Genet 88:509–519PubMedCrossRefGoogle Scholar
  78. Spangenberg G, Wang ZY, Legris G, Montavon P, Takamizo T, Perezvicente R, Valles MP, Nagel J, Potrykus I (1995) Intergeneric symmetric and asymmetric hybridization in Festuca and Lolium. Euphytica 85:235–245CrossRefGoogle Scholar
  79. Tabaeigadeh Z, Ferl JF, Vasil IK (1986) Somatic hybridization in the Gramineae: Saccharum officinarum L. (Sugarcane) and Pennisetum americanum (L.) K. Schum. Proc Natl Acad Sci USA 83:5616–5619CrossRefGoogle Scholar
  80. Tate JA, Ni Z, Scheen A-C, Koh J, Gilbert CA, Lefkowitz D, Chen ZJ, Soltis PS, Soltis DE (2006) Evolution and expression of homeologous loci in Tragopogon miscellus (Asteraceae), a recent and reciprocally formed allopolyploid. Genetics 173:1599–1611PubMedCentralPubMedCrossRefGoogle Scholar
  81. Terada R, Kyozuka J, Nishibayashi S, Shimamoto K (1987) Plantlet regeneration from somatic hybrids of rice (Oryza sativa L.) and barnyard grass (Echinochloa oryzicola Vasing). Mol Gen Genet 210:39–43CrossRefGoogle Scholar
  82. Vasil V, Ferl RJ, Vasil IK (1988) Somatic hybridization in the Gramineae: Triticum monococcum L. (Einkorn) + Pennisetum americanum (L.) K. Schum. (Pearl Millet). J Plant Physiol 132:160–163CrossRefGoogle Scholar
  83. Waara S, Glimelius K (1995) The potential of somatic hybridization in crop breeding. Euphytica 85:217–233CrossRefGoogle Scholar
  84. Wang YP, Sonntag K, Rudloff E (2003) Development of rapeseed with high erucic acid content by asymmetric somatic hybridization between Brassica napus and Crambe abyssinica. Theor Appl Genet 106:1147–1155PubMedGoogle Scholar
  85. Wang J, Tian L, Madlung A, Lee H-S, Chen M, Lee JJ, Watson B, Kagochi T, Comai L, Chen ZJ (2004a) Stochastic and epigenetic changes of gene expression in Arabidopsis polyploids. Genetics 167:1961–1973PubMedCentralPubMedCrossRefGoogle Scholar
  86. Wang J, Xiang FN, Xia GM (2004b) Transfer of small chromosome fragments of Agropyron elongatum to wheat chromosome via asymmetric somatic hybridization. Sci China (Ser. C) 47:434–441CrossRefGoogle Scholar
  87. Wang J, Xiang FN, Xia GM (2005) Agropyron elongatum chromatin localization on the wheat chromosomes in an introgression line. Planta 221:277–286PubMedCrossRefGoogle Scholar
  88. Wang MC, Peng ZY, Li CL, Li F, Liu C, Xia GM (2008) Proteomic analysis on a high salt tolerance introgression strain of Triticum aestivum/Thinopyrum ponticum. Proteomics 8:1470–1489PubMedCrossRefGoogle Scholar
  89. Wolters AMA, Schoenmakers HCH, Koornneef M (1995) Chloroplast and mitochondrial DNA composition of triploid and tetraploid somatic hybrids between Lycopersicon esculentum and Solanum tuberosum. Theor Appl Genet 90:285–293PubMedCrossRefGoogle Scholar
  90. Xia GM (2009) Progress of chromosome engineering mediated by asymmetric somatic hybridization. J Genet Genomics 36:547–556PubMedCrossRefGoogle Scholar
  91. Xia GM, Chen HM (1996) Plant regeneration from intergeneric somatic hybridization between Triticum aestivum and Leymus chinensis (Trin.) Tzvel. Plant Sci 120:197–203CrossRefGoogle Scholar
  92. Xia GM, Li ZY, Zhou AF, Guo GQ, Chen HM (1995) Plant regeneration of wheat protoplasts prepared from different composition of suspension culture. Chin J Biotechnol 11:63–66 (In Chinese with an English abstract)Google Scholar
  93. Xia GM, Wang H, Chen HM (1996) Plant regeneration from intergeneric asymmetric hybridization between wheat (Triticum aestivum L.) and Russian wildrye (Psathyrostachys juncea (Fisch) Nevski) and wheat grass (Agropyron elongatum (Host) Nevski). Chin Sci Bull 41:1382–1386Google Scholar
  94. Xia GM, Li ZY, Wang SL, Xiang FN, Chen PD, Liu DJ (1998) Asymmetric somatic hybridization between haploid common wheat and UV irradiated Haynaldia villosa. Plant Sci 137:217–223CrossRefGoogle Scholar
  95. Xia GM, Xiang FN, Zhou AF, Wang H, Chen HM (2003) Asymmetric somatic hybridization between wheat (Triticum aestivum L.) and Agropyron elongatum (Host) Nevishi. Theor Appl Genet 107:299–305PubMedCrossRefGoogle Scholar
  96. Xiang FN, Xia GM, Zhou AF, Chen HM (1999) Asymmetric somatic hybridization between wheat (Triticum aestivum) and Bromus inermis. Acta Bot Sin 41:458–462Google Scholar
  97. Xiang FN, Xia GM, Chen HM (2003a) Asymmetric somatic hybridization between wheat (Triticum aestivum) and Avena sativa L. Sci China (Ser. C) 46:243–252CrossRefGoogle Scholar
  98. Xiang FN, Xia GM, Chen HM (2003b) Effect of UV dosage on somatic hybridization between common wheat (Triticum aestivum L.) and Avena sativa L. Plant Sci 164:697–707CrossRefGoogle Scholar
  99. Xiang FN, Xia GM, Zhi DY (2004) Hybrid plant regeneration in relation to the nuclear and cytoplasmic genomes of wheat and Setaria italica. Genome 47:680–688PubMedCrossRefGoogle Scholar
  100. Xu CH, Xia GM, Zhi DY, Xiang FN, Chen HM (2003) Integration of maize nuclear and mitochondrial DNA into the wheat genome through somatic hybridization. Plant Sci 165:1001–1008CrossRefGoogle Scholar
  101. Yamagishi H, Landgren M, Forsberg J, Glimelius K (2002) Production of asymmetric hybrids between Arabidopsis thaliana and Brassica napus utilizing an efficient protoplast culture system. Theor Appl Genet 104:959–964PubMedCrossRefGoogle Scholar
  102. Yue W, Xia GM, Zhi DY, Chen HM (2001) Transfer of salt tolerance from Aeleuropus littoralis sinensis to wheat (Triticum aestivum L.) via asymmetric somatic hybridization. Plant Sci 161:256–262Google Scholar
  103. Zhang QL, Xia GM, Quan TY, Zhang YG, Li FX, Li FM (2005) Trait variation of somatic hybrid progenies from Triticum aestivum/Agropyron elongatum Host (Nevski). J Triticeae Crop 25:11–14 (In Chinese with English Abstract)Google Scholar
  104. Zhong GZ, Mu SM, Zhang ZB (2002) Triticeal distant crossing. Science Press, Bejing, pp 92–96Google Scholar
  105. Zhou AF, Xia GM (2005) Introgression of the Haynaldia villosa genome to γ-ray induced asymmetric somatic hybrids of wheat. Plant Cell Rep 24:289–296PubMedCrossRefGoogle Scholar
  106. Zhou AF, Xia GM, Chen HM (1996) Asymmetric somatic hybridization between Triticum aestivum and Haynaldia villosa Schur. Sci China (Ser. C) 39:617–626Google Scholar
  107. Zhou AF, Xia GM, Chen HM, Hu H (2001a) Comparative study of symmetric and asymmetric somatic hybridization between common wheat and Haynaldia villosa. Sci China (Ser. C) 44:294–304CrossRefGoogle Scholar
  108. Zhou AF, Xia GM, Zhang X, Chen HM, Hu H (2001b) Analysis of chromosomal and organellar DNA of somatic hybrids between Triticum aestiuvm and Haynaldia villosa Schur. Mol Genet Genomics 265:387–393PubMedCrossRefGoogle Scholar
  109. Zhou AF, Xia GM, Chen XL, Chen HM (2002) Production of somatic hybrid plants between two types of wheat protoplasts and the protoplasts of Haynaldia villosa. Acta Bot Sin 44:1004–1008Google Scholar
  110. Zhou CE, Xia GM, Zhi DY, Chen Y (2006) Genetic characterization of asymmetric somatic hybrids between Bupleurum scorzonerifolium Willd and Triticum aestivum L.: potential application to the study of the wheat genome. Planta 223:714–724PubMedCrossRefGoogle Scholar
  111. Zhu XL, Liu SW, Meng C, Qin LM, Kong LN, Xia GM (2013) WRKY transcription factors in wheat and their induction by biotic and abiotic stress. Plant Mol Biol Rep 31:1053–1067CrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  1. 1.The Key Laboratory of Plant Cell Engineering and Germplasm InnovationMinistry of Education, School of Life Sciences, Shandong UniversityJinanChina

Personalised recommendations