摘要
小麦（Triticum aestivum L.，2n=6×=42）和大麦（Hordeum vulgare L.，2n=2×=14)是世界上两大重要的麦类作物，通过小麦和大麦的远缘杂交，可以将大麦的优良基因及其相应性状导入小麦，如早熟、耐生物和非生物胁迫及各种品质性状。小麦成熟期穗发芽是一种世界性自然灾害，穗发芽时小麦α-淀粉酶活性提高，对胚乳中淀粉分解能力增强，不仅影响小麦产量，而且还会影响小麦的营养品质和加工品质。位于大麦2H染色体长臂上的Isa基因所编码的大麦α-淀粉酶抑制蛋白BASI（bifunctional α-amylase/subtilisin inhibitor），可以通过与小麦α-淀粉酶相结合而抑制其活性，从而降低小麦穗发芽的危害，提升小麦品质。
本研究通过郑麦9023，CB037，中麦16等小麦品种与小-大麦2H染色体异代换系2H(2A)及2H(2B)的杂种幼胚组织培养，经愈伤组织诱导、继代、分化的途径最终获得522株结实的SC1代再生植株。以大麦Betzes、小麦CS、小-大麦2H染色体异附加系、一整套小-大麦2H染色体异代换系2H(2A)、2H(2B)和2H(2D)为材料共筛选出10对分别位大麦2H染色体短臂和长臂的大麦2H染色体特异SSR分子标记。通过大麦第二部分同源群特异SSR分子标记，对组培SC1代再生植株及其自交后代SC2-5代植株进行逐代筛选，以追踪和鉴定大麦2H染色体。通过以大麦Betzes基因组DNA为探针、小麦CS基因组DNA为封阻的基因组原位杂交对部分SC4及SC5植株进行进一步验证，最终获得了携带有大麦2HL染色体的杂合易位、纯合易位、单端体、双端体及单等臂体遗传材料。经大麦Isa基因特异引物进行PCR验证，以上这些材料均携带Isa基因。对小-大麦2HL染色体杂合易位系32-4-9，32-4-18及纯合易位系32-4-11花粉母细胞减数分裂I中期的染色体构型进行检测，表明杂合易位系和纯合易位系染色体配对正常，细胞学上遗传稳定。
在将外源种质转移至小麦遗传背景的过程中，会引起小麦基因组结构及基因表达的广泛遗传变异。为了了解由大麦2H染色体导入而引起的小麦基因组结构的变异，采用荧光AFLP对大麦Betzes，小麦CS,小-大麦2H异附加系，小-大麦2H异代换系2H(2A)、2H(2B)、2H(2D)这一整套材料进行分析。64对引物组合在大麦中获得了3157个条带，在其它材料中获得了4736个条带。AFLP扩增类型表明，附加系中检测到了消减和激活位点，而代换系基因组结构几乎无变化。同时还检测到了多条大麦2H染色体及小麦2A、2B、2D染色体特异的AFLP片段，经回收测序后，将AFLP分子标记转换成STS分子标记并进行验证，共获得2条大麦2H染色体特异的AFLP-STS分子标记，小麦2A、2B、2D染色体特异的AFLP-STS分子标记各1条。
为了了解由大麦2H染色体导入而引起的小麦基因组的表观遗传学变异，在荧光AFLP检测的基础上构建了荧光MSAP检测体系，对大麦Betzes，小麦CS,小-大麦2H异附加系，小-大麦2H异代换系2H(2A)、2H(2B)、2H(2D)这一整套材料的基因组甲基化模式进行分析。MSAP结果表明，大麦2H染色体导入引起了小麦基因组的甲基化变异，同时大麦2H染色体本身也检测到了甲基化的升高。
为了了解由大麦2H染色体导入所引起的基因表达差异，通过荧光cDNA-AFLP对大麦Betzes，小麦CS,小-大麦2H异附加系，小-大麦2H异代换系2H(2A)、2H(2B)、2H(2D)这一整套材料进行了基因表达差异分析，分离和克隆了大量差异表达基因，并初步确定了大麦2H染色体在小麦基因背景中的表达模式。
通过iTRAQ技术来了解由大麦2H染色体导入所引起的蛋白表达差异，尝试对以上材料进行差异蛋白质组学分析，目前为止，共鉴定出589个蛋白。
Wheat (Triticum aestivum L.,2n=6×=42) and barley (Hordeum vulgare L.,2n=2×=14)are two of the most important cereal crops worldwide. The hybridization of wheat and barleymakes it possible to transfer desirable genes and traits, such as earliness, tolerance to biotic andabiotic stresses, and various nutritional parameters, from barley to wheat.The wheat pre-harvestsprouting is a worldwide natural disaster. In the course of high activity of α-amylase, pre-harvestsprouting can not only result in a great decrease in wheat yield, but also dramatically degrade thenutritional and processing quality of the seeds. The bifunctional α-amylase/subtilisin inhibitor(BASI) encoded by the Isa gene on barley chromosome2HL could inhibit wheat α-amylaseactivity by reducing pre-harvest sprouting and improving the quality of wheat.
Regenerated plants were derived from immature embryo culture of hybrids of common wheatvarieties Zhengmai9023, CB037and Zhongmai16with the wheat-barley2H alien substituionlines2H(2A) and2H(2B) after callus induction, subculture and differentiation.10SSR molecularmarkers specific to barley chromosome2H located on short arm and long arm were selected frombarley Betzes, wheat CS, wheat-barley2H addition line, wheat-barley2H substitution line2H(A),2H(2B) and2H(2D). The presence of barley2H chromatin was detected in regenerated plants(SC1) and their selfed progenies (SC2-5) using homoeologous group2SSR markers from barley,and further identified in selected SC4and SC5lines using genomic in situ hybridization withbarley genomic DNA as probe and CS genomic DNA for blocking. The Isa gene from theidentified SC4and SC5lines was also amplified using Isa specific primers. We identifiedwheat-barley2HL chromosome heterozygous translocation lines, a homozygous translocationlines, monotelosomic lines, ditelosomic lines, and an isochromosome line carring the Isa gene.Metaphase I pairing of line32-4-9, line32-4-18, line32-4-11indicated that both the wheat-barley2HL chromosome heterozygous translocation line and homozygous translocation line shouldregularly disjoin and be stable in cytology.
During the process of alien germplasm introduced into wheat genetic backgroud, extensivegenetic variations of genome structure and gene expression could be induced in wheat. Tounderstand genetic changes associated with introduction of barely chromosome2H into wheatgenome, a set of barley Betzes, wheat CS, wheat-barley2H addition line, wheat-barley2Hsubstitution line2H(A),2H(2B) and2H(2D) were analyzed by fluorescence AFLP. Using64primer combinations,3157bands were obtained in barley and4736bands in other materials.AFLP amplification patterns indicated that elimination bands and novel bands were detected inaddition line, but on genome variation were dectected in substitution lines. Several AFLP bandsspecific to chromosome2H,2A,2B,2D were recoveried and sequenced respectively, for STS primers design. Finally,2AFLP-STS molecular markers specific to barley chromosome2H and1AFLP-STS molecular markers specific to each one of wheat chromosome2A,2B,2D wereestablished.
To understand epigenetic variations associated with introduction of barely chromosome2Hinto wheat genome, fluorescence MSAP detection system was constructed based on fluorescenceAFLP, for the methylation patterns detection of a set of barley Betzes, wheat CS, wheat-barley2Haddition line, wheat-barley2H substitution line2H(A),2H(2B) and2H(2D). The results thatmethylation variations were detected in wheat and the methylation extent increased in barleychromosome2H.
To understand gene expression characteristics associated with introduction of barelychromosome2H, we used the fluorescence cDNA-AFLP to analyse the different expression of aset of barley Betzes, wheat CS, wheat-barley2H addition line, wheat-barley2H substitution line2H(A),2H(2B) and2H(2D). A large number of differentially expressed genes were isolated andexpress patterns of barley2H chromosome were confirmed
In order to understand the protein expression differences associated with introduction ofbarely chromosome2H, iTRAQ technology was used for differential proteomics analysis forabove materials, and a total of589proteins were identified.
引文
[1]原亚萍，陈孝，肖世和.小麦穗发芽的研究进展[J].麦类作物学报，2003,23(3):136-139.
[2] Buonocore V, Petrucci T, Silano V. Wheat protein inhibitors of a-amylase[J].Phytochemistry,1977,16:811-820.
[3] Orton T J. Chromosomal variability in tissue cultures and regenerated plants ofHordeum[J]. Theoretical and Applied Genetics,56:101-112.
[4]肖世和，闫长生，张海萍，孙果忠.小麦穗发芽研究[M].北京：中国农业科学技术出版社，2004.
[5] Meera K. Management of Pre-harvest Sprout Damage in Wheat and Improvement of SoftWheat Quality in US [EB/OL]. http://www.ars.usda.gov/research/publications/publications.htm?seq_no_115=261927.
[6] Derera N F. Preharvest field sprouting in cereals[M].1989, CRC Press, Boca Raton.
[7] Depauw R M, Knox R E, Singh A K, Fox S L, Humphreys D G, Hucl P. Developingstandardized methods for breeding preharvest sprouting resistant wheat, challenges andsuccesses in Canadian wheat[J]. Euphytica,2012, DOI:10.1007/s10681-011-0611-y.
[8]张海峰，卢荣禾.小麦穗发芽抗性机理与遗传研究[J].作物学报,1993,19(6):523-530.
[9]原亚萍.大麦2H染色体上的α-淀粉酶抑制蛋白基因Isa-H1向普通小麦的导入、鉴定及表达[D].北京：中国农业科学院研究生院，2001.
[10]周凤明，解晓林，陈春，张晓慧，吕玉亮，王子明.穗上发芽对小麦种子发芽率的影响[J].江苏农业科学，2010,4:69-70.
[11]麦S.想给小麦种子管理部门一点建议-关于小麦穗发芽[EB/OL]. http://songyinming3.blog.163. om/blog/#m=0.
[12]唐建明.穗上发芽严重影响小麦生产[EB/OL]. http://blog.sciencenet.cn/blog-7170-256796.ht ml
[13] Jiang G L, Xiao S. Factorial cross analysis of pre-harvest sprouting resistance in whitewheat[J]. Field Crop Research,2005,91:63-69.
[14] Morad M M, Rubenthaler G L. Germination of soft white wheat and its effect on flourfractions, breadbaking, and crumb firmness[J]. Cereal Chemistry,60:413-417.
[15] Groos C, Gay G, Perretant M R, Gervais L, Bernard M, Dedryver F, Charmet. Study of therelationship between preharvest sprouting and grain color by quantitative trait loci analysisin a white×red grain bread-wheat cross. Theoretical and Applied Genetics,2002,104:39-47.
[16]何震天，陈秀兰，韩月澎.白皮小麦抗穗发芽研究概况[J].种子,2000,2:36-38.
[17] Mares D J. Preservation of dormancy in freshly harvested wheat grain[J]. AustrialianJouranl of Agricultural research,1983,34:33-38.
[18] King R W. Water uptake in relation to preharvest sprouting damage in wheat: graincharacteristics[J]. Austrialian Jouranl of Agricultural research,1984,35:337-345.
[19] Nielsen M T. Effects of weather variables during maturation on preharvest sprouting ofhard white winter wheat[J]. Crop Science,1984,24(2):779-782.
[20] Kihachi U. Eeffects of desiccation and a change in temperature on germination ofimmature grains of wheat (Triticum aestivum L.)[J]. Euphytica,2002,126:107-113.
[21] King R W, Richar R A. Water uptake in relation to preharvest sprouting damage in wheat:ear characteristics[J]. Austrialian Jouranl of Agricultural research,1984,35:327-336.
[22] King R W, Wettstein-Knowles Von P. Epiticular waxes and regulation of ear wetting andpre-harvest sprouting in barley and wheat[J]. Euphytica,122:157-166.
[23]秦代红.小麦抗穗发芽特性的研究[J].四川农业大学学报，1989,7(3):188-193.
[24] Derera N F, Bhatt G M. Germination inhibition of the bracts in relation to preharvestsprouting tolerance in wheat[J]. Cereal Research Communications,1980,8:199-201.
[25]沈正兴，俞世蓉，吴兆苏.小麦品种穗发芽抗性的研究[J].中国农业科学，1991,24(5):44-50.
[26] Metzger R J, Silbaugh B A. Location of Genes for Seed Coat Color in Hexaploid Wheat,Triticum aestivum L[J]. Crop Science,1970,10(5):495-496.
[27] Flinthan J E, Humphray S J. Red coat genes and wheat dormancy[J]. Aspects of AppliedBiology,1993,36:135-141.
[28]梅籍芳，曾道孝，赵德芳.麦类作物[M].北京：农业出版社，1990.
[29]蒋国梁，吴兆苏，陈兆夏.小麦穗发芽抗性鉴定与白粒抗源的筛选[J].上海农业学报，1992，8(3):9-14.
[30] Gubler F, Millar A A, Jacobsen J V. Dormancy release, ABA and pre-harvest sprouting[J].Current Opinion in Plant Biology,2005,8(2):183-187.
[31] Bewley J D. Seed germination and dormancy[J]. Plant Cell1997,9:1055-1066.
[32]张海峰，Zemetra R S, Liu C T.冬小麦穗发芽抗性及其鉴定方法研究[J].作物学报,1989,15(2):116-122.
[33]胡汉桥，原亚萍，王罡. ASI蛋白的特性及其应用研究进展[J].麦类作物学报，2000，20(4):81-85.
[34] Lazarus C M, Baulcombe D C, Martienssen R A. Alpha amylase genes of wheat are twomultigene families which are differentially expressed[J]. Plant Molecular Biology,1985,13-24.
[35] Baulcombe D C, Huttly A K, Martienssen R A, Barker R F, Jarvis MG. A novel wheatalpha-amylase gene (alpha-amy3)[J]. Molecular and General Genetics,1987,209:33-40.
[36] Mares D J, Gitta Oettler. α-amylase activity in developing tricales grains[J]. Journal ofCereal Science,1991,13:151-160.
[37] Sargeant J G. α-Amylase enzymes and starch degradation[J]. Cereal ResearchCommunications,1980,8:77-86.
[38] Jacobsen J V, Higgins T J V. Characterization of the alpha amylases synthesized byaleurone layers of Himalaya barley in response to gibberellic acid[J]. Plant Physiology,1982,70:1647-1653.
[39] Mundy J, Hejaard J, Svendsen I B. Characterization of a bifunctional wheat inhibitor ofendogenous α-Amylase and subtilisin[J]. FEBS Letters,1984,167:210-214.
[40] Mundy J, Svendsen I B, Hejaard J. Barley α-Amylase/subtilisin inhibitor. I. Isolation andcharacterization[J]. Carlsberg Research Communications,1983,48:81-90.
[41] Zawistowska U, Langstaff J, Friesen A D. Purification and characterization of twodouble-headed triticale isoinhibitors of endogenous α-Amylase and subtilisin[J]. Journal ofFood Biochemistry,1989,13:215-239.
[42] Yamagata H, Kunimatsu K, Kamasaka H, Kuramoto T, Iwasaki T. Rice bifunctionalα-amylase/subtilisin inhibitor: characterization, localization and changes in developing andgerminating seeds[J]. Bioscience Biotechnology Biochemistry,1998,62:978-985.
[43] Henry R J, Battershell V G, Brennan P S, Oono K. Control of wheat alpha-amylase usinginhibitors from cereals[J]. Journal of the Science of Food and Agriculture,1991,58:281-284.
[44] Yamasaki T, Deguchi M, Fujimoto T, Masumura T, Uno T, Kanamar K, Yamagat H. Ricebifunctional α-amylase/subtilisin Inhibitor: cloning and characterization of therecombinant inhibitor expressed in Escherichia coli[J]. Bioscience BiotechnologyBiochemistry,2006,70(5):1200-1209.
[45] Weselake R J, Macgregor A W, Hill R D. An endogenous α-amylase inhibitor in barleykernels[J]. Plant Physiology,1983,72:809–812.
[46] Svendsen I, Hejgaard J, Mundy J. Complete amino acid sequence of theα-Amylase/subtilisin inhibitor from barley[J]. Carlsberg Research Communications,1986,51:43-50.
[47] Ohtsubo K I, Richardson M. The amino acid sequence of a20kDa bifunctionalsubtilisin/alpha-amylase inhibitor from brain of rice (Oryza sativa L.) seeds[J]. FEBSLetters.1992,309:68–72.
[48] Sancho A I, Faulds C B, Svensson B, Bartolme B, Williamson G, Juge N. Cross-inhibitoryactivity of cereal protein inhibitors against α-amylases and xylanases [J]. Biochimica etBiophysica Acta,2003,1650:136-144.
[49] Abdul-Hussain S, Paulsen G M. Role of proteinaceous alpha-amylase inhibitors inpreharvest sprouting of wheat grain[J], Journal of Agriculture and Food Chemistry,1989,37:295-299.
[50] Munck L, Mundy J, Vaag P. Characterization of enzyme inhibitors in barley and theirtentative role in malting and brewing[J]. Journal of the American Society for BrewingChemistry,1985,43(1):35-38.
[51] Leah R, Mundy J, The bifunctional a-amylase/subtilisin inhibitor of barley: nucleotidesequence and patterns of seed-specific expression[J]. Plant Molecular Biology,1989,12:673–682.
[52] Mundy J, Rogers J C. Selective expression of a probable amylase/protease inhibitor inbarley (Hordeum vulgare) aleurone cells: comparison to the barley amylase/subtilisininhibitor[J].Planta,1986,169:51–63.
[53] Liu J H, Hill R D. Post-transcriptional regulation of bifunctional alpha-amylase/subtilisininhibitor expression in barley embryos by abscisic acid[J]. Plant Molecular Biology,1995,29:1087-1091.
[54] Furtado A，Henry R J，Scott K J，Meech S B. The promoter of the asi gene directsexpression in the maternal tissues of the seed in transgenic barley[J]. Plant MolecularBiology.52:787-799.
[55]肖世和，陈孝，徐惠君，林志珊，杜丽璞.大麦α-淀粉酶抑制基因对小麦α-淀粉酶的抑制作用研究[J].作物学报,1998,24(6):763-767.
[56]原亚萍，陈孝，肖世和，张伟.大麦α-淀粉酶抑制蛋白的提取及其功能检测[J].麦类作物学报,2005,25(1):40-43.
[57] Zawistowska U, Langstaff J, Bushuk W. Improving effect of a natural α-amylase inhibitoron the baking quality of wheat flour containing malted barley flour[J]. Journal of CerealScience,1988,8:207-209.
[58] Masojc P, Zawistowski J, Zawistowdka U. Polymorphism of endogenous alpha-amylaseinhibitor in barley[J]. Journal of Cereal Science,1994,20:43-49.
[59] Jarrett S J, Marschke R J, Symons M H, Gibson C E, Henry R J, Fox G P. Alpha-amylase/subtilisin inhibitor levels in Australian barleys[J]. Journal of Cereal Science,1997,25:261-266.
[60] Zawistowski J, Ansell M, Zawistowska U, Howes N K. A monoclonal antibody toendogenous alpha-amylase inhibitor of barley[J]. Jouranl of Cereal Science,1992,15(1):1-4.
[61] Bundock P C, Henry R J. Single nucleotide polymorphism, haplotype diversity andrecombination in the Isa gene of barley[J]. Theoretical and Applied Genetics,2004,109(3):543-551.
[62] Cronin J K, Bundock P C, Henry R J, Nevo E. Adaptive climatic molecular evolution inwild barley at the Isa defense locus[J]. Proceedings of the National Academy of Sciencesof the United States of America,2007,104(8):2773-2778.
[63] B nsager C B, Pr torius-Ibba M, Nielsen P K, Svensson B. Purification andcharacterization of the b-trefoil fold protein barley a-amylase/subtilisin inhibitoroverexpressed in Escherichia coli[J]. Protein Expression and Purification,2003,30:185–193.
[64] Micheelsen P A, stergaard P R, Lange L, Skj t M. High-level expression of the nativebarley α-amylase/subtilisin inhibitor in Pichia pastoris[J]. Journal of Biotechnology,2008,133:424-432.
[65] Vallee F, Kadziola A, Bourne Y, Juy M, Rodenburg K W, Svensson B, Haser, R. Barleyalpha-amylase bound to its endogenous protein inhibitor BASI: crystal structure of thecomplex at1.9resolution[J]. Structure,1998,6:649–659.
[66] Micheelsen O P, Vévodová J, Maria L D, stergaard P R, Friis E P, Wilson K, Skj t M.Structural and Mutational Analyses of the Interaction between the Barleyα-Amylase/Subtilisin Inhibitor and the Subtilisin Savinase Reveal a Novel Mode ofInhibition[J]. Journal of Molecular Biology,2008,380(4):681-690.
[67] Yuan Y P, Chen X, Xiao S H, Islam A K M R, Xin Z Y. Identification of wheat-Barley2Halien substitution lines[J]. Acta Botanica Sinica,2003,48(9):1096-1102.
[68]张艳，原亚萍，陈孝，李韬，张勇，何中虎.大麦2H染色体对小麦农艺和品质特性的影响[J].麦类作物学报,2007,27(3):402-406.
[69] Yuan Y P, Chen X, Xiao S H, Islam A K M R. The improving wheat starch quality bytransferring barley α-amylase inhibitor gene to wheat[C]. Proceedings of the10thInternational Wheat Genetics Symposium, Paestum, Italy,2003:463-466.
[70]刘淑会，陈新宏，赵继新，武军，李璋.抗穗发芽2D/2H小大麦易位系的鉴定和应用研究[J].华北农学报,2005,20(1):71-74.
[71] Molnár–Láng M, Szakács é, Linc G, Molnár I. Chromosome mediated gene transfer viaclassical hybridization techniques into wheat and detection of the alien chromosomes usingup-to-date molecular cytogenetic and genetic methods[J]. Hungarian Agricultural Research,2008,17(2-3):24-27.
[72]原亚萍,陈孝.大麦有益基因向普通小麦导入的研究进展[J].麦类作物学报,2004,24(4):129-132.
[73]陈新宏.大麦与小麦杂交及其杂种后代的创制、鉴定和应用研究[D].杨凌：西北农林科技大学，2003.
[74]陈新宏，赵继新，刘淑会，武军，陈锦刚，李酶.普通小麦与大麦杂交研究进展[J].西北农业学报,2005,14(6):38-43.
[75] von Bothmer R, Jacobsen N, Baden C, J rgensen R B, Linde-Laursen I. Anecogeographical study of the genus Hordeum. Systematic and ecogeographical studies oncrop genepools7[M].2nd ed. Rome: International Plant Genetic Resources Institute,1995.
[76] Komatsuda T, Tanno K, Salomon B, Bryngelsson T, von Bothmer R. Phylogeny in thegenus Hordeum based on nucleotide sequences closely linked to the vrs1locus (rownumber of spikelets)[J]. Genome,1999,42(5):973-981.
[77] Badr A, Müller K, Sch fer-Pregl R, El Rabey H, Effgen S, Ibrahim HH, Pozzi C, Rohde W,Salamini F. On the origin and domestication history of barley (Hordeum vulgare)[J].Molecular Biology and Evolution,2000,17(4):499-510.
[78]钟冠昌，穆素梅，张正斌.麦类远缘杂交[M].北京：科学出版社，2002.
[79]张起昌.小大麦远缘杂交种研究进展[J].小麦研究,2010,31(2):16-25.
[80] Shewry P R, Parmar S, Franklin J, Burgess S R. Analysis of a rare recombination eventwithin the multigenic Hor2locus of barley (Hordeum vulgare L.)[J]. Genetics Research,1990,55:171-176.
[81] Wettstein-Knowles V P. Barley (Hordeum vulgare)2n=14[M]. In: O’Brien S J, eds.,Genetic Maps, New York: Cold Spring Harbor Lab Press,1990:6125-6135.
[82] Tang J, Ohyama K, Kawaura K, Hashinokuchi H, Kamiya Y, Suzuki M, Muranaka T,Ogihara Y. A new insight into application for barley chromosome addition lines ofcommon wheat: achievement of stigmasterol accumulation[J]. Plant Physiology,2011,157(3):1555-1567.
[83] Kihara M, Okada Y, Iimure T Ito K. Accumulation and degradation of two functionalconstituents, GABA and β-Glucan, and their varietal differences in germinated barleygrains[J]. Breed Science,2007,57:85-89.
[84]陈朱希昭，陈孝，张文祥，杜振华.栽培大麦×普通小麦杂种发育的胚胎学观察[J].遗传学报,1984,11(5):368-373.
[85] Farrer W. Some notes on the wheat ‘Bobs’; its peculiarities, economic value and origin[J].Agric Gaz of NSW,1984,15:849-854.
[86] Kruse A. Hordeum×Triticum hybrids [J]. Hereditas,1973,73(1):157-161.
[87] Fedak G. Increased homoeologeous chromosome pairing in Hordeum vulgare×Triticumaestivum hybrids[J]. Nature,1977,266(5602):529-530.
[88] Takato K, Shigeo T, Takiko S. Isolation, identification and characterization of disomic andtranslocated barley chromosome addition lines of common wheat[J]. Euphytica,1997,96:289-296.
[89]陈孝，杜振华，张文祥，尹福玉，徐惠君，朱至清.大麦与提莫菲维小麦杂种及其再生植株[J].中国农业科学,1983,16(02):9-13.
[90]陈孝，杜振华，张文祥，尹福玉，徐惠君，朱至清.栽培大麦×普通小麦杂种及其再生植株[J].作物学报,1984,10(1):65-71.
[91]汪丽泉，朱汉如，戎均康，梁竹青，郑毅仁，管启良.苏联球茎大麦有利基因引入普通小麦的研究[J].遗传学报，1988(01):1-8+81.
[92]胡适全，戎均康，汪丽泉.普通小麦(6×)×苏联球茎大麦(4×)属间整倍体杂种后代的细胞遗传学研究[J].遗传学报，1991(01):67-72+103-104.
[93] Islam A K M R, Shepherd K W, Sparrow D H B: Addition of individual barleychromosomes to wheat[C]. Proceedings of the3rd International Barley GeneticsSymposium, Garching, B R D,1975.
[94] Islam A K M R, Shepherd K W, Sparrow D H B. Production and characterization of wheatbarley addition lines[C]. Proceedings of the5th International Wheat Genetics Symposium,New Delhi,1978.
[95]李璋，刘翠云，阎正录，王鸣.普通小麦×栽培大麦杂种植株及其回交后代的产生和鉴定[J].遗传学报,1987,14(3):188-192.
[96]刘翠云，李璋，魏小平，何一哲，阎正录.普通小麦×栽培大麦杂种体细胞无性繁殖系建立[J].西北植物学报,1991,11(1):8-14.
[97] Molnár-Láng M, Linc G, Logojan A, Sutka J. Production and meiotic pairing behaviour ofnew hybrids of winter wheat (Triticum aestivum)×winter barley (Hordeum vulgare)[J].Genome,2000,43:1045-1054.
[98] Islam A K M R, Shepherd K W, Sparrow D H B. Isolation and characterization ofeuplasmic wheat-barley chromosome addition lines[J]. Heredity,1981,46:161-174.
[99] Islam A K M R. Ditelosomic additions of barley chromosomes to wheat[C]. In: SakamotoS eds, Proceeding of the6th International Wheat Genetic Symposium, Maruzen Co. Ltd.,Kyoto,1983.
[100] Islam A K M R, Shepherd K W. Incorporation of barley chromosomes into wheat[M]. In:Bajaj Y P S eds, Biotechnology in Agriculture and Forestry, Vol13Wheat. Berlin:Springer-Verlag Publishers,1990:128-151.
[101] Islam A K M R, Shepherd K W. Isolation of a fertile wheat-barley addition line carryingthe entire barley chromosome1H[J]. Euphytica,2000,111:145-149.
[102] Islam A K M R, Shepherd K W. Substituting ability of individual barley chromosomes forwheat chromosomes.1. Substitutions involving barley chromosomes1,3and6[J]. PlantBreeding,1992,109:141-150.
[103] Islam A K M R, Shepherd K W. Substitution of barley chromosome4for group4homoeologous of wheat[C]. In: Z S Lin, Z Y Xin eds. Proceedings of the8th InternationalWheat Genetics Symposium, Beijing, China,1993:141-144.
[104] Islam A K M R, Shepherd K W. Induced pairing between wheat and barleychromosomes[C]. In: Miller T E, Koebner RMD, eds. Proceeding of the7th InternationalWheat Genetics Symposium, Cambridge, England,1988:309-314.
[105] Islam A K M R, Shepherd K W. Production of wheat barley recombinant chromosomesthrough induced homoeologous pairing.1. Isolation of recombinants involving barley arms3HL and6HL. Theoretical and Applied Genetics,1992,83:489-494.
[106]陈新宏，刘淑会，赵继新，武军，李璋，阎正禄.普通小麦-大麦异附加系的分子生物学技术鉴定[J].西北农林科技大学学报,2003,31(6):5-8.
[107]陈新宏，王小利，刘淑会，赵继新，武军，李璋，闫正禄，张改生.普通小麦×大麦杂交后代中间材料的GISH及PAGE鉴定[J].遗传学报，2004,31(7):730-740.
[108]刘淑会，陈新宏，李璋，闫正禄，蔡振虎.小麦-大麦异代换系的创制及鉴定研究[J].西北植物学报，2008，28(8):1554-1558.
[109]刘淑会，陈新宏，赵继新，武军，李璋.抗穗发芽小大麦易位系2D/2H的鉴定和应用研究[J].华北农学报，2005，20(1):71-74.
[110]刘淑会，蔡振虎，陈新宏，武军，赵继新，杨群慧，徐蕾.普通小麦-大麦杂交后代花粉母细胞减数分裂染色体行为的检测与分析[J].河南农业大学学报，2010，44(5):504-507.
[111]刘淑会，蔡振虎，陈新宏，武军，赵继新，杨群慧.小麦×大麦杂种后代早熟衍生系的选择和鉴定[J].山西农业大学学报（自然科学版），2011，31(2):142-146.
[112]刘淑会，蔡振虎，陈新宏，武军，赵继新，杨群慧.普通小麦-大麦杂种后代衍生系抗旱性鉴定[J].甘肃农业大学学报，2010，45(6):60-65.
[113] Molnár-Láng M, Galiba G, Kovács G, Sutka J. Changes in the fertility and meioticbehaviour of barley (Hordeum vulgare)×wheat (Triticum aestivum) hybrids regeneratedfrom tissue cultures[J]. Genome,1991,34(2):261-266.
[114] Szakács E, Molnár-Láng M. Development and molecular cytogenetic identification of newwinter wheat-winter barley ('Martonvásári9kr1'-'Igri') disomic addition lines[J]. Genome,2007,50(1):43-50.
[115] Szakács E, Molnár-Láng M. Identification of new winter wheat–winter barley additionlines (6HS and7H) using fluorescence in situ hybridization and the stability of the whole‘Martonvásári9kr1’–‘Igri’ addition set[J]. Genome,2010,53(1):35-44.
[116] Molnár-Láng M, Novotny C, Linc G, Nagy E D. Changes in the meiotic pairing behaviourof a winter wheat-winter barley hybrid maintained for a long term intissue culture, andtracing the barley chromatin in the progenies using GISH and SSR markers[J]. PlantBreeding,2005,124(3):247-252.
[117] Molnár-Láng M, Kruppa K, Cseh A, Bucsi J, Linc G. Identification and phenotypicdescription of new wheat–six-rowed winter barley disomic additions[J]. Genome,2012,55:302-311.
[118] Molnár I, Linc G, Dulai S, Nagy E D, Molnár-Láng M. Ability of chromosome4H tocompensate for4D in response to drought stress in a newly developed and identifiedwheat-barley4H(4D) disomic substitution line[J]. Plant Breeding,2007,126:369-374.
[119] Kruppa K, Türk si E, Szakács é, Cseh A, Molnár-Láng M. Development and identificationof a4HL.5DL wheat/barley centric fusion using GISH, FISH and SSR markers[J]. CerealResearch Communications,2012, DOI:10.1556/RC.2012.0038.
[120] Molnár-Láng M, Linc G, Friebe B R, Sutka J. Detection of wheat-barley translocations bygenomic in situ hybridization in derivatives of hybrids multiplied in vitro[J]. Euphytic,2000,112:117-123.
[121] Nagy E D, Molnár-Láng M, Linc G, Láng L. Identification of wheat-barley translocationsby sequential GISH and two-colour FISH in combination with the use of geneticallymapped barley SSR markers[J]. Genome,2002,45(6):1238-1247.
[122] Dulai S, Molnár I, Haló B, Molnár-Láng M. Photosynthesis in the7H Asakazekomugi/Manas wheat/barley addition line during salt stress[J]. Acta AgronomicaHungarica,2010,58:367-376.
[123] Darko E, Barnabás B, Molnár-Láng M. Characterization of newly developed wheat/barleyintrogression lines in respect of aluminium tolerance[J]. American Journal of PlantSciences,2012,3:1462-1469
[124] Cseh A, Kruppa K, Molnár I, Rakszegi M, Dole el J, Molnár-Láng M. Characterization ofa new4BS.7HL wheat–barley translocation line using GISH, FISH, and SSR markers andits effect on the β-glucan content of wheat[J]. Genome,2011,54(10):795-804.
[125] Suchánková P, Kubaláková M, Koválová P, Barton J, Bíhalíková J, Molnár-Láng M, EndoT R. Dissection of the nuclear genome of barley by chromosome flow sorting[J].Theoretical and Applied Genetics,2006,113:651-659.
[126] Linc G, Molnár-Láng M. Two methods for the development of wheat/barley disomicadditions in winter wheat varieties, and identification using molecular cytogenetictechniques (C-banding, GISH)[J]. Novenytermeles Hungary,2003,521:3-13.
[127] Sepsi Adél, Bucsi J. Physical mapping of the7D chromosome using a wheat/barleytranslocation line (5HS.7DL) produced in a Martonvásári wheat background usingmicrosatellite markers[J]. Cereal Research Communications,2009,37(Suppl.):297-300.
[128] McGuire P E, Qualset C O. Transfer of the Yd2barley yellow dwarf virus resistance genefrom barley to wheat[M]. In: Burnett P A eds. World perspectives on barley yellow dwarf.CIMMYT, Mexico,1990:476-481.
[129]阎新甫，刘文轩.大麦DNA导入小麦诱导抗白粉病变异的研究[J].华北农学报，1992，7(2):40-45.
[130]原亚萍,陈孝,肖世和,辛志勇,张增艳,林志珊,马有志,胡汉桥.应用基因组原位杂交及RFLP标记鉴定小麦中的大麦染色体[J].遗传学报，2000,27(12):1080-1083.
[131] Bernardo R. Prediction of maize single-cross performance using RFLPs and informationfrom related hybrids[J]. Crop Science,1994,34:20-25.
[132] Sharp P J, Shao S, Desai S, Gale M D. The isolation, characterization and application inthe Triticeae of a set wheat RFLP probes identifying each homoeologous chromosomearm[J]. Theoretical and Applied Genetics,1989,78(3):342-348.
[133] Blake T K, Kadyrzhanova D, Shepherd K W, Islam A K M R, Langridge P L, McDonald CL, Erpelding J, Larson S, Blake N K, Talbert L E. STS-PCR markers appropriate forwheat-barley introgression[J]. Theoretical and Applied Genetics,1996,93(5-6):826-832.
[134] Murai K, Taketa S, Islam A K M R, Shepherd K W. Barley allele-specific amplicons usefulfor identifying wheat-barley recombinant chromosomes[J]. Genes&Genetic Systems,2000,75:131-139.
[135] Williams J G K, Kubelik A R, Livak K J, Rafalski J A, Tingey S V. DNA polymorphismsamplified by arbitrary primers are useful as genetic markers[J]. Nucleic Acids Research,1990,18:6531-6535.
[136] Welsh J, McClelland M. Fingerprinting genomes using PCR with arbitrary primers.Nucleic Acids Research,1990,18:7231-7238.
[137] Koebner R M D. Generation of PCR-based markers for the detection of rye chromatin in awheat background[J]. Theoretical and Applied Genetics,1995,90(5):740-745.
[138] Talbert L E, Bruckner P L, Smith L Y, Sears R, Martin T J. Development of PCR markerslinked to resitance to wheat streak mosaic virus in wheat[J]. Theoretical and AppliedGenetics,1996,93(3):463-467.
[139] Paran I, Michelmore R W. Development of reliable PCR-based markers linked to downymildew resistance genes in lettuce[J]. Theoretical and Applied Genetics,1993,85:985-993.
[140]黄朝峰，张文俊.大麦6H染色体特异性标记的筛选和鉴定[J].遗传学报,2000,27(8):713-718.
[141] Tautz D, Renz M. Simple sequences are ubiquitous repetitive components of eukaryoticgenomes[J]. Nucleic acids research,1984,12(10):4127-4138.
[142] Malysheva L, Sjakste T, Matzk F, Roder M, Ganal M. Molecular cylogenetic analysis ofwheat-barley hybrids using genomic in silu hybridization and barley microsatellitemarkers[J].Genome,2003,46:314-322.
[143] Zabeau M, Vos P. European Patent Application, EP0534858[P].1993.
[144] Vos P, Hogers R, Bleeker M, Reijans M, van de Lee T, Hornes M, Frijters A, Pot J,Peleman J, Kuiper M, Zabeau M. AFLP: a new technique for DNA fingerpring[J]. NucleicAcids Research,1995,23(21):4407-4414.
[145] Serizawa N, Nasuda S, Endo T R. Barley chromosome addition lines of wheat forscreening of AFLP markers on barley chromosomes[J] Genes&Genetic Systems,2001,76:107-110.
[146] Ma X F, Fang P, Gustafson P J. Polyploidization-induced genome variation in triticale[J].Genome,2004,47:839-848.
[147] Ma X F, Gustafson J P. Timing and rate of genome variation in Triticale followingallpolypoidization[J]. Genome,2004,47:839-848.
[148]张勇，刘朝辉，刘成，杨足君，邓科君，彭金华，周建平，李光蓉，唐宗祥，任正隆.外源种质导入引发小麦表观遗传变异的MSAP分析[J].科学通报,2007,52(24):2857-2865.
[149]李锁平，张大乐，王秀娥，亓增军，刘大钧.节节麦和黑麦杂种F1及双二倍体中基因组变异分析[J].作物学报，2012，38(6):996-1002.
[150]王立新，常利芳，黄岚，王晓维，赵昌平.小麦AFLP片段序列多态性分析和AFLP-SCAR标记的研究[J].麦类作物学报,2007,27(6):943-951.
[151]张丽，颜泽洪，郑有良，刘登才，代寿芬，张连全，魏育明.小麦中国春背景下长穗偃麦草Ee染色体组特异AFLP及STS标记的建立[J].农业生物技术学报,2008,16(3):465-473.
[152] Wolf A P, Matzke M A. Epigenetics: regulation throuth repression[J]. Science,1999,286(5439):481-486.
[153] Wu Ct, Morris J R. Genes, genetics, and epigenetics: a correspondence[J]. Science,2001,293(5532):1103-1105.
[154] Feng S H, Jacobsen S E, Reik W. Epigenetic reprogramming in plant and animaldevelopment[J]. Science,2010,330(6604):622-627.
[155] Saze H, Mittelsten Scheid O, Paszkowski J. Maintenance of CpG methylation is essentialfor epigenetic inheritance during plant gametogenesis[J]. Nature Genetics,2003,34(1):65-69.
[156] Flavell R B. Inactivation of gene expression in plants as a consequence of specificsequence duplication[J]. Proceedings of the National Academy of Sciences of the UnitedStates of America,1994,91(9):3490-3496.
[157] Bezd k M, Koukalová B, Kuhrová V, Vyskot B. Differential sensitivity of CG and CCGDNA sequences to ethionine-induced hypomethylation of the Nicotiana tabacumgenome[J]. FEBS Letters,1992,300(3):268-270.
[158] Wagner I, Capesius I. Determination of5-methylcytosine from plant DNA byhigh-performance liquid chromatography[J]. Acta Biochimica et Biophysica Sinica,1981,654(1):52-56.
[159] Chan S W, Henderson I R, Jacobsen S E. Gardening the genome: DNA methylation inArabidopsis thaliana[J]. Nature Reviews Genetics,2005,6(5):351-360.
[160] Plant Epigenetics[M]. In: Kovalchuk I, Zemp F J, eds., Humana Press,2010.
[161] Xiong L Z, Xu C G, Saghai Maroof M A, Zhang Q F. Patterns of cytosine methylation inan elite rice hybrid and its parental lines, detected by a methylation-sensitive amplificationpolymorphism technique[J]. Molecular and General Genetics,1999,261(3):439-446.
[162] Nelson M, McClelland M. Effect of site-specific methylation on DNA modificationmethyltransferases and restriction endonucleases[J]. Nucleic Acids Research,1989,17(suppl):389-415.
[163] Sherman J D, Talbert L E. Vernalization-induced changes of the DNA methylation patternin winter wheat[J]. Genome,2002,45:253-260.
[164] Meng F R, Li Y C, Yin J, Liu H, Chen X J, Ni Z F, Sun Q X. Analysis of DNA methylationduring the germination of wheat seeds[J]. Biologia Plantarum,2012,56(2):269-275.
[165] Qi B, Zhong X F, Zhu Bo, Zhao Na, Xu L Y, Zhang H K, Yu Xiaoming, Liu B. Generalityand characteristics of genetic and epigenetic changes in newly synthesized allotetraploidwheat lines [J]. Journal of Genetics and Genomics,2010,37:737-748.
[166] Adams M D, Kelley J M, Gocayne J D, Dubnick M, Polymeropoulos M H, Xiao H, MerrilC R, Wu A, Olde B, Moreno R F, Kerlavage A R, McCombie W R, Venter J C.Complementary DNA sequenceing: expressed sequence tags and human genome project[J].Science,1991,252:1651-1656.
[167] Nasuda S, Kikkawa Y, Ashida T, Islam A K M R, Sato K, Endo T R. Chromosomalassignment and deletion mapping of barley EST markers[J]. Genes&Genetic Systems,2005,80(5):357-366.
[168] Endo T R. Induction of chromosomal structural changes by a chromosome of Aegilopscylindrica L. in common wheat[J]. Journal of Heredity,1988,79:366-370.
[169] Endo T R. Gametocidal chromosomes and their induction of chromosome mutations inwheat[J]. The Japanese Journal of Genetics,1990,65,3:135-152.
[170] Ashida T, Nasuda S, Sato K, Endo T R. Dissection of barley chromosome5H in commonwheat[J]. Genes&Genetic Systems,2007,82:123-133.
[171] Sakai Kazuhiko, Nasuda S, Sato K,Endo T R. Dissection of barley chromosome3H incommon wheat and a comparison of3H physical and genetic maps[J]. Genes&GeneticSystems,2009,84(1):25-34.
[172] Sakata M, Nasuda S, Endo T R. Dissection of barley chromosome4H in common wheatby the gametocidal system and cytological mapping of chromosome4H with ESTmarkers[J]. Genes&Genetic Systems,2010,85(1):19-29.
[173] Joshi G P, Nasuda S, Endo T R. Dissection and cytological mapping of barley chromosome2H in the genetic background of common wheat[J]. Genes&Genetic Systems,2011,86(4):231-248.
[174] Wang M J, Zou H D, Lin Z S, Wu Y, Chen X, Yuan Y P. Expressed sequence tag-PCRmarkers for identification of alien barley chromosome2H in wheat[J]. Genetics andMolecular Research,2012,11(3):3452-3463.
[175] Schaffer R, Landgraf J, Accerbi M, Simon V, Larson M, Wisman E. Microarray analysis ofdiurnal and circadian-regulated genes in Arabidopsis[J]. Plant Cell,2001,13(1):113-123.
[176] Cho S, Garvin D F, Muehlbauer GJ. Transcriptome analysis and physical mapping ofbarley genes in wheat–barley chromosome addition lines[J]. Genetics,2006,172(2):1277-1285.
[177] Bilgic H, Cho S, Garvin D F, Muehlbauer G J. Mapping barley genes to chromosome armsby transcript profiling of wheat–barley ditelosomic chromosome addition lines[J]. Genome,2007,50:898-906.
[178] Bhat P R, Lukaszewski Adam, Cui Xinping, Xu Jin, Svensson Jan T, Wanamaker S,Waines J G, Close T J. Mapping translocation breakpoints using wheat microarray[J].Nucleic Acids Research,2007,35(9):2936-2943.
[179] Aversan R, Ercolano M R, Caruso I, Fasano Carlo, Rosellini Daniele, Carputo D.Molecular tools for exploring polyploid genomes in plants[J]. International Journal ofMolecular Sciences,2012,13(8):10316-10335.
[180] Bachem C W B, Vanderhoeven R S, Debrujin S M, Vreugdenhil D, Zabeau M.Visualization of differential gene expression using a novel method of RNA fingerpritingbased on AFLP-analysis of gene expression during potato tuber development[J]. The PlantJournal,1996,9(5):745-753.
[181] He P, Friebe B R, Gill B S, Zhou J M. Allopolyploidy alters gene expression in the highlystable hexaploid wheat[J]. Plant Molecular Biology,2003,52:401-414.
[182] Adams K L, Cronn R, Percifield R, Wendel J F. Genes duplicated by polyploidy showunequal contributions to the transcriptome and organ-specific reciprocal silencing[J].Proceedings of the National Academy of Sciences of the United States of America,2003,100:4649-4654.
[183] Kashkush K, Feldman M, Levy A A. Gene loss, silencing and activation in a newlysynthesized wheat allotetraploid[J]. Genetics,2000,160:1651-1659.
[184] Osborne T B. The proteins of the wheat kerne[M]. Washingto D C: Garnegie institution ofWashington,1907.
[185] Branlard G, Dardevet M, Saccomano R, Lagoutte F, Gourdon. Genetic diversity of wheatstorage proteins and bread wheat quality[J]. Euphytica,2001,119:59-67.
[186] Hart G E, Islam A K M R, Shepherd K W. Use of isozymes as chromosome markers in theisolation of wheat-barley chromosome addition lines[J]. Genetical Research,1980,36(3):311-325.
[187] Hart G E, Tuleen A. Introduction and characterization of alien genetic material[A].Isozymes in Pant Genetics and Breeding. part A [C]. In: Tanksley S, Orton T J, eds.,Amsterdam: Elsevier Science Publishers,1983, pp339-362.
[188] Powling A, Islam A K M R, Shepherd K W. Isozymes in wheat-barley hybrid derivativelines[J]. Biochemical Genetics,1981,19(3-4):237-253.
[189] Islam N, Woo S H, Tsujimoto H, Kawasaki H, Hirano H. Proteome approaches tocharacterize seed storage proteins related to ditelocentric chromosomes in common wheat(Triticum aestivum L.)[J]. Proteomics,2002,2:1146-1155.
[190] Islam N, Tsujimoto H, Hirano H. Proteome analysis of diploid, tetraploid and hexaploidwheat: Towards understanding genome interaction in protein expression[J]. Proteomics,2003,3:549-557.
[191] Islam N, Tsujimoto H, Hirano H. Wheat proteomics: Relationship between finechromosome deletion and protein expression[J]. Proteomics,2003,3:307-316.
[192] Kim K H, Kamal A H M, Shin K H, Choi J S, Heo H Y, Woo S H. Large-scale proteomeinvestigation in wild relatives(A, B, and D genomes) of wheat[J]. Acta Biochimica etBiophysica Sinica,2010,42(10):709-716.
[193] Kim A, Kamal A H M, Seo Y W, Park C S, Nam J, Kim S II, Choi J S, Woo S H. Leafproteome analysis of wheat-rye translocation lines[J]. Australian Journal of Crop Science,2011,5(12):1670-1677.
[194]崔会肖.小麦-簇毛麦染色体代换系、易位系SSR、AFLP和双向电泳比较[D].天津：天津师范大学，2005.
[195]李纯正.簇毛麦V染色体对小麦-簇毛麦染色体代换系（6A/6V）、易位系（6DL/6VS）蛋白质组的影响及与白粉病抗性关系探讨[D].天津：天津师范大学,2008.
[196] Yanaka M, Takata K, Tersawa Y, Ikeda T M. Chromosome5H of Hordeum speciesinvolved in reduction in grain hardness in wheat genetic background[J]. Theoretical andApplied Genetics,2011,123:1013-1018.
[197] Smolka M B, Zhou H, Purkayastha S, Aebersold R. Optimization of the isotope-codedaffinity tag-labeling procedure for quantitative proteome analysis[J]. AnalyticalBiochemisty,2001,297(1):25-31.
[198] Oeljeklaus S, Meyer H E,Warscheid B. Advancements in plant proteomics usingquantitative mass spectrometry[J]. Journal of Proteomics,2009,72:545-54.
[199] Xiong L Z, Xu C G, Saghai Maroof M A, Zhang Q F. Patterns of cytosine methylation inan elite rice hybrid and its parental lines, detected by a methylation-sensitive amplificationpolymorphism technique[J]. Molecular and General Genetics,1999,261(3):439-446.
[200] ng D W, Zhang C, Miller M, Shen Z, Briggs S P, Chen Z J. Proteomic divergence inArabidopsis autopolyploids and allopolyploids and their progenitors[J]. Heredity,2012,108(4):419-430.
[201] Ford K L, Cassin Andrew, Bacic A. Quantitative proteomic analysis of wheat cultivarswith differing drought stress tolerance[J]. Frontiers in Plant Science,2011, doi:10.3389/fpls.2011.00044
[202] Chen Q, Jahier J, Cauderon Y. Production and cytogenetic analysis of BC1, BC2, and BC3progenies of an intergeneric hybrid between Triticum aestivum (L.) Thell. and tetraploidAgropyron cristatum (L.) Gaertn[J]. Theoretical and Applied Genetics,1992,83:698-703.
[203]王关林，方宏筠.植物基因工程[M].北京：科学出版社，2002.
[204] Varshney R K, Marcel T C, Ramsay L, Russell J, R der M S, Stein N, Waugh R,Langridge P, Niks R E, Graner A. A high density barley microsatellite consensus map with775SSR loci[J]. Theoretical and Applied Genetics,2007,114:1091-1103.
[205] Liu Z W, Biyashev R M, Saghai Maroof M A. Development of simple sequence repeatDNA markers and their integration into a barley linkage map[J]. Theoretical and AppliedGenetics,1996,93:866-876.
[206] Ramsay L, Macaulay M, degli Ivanissevich S, MacLean K, Cardle L, Fuller J, Edwards KJ,Tuvesson S, Morgante M, Massari A, Maestri E, Marmiroli N, Sjakste T, Ganal M, PowellW, Waugh R. A simple sequence repeat-based linkage map of barley[J]. Genetics,2000,156:1997-2005.
[207] Islam A K M R, Shepherd K W. Meiotic restitution in wheat-barley hybrids[J].Chromosoma,1980,78:363-372.
[208] Sears E R. Relationships of chromosomes2A,2B and2D with their rye homoeologue[C].In: Finalay K W, Shepherd K W eds., Proceeding of the3rd International Wheat GeneticsSymposium, Canberra, Austrilia,1968.
[209] Sears E R. The transfer of leaf-rust resistance from Aegilops umbellulata to wheat[M].Book Genetics in plant breeding. Brook-haven Symposia in Biology,1956.
[210] Sears E R. Agropyron-wheat transfers induced by homoeologous pairing[C]. Proceedingof the4th International Wheat Genetics Symposium, Columbia, Mo.,1986.
[211] Knott D R, Dvo ák J. Agronomic and quality characteristics of wheat lines with leaf rustresistance derived from Triticum speltoides[J]. Canadian Journal of Genetics and Cytology,1981,23(3):475-480.
[212] Endo T R. Induction of chromosomal structural changes by a chromosome of Aegilopscylindrica L. in common wheat[J]. Journal of Heredity,1988,79(5):366-370.
[213] Lapitan N L V, Sears R G, Gill BS. Translocations and other karyotypic structural changesin wheat x rye hybrids regenerated from tissue culture[J]. Theoretical and Applied Genetics,1984,68(6):547-554.
[214] Friebe B, Mukai Y, Gill B S, Cauderon Y. C-banding and in-situ hybridization analyses ofAgropyron intermedium, a partial wheat x Ag. intermedium amphiploid, and six derivedchromosome addition lines[J]. Theoretical and Applied Genetics,1992,84:889-905.
[215] Bayliss M W. Chromosomal variation in plant tissues inculture[J]. International Review ofCytology,1980,11:113-144.
[216] Larkin P J, Scowcroft W R. Somaclonal variation–A novel source of variability from cellcultures for plant improvement[J]. Theoretical and Applied Genetics,1980,60:197-204.
[217] Fedak G, Grainger J. Chromosome instability in somaclones of a Triticum aestivum×Hordeum vulgare hybrid[J]. Canadian Journal of Genetics and Cytology,1986,28:618-623.
[218] Jiang J, Friebe B, Gill B S. Recent advances in alien gene transfer in wheat[J]. Euphytica,1994,73:199-212.
[219]辛志勇，徐惠君，陈孝，林志珊，周广和，钱幼亭，成卓敏，Larkin P J, Banks P,Appels R, Clarke B, Brettell R I S.应用生物技术向小麦导入黄矮病抗性的研究[J].中国科学(B辑),1991,21:36-42.
[220] Banks P M, Larkin P J, Bariana H S, Lagudah E S, Apples R, Waterhous P M, Brettell R IS, Chen X, Xu H J, Xin Z Y, Qian T, Zhou X M, Cheng Z M, Zhou G H. The use of cellculture for subchromosomal introgressions of barley yellow dwarf virus resistance fromThinopyrum intermedium to wheat[J]. Genome,1995,38(2):395-405
[221]李洪杰，李义文，张艳敏，李辉，郭北海，王子宁，温之雨，刘志勇，朱至清，贾旭.组织培养创造抗白粉病小麦-簇毛麦染色体易位及分子标记辅助选择[J].遗传学报，2000，27(7):608-613.
[222]李辉，陈孝，辛志勇，徐惠君，马有志.组织培养诱导普通小麦-簇毛麦抗白粉易位系研究[J].河北农业科学,2005,9(1):1-5.
[223]崔志富.小麦-中间偃麦草抗黄矮病易位系的细胞与分子水平鉴定[D].福建：福建农林大学，2006.
[224] Wang M J, Zhang Y, Lin Z S, Ye X G, Yuan Y P, Ma W, Xin Z Y. Development ofEST-PCR markers for Thinopyrum intermedium chromosome2Ai#2and their applicationin characterization of novel wheat-grass recombinants[J]. Theoretical and AppliedGenetics,2010,121:1369-1380.
[225]原亚萍，陈孝.大麦有益基因向普通小麦导入的研究进展[J].麦类作物学报，2004，24(4):129-132.
[226]曲鲁江，李显耀，杜志强，张龙超，杨宁.微卫星PCR产物变性与非变性PAGE-银染检测方法的比较[J].遗传,2004,26(4):522-524.
[227] Warburton M. Laboratory protocols: CIMMYT Applied Genetic EngineeringLaboratory[M]. Mexico D.F.: CIMMYT,2005.
[228] IRDye Fluorescent AFLP Protocol for Large Plant Genome Anaylsis[EB/OL].http://biosupport.licor.com.
[229]4300DNA Analyzer Applications Manual[EB/OL]. http://www.licor.com/bio/applications/aflp/aflpSupportMaterials.jsp
[230] SagaMX AFLP V.3.3Analysis User Guide v1.0n (9/2008)[EB/OL].http://www.licor.com/bio/applications/aflp/aflpSupportMaterials.jsp
[231] AFLP protocol[EB/OL]. www.css.msu.edu/bean/_pdf/AFLP_protocol.pdf
[232] Hazen S P, Leroy P, Ward R. AFLP in Triticum aestivum L.: patterns of genetic diversityand genome distribution[J]. Euphytica,2002,125(1):89-102.
[233] Xu D H, Ban T. Conversion of AFLP markers associated with FHB resistance in wheatinto STS markers with an extension-AFLP method[J]. Genome,2004,47(4):660-665.
[234]方宣钧，吴为人，唐纪良.作物DNA标记辅助育种[M].北京：科学出版社，2001.
[235] Bassam J B, Caetano-Anollés G, Gresshoff P M. Fast and sensitive silverstaining of DNAin polyacrylamide gels[J]. Analytical Biochemistry,1991,196(1):80-83.
[236] Srinivas G, Huang Y, Carver B F, Mornhinweg D. AFLP genetic diversity analysis inRussian wheat aphid resistant wheat accessions[J].2012,185(1):27-35.
[237] Mohan A, Kulwal P, Singh R, Kumar V, Mir R R, Kumar J, Prasad M, Balyan H S, GuptaP K. Genome-wide QTL analysis for pre-harvest sprouting tolerance in bread wheat[J].2009,168(3):319-329.
[238] Shan X, Blake T K, Talbert L E. Conversion of AFLP markers to sequence-specific PCRmarkers in barley and wheat[J]. Theoretical and Applied Genetics,1999,98:1072-1078.
[239]彭金华.黑麦染色体附加对小麦基因组表观遗传影响的研究[D].四川：电子科技大学，2010.
[240] Seo Y W, Jang C S, Johnson J W. Development of AFLP and STS markers for identifyingwheat-rye translocations possessing2RL[J]. Euphytica,2001,121:279-287.
[241] Dong Z Y, Wang Y M, Zhang Z J, Shen Y, Lin X Y, Ou X F, Han F P, Liu Bao. Extent andpattern of DNA methylation alteration in rice lines derived from introgressivehybridization of rice and Zizania latifolia Griseb[J]. Theoretical and Applied Genetics,2006,113(2):196-205.
[242] Liu B, Vega J M, Segal G, Abbo S, Rodova M, Feldman M. Rapid genomic changes innewly synthesized amphiploids of Triticum and Aegilops. I. Changes in low-copynoncoding DNA sequences[J]. Genome,1998,41(2):272-277.
[243] Liu B, Vega J M, Feldman M. Rapid genomic changes in newly synthesized amphiploidsof Triticum and Aegilops. II. Changes in low-copy coding DNA sequences[J]. Genome,1998,41(4):535-542.
[244] Liu B, Piao H M, Zhao F S, Zhao J H, Liu Zhenlan, Huang B Q. DNA MethylationChanges in Rice Induced by Zizania Latifolia (Griseb.) DNA Introgression[J]. Hereditas,1999,131:75-78．
[245] Liu B, Xu C M, Zhao N, Qi B, Kimatu N J, Pang J S, Han F P. Rapid genomic changes inpolyploid wheat and related species: implications for genome evolution and geneticimprovement[J]. Journal of Genetics and Genomics,2009,36(9):519-528.
[246] Cho S, Garvin D F, Muehlbauer GJ. Transcriptome analysis and physical mapping ofbarley genes in wheat–barley chromosome addition lines[J]. Genetics,2006,172(2):1277-1285.
[247]张敏.利用cDNA-AFLP技术研究附加/丢失异源染色体小麦的基因表达差异[D].吉林：东北师范大学，2007.
[248] Vuylsteke M, Peleman J D, Eijk van M J T. AFLP-based transcript profiling (cDNA-AFLP)for genome-wide expression analysis[J]. Nature Protocols,2007,2:1399-1413.
[249] Dang J, Copeland L. Studies of the fracture surface of rice grains using environmentalscanning electron microscopy[J]. Journal of the Science of Food and Agricultrue,2004,84(7):707-713.
[250] Bradford M M. A rapid and sensitive method for the quantitation of microgram quantitiesof protein utilizing the principle of protein-dye binding[J]. Analytical Biochemistry,1976,72:248-254.
[251] Ye J, Fang L, Zheng H, Zhang H, Zhang Y, Chen J, Zhang Z, Wang J, Li S, Li R, Bolund L,Wang J. WEGO: a web tool for plotting GO annotations[J]. Nucleic Acids Research,2006,34(2):293-297.
[252]张瑞奇，荣曼，张守忠，胡琳，许为钢，陈佩度.普通小麦籽粒硬度与胚乳组成及显微结构的关系[J].中国农业科学，2011，44(9):1753-1762.