抗源望水白赤霉病抗性遗传分析和QTL定位研究
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摘要
以小麦赤霉病抗源望水白与感病品种安农8455和Alondra构建的重组自交系为材料，运用主基因+多基因模型进行抗性遗传分析，并以望水白/安农8455的RIL群体进行与赤霉病抗性连锁的SSR(简单重复序列)分子标记筛选及QTL（数量性状位点）定位，获得如下结果：
抗源望水白赤霉病抗性遗传分析
抗源望水白抗性由2对主基因控制，且符合两对主基因+多基因模型。2001年、03年、04年望水白/安农8455抗性分别符合E-2-6模型（两对主基因+多基因的加性－加性模型）、E-1-8模型（两对主基因+多基因,主基因间为重叠作用）、E-1-8模型（两对主基因+多基因，主基因间为重叠作用）。主基因的遗传率较高分别为63.8%、69.02％、73.66％，多基因的遗传率分别为13.94%、21.38%、16.80%。2003年望水白/Alondra抗性符合E-1-9模型（两对主基因+多基因，主基因间为相互抑制作用）。主基因的遗传率为85.18％，多基因的遗传率较低为2.74%。
望水白/安农8455群体的SSR标记
在望水白/安农8455群体的抗感亲本间的引物筛选表明，多态性为33%。根据群体的SSR标记，在2001年、03年和04年3年中至少有2年资料表明与抗赤性连锁的SSR标记有14个，定位在3B、2A、2B和5D染色体上。已经发表的资料表明，在望水白/Alondra群体中至少两次鉴定资料中都出现的与抗赤基因连锁的SSR标记共有8个，定位在3B、4B、2D、4D染色体上。
3. 抗源望水白的QTL定位
通过构建望水白/安农8455群体的连锁图，把25个SSR标记定位在5条染色体上。2001年、03年和04年的抗性资料表明在抗源的3B染色体上存在一个QTL。2001年和03年的抗性资料表明在2A上存在一个抗性QTL。根据2001年、03年和04年的抗性资料分析3B上的QTL分别位于Xgwm533.1-Barc133、 Xgwm389-Xgwm533.1、 Xgwm533.1-Barc133之间，LOD值为2.47、5.57、2.52，能解释10.56%、21.97%、8.37%的表型变异。2001年和03年的抗性资料分析2A上抗性QTL，都位于Xgwm425-Xgwm372之间，LOD值分别为2.5、2.80，能解释9.76%、11.04%。已发表的望水白/Alondra群体中抗源望水白的3B染色体上存在抗性QTL，位于Barc147-Xgwm493之间，能解释13.7%的表型变异。
对抗源望水白的抗性进行遗传分析，得出其抗赤性为两对主基因控制，有助于对抗源抗性机制的理解。运用SSR标记技术，获得与抗赤性连锁的遗传标记，在抗源望水白3B染色体上存在抗性QTL，为抗赤性分子标记辅助育种及最终克隆抗病基因奠定基础。
The RIL populations of Wangshuibai /annong8455 and Wangshuibai/Alondra to wheat scab were analysised with major gene plus ploygenes. The SSR（simple sequence repeat） marker analysis and QTL(quantitative trait loci) tagging of wheat scab resistance were carried out with the population of Wangshuibai/Annong8455.The results showed as follows:
the genetic analysis of Wangshuibai to scab
The scab resisitance of Wangshubai fits the genetic model of two major genes plus polygenes. The RIL populations of Wangshuibai /annong8455 to scab in 2001,2003 and 2004 fits the model of E-2-6(two major genes plus polygenes with additive –epistatic effect), the model of E-1-8(two major genes plus polygenes with effect) and the model of E-1-8. The heritability of major genes is 63.8%,69.02% and 73.66%.The heritability of polygenes is 13.94%,21.38% and 16.08%. The RIL populations of Wangshuibai /Alondra to scab in 2003 fits the model of E-1-9(two major genes plus polygenes with effect).The heritability of major genes is 85.18%.The heritability of polygenes is 2.74%.
the SSR marker linked to scab in the RIL population of Wangshuibai/annong8455
78 SSR primers have polymorphism between Wangshuibai and Annong8455 when the 236 microsatellite primers are screened. The polymorphism is 33%.There are 14 SSR markers linked to scab in the two of three experimental years(2001,2003 and 2004).They are locused on chromosome 3B,2A,2B and 5D. There are 8 SSR markers linked to scab in the RIL population of Wangshuibai /Alondra. This is published. They are at least in the two of all phenotypes and are locused on chromosome 3B,4B,2D and 4D.
the tagging of QTL for Wangshuibai for wheat scab
The linked map of the RIL population of Wangshuibai/annong8455 is constructed. There are 25 markers in the linked map and are locused on 5 chromosomes. There is respectively one QTL on chromosome 3B in 2001 ,2003 and 2004.The QTLs on chromosome 3B are respectively between the primer Xgwm533.1 and Barc133, the
primer Xgwm389 and Xgwm533.1 and the primer Xgwm533.1 and Barc133.
The LOD values are respectively 2.47, 5.57and 2.52.They can explain 10.56% , 21.97% and 8.37%of the phenotypic variations of the scab. There is respectively one QTL on chromosome2A.The QTL on chromosome 2A is both between the primer Xgwm425 and Xgwm372.The LOD values are 2.5 and 2.80.They can explain 9.76% and 11.04% of the phenotypic variations of the scab. There is one QTL on chromosome 3B in the RIL population of Wangshuibai/Alondra. It has been published.The QTL is between Barc147 and Xgwm493 and can explain 13.7 % of the phenotypic variations.
The resistance to scab of Wangshuibai fits the model of two major genes and plus polygene. This helps to know the resistant to scab of Wangshuibai. The linked markers of scab-resistance is searched .There is one QTL on chromosome 3B of Wangshuibai. This is benefits to molecular breeding and is foundation to the clone of the genes to resistant to disease.
抗源望水白赤霉病抗性遗传分析
抗源望水白抗性由2对主基因控制，且符合两对主基因+多基因模型。2001年、03年、04年望水白/安农8455抗性分别符合E-2-6模型（两对主基因+多基因的加性－加性模型）、E-1-8模型（两对主基因+多基因,主基因间为重叠作用）、E-1-8模型（两对主基因+多基因，主基因间为重叠作用）。主基因的遗传率较高分别为63.8%、69.02％、73.66％，多基因的遗传率分别为13.94%、21.38%、16.80%。2003年望水白/Alondra抗性符合E-1-9模型（两对主基因+多基因，主基因间为相互抑制作用）。主基因的遗传率为85.18％，多基因的遗传率较低为2.74%。
望水白/安农8455群体的SSR标记
在望水白/安农8455群体的抗感亲本间的引物筛选表明，多态性为33%。根据群体的SSR标记，在2001年、03年和04年3年中至少有2年资料表明与抗赤性连锁的SSR标记有14个，定位在3B、2A、2B和5D染色体上。已经发表的资料表明，在望水白/Alondra群体中至少两次鉴定资料中都出现的与抗赤基因连锁的SSR标记共有8个，定位在3B、4B、2D、4D染色体上。
3. 抗源望水白的QTL定位
通过构建望水白/安农8455群体的连锁图，把25个SSR标记定位在5条染色体上。2001年、03年和04年的抗性资料表明在抗源的3B染色体上存在一个QTL。2001年和03年的抗性资料表明在2A上存在一个抗性QTL。根据2001年、03年和04年的抗性资料分析3B上的QTL分别位于Xgwm533.1-Barc133、 Xgwm389-Xgwm533.1、 Xgwm533.1-Barc133之间，LOD值为2.47、5.57、2.52，能解释10.56%、21.97%、8.37%的表型变异。2001年和03年的抗性资料分析2A上抗性QTL，都位于Xgwm425-Xgwm372之间，LOD值分别为2.5、2.80，能解释9.76%、11.04%。已发表的望水白/Alondra群体中抗源望水白的3B染色体上存在抗性QTL，位于Barc147-Xgwm493之间，能解释13.7%的表型变异。
对抗源望水白的抗性进行遗传分析，得出其抗赤性为两对主基因控制，有助于对抗源抗性机制的理解。运用SSR标记技术，获得与抗赤性连锁的遗传标记，在抗源望水白3B染色体上存在抗性QTL，为抗赤性分子标记辅助育种及最终克隆抗病基因奠定基础。
The RIL populations of Wangshuibai /annong8455 and Wangshuibai/Alondra to wheat scab were analysised with major gene plus ploygenes. The SSR（simple sequence repeat） marker analysis and QTL(quantitative trait loci) tagging of wheat scab resistance were carried out with the population of Wangshuibai/Annong8455.The results showed as follows:
the genetic analysis of Wangshuibai to scab
The scab resisitance of Wangshubai fits the genetic model of two major genes plus polygenes. The RIL populations of Wangshuibai /annong8455 to scab in 2001,2003 and 2004 fits the model of E-2-6(two major genes plus polygenes with additive –epistatic effect), the model of E-1-8(two major genes plus polygenes with effect) and the model of E-1-8. The heritability of major genes is 63.8%,69.02% and 73.66%.The heritability of polygenes is 13.94%,21.38% and 16.08%. The RIL populations of Wangshuibai /Alondra to scab in 2003 fits the model of E-1-9(two major genes plus polygenes with effect).The heritability of major genes is 85.18%.The heritability of polygenes is 2.74%.
the SSR marker linked to scab in the RIL population of Wangshuibai/annong8455
78 SSR primers have polymorphism between Wangshuibai and Annong8455 when the 236 microsatellite primers are screened. The polymorphism is 33%.There are 14 SSR markers linked to scab in the two of three experimental years(2001,2003 and 2004).They are locused on chromosome 3B,2A,2B and 5D. There are 8 SSR markers linked to scab in the RIL population of Wangshuibai /Alondra. This is published. They are at least in the two of all phenotypes and are locused on chromosome 3B,4B,2D and 4D.
the tagging of QTL for Wangshuibai for wheat scab
The linked map of the RIL population of Wangshuibai/annong8455 is constructed. There are 25 markers in the linked map and are locused on 5 chromosomes. There is respectively one QTL on chromosome 3B in 2001 ,2003 and 2004.The QTLs on chromosome 3B are respectively between the primer Xgwm533.1 and Barc133, the
primer Xgwm389 and Xgwm533.1 and the primer Xgwm533.1 and Barc133.
The LOD values are respectively 2.47, 5.57and 2.52.They can explain 10.56% , 21.97% and 8.37%of the phenotypic variations of the scab. There is respectively one QTL on chromosome2A.The QTL on chromosome 2A is both between the primer Xgwm425 and Xgwm372.The LOD values are 2.5 and 2.80.They can explain 9.76% and 11.04% of the phenotypic variations of the scab. There is one QTL on chromosome 3B in the RIL population of Wangshuibai/Alondra. It has been published.The QTL is between Barc147 and Xgwm493 and can explain 13.7 % of the phenotypic variations.
The resistance to scab of Wangshuibai fits the model of two major genes and plus polygene. This helps to know the resistant to scab of Wangshuibai. The linked markers of scab-resistance is searched .There is one QTL on chromosome 3B of Wangshuibai. This is benefits to molecular breeding and is foundation to the clone of the genes to resistant to disease.
引文
[1]Reitz.L.P. Wheat and wheat improvement.USA. 1967，P77.
[2]H..W.Schroederand .J.Christensen, pphytopathology, 1963, 53(7):831-838.
[3]张乐庆，潘雪萍.小麦品种对赤霉病的抗扩展性的遗传研究.华南农学院学报.1982,3(4):21-29.
[4]Snijders CHA..The inheritance of resistance to head blight caused by Fusarium culmorum in weinter wheat.Euphytica ,1990,50:11-18.
[5] Snijders CHA..Diallel analysis of resistance to head blight caused by Fusarium culmorum in weinter wheat.Euphytica,1990, 50:1-9.
[6]Bai GH,ZhouCF.QianCM,Ge YF..Genetic analysis of genes for resistance to scab spread in wheat cultivars.In Advances in Genetic Researches in Resistance to Diseases in Major Crops.ed. LHZhu.pp.,1990,171-77.Nanjing,Chiana: Jiang su Sci.Tech.Pub1.House.
[7]Singh RP, Ma H&Rajaram .Genetic analysis of resistance to scab in spring wheat cultivar.Frontana Plant Disease, 1995,79:238-240.
[8]Nicholson P,Mentewab A,Rezanoor HN&Gosman N.2000.Chromosamal location of Fusarium head blight resistance genes and analysis of the relationship between resistance to head blight and brown foot rot.In:John Raupp W,Ma ZQ Chen PD&Lin DJ eds .Proceedings of the international symposium on wheat improvement for scab resistance,140.
[9]王雅平，王先进.小麦品种对赤霉病抗扩展性的遗传研究.吉林农业科学，1991（1）：21－28.
[10]陈楚和.小麦抗赤霉病遗传的研究.浙江农业大学学报.1983,9(2):115-126.
[11] 余毓君.小麦品种苏麦3号抗赤霉病性及产量因素的单体分析.华中农学院学报.1982(6):70-72.
[12]姚金保.小麦品种苏麦3号抗赤霉病基因的染色体定位研究.作物学报，1997,23：450－453。
[13]夏穗生，周朝飞，钱存鸣，姚国才.苏麦3号、望水白赤霉病抗性遗传初步研
究.江苏农业科学.1984（8）：5-8.
[14]廖玉才，余毓君.小麦地方品种望水白抗赤霉病性的遗传分析.华中农学院学报.1985,4（2）：6-14.
[15]柏贵华，周朝飞，钱存鸣等.小麦品种抗扩展基因的遗传分析.朱立宏主编.主要农作物抗病性遗传研究进展.南京：江苏科学技术出版社，1990，171－175.
[16]林一波,杨竹平，吴兆苏.不同地理来源抗赤霉病小麦品种的抗性遗传分析.上海农业学报，1992,8（1）：31－36.
[17]Bostein,D.,White,R.L.,Skolnik,M.,Davis,R.W.,1980.Construction of a genetic linkage map in man using restriction fragment length polymorphisms.Am.H.Jim.Genet.32:314-331.
[18]Donis Keller,H.,Stephens,B.,et al.A genetic linkage map of the human genome.Cell.1987, 51:319-337.
[19]赵淑清，武维华.DNA分子标记和基因定位.生物技术通报，2000，16（6）:1-3.
[20]Diffenbach CW et al.PCR技术实验指南.北京：科学出版社，1998，111-113.
[21]Bennett,M.D.,Smith,J.B..Nuclear DNA amounts in angiosperms.Phil Trans RS London 1976,274:227-274.
[22] 郑敏,罗玉萍.真核生物基因组多态性分析的DNA指纹技术.生物技术,1999,9(3):35-38.
[23]Williams, G.K., kubelid, A.R.et al.DNA polymorphisms amplified by arbitry primers are useful as genetic markers. Nucleic Acids Research 1990, 18: 6531-6535.
[24]黎裕.分子标记的种类及其发展.生物技术通报，1999，15（4）：19－22.
[25]Paran,I.,Michelmore,R.W..Development of reliable PCR-based markers linked to downy mildew resistance genes in lettuce.Theor Appl.Genet,1993,85:985-993.
[26]Zabeau,M.,Vos,P.,Selective restriction fragment amplification: a general method for DNA fingerprinting. European Patent Application 92402629.7 ,1993.(Publication number: 05348-58A1).
[27]Olson,M.,Hood,L.,et al.A common language for physical mapping of the Human Genome.Science(Washington D.C.), 1989,254:1434-1435.
[28]Akkaya,M.S.,Bhagwat,A.A.,et al.Length polymorphisms of simple sequence repeat DNA in soybean.Genetics 1992,132:1131-1139.
[29]Morgante.M.,Olivieri,A.m.PCR-based amplified microsatellites as markers in plant
genetics.Plant J， 1993,3:175-182.
[30]徐云碧，朱立煌.《分子数量遗传学》.北京：中国农业出版社，1994，P85.
[31]Cheung W Y, Gugel R K, Landry B S.A RFLP-based linkage map of mustard Brassic juncea L Cross[J].Theor Appl Genet,1997,94:841-851.
[32]陈洪，朱立煌，徐吉臣等.RAPD标记构建水稻分子连锁图谱.植物学报，1995，677－684.
[33]Roder M S, Korzum V, Wendehake K, et al. A microsatellite map of wheat. Genetics ,1998,149:2007-2023.
[34]Van Degnze,A.E.,Gill,K.S.,et al.Molecular-genetic maps for group 1 chromosomes of Triticeae species and their relation to chromosome 1R.Theor.Appl.Genet, 1995,84:794-801.
[35]Nelson,J.C.,et al.Molecular mapping of wheat homoeologous group 2.Genome, 1995a 38:516-524.
[36]Devos,K.M.,Gale,M.D..The gentic maps of wheat and their potential in plant breeding.Outlook on Agricuture.1993，22:93-99.
[37]Nelson,J.C.et al.,.Molecular mapping of wheat homoeologous group3.Genome,
1995b ，38:525-533.
[38]Nelosn, J.C.et al.Molecular mapping of wheat major genes and rearrangments in homoeologous group4,5 and 7.Geonme ,1995b, 73:836-845.
[39]Xie,D.X.,Gale,M.D.,et al.,RFLP-based genetic maps of the homoeologous group 5 chromosomes of bread wheat(Triticum aestivum L.)Theor. Appl.Genet, 1993,87:70-74.
[40]Marino,C.L.,Nelson,J.C.,et al..Molecular genetic maps of the group 6 chromosomes of hexaploid wheat(Triticum aestivum L.Em.Thell). Genome, 1996, 39:359-366.
[41]Chao,S.,Sharp,P.J.,Gale,M.D..RFLP-based genetic maps of wheat homoeologous group 7 chromosomes.Theor.Appl.Genet, 1989,78:495-504.
[42]Kota,R.S.,Gill,K.S.,et al.A cytogenetically based physical map of chromosome of wheat.Theor.Appl.Genet,199391:568-573.
[43] Delaney,D.E.,Gill,B.S.,et al.Cytologically based physical map of the group 2 chromosome of wheat.Theor.Appl.Genet,1995b 91:780-782.
[44]Delaney,D.E.,Gill,B.S.,et al..Cytologically based physical map of the group 3
chromosome of wheat.Theor.Appl.Genet, 1995b ,91:780-782.
[45]Mickelosn,Y.L.,Endo,T.R.,and Gill,B.S. A cytogenetic ladder-map of the wheat homoeologous group-4 chromosome.Theor.Appl.Genet ,1995,90:1007-1011.
[46]Ogihara,Y.,et al.Fine cytological mapping of the long arm of chromosome 5A in common wheat by use of a series deletion line induced by gametocidal(Gc)genesofAegilops peltoides.Mol.Gen.Genet, 1994,244:253-259.
[47]Gill,K.S.,Boyko,E.V.,et al. Identification and highdensity mapping of
gene-rich regions in chromosome group 5 of wheat.Genetics , 1996.143 :1001-1002..
[48]Gill,K.S.,Gill,B.S.,et al.Agenetic linkage map of Triticum Tauschii
(DD)and its relationship to the D genome of bread wheat (AABBDD). Genome ,
1991,34:362-374.
[49]Hohmann,U.,Gill,B.S.,et al..Comparison of genetic and physical maps of group 7 chromosomes from Triticum aestivum.Mol.Gen.Genet ,1994,2450:644-653.
[50]Hohmann,U.,Gill,B.S..Characterization of deletions in common wheat induced by and Aegilops cylindrical Chromosome detection of multiple chromosome rearrangements. Theor. Appl.Genet, 1995a ,91:611-617.
[51]Hohmann,U.,Gill,B.S..Comparison of wheat physical maps with barley linkage maps for group 7 chromosoems.Theor.Appl.Genet, 1995b ,91:618-626.
[52]Huang N,Angeles E R,Domingo J,et al.Pyramiding of bacterial blight resistance genes in rice:sisted selection using PFLP and PCR[J].Theor Appl Genet ,1996,95:313-320.
[53]刘金元，刘大均，陈佩度等.分子标记辅助育种新尝试－与pm2及pm4a基因紧密连锁RFLP标记在小麦抗白粉病中的应用.南京农业大学学报，1997，20：1－5.
[54]Mora D,Parini C,Fortina MG,Manachini PL.Development of molecular RAPD marker for the identification of pediococcus acidilactici strains.Syst Appl Microbiol,2000:23(3):
400-408.
[55]Liang C Z, Gu M J, Pan X B.PFLP tagging of a new semidwarfing gene in rice[J]Genetics,1989,124:185-199.
[56]Michelmore R，Paranand I Kessali R V. Identification of markers linked to disease
resistance gene by bulked segregant analysis: a rapidmethod to detect markers in specifif genomic regions using segregating populations[J].Proc Natl Acad Sci USA, 1991,88: 9829-9832..
[57]Schachermayr G, Feuillet C, keller B. Molecular markers for the detection of the wheat leaf rust resistance gene Lr10 in diverse genetic backgrounds [J].Theor Appl Genet ,1990,80:833-840.
[58]ZhangQ-F, Shen B-Z, Dai X-K., Using bulked extremes and recessive class to map genes for photoperiod-sensitive genic male sterility in rice. PNAS USA,1994,8675-8679.
[59]Deal K K. Abstract of plant GenomeⅢ[M].San Diego.USA.1995:41.
[60]Tanhuanpaa P K. Association of a RAPD markers with Linoleic acid concentration in the seed oil [J].Genome ,1995,38(2):414-416.
[61]Stuber C W, Lincoln S E, Wolff D W, et al. Identification of genetic factors contributing to heterosis in a hybrid from two elite maize inbred lines using molecular markers[J].Genetics,1992,132:823-839.
[62]Lee M, Godshalk E B, Lamkey K R. Association of restriction fragment length polymorphism maize inbreds with agronomic performance of their crosses[J]Crop Science,1989,29:1067-1071.
[63]Brisses S.Dujardin HC Tibayrenc M., Identification of six Trypanosoma Cruzi lineages by sequence-characteerzed amplifified region markers.Mol Biochem Parasitol ,2000,111(1): 95- 105.
[64]种康，谭克辉，黄华梁等.冬小麦春化作用相关基因的DNA分子克隆研究.中国科学（B辑），1998.43:2419-3422.
[65]Ban, T. and Suenaga k.1997.Inheritance of resistance to Fusarium head blight caused by F. graminearum in wheat. Cereal Res.Comm.25:727-728.
[66]Otto, C.D, S.F. Kianian E.M., Elias E.M.,Stack R.W. and Joppa L.R. Molecular mapping for FHB resistance in a RICL population of tetraploid wheat, 1999,p.159. Agron.Abstr.ASA,Madison,WI.
[67] B.L.Waldron et al. RFLP Mapping of QTL for Fusarium Head Blight Resistance in Wheat. Crop Sci. 1999,39:805-811.
[68]Bai GH.Kolb FL.Shaner GE.Domier LL.Amplified fragment length polymorphism
markers linked to a major quantitative trait locus controlling scab resistance in wheat ,Phytopathology ,1999,89:343-48.
[69]Gilbert J.Fedak G.Procunier JD.Aung T.Tekauz A.Strategies for breeding for resistance to Fusarium head blight in Canadian spring wheat .See Ref,1997,44.pp47-51.
[70]Zhou WC,Kolb FL,Yu JB,BaiGH,Boze LK,Domier LL.Genetic analysis of scab resistance in wheat with microsatellite and markers,Genome, 2002,45:719-27.
[71] Zhou WC,Kolb FL,Yu JB,BaiGH,Boze LK,Domier LL.Molecular mapping of scab
resistance QTL in Wangshuibai.Natl.Fasarium Head Blight Forum Proc. Bloomington,MN,.Okemos,MI:Kinko’s. 2003,p.55.
[72]任丽娟,许雯，沈晓蓉等.中国春-苏麦3号7A代换系抗赤霉病基因的RAPD分析.农业生物技术学报，2000，8（3）：216
[73]姚金保.小麦品种苏麦3号抗赤霉病基因的染色体定位研究。作物学报，1997，23：450-453.
[74]赵涵，陈佩度，赵寅槐等.苏麦3号感赤霉病近等基因系的选育及分子标记，遗传学报，2002，29（10）：894－898.
[75]任丽娟，沈晓蓉，张旭等.三个小麦重组自交系群体抗赤霉病QTLs的SSR分析，中国农业科学，2003，36（10）：1150－1155.
[76]沈晓蓉,陆维忠.小麦体细胞无性系895004与供体亲本的抗赤性、农艺性比较和RAPD分析.江苏农业学报，1996，12（1）：7-10.
[77]Z.Q.Ma,F.Lin ,ZX kong et al.Mapping QTLs associated with FJB resistance in a Wangshuibai ×Nanda2419 population. Tenth interational wheat genetics symposium. Italy.2003,320-323.
[78]向阳海,陆维忠，朱作为等.小麦赤霉病抗性的RAPD标记筛选与分析，农业生物技术学报，2001,9（3）：247-250.
[79] Xu Zhang et al. Molecular characterization of FHB Resistance from wheat wangshuibai,2004,(待发表).
[80]周淼平，张旭，任丽娟等.用JoinMap3.0初步构建了小麦遗传连锁图，江苏农业学报，2003，19（3）：133-138..
[81]Shen X. et al., inberitance and QTL detection of FHB resistance in wheat population Ning894037/Alondra,2002,TAG.10.1007/s00122-113308.
[82] 马鸿翔，张旭，任丽娟等.?小麦赤霉病抗性QTL分子标记及辅助选择研究进
展，植物细胞工程与分子育种技术研究.中国农业科学技术出版社, 2003,186-193.
[83]邓怀，余懋群.?小麦分子标记及其在遗传育种研究中的应用.科技前沿与学术评论北京：科学出版社,2002，24（2）37-43.
[84] 盖均镒，王健康.利用回交或F2:3家系世代鉴定主基因－多基因的混合遗传模型[J].作物学报，1998,24(4):402-409.
[85] 王健康,盖均镒.利用杂种F2世代鉴定数量性状主基因－多基因混合遗传模型并估计其遗传效应[J].遗传学报,1997,24(5):432-440.
[86]Gai J Y,Wang J K.Identification and estimation of a QTL model and its effects[J].Theor.Appl.Genet.1998,97:1162-1168.
[87]章元明，盖均镒.利用DH或RIL群体检测QTL体系并估计其遗传效应[J].遗传学报，2000，27（7）：634－640.
[88] 章元明，盖均镒，王永军.利用P1、P2和DH或RIL群体联合分离分析的拓展[J].遗传，2001，23（5）：567－470.
[89]Saghai-Mmaroof M A, Soliman K, Jorgensen R A, Allard R W. Ribosomal DNA spacer-length polymorphisms in barley: mendelian inheritance, chromosomal location and population dynamics.PNAS,1984,81:8104-8018.
[90] 陆维忠，程顺和，王裕中.小麦赤霉病研究.北京：科学出版社,P29.
[91]陆维忠，张旭，黄昌等，两个小麦遗传群体不同年份和地点赤霉病抗性鉴定与比较，江苏农业学报，2002，18（1）：23-28.
[92]Elston R C and J Steart.The annlysis of quantitative traits for simple genetic models from parental,F1 and backcross data.Genetics,197373:695-711.
[93]Morton N E and C J Maclean. Analysis of family resemblance.Ⅲ.Complex segregation of quantitative traits.Am j Hum Genet, 1974.26:489-503.
[94]Elkind Y and A Cahaner. A mixed model for the effects of single gene,polygenes and their interaction on quantitative traits.1.The model and experimental design.Theor Appl Genet, 1986,72:377-383.
[95] Elkind Y and A Cahaner. A mixed model for the effects of single gene,polygenes and their interaction on quantitative traits.2. The effects of the major gene and polygenes on tomato fruit softness.Heredity, 1990, 64:205-213.
[96]莫惠栋.质量-数量性状的遗传分析 Ⅱ.世代平均数和遗传方差.作物学报，1993b，19（3）：193-200.
[97]姜长鉴，莫惠栋.1995.质量-数量性状的遗传分析 Ⅳ.极大似然法的应用.作物学报，21(6):641-648.
[98]张勇.小麦赤霉病新抗源S42、 H35、 N553的遗传分析及望水白抗赤霉病QTLs
的SSR标记筛选:[硕士学位论文].扬州：扬州大学，2002.
[99]Liu ZY,Sun QX,Li HJ,Ni ZF,Yang ZM,Tang BR,Yang AD,Jia X.,Molecular identigication and marker-assisted selection of Pm21 gene conferring resistance to powdery mildew in wheat.,YiChuan Xue Bao.,1999,26(6):673-682.
[100]Xu Y,Zhang JY,Kang GB,Wang YJ,Chen H.,Studies of molecular marker-assisted
-selection for resistance to Fusarium wilt in watermelon (Citrullus lanatus) breeding.,Yi Chuan Xue Bao 2000,27(2):151-157.
[101]吴建宇，汤继华，夏宗良等,玉米矮花叶病毒一个新的抗性基因位点的微卫星标记.植物学报，2002,（2）：177-180.
[102] 盖均镒，章元明，王健康.植物数量性状遗传体系.北京：科学出版社，2003,340-3412003,340-341.
[2]H..W.Schroederand .J.Christensen, pphytopathology, 1963, 53(7):831-838.
[3]张乐庆，潘雪萍.小麦品种对赤霉病的抗扩展性的遗传研究.华南农学院学报.1982,3(4):21-29.
[4]Snijders CHA..The inheritance of resistance to head blight caused by Fusarium culmorum in weinter wheat.Euphytica ,1990,50:11-18.
[5] Snijders CHA..Diallel analysis of resistance to head blight caused by Fusarium culmorum in weinter wheat.Euphytica,1990, 50:1-9.
[6]Bai GH,ZhouCF.QianCM,Ge YF..Genetic analysis of genes for resistance to scab spread in wheat cultivars.In Advances in Genetic Researches in Resistance to Diseases in Major Crops.ed. LHZhu.pp.,1990,171-77.Nanjing,Chiana: Jiang su Sci.Tech.Pub1.House.
[7]Singh RP, Ma H&Rajaram .Genetic analysis of resistance to scab in spring wheat cultivar.Frontana Plant Disease, 1995,79:238-240.
[8]Nicholson P,Mentewab A,Rezanoor HN&Gosman N.2000.Chromosamal location of Fusarium head blight resistance genes and analysis of the relationship between resistance to head blight and brown foot rot.In:John Raupp W,Ma ZQ Chen PD&Lin DJ eds .Proceedings of the international symposium on wheat improvement for scab resistance,140.
[9]王雅平，王先进.小麦品种对赤霉病抗扩展性的遗传研究.吉林农业科学，1991（1）：21－28.
[10]陈楚和.小麦抗赤霉病遗传的研究.浙江农业大学学报.1983,9(2):115-126.
[11] 余毓君.小麦品种苏麦3号抗赤霉病性及产量因素的单体分析.华中农学院学报.1982(6):70-72.
[12]姚金保.小麦品种苏麦3号抗赤霉病基因的染色体定位研究.作物学报，1997,23：450－453。
[13]夏穗生，周朝飞，钱存鸣，姚国才.苏麦3号、望水白赤霉病抗性遗传初步研
究.江苏农业科学.1984（8）：5-8.
[14]廖玉才，余毓君.小麦地方品种望水白抗赤霉病性的遗传分析.华中农学院学报.1985,4（2）：6-14.
[15]柏贵华，周朝飞，钱存鸣等.小麦品种抗扩展基因的遗传分析.朱立宏主编.主要农作物抗病性遗传研究进展.南京：江苏科学技术出版社，1990，171－175.
[16]林一波,杨竹平，吴兆苏.不同地理来源抗赤霉病小麦品种的抗性遗传分析.上海农业学报，1992,8（1）：31－36.
[17]Bostein,D.,White,R.L.,Skolnik,M.,Davis,R.W.,1980.Construction of a genetic linkage map in man using restriction fragment length polymorphisms.Am.H.Jim.Genet.32:314-331.
[18]Donis Keller,H.,Stephens,B.,et al.A genetic linkage map of the human genome.Cell.1987, 51:319-337.
[19]赵淑清，武维华.DNA分子标记和基因定位.生物技术通报，2000，16（6）:1-3.
[20]Diffenbach CW et al.PCR技术实验指南.北京：科学出版社，1998，111-113.
[21]Bennett,M.D.,Smith,J.B..Nuclear DNA amounts in angiosperms.Phil Trans RS London 1976,274:227-274.
[22] 郑敏,罗玉萍.真核生物基因组多态性分析的DNA指纹技术.生物技术,1999,9(3):35-38.
[23]Williams, G.K., kubelid, A.R.et al.DNA polymorphisms amplified by arbitry primers are useful as genetic markers. Nucleic Acids Research 1990, 18: 6531-6535.
[24]黎裕.分子标记的种类及其发展.生物技术通报，1999，15（4）：19－22.
[25]Paran,I.,Michelmore,R.W..Development of reliable PCR-based markers linked to downy mildew resistance genes in lettuce.Theor Appl.Genet,1993,85:985-993.
[26]Zabeau,M.,Vos,P.,Selective restriction fragment amplification: a general method for DNA fingerprinting. European Patent Application 92402629.7 ,1993.(Publication number: 05348-58A1).
[27]Olson,M.,Hood,L.,et al.A common language for physical mapping of the Human Genome.Science(Washington D.C.), 1989,254:1434-1435.
[28]Akkaya,M.S.,Bhagwat,A.A.,et al.Length polymorphisms of simple sequence repeat DNA in soybean.Genetics 1992,132:1131-1139.
[29]Morgante.M.,Olivieri,A.m.PCR-based amplified microsatellites as markers in plant
genetics.Plant J， 1993,3:175-182.
[30]徐云碧，朱立煌.《分子数量遗传学》.北京：中国农业出版社，1994，P85.
[31]Cheung W Y, Gugel R K, Landry B S.A RFLP-based linkage map of mustard Brassic juncea L Cross[J].Theor Appl Genet,1997,94:841-851.
[32]陈洪，朱立煌，徐吉臣等.RAPD标记构建水稻分子连锁图谱.植物学报，1995，677－684.
[33]Roder M S, Korzum V, Wendehake K, et al. A microsatellite map of wheat. Genetics ,1998,149:2007-2023.
[34]Van Degnze,A.E.,Gill,K.S.,et al.Molecular-genetic maps for group 1 chromosomes of Triticeae species and their relation to chromosome 1R.Theor.Appl.Genet, 1995,84:794-801.
[35]Nelson,J.C.,et al.Molecular mapping of wheat homoeologous group 2.Genome, 1995a 38:516-524.
[36]Devos,K.M.,Gale,M.D..The gentic maps of wheat and their potential in plant breeding.Outlook on Agricuture.1993，22:93-99.
[37]Nelson,J.C.et al.,.Molecular mapping of wheat homoeologous group3.Genome,
1995b ，38:525-533.
[38]Nelosn, J.C.et al.Molecular mapping of wheat major genes and rearrangments in homoeologous group4,5 and 7.Geonme ,1995b, 73:836-845.
[39]Xie,D.X.,Gale,M.D.,et al.,RFLP-based genetic maps of the homoeologous group 5 chromosomes of bread wheat(Triticum aestivum L.)Theor. Appl.Genet, 1993,87:70-74.
[40]Marino,C.L.,Nelson,J.C.,et al..Molecular genetic maps of the group 6 chromosomes of hexaploid wheat(Triticum aestivum L.Em.Thell). Genome, 1996, 39:359-366.
[41]Chao,S.,Sharp,P.J.,Gale,M.D..RFLP-based genetic maps of wheat homoeologous group 7 chromosomes.Theor.Appl.Genet, 1989,78:495-504.
[42]Kota,R.S.,Gill,K.S.,et al.A cytogenetically based physical map of chromosome of wheat.Theor.Appl.Genet,199391:568-573.
[43] Delaney,D.E.,Gill,B.S.,et al.Cytologically based physical map of the group 2 chromosome of wheat.Theor.Appl.Genet,1995b 91:780-782.
[44]Delaney,D.E.,Gill,B.S.,et al..Cytologically based physical map of the group 3
chromosome of wheat.Theor.Appl.Genet, 1995b ,91:780-782.
[45]Mickelosn,Y.L.,Endo,T.R.,and Gill,B.S. A cytogenetic ladder-map of the wheat homoeologous group-4 chromosome.Theor.Appl.Genet ,1995,90:1007-1011.
[46]Ogihara,Y.,et al.Fine cytological mapping of the long arm of chromosome 5A in common wheat by use of a series deletion line induced by gametocidal(Gc)genesofAegilops peltoides.Mol.Gen.Genet, 1994,244:253-259.
[47]Gill,K.S.,Boyko,E.V.,et al. Identification and highdensity mapping of
gene-rich regions in chromosome group 5 of wheat.Genetics , 1996.143 :1001-1002..
[48]Gill,K.S.,Gill,B.S.,et al.Agenetic linkage map of Triticum Tauschii
(DD)and its relationship to the D genome of bread wheat (AABBDD). Genome ,
1991,34:362-374.
[49]Hohmann,U.,Gill,B.S.,et al..Comparison of genetic and physical maps of group 7 chromosomes from Triticum aestivum.Mol.Gen.Genet ,1994,2450:644-653.
[50]Hohmann,U.,Gill,B.S..Characterization of deletions in common wheat induced by and Aegilops cylindrical Chromosome detection of multiple chromosome rearrangements. Theor. Appl.Genet, 1995a ,91:611-617.
[51]Hohmann,U.,Gill,B.S..Comparison of wheat physical maps with barley linkage maps for group 7 chromosoems.Theor.Appl.Genet, 1995b ,91:618-626.
[52]Huang N,Angeles E R,Domingo J,et al.Pyramiding of bacterial blight resistance genes in rice:sisted selection using PFLP and PCR[J].Theor Appl Genet ,1996,95:313-320.
[53]刘金元，刘大均，陈佩度等.分子标记辅助育种新尝试－与pm2及pm4a基因紧密连锁RFLP标记在小麦抗白粉病中的应用.南京农业大学学报，1997，20：1－5.
[54]Mora D,Parini C,Fortina MG,Manachini PL.Development of molecular RAPD marker for the identification of pediococcus acidilactici strains.Syst Appl Microbiol,2000:23(3):
400-408.
[55]Liang C Z, Gu M J, Pan X B.PFLP tagging of a new semidwarfing gene in rice[J]Genetics,1989,124:185-199.
[56]Michelmore R，Paranand I Kessali R V. Identification of markers linked to disease
resistance gene by bulked segregant analysis: a rapidmethod to detect markers in specifif genomic regions using segregating populations[J].Proc Natl Acad Sci USA, 1991,88: 9829-9832..
[57]Schachermayr G, Feuillet C, keller B. Molecular markers for the detection of the wheat leaf rust resistance gene Lr10 in diverse genetic backgrounds [J].Theor Appl Genet ,1990,80:833-840.
[58]ZhangQ-F, Shen B-Z, Dai X-K., Using bulked extremes and recessive class to map genes for photoperiod-sensitive genic male sterility in rice. PNAS USA,1994,8675-8679.
[59]Deal K K. Abstract of plant GenomeⅢ[M].San Diego.USA.1995:41.
[60]Tanhuanpaa P K. Association of a RAPD markers with Linoleic acid concentration in the seed oil [J].Genome ,1995,38(2):414-416.
[61]Stuber C W, Lincoln S E, Wolff D W, et al. Identification of genetic factors contributing to heterosis in a hybrid from two elite maize inbred lines using molecular markers[J].Genetics,1992,132:823-839.
[62]Lee M, Godshalk E B, Lamkey K R. Association of restriction fragment length polymorphism maize inbreds with agronomic performance of their crosses[J]Crop Science,1989,29:1067-1071.
[63]Brisses S.Dujardin HC Tibayrenc M., Identification of six Trypanosoma Cruzi lineages by sequence-characteerzed amplifified region markers.Mol Biochem Parasitol ,2000,111(1): 95- 105.
[64]种康，谭克辉，黄华梁等.冬小麦春化作用相关基因的DNA分子克隆研究.中国科学（B辑），1998.43:2419-3422.
[65]Ban, T. and Suenaga k.1997.Inheritance of resistance to Fusarium head blight caused by F. graminearum in wheat. Cereal Res.Comm.25:727-728.
[66]Otto, C.D, S.F. Kianian E.M., Elias E.M.,Stack R.W. and Joppa L.R. Molecular mapping for FHB resistance in a RICL population of tetraploid wheat, 1999,p.159. Agron.Abstr.ASA,Madison,WI.
[67] B.L.Waldron et al. RFLP Mapping of QTL for Fusarium Head Blight Resistance in Wheat. Crop Sci. 1999,39:805-811.
[68]Bai GH.Kolb FL.Shaner GE.Domier LL.Amplified fragment length polymorphism
markers linked to a major quantitative trait locus controlling scab resistance in wheat ,Phytopathology ,1999,89:343-48.
[69]Gilbert J.Fedak G.Procunier JD.Aung T.Tekauz A.Strategies for breeding for resistance to Fusarium head blight in Canadian spring wheat .See Ref,1997,44.pp47-51.
[70]Zhou WC,Kolb FL,Yu JB,BaiGH,Boze LK,Domier LL.Genetic analysis of scab resistance in wheat with microsatellite and markers,Genome, 2002,45:719-27.
[71] Zhou WC,Kolb FL,Yu JB,BaiGH,Boze LK,Domier LL.Molecular mapping of scab
resistance QTL in Wangshuibai.Natl.Fasarium Head Blight Forum Proc. Bloomington,MN,.Okemos,MI:Kinko’s. 2003,p.55.
[72]任丽娟,许雯，沈晓蓉等.中国春-苏麦3号7A代换系抗赤霉病基因的RAPD分析.农业生物技术学报，2000，8（3）：216
[73]姚金保.小麦品种苏麦3号抗赤霉病基因的染色体定位研究。作物学报，1997，23：450-453.
[74]赵涵，陈佩度，赵寅槐等.苏麦3号感赤霉病近等基因系的选育及分子标记，遗传学报，2002，29（10）：894－898.
[75]任丽娟，沈晓蓉，张旭等.三个小麦重组自交系群体抗赤霉病QTLs的SSR分析，中国农业科学，2003，36（10）：1150－1155.
[76]沈晓蓉,陆维忠.小麦体细胞无性系895004与供体亲本的抗赤性、农艺性比较和RAPD分析.江苏农业学报，1996，12（1）：7-10.
[77]Z.Q.Ma,F.Lin ,ZX kong et al.Mapping QTLs associated with FJB resistance in a Wangshuibai ×Nanda2419 population. Tenth interational wheat genetics symposium. Italy.2003,320-323.
[78]向阳海,陆维忠，朱作为等.小麦赤霉病抗性的RAPD标记筛选与分析，农业生物技术学报，2001,9（3）：247-250.
[79] Xu Zhang et al. Molecular characterization of FHB Resistance from wheat wangshuibai,2004,(待发表).
[80]周淼平，张旭，任丽娟等.用JoinMap3.0初步构建了小麦遗传连锁图，江苏农业学报，2003，19（3）：133-138..
[81]Shen X. et al., inberitance and QTL detection of FHB resistance in wheat population Ning894037/Alondra,2002,TAG.10.1007/s00122-113308.
[82] 马鸿翔，张旭，任丽娟等.?小麦赤霉病抗性QTL分子标记及辅助选择研究进
展，植物细胞工程与分子育种技术研究.中国农业科学技术出版社, 2003,186-193.
[83]邓怀，余懋群.?小麦分子标记及其在遗传育种研究中的应用.科技前沿与学术评论北京：科学出版社,2002，24（2）37-43.
[84] 盖均镒，王健康.利用回交或F2:3家系世代鉴定主基因－多基因的混合遗传模型[J].作物学报，1998,24(4):402-409.
[85] 王健康,盖均镒.利用杂种F2世代鉴定数量性状主基因－多基因混合遗传模型并估计其遗传效应[J].遗传学报,1997,24(5):432-440.
[86]Gai J Y,Wang J K.Identification and estimation of a QTL model and its effects[J].Theor.Appl.Genet.1998,97:1162-1168.
[87]章元明，盖均镒.利用DH或RIL群体检测QTL体系并估计其遗传效应[J].遗传学报，2000，27（7）：634－640.
[88] 章元明，盖均镒，王永军.利用P1、P2和DH或RIL群体联合分离分析的拓展[J].遗传，2001，23（5）：567－470.
[89]Saghai-Mmaroof M A, Soliman K, Jorgensen R A, Allard R W. Ribosomal DNA spacer-length polymorphisms in barley: mendelian inheritance, chromosomal location and population dynamics.PNAS,1984,81:8104-8018.
[90] 陆维忠，程顺和，王裕中.小麦赤霉病研究.北京：科学出版社,P29.
[91]陆维忠，张旭，黄昌等，两个小麦遗传群体不同年份和地点赤霉病抗性鉴定与比较，江苏农业学报，2002，18（1）：23-28.
[92]Elston R C and J Steart.The annlysis of quantitative traits for simple genetic models from parental,F1 and backcross data.Genetics,197373:695-711.
[93]Morton N E and C J Maclean. Analysis of family resemblance.Ⅲ.Complex segregation of quantitative traits.Am j Hum Genet, 1974.26:489-503.
[94]Elkind Y and A Cahaner. A mixed model for the effects of single gene,polygenes and their interaction on quantitative traits.1.The model and experimental design.Theor Appl Genet, 1986,72:377-383.
[95] Elkind Y and A Cahaner. A mixed model for the effects of single gene,polygenes and their interaction on quantitative traits.2. The effects of the major gene and polygenes on tomato fruit softness.Heredity, 1990, 64:205-213.
[96]莫惠栋.质量-数量性状的遗传分析 Ⅱ.世代平均数和遗传方差.作物学报，1993b，19（3）：193-200.
[97]姜长鉴，莫惠栋.1995.质量-数量性状的遗传分析 Ⅳ.极大似然法的应用.作物学报，21(6):641-648.
[98]张勇.小麦赤霉病新抗源S42、 H35、 N553的遗传分析及望水白抗赤霉病QTLs
的SSR标记筛选:[硕士学位论文].扬州：扬州大学，2002.
[99]Liu ZY,Sun QX,Li HJ,Ni ZF,Yang ZM,Tang BR,Yang AD,Jia X.,Molecular identigication and marker-assisted selection of Pm21 gene conferring resistance to powdery mildew in wheat.,YiChuan Xue Bao.,1999,26(6):673-682.
[100]Xu Y,Zhang JY,Kang GB,Wang YJ,Chen H.,Studies of molecular marker-assisted
-selection for resistance to Fusarium wilt in watermelon (Citrullus lanatus) breeding.,Yi Chuan Xue Bao 2000,27(2):151-157.
[101]吴建宇，汤继华，夏宗良等,玉米矮花叶病毒一个新的抗性基因位点的微卫星标记.植物学报，2002,（2）：177-180.
[102] 盖均镒，章元明，王健康.植物数量性状遗传体系.北京：科学出版社，2003,340-3412003,340-341.