小麦抗叶锈病近等基因系TcLr35的RGA分析
|
摘要
由小麦叶锈菌(Puccinia triticina)引起的小麦叶锈病是影响世界小麦稳产高产的重要病害之一,也是影响我国小麦生产的重要因素。持续筛选抗病基因和培育抗病品种是防治小麦叶锈病发生的主要手段。小麦抗叶锈病基因Lr35最初来源于拟斯卑尔脱山羊草(Aegilops speltoides),通过与Triticina.monococcum的二倍体回交将其转移到小麦品种马奎斯(Marquis)中。该基因位于2B染色体上,与Sr39紧密连锁。其抗性在二叶期开始表达,六叶期完全表达,是一个应用潜力很大的由主效基因控制的抗病基因。
抗病基因类似序列(Reistance gene analogys,RGAs)技术是利用抗病基因具有保守结构域的特性,通过人工合成引物对基因组DNA进行定点扩增,在全基因组水平上检测DNA变异。本试验以含小麦抗叶锈病基因Lr35的近等基因系材料TcLr35和感病亲本Thatcher以及其它小麦抗叶锈病近等基因系材料进行RGA分析。主要结果如下:
根据已克隆的植物抗病基因普遍具有的NBS和LRR结构域设计49条RGA引物,组成260对引物组合,从近等基因系TcLr35中扩增获得4条特异性片段。其中引物对Pto-kin1IN/XLRR-INV1从近等基因系TcLr35中扩增获得一条747bp的稳定特异性片段。以本实验室保存的49个小麦抗叶锈病近等基因系材料进行扩增,发现此条带仅在TcLr35中存在,而在其他48个小麦抗叶锈病近等基因系材料及感病亲本Thatcher中均未出现,说明该片段特异性很强。对该特异性片段进行序列测定与分析,经BLAST比较,表明该片段与许多已克隆出来的小麦基因序列相似:与Triticummonococcum DV92的BAC克隆231A16有84%的同源性,与Triticum urartu clone BAC210J24有96%同源性;与Triticum turgidum subsp.durum clone BAC 221H19有90%同源性;与Triticum urartu clone BAC 41C8有89%同源性;与Triticum urartu clone BAC292N12有88%同源性等。对该特异性片段进行蛋白质序列比对,未发现与其于已知功能小麦片段有较高同源性。
Wheat leaf rust caused by Puccinia triticina is one of the most important diseases affected the wheat production world widely. Utilization of genetic resistance is the most economical and environment friendly strategy for disease control. Wheat leaf rust resistance gene Lr35 mapped on chromosome 2B was derived from Aegilops speltoides and was transferred to the wheat cultivar Marquis through baekcrossing with diploid Triticina monococcum. The gene closely links with the wheat stem rust resistance gene Sr39. Its resistance to the disease first appears at the growth stage with two leaves, and completely expresses with six leaves. It is a very effective leaf rust resistance gene which has great potential to be used on wheat production
Reistance gene analogys (RGAs) is a technique which can detect the DNA mutation from the genome scale by utilizing the character of the resistance gene conserved region and amplify the genome DNA specifically through man-made synthesing primers. In this experiment, near-isogenic line (NIL) TcLr35 carrying Lr35, its susceptible parent Thatcher and the other NILs were served as material to perform RGA analysis. The main results were as follows:
Using the 260 primer pairs from 49 individual RGA primers which based on the conserved domains (NBS、LRR etc) of the known resistance genes, 4 specific fragments were found in TcLr35. A steady and specific747bp fragment was found in TcLr35 by primer pair Pto-kin1IN/XLRR-INV1. The primer pair then was used to amplify resistant homologous sequences in the 49 near isogenic lines (NILs) of wheat leaf rust resistance. The specific fragment was only found in TcLr35, but never in the other 48 near isogenic lines and Thatcher. The result indicates that the specific fragment is high differential. The specific fragment was isolated from the polyacrylamide gels, re-amplified, cloned and sequenced. It has the high homology with other wheat gene sequences cloned: 84% identical with Triticum monococcum DV92 clone BAC 231 A16, 96% identical with Triticum urartu clone BAC 210J24, 90% identical with Triticum turgidum subsp. durum clone BAC 221H19, 90% identical with Triticum urartu clone BAC 41C8, 88% identical with Triticum urartu clone BAC 292N12 etc. The BLASTx analysis indicates that there are no wheat DNA fragments showing high homologous with the specific fragment.
抗病基因类似序列(Reistance gene analogys,RGAs)技术是利用抗病基因具有保守结构域的特性,通过人工合成引物对基因组DNA进行定点扩增,在全基因组水平上检测DNA变异。本试验以含小麦抗叶锈病基因Lr35的近等基因系材料TcLr35和感病亲本Thatcher以及其它小麦抗叶锈病近等基因系材料进行RGA分析。主要结果如下:
根据已克隆的植物抗病基因普遍具有的NBS和LRR结构域设计49条RGA引物,组成260对引物组合,从近等基因系TcLr35中扩增获得4条特异性片段。其中引物对Pto-kin1IN/XLRR-INV1从近等基因系TcLr35中扩增获得一条747bp的稳定特异性片段。以本实验室保存的49个小麦抗叶锈病近等基因系材料进行扩增,发现此条带仅在TcLr35中存在,而在其他48个小麦抗叶锈病近等基因系材料及感病亲本Thatcher中均未出现,说明该片段特异性很强。对该特异性片段进行序列测定与分析,经BLAST比较,表明该片段与许多已克隆出来的小麦基因序列相似:与Triticummonococcum DV92的BAC克隆231A16有84%的同源性,与Triticum urartu clone BAC210J24有96%同源性;与Triticum turgidum subsp.durum clone BAC 221H19有90%同源性;与Triticum urartu clone BAC 41C8有89%同源性;与Triticum urartu clone BAC292N12有88%同源性等。对该特异性片段进行蛋白质序列比对,未发现与其于已知功能小麦片段有较高同源性。
Wheat leaf rust caused by Puccinia triticina is one of the most important diseases affected the wheat production world widely. Utilization of genetic resistance is the most economical and environment friendly strategy for disease control. Wheat leaf rust resistance gene Lr35 mapped on chromosome 2B was derived from Aegilops speltoides and was transferred to the wheat cultivar Marquis through baekcrossing with diploid Triticina monococcum. The gene closely links with the wheat stem rust resistance gene Sr39. Its resistance to the disease first appears at the growth stage with two leaves, and completely expresses with six leaves. It is a very effective leaf rust resistance gene which has great potential to be used on wheat production
Reistance gene analogys (RGAs) is a technique which can detect the DNA mutation from the genome scale by utilizing the character of the resistance gene conserved region and amplify the genome DNA specifically through man-made synthesing primers. In this experiment, near-isogenic line (NIL) TcLr35 carrying Lr35, its susceptible parent Thatcher and the other NILs were served as material to perform RGA analysis. The main results were as follows:
Using the 260 primer pairs from 49 individual RGA primers which based on the conserved domains (NBS、LRR etc) of the known resistance genes, 4 specific fragments were found in TcLr35. A steady and specific747bp fragment was found in TcLr35 by primer pair Pto-kin1IN/XLRR-INV1. The primer pair then was used to amplify resistant homologous sequences in the 49 near isogenic lines (NILs) of wheat leaf rust resistance. The specific fragment was only found in TcLr35, but never in the other 48 near isogenic lines and Thatcher. The result indicates that the specific fragment is high differential. The specific fragment was isolated from the polyacrylamide gels, re-amplified, cloned and sequenced. It has the high homology with other wheat gene sequences cloned: 84% identical with Triticum monococcum DV92 clone BAC 231 A16, 96% identical with Triticum urartu clone BAC 210J24, 90% identical with Triticum turgidum subsp. durum clone BAC 221H19, 90% identical with Triticum urartu clone BAC 41C8, 88% identical with Triticum urartu clone BAC 292N12 etc. The BLASTx analysis indicates that there are no wheat DNA fragments showing high homologous with the specific fragment.
引文
[1]Komer J A.Genetics of resistance to wheat leaf rust[J].Annual Review Phytopathology,1996,34:435-455.
[2]Kloppers F J,Pretorius Z A.Effects of combinations amongst genes Lr13,Lr34 and Lr37 on components of resistance in wheat to leaf rust[J].Plant Pathology,1997,46:737-750.
[3]Knott D R.The wheat rust breeding for resistance[J].TheorAppl Genet,1989,191-198.
[4]Yu J,Hu S N,Wang J,et al.A draft sequence of the rice genome(Oryza sativa L.ssp.indic a)[J].Science,2002,296:79-91.
[5]Deslandes L,Olivier J,Theulieres F.Resistance to Ralstonia solanacearum in Arabidopsis thaliana is conferred by the recessive RRSl-R gene,a member of a novel family of resistance genes[J].Proc Natl Acad SciUSA,2002,99:2404-2409.
[6]Bryan G T,Wu K S,Farrall L.A single amino acid difference distinguishes resistant and susceptible alleles of the rice blast resistance genePi-to[J].Plant Cell,2000,12:2033-2046.
[7]Buschges R,Hollrichner K,Panstrnga R.The barely Mlo gene:A novel control element of plant pathogen resistance[J].Cell,1997,88:695-705.
[8]Brandwagt B F,Mesbah L A,Takken F L,et al.A longevity assurance gene homolog of tomato mediates resistance to Altemaria alternata f.sp.lycopersiei toxins and fumonisin B1[J].Proc Natl Acad Sci USA,2000,97:4961-4966.
[9]Kawchuk L M,Hachey J,Lynch D R,et al.Tomato Ve disease resistance genes encode cell surface-like receptors[J].Proc Natl Acad Sci USA,2001,98:6511-6515.
[10]Xiao S,Ellwood S,Calis O,et al.Broad-spectrum mildew resistance in Arabidopsis thaliana mediated byRPW8[J].Science,2001,291:118-120.
[11]Boyes D C,Nam J,Dangl J,et al.The Arabidopsis thaliana RPMl disease resistance gene product is a peripheral plasma membrane protein that is degrade coincident with the hypersensitive response[J].Proc Natl Acad Sci USA,1998,95:15849-15854.
[12]Bent A F.Plant disease resistance genes:function meets structure[J].Plant Cell,1996,8:1757-1771.
[13]Jones D A,Thomas C M,Hammond-Kosack K E,et al.Isolation of the tomato Cf-9 gene for resistance to Cladosporium fulvum by transposon tagging[J].Science,1994,266:789-793.
[14]Martin G B,Brommonschenkel S H,Chunwongse J,et al.Map-based cloning of a protein kinase gene conferring disease resistance in tomato[J].Science,1993,262:1432-1436.
[15]Whitham S,Dinesh-kumar S P,Chol D,et al.The product of the tobacco mosaic virus resistance gene N:similarity to Toll and the lnterleukin-1 receptor[J].Cell,1994,78:1101-1115.
[16]罗敏,末友林,余潮等.植物抗病基因的克隆及其结构与功能[J].遗传,2000,6:429-34.
[17]Song W Y,Wang G L,Chen L L,et al.A receptor kinase-like protein encoded by the rice disease resistance gene,Xa21[J].Science,1995,270:1804-1806.
[18] Meyers B C, Dickerman A W, Michelmore R W, et al. Plant disease resistance gene encode members of an ancient and diverse protein family within the nucleotide binding superfamily [J]. The Plant J, 1999, 20(3): 317-332.
[19] Brightbill H D, Libraty D H, Krutzik S R, et al. Host defense mechanisms triggered by microbial lipoprotein through Toll-like receptors [J]. Science, 1999, 285: 732-739.
[20] Lawrence G J, Finnegan E J, Ayliffe M A, et al. The L6 Gene for Flax Rust Resistance 1s Related to the Arabidopsis Bacterial Resistance Gene RPSP and the Tobacco Viral Resistance Gene N [J]. The Plant Cell, 1995, 7: 1195-1206.
[21] Aarts G M, Lintel Hekkert B, Holub E B. et al. Identification of R-gene homologous DNA fragments genetically linked to disease resistance loci in Arabidopsis thaliana [J]. Mol. Plant Microbe Interact, 1998, 11(4): 251-258.
[22] Dodds P N. Lawrence G J, Ellis J G. Contrasting modes of evolution acting on the complex N locus for rust resistance in flax [J]. The Plant J, 2001, 27(5): 439-453.
[23] Yu Y G, Buss G R, Maroof M A S. Isolation of a superfamily of candidate disease-resistance genes in soybean based on a conserved nucleotide-binding site [J]. Proc Natl Acad Sci, 1996, 93(21): 11751-11756.
[24] Chen X M, Line R F, Leung H. Genome scanning for resistance-gene analogs in rice, barley, and wheat by high-resolution electrophoresis [J]. Theor Appl Genet, 1998, 97(3): 345-355.
[25] Madsen L H, Collins N C, Rakwalska M, et al. Barley disease resisitance gene analogs of the NBS-LRR class: identification and mapping [J]. Mol Genet Genomics, 2003, 269(1): 150-161.
[26] Mohler V, Klahr A, Wenzel G, et al. A resistance gene analog useful for targeting disease resistance genes against different pathogens on group 1S chromosomes of barley, wheat and rye [J]. Theor Appl Genet, 2002, 105(2-3): 364-368.
[27] Shi Z X, Chen X M, Line R F, et al. Development of resistance gene analog polymorphism markers for the Yr9 gene resistance to wheat stripe rust [J]. Genome, 2001, 44(4): 509-516.
[28] Collins N C, Webb C A, Seah S, et al. The isolation and mapping of disease resistance gene analogs in maize [J]. Mol Plant Microb Interact, 1998, 11(10): 968-978.
[29] Quint M, Mihaljevic R, Dussle M, et al. Development of RGA-CAPS markers and genetic mapping of candidate genes for sugarcane mosaic virus resistance in maize [J]. Theor Appl Genet, 2002, 105(2-3): 355-363.
[30] Kalavacharia V, Stavely J R, Myers J R, et al. Crg, a gene required for Ur-3-mediatd rust resistance in common bean, maps to a resistance gene analog cluster [J]. Mol Plant Microbe Interact, 2000, 13(11): 1237-1242.
[31] Meyers B C, Shen K A, Rohani P, et al. Receptor-like genes in the major resistance locus of lettuce are subject to divergent selection[J]. Plant Cell, 1998, 10(11): 1833-1846.
[32] Noel L, Moore T L, Biezen E A, et al. Pronounced intraspecific haplotype divergence at the RPP5 complex disease resistance locus of Arabidopsis [J]. Plant Cell, 1999, 11(11): 2099-2112.
[33] Meyers B C, Kozik A, Griego A, et al. Genome-wide analysis of NBS-LRR-encoding genes in Arabidopsis [J]. Plant Cell, 2003, 15(4): 809-834.
[34] Collins N, Drake J, Aycliffe M, et al. Molecular characterization of the maize Rpl-D rust resistance haplotype and its mutants [J]. Plant Cell, 1999, 11: 1365-1376.
[35] Graham M A, Marek L F, Lohnes D, et al. Expression and genome organization of resistance gene analogs in soybean [J]. Genome, 2000, 43(1): 86-93.
[36] Ramalingam J, vera Cruz C M, Kukreja K, et al. Candidate defense genes from rice, barley, and maize and their association with qualitative and quantitative resistance in rice [J]. Mol Plant Microbe Interact, 2003, 16(1): 14-24.
[37] Di Gaspero G, Cipriani G. Nucleotide binding site/leucine-rich repeats, pto-like and receptor-like kinases related to disease resistance in grapevine [J]. Mol Genet Genomics, 2003, 269(5): 612-623.
[38] Donald TM, PelleroneF, Adam-Blondon AF, et al. Indentification of resistance gene analogs linked to a powdery mildew resistance locus in grapevine [J]. Theor Appl Genet, 2002, 104(4): 610-618.
[39] Leister D, Ballvora A, Salamin F, et al. A PCR-based approach for isolating pathogen resistance genes from potato with potential for wide application in plants [J]. Nature Genetics, 1996, 14: 421-429.
[40] Huang L, Brooks S A, Li W L, et al. Map-based cloning of leaf rust resistance gene Lr21 from the large and polyploid genome of bread wheat [J]. Genetics, 2003, 164: 655-664.
[41 ] Radwan O, Bouzidi M F, Vear F, et al. Indentification of non-TIR-NBS-LRR markers linked to the PI5/PI8 locus for resistance to downy mildew in sunflower [J]. Theor Appl Genet, 2003, 106(8): 1438-1446.
[42] Byrne PF, McMullenMD, Snook ME, et al. Quantitative Trait loci and metabolic pathways: genetic control of the concentration of maysin, a corn earworm resistance factor in maize siks [J]. Proc Natl Acad Sci, 1996, 93(17): 8820-8825.
[43]Geffroy V, Sevignac M, De Oliveira J C, et al. Inheritance of partial resistance against Colletotrichum lindemuthianum in Phaseolus vulgaris and co-localization of quantitative trait loci with genes involved in specific resistance [J]. Mol Plant Microbe Interact, 2000, 13(3): 287-296.
[44] Zhang L P, Khan A, Nino-Liu D, et al. A molecular linkage map of tomato displaying chromosomal locations of resistance gene analogs based on a Lycopersicon esculentum Lycopersicon hirsutum cross [J]. Genome, 2002, 45(1): 133-146.
[45] Flandez Galvez H, Ford R, Pang E C, et al. An intraspecific linkage map of the chickpea (Cicer arietinum L. ) genome based on sequence tagged microsatellite site and resistance gene analog markers [J]. Theor Appl Genet, 2003, 106(8): 1447-1456.
[46] Nior S, Combes M C, Anthony F, et al. Origin, diversity and evolution of NBS-type disease-resistance gene homologues in coffee trees (Coffea L) [J]. Mol Genet Genomics, 2001, .265(4): 654-662.
[47]Graham M A, Marek L F, Shoemaker R C. Organization, expression and evolution of a disease resistance gene cluster in soybean[J].Genetics,2002,162(4):1961-1977.
[48]Lee S Y,Seo J S,Rodriguez-Lanetty M,et al.Comparative analysis of superfamilies of NBS-encoding disease resistance gene analogs in cultivated and wild apple species[J].Mol Genet Genomics,2003,269(1):101-108.
[49]Liu J J,Ekramoddoullah A K.Isolation,genetic variation and expression of TIR-NBS-LRR resistance gene analogs from western white pine(Pinus monticola Dougl ex D Don)[J].Mol Genet Genomics,2003,270(5):432-441.
[50]Rossi M,Araujo P G,Paulet F,et al.Genomic distribution and characterization of EST-derived resistance gene analogs(RGAs)in sugarance[J].Mol Genet Gen,2003,269(3):406-419.
[51]Sun Y,Wang Y Y,He Y Q,et al.Genetic diversity and identification among rice varieties based on resistance gene analogues[J].Seed,2001,114(2):1-2,6.(孙雁,王云月,何月秋,等.水稻抗病基因同源序列多态性与品种鉴定[J].种子,2001,114(2):1-2,6.)
[52]王石平,刘王德,王江,等.用同源序列的染色体定位寻找水稻抗病基因DNA片段[J].植物学报,1998,40:42-50.
[53]曹张军,王美南,井金学.小麦抗条锈性遗传研究进展.麦类作物学报[J].2001,21(3):80-83.
[54]杨勤忠,杨佩文,王群,等.水稻抗病基因同源序列的克隆及测序分析[J].中国水稻科学,2001,15(4):241-247.
[55]吴建利,庄杰云,柴荣耀,等.水稻抗穗瘟基因的分子定位[J].植物病理学报.2000,30(2):111-115.
[56]李子银,陈受宜.水稻抗病基冈同源序列的克隆、定位及其表达[J].科学通报,1999,44(7):727-734.
[57]孙雁,王云月,万瑞亭,等.黑龙江水稻品种抗病基冈同源序列聚类分析[J].云南农业人学学报,2001,16(2):107-109.
[58]Kanazin V,Marek L F,Shoemaker R.Resistance gene analogues are conserved and clustered in soybean[J].ProcNatlAcad Sci USA,1996,93:11746-11750.
[59]Leister D,Kurth J,Schulze P,et al.RFLP and physical mapping of resistance gene homologues in rice(O.sative)and Barley(H.vulgare)[J].Theor Appl Genet,1999,98:509-520.
[60]Leister D,Kurth J,Laurie D A,et al.Rapid reorganization of resistance gene homologues in cereal genomes[J].Plant Biology,1998,95:370-375.
[61]薛勇彪,唐定中,张燕生,等.水稻基因组中R类抗病基冈同源序列的分离.科学通报,1998,43(3):277-281.
[62]Collins N,Park R,Spiel Meyer W,et al.A high-density cytogenetic map of the Aegilops tauschii genome incorporating retrotransposons and defense-related genes:insights into cereal chromosome structure and function[J].Plant Mol Biol,2002,48(5-6):767-790.
[63]Gentzbittel L,Mouzeyar S,Badaoui S,et al.Cloning of molecular markers for disease resistance in sunflower,Helianthus annuus L[J].TheorAppl Genet,1998,96:519-525.
[64]Onmori T,Murata M,Motoyoshi F.Characterization of disease resistance gene-like sequences in near～isogenic lines of tomato[J].Theor Appl Genet,1998,96:519-525.
[65]陈爱武,赵建伟,甘霖,等.甘蓝型油菜芥酸含量的双列杂交分析[J].中国油料,1999,21(3):10-14.
[66]Plieger S,Lefebvre V,Caranta C,et al.Disease resistance gene analogs as candidates for QTLs involved in pepper-pathogen interactions[J].Genome,1999,42:1100-1110.
[67]Deng Z,Huang S,Ling P,et al.Cloning and characterization of NBS-LRR class resistance-gene candidate sequences in citrus[J].Theor Appl Genet,2001,101:814-822.
[68]Boris A,Vinatzer B A,Patocchi A,et al.Apple contains receptor～like genes homologous to the Cladosporiumfulvum resistance gene family of tomato with a cluster of genes consegregating with Vfapple scab resistance[J].Mo1 Plant Microbe Interact,2001,14(4):508-515.
[69]Feuillet C,Schachermayr G,Keller B.Molecular cloning of a new receptor-like kinase gene encoded at the Lr1O disease resistance locus of wheat[J].The Plant J,1997,11(1):45-52.
[70]王时平,刘克德,王江,等.用同源序列的染色体定位寻找水稻抗病基因DNA片段[J].植物学报,1998,40:42-50.
[71]Botella M A,Coleman M J,Hughes D E,et al.Map position of 47 Arabidopsis sequence with sequence similarity to disease resistance genes[J].The Plant J,1997,12(5):1197-1211.
[72]Speulman E,Bouchez D,Holub E B,et al.Disease resistance gene homologs correlate with disease resistance loci of Arabidopsis thaliana[J].The Plant J,1998,14(4):467-474.
[73]Mago R,Nair S,Mohan M.Resistance gene analogues from rice:cloning sequencing and mapping [J].Theor Appl Genet,1999,99:50-57.
[74]Seah S,Sivasithamparam S,Karakousis A,et al.Cloning and characterization of a family of disease resistance gene analogs from wheat and barley[J].Theor Appl Genet,1998,97:937-945.
[75]Creusot F,Macadre C,Cana E F,et al.Cloning and molecular characterization of three members of the NBS-LRR subfamily located in the vicinity of the Co-2 locus for anthracnose resistance in phaseolusvulgaris[J].Genome,1999,42:254-264.
[76]郑先武,翟文学,李晓兵,等.水稻NBS-LRR类R基冈同源序列[J].中国科学(C辑),2001,31(1):43-51.
[77]Feuillet C,Travella S,Stein N.Map-based isolation of the leaf rust disease resistance gene Lr1O from the hexaploid wheat(Triticum aestivum L)genome[J].Proc Natl Acad Sci USA,2003,100:15253-15258.
[78]Seah S,Bariana H,Jahier J,et al.The introgressed segment carry rust resistance genes Yr17,Lr37and Sr38 in wheat can be assayed by a cloned disease resistance gene-like sequence[J].Thero Appl Genet,2001,102:600-605.
[79]王海燕,杨文香,孟庆芳,等.小麦抗叶锈病基因Lr35相关基因组DNA片段的分离[J].农业生物技术学报,2006,14(5):824-825.
[80]褚栋,杨文香,闫红飞,等.小麦抗叶锈病基因Lr38的RGA分析初报[J].植物病理学报,2005,35(6)(ZK):179-180.
[81]Kerber E R,Dyck P L.Transfer to hexaploid wheat of linked genes for adult plant leaf rust and seeding stem rust resistance from an amphiploid of Aegilops speltoide×Triticum monococcum [J].Genome,1990,33:530-537.
[82]Gold J.Development of a molecular marker for rust resistance genes Sr39 and Lr35 in wheat breeding lines[J].Electronic Journal of Biotechnology,1999,2(1):35-40.
[83]Seyfarth R,Feuillet C,Schachermayr G.et al.Development of a molecular marker for the adult plant leaf rust resistance gene Lr35 in wheat[J].TheorAppl Genet,1999,99:554-560.
[84]王海燕,杨文香,刘大群.小麦NBS-LRR类抗病基因同源序列的分离与鉴定[J].中国农业科学.2006,39(8):1558-1564.
[85]王海燕,刘大群,杨文香,等.TcLr35小麦中病程相关蛋白1基因的克隆及分析[J].植物遗传资源学报.2007,8(1):16-20.
[86]Roelf A P.Race specificity and methods of study.TheCereal Rust Vol I.New York:Academic Press,1985:131-164.
[87]Roelfs A P,Singh R P,Saari E E.Rust diseases of wheat:Concepts and Methods of Disease.CIMMYT,Mexico,D F Mexico,1992.
[88]Gill K S,Lubbers E L,Gill B S,et al.A genetic linkage map of Triticum tauschii(DD)and its relationship to the D genome of bread wheat(AABBDD)[J].Genome,1991,34:362-374.
[89]Di Gaspero G,Cipriani G.Resistance gene analogs are candidate markers for disease～resistance genes in grape(Vitis spp)[J].Thero Appl Genet,2002,106:1563-1572.
[90]Baldi P,Patocchi A,Zini E,et al.Cloning and linkage mapping of resistance gene homologues in apple[J].Thero Appl Genet,2004,109(1):231-239.
[91]Palomino C,Satovic Z,Cubero J I,et al.Identification and characterization of NBS-LRR class resistance gene analogs in faba bean(Vicia faba L)and chickpea(Citer arictinum L)[J].Genome,2006,49(10):1227-1237.
[92]Arumuganathan K,Earle E D.Nuclear DNA content of some important plant species[J].Plant Molecular Biology Reporter,1991,9:208-218.
[93]Gill B S,Appels R,Botha-Oberholster A M,et al.A workshop report on wheat genome sequencing [J].Genetics,2004,168:1087-1096.
[2]Kloppers F J,Pretorius Z A.Effects of combinations amongst genes Lr13,Lr34 and Lr37 on components of resistance in wheat to leaf rust[J].Plant Pathology,1997,46:737-750.
[3]Knott D R.The wheat rust breeding for resistance[J].TheorAppl Genet,1989,191-198.
[4]Yu J,Hu S N,Wang J,et al.A draft sequence of the rice genome(Oryza sativa L.ssp.indic a)[J].Science,2002,296:79-91.
[5]Deslandes L,Olivier J,Theulieres F.Resistance to Ralstonia solanacearum in Arabidopsis thaliana is conferred by the recessive RRSl-R gene,a member of a novel family of resistance genes[J].Proc Natl Acad SciUSA,2002,99:2404-2409.
[6]Bryan G T,Wu K S,Farrall L.A single amino acid difference distinguishes resistant and susceptible alleles of the rice blast resistance genePi-to[J].Plant Cell,2000,12:2033-2046.
[7]Buschges R,Hollrichner K,Panstrnga R.The barely Mlo gene:A novel control element of plant pathogen resistance[J].Cell,1997,88:695-705.
[8]Brandwagt B F,Mesbah L A,Takken F L,et al.A longevity assurance gene homolog of tomato mediates resistance to Altemaria alternata f.sp.lycopersiei toxins and fumonisin B1[J].Proc Natl Acad Sci USA,2000,97:4961-4966.
[9]Kawchuk L M,Hachey J,Lynch D R,et al.Tomato Ve disease resistance genes encode cell surface-like receptors[J].Proc Natl Acad Sci USA,2001,98:6511-6515.
[10]Xiao S,Ellwood S,Calis O,et al.Broad-spectrum mildew resistance in Arabidopsis thaliana mediated byRPW8[J].Science,2001,291:118-120.
[11]Boyes D C,Nam J,Dangl J,et al.The Arabidopsis thaliana RPMl disease resistance gene product is a peripheral plasma membrane protein that is degrade coincident with the hypersensitive response[J].Proc Natl Acad Sci USA,1998,95:15849-15854.
[12]Bent A F.Plant disease resistance genes:function meets structure[J].Plant Cell,1996,8:1757-1771.
[13]Jones D A,Thomas C M,Hammond-Kosack K E,et al.Isolation of the tomato Cf-9 gene for resistance to Cladosporium fulvum by transposon tagging[J].Science,1994,266:789-793.
[14]Martin G B,Brommonschenkel S H,Chunwongse J,et al.Map-based cloning of a protein kinase gene conferring disease resistance in tomato[J].Science,1993,262:1432-1436.
[15]Whitham S,Dinesh-kumar S P,Chol D,et al.The product of the tobacco mosaic virus resistance gene N:similarity to Toll and the lnterleukin-1 receptor[J].Cell,1994,78:1101-1115.
[16]罗敏,末友林,余潮等.植物抗病基因的克隆及其结构与功能[J].遗传,2000,6:429-34.
[17]Song W Y,Wang G L,Chen L L,et al.A receptor kinase-like protein encoded by the rice disease resistance gene,Xa21[J].Science,1995,270:1804-1806.
[18] Meyers B C, Dickerman A W, Michelmore R W, et al. Plant disease resistance gene encode members of an ancient and diverse protein family within the nucleotide binding superfamily [J]. The Plant J, 1999, 20(3): 317-332.
[19] Brightbill H D, Libraty D H, Krutzik S R, et al. Host defense mechanisms triggered by microbial lipoprotein through Toll-like receptors [J]. Science, 1999, 285: 732-739.
[20] Lawrence G J, Finnegan E J, Ayliffe M A, et al. The L6 Gene for Flax Rust Resistance 1s Related to the Arabidopsis Bacterial Resistance Gene RPSP and the Tobacco Viral Resistance Gene N [J]. The Plant Cell, 1995, 7: 1195-1206.
[21] Aarts G M, Lintel Hekkert B, Holub E B. et al. Identification of R-gene homologous DNA fragments genetically linked to disease resistance loci in Arabidopsis thaliana [J]. Mol. Plant Microbe Interact, 1998, 11(4): 251-258.
[22] Dodds P N. Lawrence G J, Ellis J G. Contrasting modes of evolution acting on the complex N locus for rust resistance in flax [J]. The Plant J, 2001, 27(5): 439-453.
[23] Yu Y G, Buss G R, Maroof M A S. Isolation of a superfamily of candidate disease-resistance genes in soybean based on a conserved nucleotide-binding site [J]. Proc Natl Acad Sci, 1996, 93(21): 11751-11756.
[24] Chen X M, Line R F, Leung H. Genome scanning for resistance-gene analogs in rice, barley, and wheat by high-resolution electrophoresis [J]. Theor Appl Genet, 1998, 97(3): 345-355.
[25] Madsen L H, Collins N C, Rakwalska M, et al. Barley disease resisitance gene analogs of the NBS-LRR class: identification and mapping [J]. Mol Genet Genomics, 2003, 269(1): 150-161.
[26] Mohler V, Klahr A, Wenzel G, et al. A resistance gene analog useful for targeting disease resistance genes against different pathogens on group 1S chromosomes of barley, wheat and rye [J]. Theor Appl Genet, 2002, 105(2-3): 364-368.
[27] Shi Z X, Chen X M, Line R F, et al. Development of resistance gene analog polymorphism markers for the Yr9 gene resistance to wheat stripe rust [J]. Genome, 2001, 44(4): 509-516.
[28] Collins N C, Webb C A, Seah S, et al. The isolation and mapping of disease resistance gene analogs in maize [J]. Mol Plant Microb Interact, 1998, 11(10): 968-978.
[29] Quint M, Mihaljevic R, Dussle M, et al. Development of RGA-CAPS markers and genetic mapping of candidate genes for sugarcane mosaic virus resistance in maize [J]. Theor Appl Genet, 2002, 105(2-3): 355-363.
[30] Kalavacharia V, Stavely J R, Myers J R, et al. Crg, a gene required for Ur-3-mediatd rust resistance in common bean, maps to a resistance gene analog cluster [J]. Mol Plant Microbe Interact, 2000, 13(11): 1237-1242.
[31] Meyers B C, Shen K A, Rohani P, et al. Receptor-like genes in the major resistance locus of lettuce are subject to divergent selection[J]. Plant Cell, 1998, 10(11): 1833-1846.
[32] Noel L, Moore T L, Biezen E A, et al. Pronounced intraspecific haplotype divergence at the RPP5 complex disease resistance locus of Arabidopsis [J]. Plant Cell, 1999, 11(11): 2099-2112.
[33] Meyers B C, Kozik A, Griego A, et al. Genome-wide analysis of NBS-LRR-encoding genes in Arabidopsis [J]. Plant Cell, 2003, 15(4): 809-834.
[34] Collins N, Drake J, Aycliffe M, et al. Molecular characterization of the maize Rpl-D rust resistance haplotype and its mutants [J]. Plant Cell, 1999, 11: 1365-1376.
[35] Graham M A, Marek L F, Lohnes D, et al. Expression and genome organization of resistance gene analogs in soybean [J]. Genome, 2000, 43(1): 86-93.
[36] Ramalingam J, vera Cruz C M, Kukreja K, et al. Candidate defense genes from rice, barley, and maize and their association with qualitative and quantitative resistance in rice [J]. Mol Plant Microbe Interact, 2003, 16(1): 14-24.
[37] Di Gaspero G, Cipriani G. Nucleotide binding site/leucine-rich repeats, pto-like and receptor-like kinases related to disease resistance in grapevine [J]. Mol Genet Genomics, 2003, 269(5): 612-623.
[38] Donald TM, PelleroneF, Adam-Blondon AF, et al. Indentification of resistance gene analogs linked to a powdery mildew resistance locus in grapevine [J]. Theor Appl Genet, 2002, 104(4): 610-618.
[39] Leister D, Ballvora A, Salamin F, et al. A PCR-based approach for isolating pathogen resistance genes from potato with potential for wide application in plants [J]. Nature Genetics, 1996, 14: 421-429.
[40] Huang L, Brooks S A, Li W L, et al. Map-based cloning of leaf rust resistance gene Lr21 from the large and polyploid genome of bread wheat [J]. Genetics, 2003, 164: 655-664.
[41 ] Radwan O, Bouzidi M F, Vear F, et al. Indentification of non-TIR-NBS-LRR markers linked to the PI5/PI8 locus for resistance to downy mildew in sunflower [J]. Theor Appl Genet, 2003, 106(8): 1438-1446.
[42] Byrne PF, McMullenMD, Snook ME, et al. Quantitative Trait loci and metabolic pathways: genetic control of the concentration of maysin, a corn earworm resistance factor in maize siks [J]. Proc Natl Acad Sci, 1996, 93(17): 8820-8825.
[43]Geffroy V, Sevignac M, De Oliveira J C, et al. Inheritance of partial resistance against Colletotrichum lindemuthianum in Phaseolus vulgaris and co-localization of quantitative trait loci with genes involved in specific resistance [J]. Mol Plant Microbe Interact, 2000, 13(3): 287-296.
[44] Zhang L P, Khan A, Nino-Liu D, et al. A molecular linkage map of tomato displaying chromosomal locations of resistance gene analogs based on a Lycopersicon esculentum Lycopersicon hirsutum cross [J]. Genome, 2002, 45(1): 133-146.
[45] Flandez Galvez H, Ford R, Pang E C, et al. An intraspecific linkage map of the chickpea (Cicer arietinum L. ) genome based on sequence tagged microsatellite site and resistance gene analog markers [J]. Theor Appl Genet, 2003, 106(8): 1447-1456.
[46] Nior S, Combes M C, Anthony F, et al. Origin, diversity and evolution of NBS-type disease-resistance gene homologues in coffee trees (Coffea L) [J]. Mol Genet Genomics, 2001, .265(4): 654-662.
[47]Graham M A, Marek L F, Shoemaker R C. Organization, expression and evolution of a disease resistance gene cluster in soybean[J].Genetics,2002,162(4):1961-1977.
[48]Lee S Y,Seo J S,Rodriguez-Lanetty M,et al.Comparative analysis of superfamilies of NBS-encoding disease resistance gene analogs in cultivated and wild apple species[J].Mol Genet Genomics,2003,269(1):101-108.
[49]Liu J J,Ekramoddoullah A K.Isolation,genetic variation and expression of TIR-NBS-LRR resistance gene analogs from western white pine(Pinus monticola Dougl ex D Don)[J].Mol Genet Genomics,2003,270(5):432-441.
[50]Rossi M,Araujo P G,Paulet F,et al.Genomic distribution and characterization of EST-derived resistance gene analogs(RGAs)in sugarance[J].Mol Genet Gen,2003,269(3):406-419.
[51]Sun Y,Wang Y Y,He Y Q,et al.Genetic diversity and identification among rice varieties based on resistance gene analogues[J].Seed,2001,114(2):1-2,6.(孙雁,王云月,何月秋,等.水稻抗病基因同源序列多态性与品种鉴定[J].种子,2001,114(2):1-2,6.)
[52]王石平,刘王德,王江,等.用同源序列的染色体定位寻找水稻抗病基因DNA片段[J].植物学报,1998,40:42-50.
[53]曹张军,王美南,井金学.小麦抗条锈性遗传研究进展.麦类作物学报[J].2001,21(3):80-83.
[54]杨勤忠,杨佩文,王群,等.水稻抗病基因同源序列的克隆及测序分析[J].中国水稻科学,2001,15(4):241-247.
[55]吴建利,庄杰云,柴荣耀,等.水稻抗穗瘟基因的分子定位[J].植物病理学报.2000,30(2):111-115.
[56]李子银,陈受宜.水稻抗病基冈同源序列的克隆、定位及其表达[J].科学通报,1999,44(7):727-734.
[57]孙雁,王云月,万瑞亭,等.黑龙江水稻品种抗病基冈同源序列聚类分析[J].云南农业人学学报,2001,16(2):107-109.
[58]Kanazin V,Marek L F,Shoemaker R.Resistance gene analogues are conserved and clustered in soybean[J].ProcNatlAcad Sci USA,1996,93:11746-11750.
[59]Leister D,Kurth J,Schulze P,et al.RFLP and physical mapping of resistance gene homologues in rice(O.sative)and Barley(H.vulgare)[J].Theor Appl Genet,1999,98:509-520.
[60]Leister D,Kurth J,Laurie D A,et al.Rapid reorganization of resistance gene homologues in cereal genomes[J].Plant Biology,1998,95:370-375.
[61]薛勇彪,唐定中,张燕生,等.水稻基因组中R类抗病基冈同源序列的分离.科学通报,1998,43(3):277-281.
[62]Collins N,Park R,Spiel Meyer W,et al.A high-density cytogenetic map of the Aegilops tauschii genome incorporating retrotransposons and defense-related genes:insights into cereal chromosome structure and function[J].Plant Mol Biol,2002,48(5-6):767-790.
[63]Gentzbittel L,Mouzeyar S,Badaoui S,et al.Cloning of molecular markers for disease resistance in sunflower,Helianthus annuus L[J].TheorAppl Genet,1998,96:519-525.
[64]Onmori T,Murata M,Motoyoshi F.Characterization of disease resistance gene-like sequences in near～isogenic lines of tomato[J].Theor Appl Genet,1998,96:519-525.
[65]陈爱武,赵建伟,甘霖,等.甘蓝型油菜芥酸含量的双列杂交分析[J].中国油料,1999,21(3):10-14.
[66]Plieger S,Lefebvre V,Caranta C,et al.Disease resistance gene analogs as candidates for QTLs involved in pepper-pathogen interactions[J].Genome,1999,42:1100-1110.
[67]Deng Z,Huang S,Ling P,et al.Cloning and characterization of NBS-LRR class resistance-gene candidate sequences in citrus[J].Theor Appl Genet,2001,101:814-822.
[68]Boris A,Vinatzer B A,Patocchi A,et al.Apple contains receptor～like genes homologous to the Cladosporiumfulvum resistance gene family of tomato with a cluster of genes consegregating with Vfapple scab resistance[J].Mo1 Plant Microbe Interact,2001,14(4):508-515.
[69]Feuillet C,Schachermayr G,Keller B.Molecular cloning of a new receptor-like kinase gene encoded at the Lr1O disease resistance locus of wheat[J].The Plant J,1997,11(1):45-52.
[70]王时平,刘克德,王江,等.用同源序列的染色体定位寻找水稻抗病基因DNA片段[J].植物学报,1998,40:42-50.
[71]Botella M A,Coleman M J,Hughes D E,et al.Map position of 47 Arabidopsis sequence with sequence similarity to disease resistance genes[J].The Plant J,1997,12(5):1197-1211.
[72]Speulman E,Bouchez D,Holub E B,et al.Disease resistance gene homologs correlate with disease resistance loci of Arabidopsis thaliana[J].The Plant J,1998,14(4):467-474.
[73]Mago R,Nair S,Mohan M.Resistance gene analogues from rice:cloning sequencing and mapping [J].Theor Appl Genet,1999,99:50-57.
[74]Seah S,Sivasithamparam S,Karakousis A,et al.Cloning and characterization of a family of disease resistance gene analogs from wheat and barley[J].Theor Appl Genet,1998,97:937-945.
[75]Creusot F,Macadre C,Cana E F,et al.Cloning and molecular characterization of three members of the NBS-LRR subfamily located in the vicinity of the Co-2 locus for anthracnose resistance in phaseolusvulgaris[J].Genome,1999,42:254-264.
[76]郑先武,翟文学,李晓兵,等.水稻NBS-LRR类R基冈同源序列[J].中国科学(C辑),2001,31(1):43-51.
[77]Feuillet C,Travella S,Stein N.Map-based isolation of the leaf rust disease resistance gene Lr1O from the hexaploid wheat(Triticum aestivum L)genome[J].Proc Natl Acad Sci USA,2003,100:15253-15258.
[78]Seah S,Bariana H,Jahier J,et al.The introgressed segment carry rust resistance genes Yr17,Lr37and Sr38 in wheat can be assayed by a cloned disease resistance gene-like sequence[J].Thero Appl Genet,2001,102:600-605.
[79]王海燕,杨文香,孟庆芳,等.小麦抗叶锈病基因Lr35相关基因组DNA片段的分离[J].农业生物技术学报,2006,14(5):824-825.
[80]褚栋,杨文香,闫红飞,等.小麦抗叶锈病基因Lr38的RGA分析初报[J].植物病理学报,2005,35(6)(ZK):179-180.
[81]Kerber E R,Dyck P L.Transfer to hexaploid wheat of linked genes for adult plant leaf rust and seeding stem rust resistance from an amphiploid of Aegilops speltoide×Triticum monococcum [J].Genome,1990,33:530-537.
[82]Gold J.Development of a molecular marker for rust resistance genes Sr39 and Lr35 in wheat breeding lines[J].Electronic Journal of Biotechnology,1999,2(1):35-40.
[83]Seyfarth R,Feuillet C,Schachermayr G.et al.Development of a molecular marker for the adult plant leaf rust resistance gene Lr35 in wheat[J].TheorAppl Genet,1999,99:554-560.
[84]王海燕,杨文香,刘大群.小麦NBS-LRR类抗病基因同源序列的分离与鉴定[J].中国农业科学.2006,39(8):1558-1564.
[85]王海燕,刘大群,杨文香,等.TcLr35小麦中病程相关蛋白1基因的克隆及分析[J].植物遗传资源学报.2007,8(1):16-20.
[86]Roelf A P.Race specificity and methods of study.TheCereal Rust Vol I.New York:Academic Press,1985:131-164.
[87]Roelfs A P,Singh R P,Saari E E.Rust diseases of wheat:Concepts and Methods of Disease.CIMMYT,Mexico,D F Mexico,1992.
[88]Gill K S,Lubbers E L,Gill B S,et al.A genetic linkage map of Triticum tauschii(DD)and its relationship to the D genome of bread wheat(AABBDD)[J].Genome,1991,34:362-374.
[89]Di Gaspero G,Cipriani G.Resistance gene analogs are candidate markers for disease～resistance genes in grape(Vitis spp)[J].Thero Appl Genet,2002,106:1563-1572.
[90]Baldi P,Patocchi A,Zini E,et al.Cloning and linkage mapping of resistance gene homologues in apple[J].Thero Appl Genet,2004,109(1):231-239.
[91]Palomino C,Satovic Z,Cubero J I,et al.Identification and characterization of NBS-LRR class resistance gene analogs in faba bean(Vicia faba L)and chickpea(Citer arictinum L)[J].Genome,2006,49(10):1227-1237.
[92]Arumuganathan K,Earle E D.Nuclear DNA content of some important plant species[J].Plant Molecular Biology Reporter,1991,9:208-218.
[93]Gill B S,Appels R,Botha-Oberholster A M,et al.A workshop report on wheat genome sequencing [J].Genetics,2004,168:1087-1096.