马铃薯抗病同源序列系统进化分析及晚疫病抗病基因的克隆
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摘要
马铃薯是仅次于水稻、小麦、玉米的世界第四大粮食作物,在国民经济中具有重要作用。马铃薯晚疫病是威胁马铃薯生产的主要病害之一,传统防治马铃薯晚疫病方法主要是采用抗菌剂及抗性育种策略。植物基因工程技术的不断完善和应用为马铃薯晚疫病的防治研究提供了新的契机,从马铃薯抗病资源中挖掘具有较高抗性的抗病基因,并将其导入到栽培品种中,无疑是我们获得和培育抗晚疫病马铃薯品系较为快捷的手段。本研究采用抗病基因同源序列法(即RGA法),结合RACE技术克隆马铃薯晚疫病抗病基因,研究结果如下:
(1)马铃薯抗病基因同源序列(RGAs)的分离和相似性分析
以马铃薯抗晚疫病野生品种Solanum dimissum为试材,根据GenBank中已登录的植物抗病基因(R3a、R1、Rpi-blb1、Rpi-blb2、I2、I2C-1等)的NBS区的保守序列P-loop和GLPL区设计简并引物,以S.demissum的cDNA作模板进行RT-PCR扩增,共获得30条具有开放阅读框的含NBS区保守结构域的序列,利用生物信息学软件对所得序列进行同源性比较和系统进化分析,30条具开放阅读框的序列根据核酸相似性90%为界共分成9类,皆为马铃薯的抗病基因同源序列(RGAs),且全部属于nonTIR-NBS-LRR类型。
(2)马铃薯晚疫病抗病基因候选基因的确定
根据9类RGAs聚类分析结果分别设计特异引物,对S.demissum及六个对照栽培品种进行PCR和RT-PCR检测,结果表明:第Ⅳ(PB6-22)类仅在S.demissum中存在,第Ⅵ类(PB1-1)在S.demissum和大西洋中呈阳性,而其他类RGAs在所有的七个品种中都有扩增产物。初步确定第Ⅳ类和第Ⅵ类为马铃薯抗晚疫病相关的候选基因。
(3)晚疫病抗病基因的克隆
利用RACE技术克隆出了第Ⅳ类(PB6-22)的基因,该基因命名为Sdpa,基因全长为3570bp,编码1189氨基酸,该基因含有抗病基因的NBS保守结构域(P-loop、Kinase2、Kinase-3a、GLPL等),并在此基因中鉴定出了至少8个LRR结构域,其编码蛋白与已克隆的马铃薯晚疫病抗病蛋白R3a相似性达66.24%。基因表达及抗病性鉴定工作有待于进一步研究。
Potato (Solanum tuberosum. L) is one of the most important crops in the world ranking fourth after rice, wheat and maize, and plays a significant role in economy. One of the most destructive diseases of potato is the late blight, which can lead to severe yield decrease. Traditionally, people combated the late blight of potato by fungicides and conventional breeding. With the development of Genetic Engineering, a lot of R genes in wild sources have been isolated and subsequent transformation of theses genes into potato cultivars could be a fast way to obtain disease resistance cultivars. In this study, we employ the RGA method combined with RACE technology to clone the late blight resistance gene of potato, the results showed below:
(1) Isolation and Phylogenetic Analysis of Potato Resistance Gene Analogs (RGAs)
We choose the classical late blight resistant species Solanum dimissum as plant material, degenerated oligonucleotide primers were designed according to the two conservative motifs: P-loop and GLPL sequences of previously reported resistance genes (R3a, R1, Rpi-blb1, Rpi-blb2, I2, I2C-1 etc.), as a result, 30 sequences with Open Reading Frame (ORF) were acquired from the cDNA of S.demissum, which includes the NBS conservative motifs. Sequences alignment and phylogenetic analysis were performed by bioinformatics softwares, the results indicated that 30 sequences with ORF were divided into 9 types based on the 90% nucleotide similarity. All of them belong to nonTIR-NBS-LRR resistance gene analogs (RGAs).
(2) Identification of Candidate sequences of Late Blight resistance
Nine specific primers were designed based on the specific domain of these nine RGAs, respectively, PCR and RT-PCR identification were performed in S.demissum and six domestic cultivars (Long 3, Nileke, Atlantic, Zihuabai, MC-4, Favorita), results showed that groupⅣappeared uniquely in S.demissum, as well as groupⅥin S.demissum and another cultivar Atlantic, while all the other seven groups were present in all the samples tested, which indicated that the groupⅣand groupⅥmight highly relate to the late blight resistance (Candidate Genes).
(3) Cloning of Late Blight Resistance Gene
We cloned the whole sequence of GroupⅣRGA (PB6-22) by RACE technology, this gene was named Sdpa, with coding sequence is 3570 bp. The deducted open reading frame (ORF) of the Sdpa gene encodes a predicted polypeptide of 1189 amino acid. The gene includes the conservative motifs of R genes (P-loop, Kinase2, Kinase-3a, GLPL etc.), and at least eight LRR motifs were identified in this protein. This resistance protein shares sequence homology of 66.24% with reported potato late blight resistance protein R3a. Our future studies will focus on expressing this disease resistance gene for functional analysis.
(1)马铃薯抗病基因同源序列(RGAs)的分离和相似性分析
以马铃薯抗晚疫病野生品种Solanum dimissum为试材,根据GenBank中已登录的植物抗病基因(R3a、R1、Rpi-blb1、Rpi-blb2、I2、I2C-1等)的NBS区的保守序列P-loop和GLPL区设计简并引物,以S.demissum的cDNA作模板进行RT-PCR扩增,共获得30条具有开放阅读框的含NBS区保守结构域的序列,利用生物信息学软件对所得序列进行同源性比较和系统进化分析,30条具开放阅读框的序列根据核酸相似性90%为界共分成9类,皆为马铃薯的抗病基因同源序列(RGAs),且全部属于nonTIR-NBS-LRR类型。
(2)马铃薯晚疫病抗病基因候选基因的确定
根据9类RGAs聚类分析结果分别设计特异引物,对S.demissum及六个对照栽培品种进行PCR和RT-PCR检测,结果表明:第Ⅳ(PB6-22)类仅在S.demissum中存在,第Ⅵ类(PB1-1)在S.demissum和大西洋中呈阳性,而其他类RGAs在所有的七个品种中都有扩增产物。初步确定第Ⅳ类和第Ⅵ类为马铃薯抗晚疫病相关的候选基因。
(3)晚疫病抗病基因的克隆
利用RACE技术克隆出了第Ⅳ类(PB6-22)的基因,该基因命名为Sdpa,基因全长为3570bp,编码1189氨基酸,该基因含有抗病基因的NBS保守结构域(P-loop、Kinase2、Kinase-3a、GLPL等),并在此基因中鉴定出了至少8个LRR结构域,其编码蛋白与已克隆的马铃薯晚疫病抗病蛋白R3a相似性达66.24%。基因表达及抗病性鉴定工作有待于进一步研究。
Potato (Solanum tuberosum. L) is one of the most important crops in the world ranking fourth after rice, wheat and maize, and plays a significant role in economy. One of the most destructive diseases of potato is the late blight, which can lead to severe yield decrease. Traditionally, people combated the late blight of potato by fungicides and conventional breeding. With the development of Genetic Engineering, a lot of R genes in wild sources have been isolated and subsequent transformation of theses genes into potato cultivars could be a fast way to obtain disease resistance cultivars. In this study, we employ the RGA method combined with RACE technology to clone the late blight resistance gene of potato, the results showed below:
(1) Isolation and Phylogenetic Analysis of Potato Resistance Gene Analogs (RGAs)
We choose the classical late blight resistant species Solanum dimissum as plant material, degenerated oligonucleotide primers were designed according to the two conservative motifs: P-loop and GLPL sequences of previously reported resistance genes (R3a, R1, Rpi-blb1, Rpi-blb2, I2, I2C-1 etc.), as a result, 30 sequences with Open Reading Frame (ORF) were acquired from the cDNA of S.demissum, which includes the NBS conservative motifs. Sequences alignment and phylogenetic analysis were performed by bioinformatics softwares, the results indicated that 30 sequences with ORF were divided into 9 types based on the 90% nucleotide similarity. All of them belong to nonTIR-NBS-LRR resistance gene analogs (RGAs).
(2) Identification of Candidate sequences of Late Blight resistance
Nine specific primers were designed based on the specific domain of these nine RGAs, respectively, PCR and RT-PCR identification were performed in S.demissum and six domestic cultivars (Long 3, Nileke, Atlantic, Zihuabai, MC-4, Favorita), results showed that groupⅣappeared uniquely in S.demissum, as well as groupⅥin S.demissum and another cultivar Atlantic, while all the other seven groups were present in all the samples tested, which indicated that the groupⅣand groupⅥmight highly relate to the late blight resistance (Candidate Genes).
(3) Cloning of Late Blight Resistance Gene
We cloned the whole sequence of GroupⅣRGA (PB6-22) by RACE technology, this gene was named Sdpa, with coding sequence is 3570 bp. The deducted open reading frame (ORF) of the Sdpa gene encodes a predicted polypeptide of 1189 amino acid. The gene includes the conservative motifs of R genes (P-loop, Kinase2, Kinase-3a, GLPL etc.), and at least eight LRR motifs were identified in this protein. This resistance protein shares sequence homology of 66.24% with reported potato late blight resistance protein R3a. Our future studies will focus on expressing this disease resistance gene for functional analysis.
引文
1. 陈观水,周以飞,林生,潘大生.甘薯 NBS 类抗病基因类似物的分离与序列分析. 热带亚热带植物学报,2006,14(5):359-365.
2. 谌谋华,邓秀新. 利用同源序列法克隆柑桔抗病基因类似物及其初步分析.[硕士研究生学位论文]. 武汉:华中农业大学,2003.
3. 丁国华,秦智伟,刘宏宇,周秀艳,池春玉,王志坤. 黄瓜 NBS 类型抗病基因同源序列的克隆与分析. 园艺学报,2005,32(4):638-642.
4. 杲修杰,王正华. 植物抗病基因研究进展. 长江大学学报,2006, 3(4):205-210.
5. 郭军,屈冬玉,王晓武 .cDNA-AFLP 结合 BSA 研究马铃薯晚疫病菌小种特异无毒基因差异表达片段. 中国马铃薯, 2004, 18(1):1-3.
6. 黄萱,徐子勤,陈立余,王健.小麦 NBS-LRR 类抗病基因同源序列的分离与鉴定. 分子细胞生物学报,2006, 39(2):91-96.
7. 蒋敏华,杨清,李丽,黄先群. 马铃薯抗晚疫病转基因研究进展. 种子,2006,25(12):46-50.
8. 孔巍,方宣钧. 海岛棉 NBS-LRR 类抗病基因同源序列的克隆与定位. [硕士研究生学位论文]. 北京:中国农业科学院,2003.
9. 李春来,张怀渝,植物抗病基因同源序列(RGA)研究进展,分子植物育种,2004,2(6):
853-860.
10. 李金玉,李冠,赵惠新,王贤雷,杜钰. 植物抗病分子机制研究进展. 种子,2006, 25(2):
45-50.
11. 李先平,何云昆,赵志坚,孙茂林,隋启君. 马铃薯抗晚疫病育种研究进展. 中国马铃薯, 2001,
15(5):290-295.
12. 梁凤山,周春江,孔凡娜,王斌. 桃基因组中 R 类抗病基因同源序列的克隆与序列分析. 河北农业大学学报,2005,28(1):44-48.
13. 刘晓鹏, 姜宁, 何礼远. 马铃薯晚疫病菌基因文库的构建. 东北农业大学学报, 1998, 29 (3)::228-235.
14. 卢志国,韩建民,董金皋. 植物抗病基因研究现状. 河北农业大学学报,2002,25:160-163.
15. 马媛媛,甘睿,王宁宁. 植物富含亮氨酸重复序列型受体蛋白激酶的生物学功能. 植物生理与分子生物学学报. 2005, 31(4):331-339.
16. 盛刚,李付广,王涛. 海岛棉 NBS-LRR 抗性基因类似序列的分离及 Gbarvin 基因的鉴定.[硕士研究生学位论文]. 北京:中国农业科学院,2005.
17. 孙雁,王云月,何月秋,朱有勇. 水稻抗病基因同源序列多态性与品种鉴定,种子,2001,114(2):1-2,6.
18. 田振东,柳俊,谢从华. cDNA 文库与 RACE 方法结合克隆一个马铃薯病程相关蛋白基因cDNA. 遗传学报,2003, 30(11):96-1002.
19. 万里红,周亦华,陈正华. 植物防御系统中抗病相关基因的研究进展. 遗传,2002, 24(4):486-492.
20. 王海燕,刘大群. TcLr35 小麦抗病相关基因的克隆及分析.[博士学位论文]. 保定:河北农业大学,2006.
21. 王友红,张鹏飞,陈建群.植物抗病基因及其作用机理. 植物学通报,2005,22(1):92-99.
22. 徐兵强,杜中军,黄俊生. RGA 法克隆候选抗病基因的研究进展. 分子植物育种,2004,2(3):421-423.
23. 徐建飞,辛翠花, 黄三文. 利用田间抗病基因近等混合系防治马铃薯晚疫病. 中国农业科技导报,2006, 8(4):8-13.
24. 许丽,李玥莹,林凤. 植物抗病的分子基础与研究进展. 杂粮作物,2006, 26(6):428-432.
25. 杨秀红,陈庆山,杨庆凯,李文滨. 大豆 NBS 类抗病相关基因的克隆与序列分析. 高技术通讯,2006,5(2):71-78.
26. 俞志华,祝水金,夏英武. 从植物抗病基因的克隆看其基因结构、功能和进化. 浙江大学学报,2002,28(1):107-113.
27. 左示敏,殷跃军,张亚芳,陈宗祥,潘学彪. 植物数量抗病基因克隆及其抗性机制的研究进展. 分子植物育种,2006,4(5):603-613.
28. Aarts M.G.M. te Lintel Hekkert B., Holub E.B., Beynon J.L. Identification of R-gene homologous DNA fragments genetically linked to disease resistance loci in Arabidopsis thaliana. Mol. Plant Microbe Interact., 1998, 11(4):251-258.
29. Alber T. Structure of the leucine zipper. Curr. Opin. Genet. Dev., 1992, 2: 205-210.
30. Ananieva K.I. Effect of methyl ester of jasmonic acid and benzyl-aminopurine on growth and protein profile of excised cotyledons of Cucurbita pepo. Zucchini. Biologia, lantarium., 1999,42: 549-557.
31. Anderson P.A., Lawrence G.J., Morrish B.C., Aylifee M.A., Finnegan E.J., and Ellis J.G. Inactivation of the flax rust resistance gene M associated with loss of a repeated unit within the leucine-rich repeat coding region. Plant Cell, 1997, 9(4): 641-651.
32. Ballvora A, Ercolano MR, WeiB J, Meksem K, Bormann CA, Oberhagemann P, Salamini F, Gebhardt C .The R1 gene for potato resistance to late blight (Phytophthora infestans) belongs to the leucine zipper/NBS/LRR class of plant resistance genes. Plant Journal, 2002, 30: 361–371.
33. Bendahmane A, Kanyuka K, Baulcombe DC. The Rx gene from potato controls separate virus resistance and cell death response. Plant Cell, 1999, 11:781–791.
34. Bent A.F., Kunkel B.N., Dahlbeck D., et al. RPS2 of Arabidopsis thaliana: a leucine-rich repeat class of plant disease resistance genes. Science, 1994, 265:1856-1860.
35. Buschges R., Hollricher K., Panstruga R., Simons G. Wolter M., Frijters A., van Daelen R., and Schulze-Lefert. The barley Mlo gene: a novel control element of plant pathogen resistance. Cell, 1997, 88(5): 695-705.
36. Byrne P.F., McMullen M.D., Snook M.E., Musket T.A. Quantitative trait loci and metabolic pathways: genetic control of the concentration of maysin, a corn earworm resistance factor in maize silks. Proc.Natl.Acad.Sci.USA, 1996, 93(17):8820-8825.
37. Cai D., Kleine M., Kifle S., Harloff H.J., Sandal N.N., Marcker K.A., Klein-Lankhorst R.M., Salentijn E.M.J., Lange W. Positional cloning of a gene for nematode resistance in sugar beet. Science, 1997, 275(5301):832-834.
38. Catherine F, Gabrieles S, Bent K. Molecular cloning of a new receptor-like kinase gene encoded the Lr10 disease resistance locus wheat. The Plant Journal, 1997, 11 (1): 45-52.
39. Chen X.M., Line R.F., Leung H. Genome scanning for resistance-gene analogs in rice, barley, and wheat by high-resolution electrophoresis. Theor. Appl. Genet., 1998, 97(3):345-355.
40. Chunwongse J., Chunwongse C., Black L., Hanson P. Molecular mapping of the Ph-3 gene for late blight resistance in tomato. J Hort. Sco&Biotechnol., 2002, 77:281-286.
41. Collins N C, W ebb C A, Seah S, Pryor. The isolation and mapping of disease resistance gene analogs in maize. Mol. Plant Microbe. Interact. , 1998, 11: 968-978.
42. Dangl, J.L and Jones, J.D.G. Plant pathogens and integrated defence responses to infection. Nature, 2001, 411: 826-833.
43. David A.Jones, Clowyn M. Thomas, Kim. E. Hammond Kosack. Isolation of the tomato Cf-9 gene for resistance Cladosporium fulvum by transposon tagging. Science, 1994, 266: 789-793.
44. De Jong W., Forsyth A., Lister D., Hebhardt C. and Baulcombe D.C. A potato hypersensitive resistance gene cluster on chromosome Ⅴ. Theor. Appl. Genet., 1995: 153-162.
45. Dilbirligi M., Erayman M., Sandhu D. Identification of wheat chromosomal regions containing expressed resistance genes. Genetics, 2004, 166: 461- 481.
46. Dixon M.S., Hatzixanthis K., Jones D.A. The tomato Cf-5 disease resistance gene and six homologs show pronounced allelic variation in leucine-rich repeat copy number. Plant Cell,1998,10:1915～1925.
47. Donald T.M., Pellerone F., Adam-Blondon A.F., Bouquet A., Thomas M.R., and Dry I.B. Identification of resistance gene analogs linked to a powdery mildew resistance locus in grapevine. Theor. Appl. Genet., 2002,104 (4):610-618.
48. Edwin A.G. van der Vossen, Anne Sikkema, Bas te Lintel Hekkert. An ancient R gene from the wild potato species Solanum bulbocastanum confers broad-spectrum resistance to Phytophthora infestans in cultivated potato and tomato. The Plant Journal, 2003, 36 (6): 867-882.
49. Edwin A.G. van der Vossen, Jack Gros, Anne Sikkema. The Rpi-blb2 gene from Solanum bulbocastanum is an Mi-1 gene homolog conferring broad-spectrum late blight resistance in potato. The Plant Journal, 2005, 44(2):208-222.
50. Edwin A.G. van der Vossen, van der Voort J.N., Kanyuka K., Bendahmane A., Sandbrink H., Baulcombe D.C., Bakker J., Stiekema W. J. and Klein-Lankhorst R. M. Homologues of a single resistance-gene cluster in potato confer resistance to distinct pathogens: A virus and a nematode. Plant J. 2000, 23:567-576.
51. El-Kharbotly A, Palomino-Sanchez C, Salamini F, Jacobsen E, Gebhardt C. R6 and R7 alleles of potato conferring race-specific resistance to Phytophthora infestans (Mont.) de Bary identified genetic loci clustering with the R3 locus on chromosome Ⅺ. Theor. Appl. Genet., 1996,92:880–884.
52. Ellis J.G.,Doods P, Pryor T. The generation of plant disease resistance specificities. Trends in Plant Science, 2000, 5: 373-379.
53. Ellis, J.G., Lawrence, G.J., Luck, J.E. and Dodds, P.N. Identification of regions in alleles of the flax rust resistance gene L that determine differences in gene-for-gene specificity. Plant Cell, 1999, 11: 495–506.
54. Ewing, E.E., Simko, I., Smart, C.D., Bonierbale, M. W., Mizubuti, E. S. G., May, G. D., and Fry, W. E. Genetic mapping of qualitative and quantitative field resistance to Phytophthora infestans in a population derived from Solanum tuberosum and Solanum Berthaulti. Molecular Breeding, 2000, 6:25-36.
55. Feuillet C, Schachermagr G, Keller B. Molecular cloning of a new receptor-like kinase gene encoding the Lr10 disease resistance locus of wheat. The Plant Journal, 1997, 11 (1):45-52.
56. Flandez-Galvez H., Ford R., Pang E.C Taylor P.W. An intraspecific linkage map of the chickpea (Cicer arietinum L.) genomes based on sequence tagged microsatellite site and resistance gene analog markers, Theor. Appl. Genet., 2003, 106(8):1447-1456.
57. Flor H H. Current Status of the Gene- for - gene Concept. Annual Review of Phytopathology, 1971, 9: 275-296.
58. Fourmann M., Chariot F., Froger N., Delourme R., and Brunel D. Expression, mapping, and genetic variability of Brassica napus disease resistance gene analogues, Genome, 2001, 44(6):1083-1099.
59. Fry, W.E. and Goodwin, S.B. Resurgence of the Irish potato famine fungus. Bioscience, 1997, 47:363–371.
60. Rauscher G. M., Smart C. D., Simko I. Characterization and mapping of RPi-ber, a novel potato late blight resistance gene from Solanum berthaultii. Theor. Appl. Genet., 2006, 112: 674-687.
61. Gebhardt C., Valkonen J.P.T. Organization of genes controlling disease resistance in the potato genome. Annual Review of Phytopathology, 2001, 39:79–102.
62. Graham M.A., Marek L.F. Shoemaker R.C. Organization, expression and evolution of a disease resistance gene cluster in soybean, Genetics, 2002, 162(4):1961-1977.
63. Grant M.R., Godiard L., Straube E., Ashfield T., Lewald J., Sattler A., Innes R.W., and Dangl J.L. Structure of the Arabidopsis RPM1 gene enabling dual specificity disease resistance, Science, 1995, 269: 843-846.
64. Halterman D.A., Kramer L.C., Wielgus S., Jiang J. Performance of transgenic potato containing the late blight resistance gene RB. Plant Dis., 2008, 92: 339-343.
65. Hammond-Kosack K. E. Plant disease resistance genes. Annual Review of Plant Physiology and Plant Molecular Biology, 1997, 48: 575-607.
66. Hanhui Kuang, Fusheng Wei, Maria Rosa Marano, Uwe Wirtz, Xiaoxue Wang. The R1 resistance gene cluster contains three groups of independently evolving, type I R1 homologues and shows substantial structural variation among haplotypes of Solanum demissum. The Plant J., 2005,44:37-50.
67. Huang Sanwei, Edwin A.G.van der Vossen, Hanhui Kuang .Comparative genomics enabled the isolation of the R3a late blight resistance gene in potato. The Plant J., 2005, 42: 251-261.
68. Huang Sanwei, Vivianne G.A.A. Vleeshouwers, Jeroen S. Werij, Ronald C.B. Hutten, Herman J. van Eck, Richard G.F.Visser, and Evert Jacobsen. The R3 resistance to Phytophthora infestans in potato is conferred by two closely linked R genes with distinct specificities. Mol. Plant Micro. Inter., 2004, 17(4): 428-435.
69. Hulbert, S.H., Webb, C.A., Smith, S.M. and Sun, Q. Resistance gene complexes: evolution and utilization. Annu. Rev. Phytopathol., 2001, 39, 285–312.
70. Hunger S., di Gaspero G., Mohring S. Isolation and linkage analysis of expressed disease- resistance gene analogues of sugar beet( Beta vulgaris L.) Genome, 2003, 46(1):70-82.
71. Jia Y., McAdams S.A., Bryan G.T., et al. Direct interaction of resistance gene and a virulence gene product confers rice blast resistance. EMBO J, 2000, 19: 4004-4014.
72. Johal G. S., and Briggs S.P. Reductase activity encoded by the HM1 disease resistance gene in maize, Science, 1992, 258(5084):985-987.
73. John E. Bradshaw, Glenn J. Bryan, Alison K. Lees. Mapping the R10 and R11 genes for resistance to late blight (Phytophthora infestans) present in the potato (Solanum tuberosum) R-gene differentials of Black. Theor. Appl. Genet., 2006, 112(4):744-751.
74. Johnaton S.A., den Nijs, T.P.M. Peloquin S.J. and Hanneman R.E. The significance of genic balance to endosperm development in interspecific crosses. Theor. Appl. Genet., 1980, 57: 5-9.
75. Jones D.A., Thomas C.M., Hammond-Kosack K.E., Balint-Kurti P.J., and Jones J.D. Isolation of the tomato Cf-9 gene for resistance to Cladosporium fulvum by transposon tagging. Science, 1994, 165(5186):789-793.
76. Kobe B, Deisenhofer J. The leucine-rich repeat: a versatile binding motif, Trends. Biochem. Sci., 1994, 19(10):415-421.
77. Kuhl JC, Hanneman RE Jr, Havey MJ. Characterization and mapping of Rpi1, a late-blight resistance locus from diploid (1EBN) Mexican Solanum pinnatisectum. Mol. Genet. Genomics, 2001, 265(6):977-85.
78. McHale L., Tan X. P., Koehl P., Michelmore R.W. Plant NBS-LRR proteins: adaptable guards. Genome Biology, 2006, 7:212.
79. Lawrence G.J., Finnegan E.J., Ayliffe M.A., and Eliis J.G. The L6 gene for flax rust resistance is related to the Arabidopsis bacterial resistance gene RPS2 and the tobacco viral resistance gene N, Plant Cell, 1995, 7(8):1195-1206.
80. Lee S.Y., Seo J.S., Rodriguez-Lanetty M., Lee D.H. Comparative analysis of superfamilies of NBS-encoding disease resistance gene analogs in cultivated and wild apple specis, Mol. Genet.Genomics, 2003, 269 (1):101-108.
81. Leister D, Ballvora A, Salamini F, et al. A PCR-based approach for isolating pathogen resistance genes from potato with potential for wide application in plants. Proc. Natl. Acad. Sci. USA., 1996,14:421~429.
82. Leister D, Kurth T, Schulze P, RFLP and physical mapping of resistance gene homologues in rice and Barley. Theoretical and Applied Genetics, 1999, 98:509-520.
83. Leister D. Tandem and segmental gene duplication and recombination in the evolution of plant disease resistance gene. Trends in Genetics, 2004, 3: 116-122.
84. Leonards-Schippers C, Gieffers W, Schafer-Pregl R, Ritter E, Knapp S, Salamini F, Gebhardt C. Quantitative resistance to Phytophthora infestans in potato: a case study for QTL mapping in an allogamous plant species. Genetics, 1994, 137:67–77.
85. Li X, van Eck HJ, Rouppe van der Voort JNAM, Huigen DJ, Stam P, Jacobsen E. Autotetraploids and genetic mapping using common AFLP markers: the R2 allele conferring resistance to Phytophthora infestans mapped on potato chromosome 4. Theor Appl Genet., 1998, 96:1121–1128.
86. Madsen L.H., Collins N.C., Rakwalska M., Backes G., Sandal N., Krusell L., Jensen J., Waterman E.H., Jahoor A., Ayliffe M., Pryor A.J. Langridge P., Schulze-Lefert P., and Stougaard J. Barley disease resistance gene analogs of the NBS-LRR class: identification and mapping, Mol. Genet. Genomics, 2003, 269(1):150-161.
87. Mark S., Dixon, Jones D.A., Keddie J.S.The tomato Cf-2 disease resistance locus comprises two functional genes encoding leucine-rich repeat proteins.Cell,1996,84:451～459.
88. Martin G.B., Brommonschenkel S.H.Chunwongse J., Frary A., Ganal M.W., Spivey R., Wu T., Earle E.D. and Tanksley S.D., , Map-based cloning of a protein kinase gene conferring disease resistance in tomato, Science, 1993, 262(5138): 1432-1436.
89. McDowell J.M., Dangl J.L. Signal transduction in the plant immune response. Trends Biochem Sci, 2000, 25 (2): 79-82.
90. McDowell J.M., Woffenden B.J. Plant disease resistance genes: recent insights and potential applications. Trends in Biotechnology, 2003, 21: 178-183.
91. Mehar H.A simple procedure for the isolation of high quality RNA from ripening banana fruit. Plant Molecular Biology Reporter.2000, 18:109-115.
92. Meyers B.C., Dickerman A.W., Michelmore R.W., Sivaramakrishnan S., Sobral B.W., Young N.D. Plant disease resistance genes encode members of an ancient and diverse protein family within the nucleotide-binding superfamily. Plant J., 1999, 20:317- 332.
93. Meyers B.C., Kozik A., Griego A., Kuang H.H., Michelmore R.W. Genome-wide analysis of NBS-LRR-encoding genes in Arabidopsis. Plant Cell, 2003, 15: 809-834.
94. Michelmore R.W., Meyers B.C. Clusters of resistance genes in plants evolve by divergent selection and a birth-and-death process.Genome Res., 1998, 8: 1113～1130.
95. Michelmore R.W., Paran I., Kesseli R.V. Identification of markers linked to disease-resistance genes by bulked segregant analysis: a rapid method to detect markers in specific genomic regions by using segregating populations. Proc. Natl. Acad. Sci. USA, 1991, 88:9828–9832.
96. Mindrinos M., Katagiri F., Yu G.L., and Ausubel F.M. The A. thaliana disease-resistance gene RPS2 encodes a protein containing a nucleotide-binding site and leucine-rich repeats, Cell, 1994,78(6):1089-1099.
97. Mo Tinghui, Huang Junsheng, Du Zhongjun, Xu L., Chen Yeyuan. Cloning and Characterization of One NBS-LRR like Resistance Gene Analog from Banana. Chinese Journal of Tropical Crops, 2006, 27(3): 46-50.
98. Mohler V., Klahr A., Wenzel G. A resistance gene analoge useful for targeting disease resistance genes against different pathogens on group 1S chromosomes of barley, wheat and rye, Theor. Appl. Genet, 2002, 105(2-3):364-368.
99. Moreau P, Thoquet P, Olivier J, Laterrot H, Grimsley N. Genetic mapping of Ph-2, a single locus controlling partial resistance to Phytophthora infestans in tomato. Mol. Plant-Microbe Interact., 1998, 11:259–269
100. Naess S.K., Bradeen J.M., Wielgus S.M., Haberlach G.T., McGrath J.M. and Helgeson J.P. Resistance to late blight in Solanum bulbocastanum is mapped to chromosome 8. Theor. Appl. Genet. 2000, 101: 697-704.
101. Niklaus J.Grunwald1 Wilbert G. Flier. The Biology of Phytophthora infestans at its Center of Origin. Annu. Rev. Phytopathol., 2005, 43:171–90.
102. Nimchuk Z., Rohmer L., Chang J.H., Dangl J.L. Knowing the dancer from the dance: R-gene products and their interactions with other proteins from host and pathogen. Current Opinion in Plant Biology, 2000, 4:288-294.
103. Noir S., Combes M.C., Antony F., Lashermes P. Origin, diversity and evolution of NBS-type disease-resistance gene homologues in coffee trees (Coffea L.),Mol. Genet. Genomics, 2001, 265(4): 654-662.
104. Ohmori T, Murata M, Motoyoshi F. Characterization of disease resistanc gene-like sequences in near-isogenic lines of tomato. Theoretical and Applied Genetics, 1998, 96:331-338.
105. Ori N., Eshed Y., Paran I., Presting G., Aviv D., Tanksley S., Zamir D., and Fluhr R. The I2C family from the wilt disease resistance locus I2 belongs to the nucleotide binding, leucine-rich repeat superfamily of plant resistance genes, Plant Cell, 1997, 9(4): 521-532.
106. Pan Q., Wendel J., Fluhr R. Divergent evolution of plant NBS-LRR resistance gene homologues in dicot and cereal genomes. Journal of Molecular Evolution, 2000, 50: 203-213.
107. Park TH, Gros J, Sikkema A. The late blight resistance locus Rpi-bib3 from Solanum bulbocastanum belongs to a major late blight R gene cluster on chromosome 4 of potato. Mol. Plant Microbe. Interact. 2005, 18(7):722-9.
108. Park T-H, Vleeshouwers VGAA, van der Vossen E, Visser RF, van Eck HJ. A novel resistance gene against Phytophthora infestans derived from Solanum bulbocastanum. Poster S10-S30 In: 7th International congress on plant molecular biology, ISPMP, Barcelona, 2003.
109. Parker J.E., Coleman E.A., Moores T., Dean C., Daniels M.J., and Jones J.D.G. The Arabidopsis downy mildew resistance gene RPP5 shares similarity to the Toll and Interleukon-1 receptors with N and L6, Plant Cell, 1999: 879-894.
110. Pei X. Isolation characterization and phylogenetic analysis of the resistance gene analogues(RGAs)in banana (Musa spp.).Plant Science, 2007, 172(6):1166-1174.
111. Piffanelli P, Devoto A, Schulze-Lefert P. Defence signaling pathways in cereals. Curr. Opin. Plant Biol., 1999, 2(4):295-300.
112. Quint M., Mihaljevic R., Dussle M., Xu L. Development of RGA-CAPS markers and genetic mapping of candidate genes for sugarcane mosaic virus resistance in maize, Theor. Appl. Genet., 2002, 105(2-3):355-363.
113. Radwan O., Bouzidi M.F., Vear F., Philippon. Identification of nonTIR-NBS-LRR markers linked to the Pl5/Pl8 locus for resistance to downy mildew in sunflower, Theor. Appl. Genet., 2003, 106(8):1438-1446.
114. Ramaligam J., vera Cruz C.M., Kulkreja K., Chittoor J.M. Candidate defense genes from rice, barley, and maize and their association with qualitative and quantitative resistance in rice, Mol. Plant Microbe Interact., 2003, 16(1):14-24.
115. Ren T., Qu F., Morris T.J. HRT gene function requires interaction between a NAC protein and viral capsid protein to confer resistance to turnip crinkle virus. Plant Cell, 2000, 12: 1917-1925.
116. Robert B.Meeley, Gurmukh S.Johal, Steven P. Briggs. A biochemical Phenotype for a Disease Resistance Gene of Maize. The Plant Cell, 1992, 4:71-77.
117. Rock F.L., Hardiman G., Timans J.G., Kastelein R.A., and Bazan J.F. A family of human receptors structurally related to drosophila Toll, Proc. Natl. Acad. Sci., 1998, 95(2): 588-593.
118. Rossi M, Araujo P G., Paulet F, Garsmeur O. A Genomic distribution and characterization of EST-derived resistance gene analogs (RGAs) in sugarcane. Molecular Genetics, 2003, 269(3):406-419.
119. Salmeron J.M., Oldroyd E.D.G., Rommens C.M.T., et al. Tomato Prf is a member of the leucine-rich repeat class of plant disease resistance genes and lies embedded within the Pto kinase gene cluster. Cell, 1996, 86:123-133.
120. Sessa G., D'Ascenzo M., Loh Y.T., Martin G.B. Biochemical properties of two protein kinases involved in disease resistance signaling in tomato. J. Biol. Chem., 1998, 273(25): 15860-15865.
121. Shi Z.X., Chen X.M., Line R.F., Leung H., and Wellings C.R. Development of resistance gene analog polymorphism markers for the Yr9 gene resistance to wheat stripe rust, Genome, 2001, 44(4):509-516.
122. Smilde W.D., Brigneti G., Jagger L, Perkins S, Jones JD. Solanum mochiquense chromosome Ⅳ carries a novel late blight resistance gene Rpi-moc1. Theor Appl Genet., 2005, 110(2): 252-258.
123. Song J, Bradeen JM, Naess SK. Gene RB cloned from Solanum bulbocastanum confers broad spectrum resistance to potato late blight. Proc.Natl.Acad.Sci.USA, 2003, 100(16):9128-33
124. Song W.Y., Wang G.L., Chen L.L., Kim H.S., Pi L. Y., Holsten T., Gardner J., Wang B., Zhai W. X., Zhu L.H., Fauquet C., and Ronald P. A receptor kinase-like protein encoded by the rice disease resistance gene, Xa21, Science, 1995, 270(5243): 1804-1806.
125. Stewart C.N.J., Via L.E., A rapid CTAB DNA isolation technique for RAPD fingerprint and other PCR application. Biotechniques, 1993, 14: 748-750.
126. Tao Y., Yuan F.H., Leister R.T., Ausubel R.M., Katagiri F. Mutational analysis of the Arabidopsis is nucleotide binding site-leucine-rich repeats resistance gene RPS2. Plant Cell, 2000, 12: 2541- 2554.
127. Thomas C.M., Jones D.A., Parniske M. Characterization of the tomato Cf-4 gene for resistance to Cladosporium fulvum identifies sequences that determine recognition specificity in Cf-4 and Cf-9. Plant Cell, 1997, 9:2209~2224.
128. Tian Y.Y., Fan L., Thurau T., Jung C., Cai D. The absence of TIR-type resistance gene analogues in the sugar beet (Beta vulgaris L.) genome. J. Mol. Evol., 2004, 58:40-53.
129. Torri K.U., McNellis T.W., and Deng X.W. Functional dissection of Arabidopsis COP1 reveals specific roles of its three structural modules in light control of seedling development, EMBO, 1998, 17(19):5577-5587.
130. Van der Biezen E.A., Jones J.D.G. Plant disease-resistance proteins and the gene-for-gene concept. Trends in Biochemical Sciences, 1998, 23:454-456.
131. Vander plank J. E. Plant disease: epidemics and control, Nature, 1963, 360:121-125.
132. Villamon F. G., Spooner D. M., Orrillo M., Mihovilovich E., Perez W., Bonierbale M. Late blight resistance linkages in a novel cross of the wild potato species Solanum paucissectum (series Piurana). Theor. Appl. Genet., 2005, 111:1201-1214.
133. Wastie R.L. Breeding for resistance. Adv. Plant Pathol. , 1991, 7:193–223.
134. Whitham S., Dinesh-Kumar S.P., Choi D., Hehl R., Corr C., and Baker B. The product of the tobacco mosaic virus resistance gene N: similarity to toll and the interleukin-1 receptor, Cell, 1994, 78(6): 1101-1115.
135. Xiao S., Ellwood S., Calis O. Broad-spectrum mildew resistance in Arabidopsis thaliana mediated by RPW8. Science, 2001, 291(5501):118-120.
136. Yoshimura S., Umehara Y., Kurata N. Identification of a YAC clone carrying the Xa-1 allele, a bacterial blight resistance gene in rice. Theor. Appl. Genet., 1996, 93:117-122.
137. Yu Y. G., Buss G., Saghaih M. A. Isolation of a super family of candidate disease-resistance gene in soybean based on a conserved nucleotide-binding site. Proc.Natl.Acad.Sci.USA, 1996, 93: 11751~11756.
138. Zhang L. P., Khan A., Nino-liu D., Foolad M. R. A molecular linkage map of tomato displaying chromosomal locations of resistance gene analogs based on a Lycopersicon esculentum × Lycopersicon hisuturn cross. Genome, 2002, 45(1):133-146.
139. Zhou T., Wang Y., Chen J.Q., Araki H., Jing Z., Jiang K., Shen J., Tian D.. Genome-wide identification of NBS genes in Japonica rice reveals significant expansion of divergent non-TIR NBS-LRR genes. Mol. Gen. Genomics., 2004, 271: 402-415.
2. 谌谋华,邓秀新. 利用同源序列法克隆柑桔抗病基因类似物及其初步分析.[硕士研究生学位论文]. 武汉:华中农业大学,2003.
3. 丁国华,秦智伟,刘宏宇,周秀艳,池春玉,王志坤. 黄瓜 NBS 类型抗病基因同源序列的克隆与分析. 园艺学报,2005,32(4):638-642.
4. 杲修杰,王正华. 植物抗病基因研究进展. 长江大学学报,2006, 3(4):205-210.
5. 郭军,屈冬玉,王晓武 .cDNA-AFLP 结合 BSA 研究马铃薯晚疫病菌小种特异无毒基因差异表达片段. 中国马铃薯, 2004, 18(1):1-3.
6. 黄萱,徐子勤,陈立余,王健.小麦 NBS-LRR 类抗病基因同源序列的分离与鉴定. 分子细胞生物学报,2006, 39(2):91-96.
7. 蒋敏华,杨清,李丽,黄先群. 马铃薯抗晚疫病转基因研究进展. 种子,2006,25(12):46-50.
8. 孔巍,方宣钧. 海岛棉 NBS-LRR 类抗病基因同源序列的克隆与定位. [硕士研究生学位论文]. 北京:中国农业科学院,2003.
9. 李春来,张怀渝,植物抗病基因同源序列(RGA)研究进展,分子植物育种,2004,2(6):
853-860.
10. 李金玉,李冠,赵惠新,王贤雷,杜钰. 植物抗病分子机制研究进展. 种子,2006, 25(2):
45-50.
11. 李先平,何云昆,赵志坚,孙茂林,隋启君. 马铃薯抗晚疫病育种研究进展. 中国马铃薯, 2001,
15(5):290-295.
12. 梁凤山,周春江,孔凡娜,王斌. 桃基因组中 R 类抗病基因同源序列的克隆与序列分析. 河北农业大学学报,2005,28(1):44-48.
13. 刘晓鹏, 姜宁, 何礼远. 马铃薯晚疫病菌基因文库的构建. 东北农业大学学报, 1998, 29 (3)::228-235.
14. 卢志国,韩建民,董金皋. 植物抗病基因研究现状. 河北农业大学学报,2002,25:160-163.
15. 马媛媛,甘睿,王宁宁. 植物富含亮氨酸重复序列型受体蛋白激酶的生物学功能. 植物生理与分子生物学学报. 2005, 31(4):331-339.
16. 盛刚,李付广,王涛. 海岛棉 NBS-LRR 抗性基因类似序列的分离及 Gbarvin 基因的鉴定.[硕士研究生学位论文]. 北京:中国农业科学院,2005.
17. 孙雁,王云月,何月秋,朱有勇. 水稻抗病基因同源序列多态性与品种鉴定,种子,2001,114(2):1-2,6.
18. 田振东,柳俊,谢从华. cDNA 文库与 RACE 方法结合克隆一个马铃薯病程相关蛋白基因cDNA. 遗传学报,2003, 30(11):96-1002.
19. 万里红,周亦华,陈正华. 植物防御系统中抗病相关基因的研究进展. 遗传,2002, 24(4):486-492.
20. 王海燕,刘大群. TcLr35 小麦抗病相关基因的克隆及分析.[博士学位论文]. 保定:河北农业大学,2006.
21. 王友红,张鹏飞,陈建群.植物抗病基因及其作用机理. 植物学通报,2005,22(1):92-99.
22. 徐兵强,杜中军,黄俊生. RGA 法克隆候选抗病基因的研究进展. 分子植物育种,2004,2(3):421-423.
23. 徐建飞,辛翠花, 黄三文. 利用田间抗病基因近等混合系防治马铃薯晚疫病. 中国农业科技导报,2006, 8(4):8-13.
24. 许丽,李玥莹,林凤. 植物抗病的分子基础与研究进展. 杂粮作物,2006, 26(6):428-432.
25. 杨秀红,陈庆山,杨庆凯,李文滨. 大豆 NBS 类抗病相关基因的克隆与序列分析. 高技术通讯,2006,5(2):71-78.
26. 俞志华,祝水金,夏英武. 从植物抗病基因的克隆看其基因结构、功能和进化. 浙江大学学报,2002,28(1):107-113.
27. 左示敏,殷跃军,张亚芳,陈宗祥,潘学彪. 植物数量抗病基因克隆及其抗性机制的研究进展. 分子植物育种,2006,4(5):603-613.
28. Aarts M.G.M. te Lintel Hekkert B., Holub E.B., Beynon J.L. Identification of R-gene homologous DNA fragments genetically linked to disease resistance loci in Arabidopsis thaliana. Mol. Plant Microbe Interact., 1998, 11(4):251-258.
29. Alber T. Structure of the leucine zipper. Curr. Opin. Genet. Dev., 1992, 2: 205-210.
30. Ananieva K.I. Effect of methyl ester of jasmonic acid and benzyl-aminopurine on growth and protein profile of excised cotyledons of Cucurbita pepo. Zucchini. Biologia, lantarium., 1999,42: 549-557.
31. Anderson P.A., Lawrence G.J., Morrish B.C., Aylifee M.A., Finnegan E.J., and Ellis J.G. Inactivation of the flax rust resistance gene M associated with loss of a repeated unit within the leucine-rich repeat coding region. Plant Cell, 1997, 9(4): 641-651.
32. Ballvora A, Ercolano MR, WeiB J, Meksem K, Bormann CA, Oberhagemann P, Salamini F, Gebhardt C .The R1 gene for potato resistance to late blight (Phytophthora infestans) belongs to the leucine zipper/NBS/LRR class of plant resistance genes. Plant Journal, 2002, 30: 361–371.
33. Bendahmane A, Kanyuka K, Baulcombe DC. The Rx gene from potato controls separate virus resistance and cell death response. Plant Cell, 1999, 11:781–791.
34. Bent A.F., Kunkel B.N., Dahlbeck D., et al. RPS2 of Arabidopsis thaliana: a leucine-rich repeat class of plant disease resistance genes. Science, 1994, 265:1856-1860.
35. Buschges R., Hollricher K., Panstruga R., Simons G. Wolter M., Frijters A., van Daelen R., and Schulze-Lefert. The barley Mlo gene: a novel control element of plant pathogen resistance. Cell, 1997, 88(5): 695-705.
36. Byrne P.F., McMullen M.D., Snook M.E., Musket T.A. Quantitative trait loci and metabolic pathways: genetic control of the concentration of maysin, a corn earworm resistance factor in maize silks. Proc.Natl.Acad.Sci.USA, 1996, 93(17):8820-8825.
37. Cai D., Kleine M., Kifle S., Harloff H.J., Sandal N.N., Marcker K.A., Klein-Lankhorst R.M., Salentijn E.M.J., Lange W. Positional cloning of a gene for nematode resistance in sugar beet. Science, 1997, 275(5301):832-834.
38. Catherine F, Gabrieles S, Bent K. Molecular cloning of a new receptor-like kinase gene encoded the Lr10 disease resistance locus wheat. The Plant Journal, 1997, 11 (1): 45-52.
39. Chen X.M., Line R.F., Leung H. Genome scanning for resistance-gene analogs in rice, barley, and wheat by high-resolution electrophoresis. Theor. Appl. Genet., 1998, 97(3):345-355.
40. Chunwongse J., Chunwongse C., Black L., Hanson P. Molecular mapping of the Ph-3 gene for late blight resistance in tomato. J Hort. Sco&Biotechnol., 2002, 77:281-286.
41. Collins N C, W ebb C A, Seah S, Pryor. The isolation and mapping of disease resistance gene analogs in maize. Mol. Plant Microbe. Interact. , 1998, 11: 968-978.
42. Dangl, J.L and Jones, J.D.G. Plant pathogens and integrated defence responses to infection. Nature, 2001, 411: 826-833.
43. David A.Jones, Clowyn M. Thomas, Kim. E. Hammond Kosack. Isolation of the tomato Cf-9 gene for resistance Cladosporium fulvum by transposon tagging. Science, 1994, 266: 789-793.
44. De Jong W., Forsyth A., Lister D., Hebhardt C. and Baulcombe D.C. A potato hypersensitive resistance gene cluster on chromosome Ⅴ. Theor. Appl. Genet., 1995: 153-162.
45. Dilbirligi M., Erayman M., Sandhu D. Identification of wheat chromosomal regions containing expressed resistance genes. Genetics, 2004, 166: 461- 481.
46. Dixon M.S., Hatzixanthis K., Jones D.A. The tomato Cf-5 disease resistance gene and six homologs show pronounced allelic variation in leucine-rich repeat copy number. Plant Cell,1998,10:1915～1925.
47. Donald T.M., Pellerone F., Adam-Blondon A.F., Bouquet A., Thomas M.R., and Dry I.B. Identification of resistance gene analogs linked to a powdery mildew resistance locus in grapevine. Theor. Appl. Genet., 2002,104 (4):610-618.
48. Edwin A.G. van der Vossen, Anne Sikkema, Bas te Lintel Hekkert. An ancient R gene from the wild potato species Solanum bulbocastanum confers broad-spectrum resistance to Phytophthora infestans in cultivated potato and tomato. The Plant Journal, 2003, 36 (6): 867-882.
49. Edwin A.G. van der Vossen, Jack Gros, Anne Sikkema. The Rpi-blb2 gene from Solanum bulbocastanum is an Mi-1 gene homolog conferring broad-spectrum late blight resistance in potato. The Plant Journal, 2005, 44(2):208-222.
50. Edwin A.G. van der Vossen, van der Voort J.N., Kanyuka K., Bendahmane A., Sandbrink H., Baulcombe D.C., Bakker J., Stiekema W. J. and Klein-Lankhorst R. M. Homologues of a single resistance-gene cluster in potato confer resistance to distinct pathogens: A virus and a nematode. Plant J. 2000, 23:567-576.
51. El-Kharbotly A, Palomino-Sanchez C, Salamini F, Jacobsen E, Gebhardt C. R6 and R7 alleles of potato conferring race-specific resistance to Phytophthora infestans (Mont.) de Bary identified genetic loci clustering with the R3 locus on chromosome Ⅺ. Theor. Appl. Genet., 1996,92:880–884.
52. Ellis J.G.,Doods P, Pryor T. The generation of plant disease resistance specificities. Trends in Plant Science, 2000, 5: 373-379.
53. Ellis, J.G., Lawrence, G.J., Luck, J.E. and Dodds, P.N. Identification of regions in alleles of the flax rust resistance gene L that determine differences in gene-for-gene specificity. Plant Cell, 1999, 11: 495–506.
54. Ewing, E.E., Simko, I., Smart, C.D., Bonierbale, M. W., Mizubuti, E. S. G., May, G. D., and Fry, W. E. Genetic mapping of qualitative and quantitative field resistance to Phytophthora infestans in a population derived from Solanum tuberosum and Solanum Berthaulti. Molecular Breeding, 2000, 6:25-36.
55. Feuillet C, Schachermagr G, Keller B. Molecular cloning of a new receptor-like kinase gene encoding the Lr10 disease resistance locus of wheat. The Plant Journal, 1997, 11 (1):45-52.
56. Flandez-Galvez H., Ford R., Pang E.C Taylor P.W. An intraspecific linkage map of the chickpea (Cicer arietinum L.) genomes based on sequence tagged microsatellite site and resistance gene analog markers, Theor. Appl. Genet., 2003, 106(8):1447-1456.
57. Flor H H. Current Status of the Gene- for - gene Concept. Annual Review of Phytopathology, 1971, 9: 275-296.
58. Fourmann M., Chariot F., Froger N., Delourme R., and Brunel D. Expression, mapping, and genetic variability of Brassica napus disease resistance gene analogues, Genome, 2001, 44(6):1083-1099.
59. Fry, W.E. and Goodwin, S.B. Resurgence of the Irish potato famine fungus. Bioscience, 1997, 47:363–371.
60. Rauscher G. M., Smart C. D., Simko I. Characterization and mapping of RPi-ber, a novel potato late blight resistance gene from Solanum berthaultii. Theor. Appl. Genet., 2006, 112: 674-687.
61. Gebhardt C., Valkonen J.P.T. Organization of genes controlling disease resistance in the potato genome. Annual Review of Phytopathology, 2001, 39:79–102.
62. Graham M.A., Marek L.F. Shoemaker R.C. Organization, expression and evolution of a disease resistance gene cluster in soybean, Genetics, 2002, 162(4):1961-1977.
63. Grant M.R., Godiard L., Straube E., Ashfield T., Lewald J., Sattler A., Innes R.W., and Dangl J.L. Structure of the Arabidopsis RPM1 gene enabling dual specificity disease resistance, Science, 1995, 269: 843-846.
64. Halterman D.A., Kramer L.C., Wielgus S., Jiang J. Performance of transgenic potato containing the late blight resistance gene RB. Plant Dis., 2008, 92: 339-343.
65. Hammond-Kosack K. E. Plant disease resistance genes. Annual Review of Plant Physiology and Plant Molecular Biology, 1997, 48: 575-607.
66. Hanhui Kuang, Fusheng Wei, Maria Rosa Marano, Uwe Wirtz, Xiaoxue Wang. The R1 resistance gene cluster contains three groups of independently evolving, type I R1 homologues and shows substantial structural variation among haplotypes of Solanum demissum. The Plant J., 2005,44:37-50.
67. Huang Sanwei, Edwin A.G.van der Vossen, Hanhui Kuang .Comparative genomics enabled the isolation of the R3a late blight resistance gene in potato. The Plant J., 2005, 42: 251-261.
68. Huang Sanwei, Vivianne G.A.A. Vleeshouwers, Jeroen S. Werij, Ronald C.B. Hutten, Herman J. van Eck, Richard G.F.Visser, and Evert Jacobsen. The R3 resistance to Phytophthora infestans in potato is conferred by two closely linked R genes with distinct specificities. Mol. Plant Micro. Inter., 2004, 17(4): 428-435.
69. Hulbert, S.H., Webb, C.A., Smith, S.M. and Sun, Q. Resistance gene complexes: evolution and utilization. Annu. Rev. Phytopathol., 2001, 39, 285–312.
70. Hunger S., di Gaspero G., Mohring S. Isolation and linkage analysis of expressed disease- resistance gene analogues of sugar beet( Beta vulgaris L.) Genome, 2003, 46(1):70-82.
71. Jia Y., McAdams S.A., Bryan G.T., et al. Direct interaction of resistance gene and a virulence gene product confers rice blast resistance. EMBO J, 2000, 19: 4004-4014.
72. Johal G. S., and Briggs S.P. Reductase activity encoded by the HM1 disease resistance gene in maize, Science, 1992, 258(5084):985-987.
73. John E. Bradshaw, Glenn J. Bryan, Alison K. Lees. Mapping the R10 and R11 genes for resistance to late blight (Phytophthora infestans) present in the potato (Solanum tuberosum) R-gene differentials of Black. Theor. Appl. Genet., 2006, 112(4):744-751.
74. Johnaton S.A., den Nijs, T.P.M. Peloquin S.J. and Hanneman R.E. The significance of genic balance to endosperm development in interspecific crosses. Theor. Appl. Genet., 1980, 57: 5-9.
75. Jones D.A., Thomas C.M., Hammond-Kosack K.E., Balint-Kurti P.J., and Jones J.D. Isolation of the tomato Cf-9 gene for resistance to Cladosporium fulvum by transposon tagging. Science, 1994, 165(5186):789-793.
76. Kobe B, Deisenhofer J. The leucine-rich repeat: a versatile binding motif, Trends. Biochem. Sci., 1994, 19(10):415-421.
77. Kuhl JC, Hanneman RE Jr, Havey MJ. Characterization and mapping of Rpi1, a late-blight resistance locus from diploid (1EBN) Mexican Solanum pinnatisectum. Mol. Genet. Genomics, 2001, 265(6):977-85.
78. McHale L., Tan X. P., Koehl P., Michelmore R.W. Plant NBS-LRR proteins: adaptable guards. Genome Biology, 2006, 7:212.
79. Lawrence G.J., Finnegan E.J., Ayliffe M.A., and Eliis J.G. The L6 gene for flax rust resistance is related to the Arabidopsis bacterial resistance gene RPS2 and the tobacco viral resistance gene N, Plant Cell, 1995, 7(8):1195-1206.
80. Lee S.Y., Seo J.S., Rodriguez-Lanetty M., Lee D.H. Comparative analysis of superfamilies of NBS-encoding disease resistance gene analogs in cultivated and wild apple specis, Mol. Genet.Genomics, 2003, 269 (1):101-108.
81. Leister D, Ballvora A, Salamini F, et al. A PCR-based approach for isolating pathogen resistance genes from potato with potential for wide application in plants. Proc. Natl. Acad. Sci. USA., 1996,14:421~429.
82. Leister D, Kurth T, Schulze P, RFLP and physical mapping of resistance gene homologues in rice and Barley. Theoretical and Applied Genetics, 1999, 98:509-520.
83. Leister D. Tandem and segmental gene duplication and recombination in the evolution of plant disease resistance gene. Trends in Genetics, 2004, 3: 116-122.
84. Leonards-Schippers C, Gieffers W, Schafer-Pregl R, Ritter E, Knapp S, Salamini F, Gebhardt C. Quantitative resistance to Phytophthora infestans in potato: a case study for QTL mapping in an allogamous plant species. Genetics, 1994, 137:67–77.
85. Li X, van Eck HJ, Rouppe van der Voort JNAM, Huigen DJ, Stam P, Jacobsen E. Autotetraploids and genetic mapping using common AFLP markers: the R2 allele conferring resistance to Phytophthora infestans mapped on potato chromosome 4. Theor Appl Genet., 1998, 96:1121–1128.
86. Madsen L.H., Collins N.C., Rakwalska M., Backes G., Sandal N., Krusell L., Jensen J., Waterman E.H., Jahoor A., Ayliffe M., Pryor A.J. Langridge P., Schulze-Lefert P., and Stougaard J. Barley disease resistance gene analogs of the NBS-LRR class: identification and mapping, Mol. Genet. Genomics, 2003, 269(1):150-161.
87. Mark S., Dixon, Jones D.A., Keddie J.S.The tomato Cf-2 disease resistance locus comprises two functional genes encoding leucine-rich repeat proteins.Cell,1996,84:451～459.
88. Martin G.B., Brommonschenkel S.H.Chunwongse J., Frary A., Ganal M.W., Spivey R., Wu T., Earle E.D. and Tanksley S.D., , Map-based cloning of a protein kinase gene conferring disease resistance in tomato, Science, 1993, 262(5138): 1432-1436.
89. McDowell J.M., Dangl J.L. Signal transduction in the plant immune response. Trends Biochem Sci, 2000, 25 (2): 79-82.
90. McDowell J.M., Woffenden B.J. Plant disease resistance genes: recent insights and potential applications. Trends in Biotechnology, 2003, 21: 178-183.
91. Mehar H.A simple procedure for the isolation of high quality RNA from ripening banana fruit. Plant Molecular Biology Reporter.2000, 18:109-115.
92. Meyers B.C., Dickerman A.W., Michelmore R.W., Sivaramakrishnan S., Sobral B.W., Young N.D. Plant disease resistance genes encode members of an ancient and diverse protein family within the nucleotide-binding superfamily. Plant J., 1999, 20:317- 332.
93. Meyers B.C., Kozik A., Griego A., Kuang H.H., Michelmore R.W. Genome-wide analysis of NBS-LRR-encoding genes in Arabidopsis. Plant Cell, 2003, 15: 809-834.
94. Michelmore R.W., Meyers B.C. Clusters of resistance genes in plants evolve by divergent selection and a birth-and-death process.Genome Res., 1998, 8: 1113～1130.
95. Michelmore R.W., Paran I., Kesseli R.V. Identification of markers linked to disease-resistance genes by bulked segregant analysis: a rapid method to detect markers in specific genomic regions by using segregating populations. Proc. Natl. Acad. Sci. USA, 1991, 88:9828–9832.
96. Mindrinos M., Katagiri F., Yu G.L., and Ausubel F.M. The A. thaliana disease-resistance gene RPS2 encodes a protein containing a nucleotide-binding site and leucine-rich repeats, Cell, 1994,78(6):1089-1099.
97. Mo Tinghui, Huang Junsheng, Du Zhongjun, Xu L., Chen Yeyuan. Cloning and Characterization of One NBS-LRR like Resistance Gene Analog from Banana. Chinese Journal of Tropical Crops, 2006, 27(3): 46-50.
98. Mohler V., Klahr A., Wenzel G. A resistance gene analoge useful for targeting disease resistance genes against different pathogens on group 1S chromosomes of barley, wheat and rye, Theor. Appl. Genet, 2002, 105(2-3):364-368.
99. Moreau P, Thoquet P, Olivier J, Laterrot H, Grimsley N. Genetic mapping of Ph-2, a single locus controlling partial resistance to Phytophthora infestans in tomato. Mol. Plant-Microbe Interact., 1998, 11:259–269
100. Naess S.K., Bradeen J.M., Wielgus S.M., Haberlach G.T., McGrath J.M. and Helgeson J.P. Resistance to late blight in Solanum bulbocastanum is mapped to chromosome 8. Theor. Appl. Genet. 2000, 101: 697-704.
101. Niklaus J.Grunwald1 Wilbert G. Flier. The Biology of Phytophthora infestans at its Center of Origin. Annu. Rev. Phytopathol., 2005, 43:171–90.
102. Nimchuk Z., Rohmer L., Chang J.H., Dangl J.L. Knowing the dancer from the dance: R-gene products and their interactions with other proteins from host and pathogen. Current Opinion in Plant Biology, 2000, 4:288-294.
103. Noir S., Combes M.C., Antony F., Lashermes P. Origin, diversity and evolution of NBS-type disease-resistance gene homologues in coffee trees (Coffea L.),Mol. Genet. Genomics, 2001, 265(4): 654-662.
104. Ohmori T, Murata M, Motoyoshi F. Characterization of disease resistanc gene-like sequences in near-isogenic lines of tomato. Theoretical and Applied Genetics, 1998, 96:331-338.
105. Ori N., Eshed Y., Paran I., Presting G., Aviv D., Tanksley S., Zamir D., and Fluhr R. The I2C family from the wilt disease resistance locus I2 belongs to the nucleotide binding, leucine-rich repeat superfamily of plant resistance genes, Plant Cell, 1997, 9(4): 521-532.
106. Pan Q., Wendel J., Fluhr R. Divergent evolution of plant NBS-LRR resistance gene homologues in dicot and cereal genomes. Journal of Molecular Evolution, 2000, 50: 203-213.
107. Park TH, Gros J, Sikkema A. The late blight resistance locus Rpi-bib3 from Solanum bulbocastanum belongs to a major late blight R gene cluster on chromosome 4 of potato. Mol. Plant Microbe. Interact. 2005, 18(7):722-9.
108. Park T-H, Vleeshouwers VGAA, van der Vossen E, Visser RF, van Eck HJ. A novel resistance gene against Phytophthora infestans derived from Solanum bulbocastanum. Poster S10-S30 In: 7th International congress on plant molecular biology, ISPMP, Barcelona, 2003.
109. Parker J.E., Coleman E.A., Moores T., Dean C., Daniels M.J., and Jones J.D.G. The Arabidopsis downy mildew resistance gene RPP5 shares similarity to the Toll and Interleukon-1 receptors with N and L6, Plant Cell, 1999: 879-894.
110. Pei X. Isolation characterization and phylogenetic analysis of the resistance gene analogues(RGAs)in banana (Musa spp.).Plant Science, 2007, 172(6):1166-1174.
111. Piffanelli P, Devoto A, Schulze-Lefert P. Defence signaling pathways in cereals. Curr. Opin. Plant Biol., 1999, 2(4):295-300.
112. Quint M., Mihaljevic R., Dussle M., Xu L. Development of RGA-CAPS markers and genetic mapping of candidate genes for sugarcane mosaic virus resistance in maize, Theor. Appl. Genet., 2002, 105(2-3):355-363.
113. Radwan O., Bouzidi M.F., Vear F., Philippon. Identification of nonTIR-NBS-LRR markers linked to the Pl5/Pl8 locus for resistance to downy mildew in sunflower, Theor. Appl. Genet., 2003, 106(8):1438-1446.
114. Ramaligam J., vera Cruz C.M., Kulkreja K., Chittoor J.M. Candidate defense genes from rice, barley, and maize and their association with qualitative and quantitative resistance in rice, Mol. Plant Microbe Interact., 2003, 16(1):14-24.
115. Ren T., Qu F., Morris T.J. HRT gene function requires interaction between a NAC protein and viral capsid protein to confer resistance to turnip crinkle virus. Plant Cell, 2000, 12: 1917-1925.
116. Robert B.Meeley, Gurmukh S.Johal, Steven P. Briggs. A biochemical Phenotype for a Disease Resistance Gene of Maize. The Plant Cell, 1992, 4:71-77.
117. Rock F.L., Hardiman G., Timans J.G., Kastelein R.A., and Bazan J.F. A family of human receptors structurally related to drosophila Toll, Proc. Natl. Acad. Sci., 1998, 95(2): 588-593.
118. Rossi M, Araujo P G., Paulet F, Garsmeur O. A Genomic distribution and characterization of EST-derived resistance gene analogs (RGAs) in sugarcane. Molecular Genetics, 2003, 269(3):406-419.
119. Salmeron J.M., Oldroyd E.D.G., Rommens C.M.T., et al. Tomato Prf is a member of the leucine-rich repeat class of plant disease resistance genes and lies embedded within the Pto kinase gene cluster. Cell, 1996, 86:123-133.
120. Sessa G., D'Ascenzo M., Loh Y.T., Martin G.B. Biochemical properties of two protein kinases involved in disease resistance signaling in tomato. J. Biol. Chem., 1998, 273(25): 15860-15865.
121. Shi Z.X., Chen X.M., Line R.F., Leung H., and Wellings C.R. Development of resistance gene analog polymorphism markers for the Yr9 gene resistance to wheat stripe rust, Genome, 2001, 44(4):509-516.
122. Smilde W.D., Brigneti G., Jagger L, Perkins S, Jones JD. Solanum mochiquense chromosome Ⅳ carries a novel late blight resistance gene Rpi-moc1. Theor Appl Genet., 2005, 110(2): 252-258.
123. Song J, Bradeen JM, Naess SK. Gene RB cloned from Solanum bulbocastanum confers broad spectrum resistance to potato late blight. Proc.Natl.Acad.Sci.USA, 2003, 100(16):9128-33
124. Song W.Y., Wang G.L., Chen L.L., Kim H.S., Pi L. Y., Holsten T., Gardner J., Wang B., Zhai W. X., Zhu L.H., Fauquet C., and Ronald P. A receptor kinase-like protein encoded by the rice disease resistance gene, Xa21, Science, 1995, 270(5243): 1804-1806.
125. Stewart C.N.J., Via L.E., A rapid CTAB DNA isolation technique for RAPD fingerprint and other PCR application. Biotechniques, 1993, 14: 748-750.
126. Tao Y., Yuan F.H., Leister R.T., Ausubel R.M., Katagiri F. Mutational analysis of the Arabidopsis is nucleotide binding site-leucine-rich repeats resistance gene RPS2. Plant Cell, 2000, 12: 2541- 2554.
127. Thomas C.M., Jones D.A., Parniske M. Characterization of the tomato Cf-4 gene for resistance to Cladosporium fulvum identifies sequences that determine recognition specificity in Cf-4 and Cf-9. Plant Cell, 1997, 9:2209~2224.
128. Tian Y.Y., Fan L., Thurau T., Jung C., Cai D. The absence of TIR-type resistance gene analogues in the sugar beet (Beta vulgaris L.) genome. J. Mol. Evol., 2004, 58:40-53.
129. Torri K.U., McNellis T.W., and Deng X.W. Functional dissection of Arabidopsis COP1 reveals specific roles of its three structural modules in light control of seedling development, EMBO, 1998, 17(19):5577-5587.
130. Van der Biezen E.A., Jones J.D.G. Plant disease-resistance proteins and the gene-for-gene concept. Trends in Biochemical Sciences, 1998, 23:454-456.
131. Vander plank J. E. Plant disease: epidemics and control, Nature, 1963, 360:121-125.
132. Villamon F. G., Spooner D. M., Orrillo M., Mihovilovich E., Perez W., Bonierbale M. Late blight resistance linkages in a novel cross of the wild potato species Solanum paucissectum (series Piurana). Theor. Appl. Genet., 2005, 111:1201-1214.
133. Wastie R.L. Breeding for resistance. Adv. Plant Pathol. , 1991, 7:193–223.
134. Whitham S., Dinesh-Kumar S.P., Choi D., Hehl R., Corr C., and Baker B. The product of the tobacco mosaic virus resistance gene N: similarity to toll and the interleukin-1 receptor, Cell, 1994, 78(6): 1101-1115.
135. Xiao S., Ellwood S., Calis O. Broad-spectrum mildew resistance in Arabidopsis thaliana mediated by RPW8. Science, 2001, 291(5501):118-120.
136. Yoshimura S., Umehara Y., Kurata N. Identification of a YAC clone carrying the Xa-1 allele, a bacterial blight resistance gene in rice. Theor. Appl. Genet., 1996, 93:117-122.
137. Yu Y. G., Buss G., Saghaih M. A. Isolation of a super family of candidate disease-resistance gene in soybean based on a conserved nucleotide-binding site. Proc.Natl.Acad.Sci.USA, 1996, 93: 11751~11756.
138. Zhang L. P., Khan A., Nino-liu D., Foolad M. R. A molecular linkage map of tomato displaying chromosomal locations of resistance gene analogs based on a Lycopersicon esculentum × Lycopersicon hisuturn cross. Genome, 2002, 45(1):133-146.
139. Zhou T., Wang Y., Chen J.Q., Araki H., Jing Z., Jiang K., Shen J., Tian D.. Genome-wide identification of NBS genes in Japonica rice reveals significant expansion of divergent non-TIR NBS-LRR genes. Mol. Gen. Genomics., 2004, 271: 402-415.