水稻基因聚合育种及cry1Ab基因在转基因水稻中的分子标记定位研究
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摘要
水稻是世界上最重要的粮食作物之一,全球约二分之一的人口以稻米为主食,水稻也是受病虫害最多的作物之一。由Xanthomonas oryzae pv. oryzae引起的水稻白叶枯病是世界范围的细菌性病害,严重影响水稻的产量和品质。水稻每年因鳞翅目害虫(二化螟、三化螟、稻纵卷叶螟、稻螟蛉、稻苞虫等)为害造成的损失占总产量的10%-30%。另外,杂交稻纯度和稻田杂草影响水稻的产量与品质,使用除草剂费工费力,增加了生产成本,已成为生产上的主要问题。抗病、抗虫、抗除草剂转基因水稻的培育为水稻的高产与稳产提供重要保证的同时,可以减少化学农药的使用,改善环境质量。实践证明,将多个抗性基因聚合在同一水稻品种中能有效提高水稻的抗性和保持抗性持久,分子标记辅助选择以其快速、准确的选择优势成为实现基因聚合育种的有效手段。
     对转基因水稻中外源转cry1Ab基因进行定位,构建转cry1Ab基因水稻的分子遗传图谱,可为转cry1Ab基因水稻的培育进行标记辅助选择提供更多的分子标记,同时为研究转cry1Ab基因水稻的基因组变化,进行转基因水稻中外源基因的位置效应奠定基础,为转cry1Ab基因水稻生物安全性评价提供更多的理论支撑。
     本论文主要开展了三个方面的研究:多抗性(抗白叶枯、螟虫、除草剂)转基因水稻恢复系的培育;水稻香味基因与抗白叶枯基因的聚合育种;转基因水稻中转cry1Ab基因的分子标记定位。主要研究结果如下:
     1.多抗性(抗白叶枯、螟虫、除草剂)转基因水稻恢复系的培育
     (1)将恢773与携有转cry1Ab和bar基因的水稻新品系中国91(cry1Ab)杂交,并以恢773作为轮回亲本,连续回交三代和自交一代,对各回交世代的单株进行cry1Ab和bar基因的PCR分析、田间抗虫性鉴定和除草剂Basta抗性鉴定,经过选择获得了转基因水稻恢复系T773-1,田间种植对螟虫及除草剂Basta表现很好的抗性。
     (2)将恢773与带有转Xa21基因的水稻新品系豫粳6(Xa21)杂交,并以恢773作为轮回亲本,连续回交三代和自交一代,经过PCR分析、GUS活性测定和白叶枯病菌接种鉴定,获得带有Xa21基因的并具有优良农艺性状的转基因水稻恢复系T773-2,田间种植表现很好的水稻白叶枯病抗性。
     (3)转cry1Ab和bar基因的水稻恢复系T773-1和转Xa21基因的恢复系T773-2进行杂交、自交,对自交世代单株进行cry1Ab、bar和Xa21基因的分子标记辅助选择,获得了带有三基因的恢复系T773,田间种植对水稻白叶枯病、鳞翅目害虫、除草剂具有很好的抗性。并利用恢复系T773与不育系早花2A进行杂交,进行配合力测定,获得了抗白叶枯病、螟虫与除草剂的多抗转基因水稻恢复系T773。T773及其杂交后代的抗谱和抗性水平达到其抗性基因供体亲本的水平,抗性改良效果明显。
     2.水稻香味基因与抗白叶枯基因的分子标记聚合育种
     通过分子标记辅助选择与传统的杂交育种技术有机结合,将香稻栽培品种香粳9407与抗白叶枯病多基因聚合系IRBB60(Xa4、xa5、xa13、Xa21)进行杂交、自交,采用田间病原菌接种鉴定和香味鉴定筛选,结合农艺性状考查进行选择,在自交F2代获得两个抗白叶枯基因和香味基因聚合系(2R+fgr)1株;在自交F3代获得四个抗白叶枯基因和香味基因聚合系(4R+fgr)1株,三个抗白叶枯基因和香味基因聚合系(3R+fgr)2株,两个抗白叶枯基因和香味基因聚合系(2R+fgr)7株,一个抗白叶枯基因和香味基因聚合系(1R+fgr)3株;在自交F4代获得四个抗白叶枯基因和香味基因聚合系(4R+fgr)10株,三个抗白叶枯基因和香味基因聚合系(3R+fgr)28株,两个抗白叶枯基因和香味基因聚合系(2R+fgr)20株。获得的多基因聚合系单株高抗水稻白叶枯病并且具有浓郁的香味,主要农艺性状优良,可直接应用于抗白叶枯病香稻育种,对水稻抗性育种和品质改良具有重要的实践意义。
     3.转基因水稻中转cry1Ab基因的分子标记定位
     (1)首先利用纯合稳定的转cry1Ab基因的中国91株系与IRBB60杂交,对其F2代分离群体的单株进行除草剂Basta抗性检测,结果显示,在F2代分离群体中cry1Ab基因的分离符合期望的3:1分离比例,遵循孟德尔分离规律,证明了cry1Ab基因在转基因株系中以单显性位点整合。
     (2)利用Tail-PCR技术获得转基因水稻中外源cry1Ab基因插入的侧翼序列,通过序列比较分析确定了转cry1Ab基因在水稻基因组插入的物理位置。根据插入位置附近的水稻基因组序列进行微卫星序列的搜索,并利用Primer Premier 5.0软件设计了24对SSR标记引物。
     (3)利用设计的24对SSR引物对亲本转cry1Ab基因的水稻中国91与IRBB60之间进行多态性筛选,获得了6对多态性引物,然后利用这6对引物以及公开引物RM1261对杂交的F2代分离群体240个单株进行分析。依据每个单株的基因型和表型数据,利用MAPMAKER/EXP Version 3.0软件进行分析,构建了外源转基因cry1Ab在水稻第12染色体的遗传图谱,外源cry1Ab基因被定位在标记CM22和RM1262之间,两个标记相距9.8cM。微卫星标记CM22、CM01、CM12、CM13位于cry1Ab基因的一侧,与cry1Ab基因的遗传距离分别为4.6cM、4.2cM、3.1cM和2.7cM;微卫星标记CM24、CM10和RM1261位于另一侧,与cry1Ab的遗传距离分别为3.1cM、3.5cM和5.2cM。
Rice is one of the most important crops in the world, providing staple food for more than half of the world’s population, but also one crop tends to be infested by many kinds of diseases and pests. Bacterial blight caused by Xanthomonas oryzae pv. oryzae, a bacterial disease of rice worldwide, affects the yield and quality of rice seriously. The losses in rice yield caused by lepidopteran insects (Chilo suppressalis walker, Tryporyza incertulas, Cnaphalocrocis medinalis Guenee, Naranga aenescens Moore, Borbo cinnara Wallace) can account for 10-30% of total yield. In addition, purity of hybrid rice and field weeds affect the yield and quality of rice,the use of herbicide to control field weeds increases the production cost, these have been the main problems in rice production. So breeding of transgenic rice with multiple resistance to diseases, insects and herbicide can ensure the quantity and quality of rice, decrease the use of pesticides and be beneficial to the environment. Many studies had indicated that pyramiding multiple resistant genes into one rice variety can significantly enhance the resistance and keep durable resistance of rice. Molecular marker-assisted selection (MAS) has been advocated as a highly efficient breeding method for pyramiding multiple resistant genes, because it can offer rapid and precise selection of the target gene.
     Molecular mapping of transgene cry1Ab and genetic linkage map construction of transgenic cry1Ab rice increases molecular markers used for marker-assisted selection in transgenic cry1Ab rice breeding. Furthermore, this research establishes the foundation to investigate the transgene location affection, and provides the theory theoretic support for transgenic rice penalty evaluation.
     The objective of this paper was to study from three aspects, respectively: Breeding of transgenic rice restorer line for multiple resistance against bacterial blight, striped stem borer and herbicide; Marker-assisted breeding of pyramiding rice fragrance gene and multiple bacterial blight resistance genes; Molecular mapping of transgene cry1Ab in transgenic rice. Main research results are as follows:
     1. Breeding of transgenic rice restorer line for multiple resistance against bacterial blight, striped stem borer and herbicide
     (1) Transgenic rice line Zhongguo91 (carrying cry1Ab and bar gene) and elite restorer line Hui773 were crossed, then backcross was carried out three times using Hui773 as recurrent parent and self-cross was carried out one time. All plants were subjected to Basta painting and PCR analysis to verify presence of the cry1Ab and the bar genes. Through selection we obtained transgenic restorer line T773-1 with good agronomic traits and obvious resistance to striped stem borer and herbicide.
     (2) Transgenic rice line Yujing6 (carrying Xa21 gene) and elite restorer line Hui773, then backcross was carried out three times using Hui773 as recurrent parent and self-cross was carried out one time. All plants were subjected to GUS activity assay and PCR analysis to verify presence of the Xa21 gene. In BC3F2 population we obtained transgenic restorer line T773-2 with good agronomic traits, showing high resistance to bacterial blight disease under field condition.
     (3) Transgenic rice line Zhongguo91 (carrying cry1Ab and bar gene) and transgenic rice line Yujing6 (carrying Xa21 gene) were crossed. Through marker-assisted selection in self-cross populations we obtained transgenic restorer line T773 carrying three genes (cry1Ab, bar and Xa21) with resistance to bacterial blight, striped stem borer and herbicide. Transgenic restorer line T773 showed high resistance to bacterial blight disease, striped stem borer and herbicide under field condition. The hybrid F1 generation produced from the cross between transgenic restorer line T773 and a corresponding male sterile line Zaohua2A maintained obvious resistance to rice bacterial blight, rice leaffolder and striped stem borer, and showed significant heterosis.
     2. Marker-assisted breeding of pyramiding rice fragrance gene and resistance gene to bacterial blight
     Through the combination of marker-assisted selection and conventional cross breeding technology, we crossed Xiangjing9407 and transgenic pyramiding rice line IRBB60 (carrying Xa4, xa5, xa13, Xa21), screened the self-cross population using inoculation appraisal technology and fragrance identification to obtained the multiple resistance gene and fragrance gene pyramiding plant, supplemented by investigation of agronomic traits. We obtained 1 individual plant of (2R+fgr) pyramiding in F2 self-cross population. And we obtained 1 individual plant of (4R+fgr) pyramiding, 2 individual plant of (3R+fgr) pyramiding, 7 individual plant of (2R+fgr) pyramiding, 3 individual plant of (1R+fgr) pyramiding in F3 self-cross population. Finally, we obtained 10 individual plant of (4R+fgr) pyramiding, 28 individual plant of (3R+fgr) pyramiding, 20 individual plant of (2R+fgr) pyramiding in F4 self-cross population. All the plants selected had rich aroma and high resistance to bacterial blight, with excellent agronomic traits, can be used to breed aroma rice with resistance to bacterial blight. This study has important significance of rice resistance breeding and quality improvement practice.
     3. Molecular mapping of transgene cry1Ab in transgenic rice
     (1) First, the F2 segregation population plants of transgenic Zhongguo91 (cry1Ab)×IRBB60 were subjected to herbicide Basta resistance detection. The result demonstrated that the segregation of cry1Ab gene was the ratio of 3:1, following Mendelian segregation law. The result proved that the cry1Ab gene was single dominant locus integrated in transgenic rice.
     (2) The flanking sequences of cry1Ab gene insertions in transgenic rice genome were rescued by Tail-PCR. Then using sequence blast analysis on NCBI website we confirmed the cry1Ab gene insertions location in chromosome 12 of rice. Using software“simple sequence repeats identification tool”and“Primer Premier 5.0”, 24 pairs of primers were designed on the basis of the rice genome sequence close to the transgene cry1Ab insertions location. (3) Polymorphism screen using parent Zhongguo91 (cry1Ab) and IRBB60 we obtained 6 pairs of primers with polymorphism, then using the 6 pairs of primers and primer RM1261 we analyze the genotype of 240 plants in F2 segregation population. By“MAPMAKER/EXP Version 3.0”software we construct the molecular genetic linkage map of exogenous cry1Ab gene in 12 chromosome of rice genome according to their genotype and phenotype. The result indicated that exogenous cry1Ab gene is located between the SSR markers CM22 and RM1261 in 12 chromosome of rice genome. SSR marker CM22, CM01, CM12, CM13 were found to be linked to cry1Ab gene with genetics distances of 4.6, 4.2, 3.1 and 2.7 respectively, close to the short arm side; SSR marker CM24, CM10, RM1261 were found to be linked to cry1Ab gene with genetics distances of 3.1, 3.5 and 5.2 respectively, close to the other side.
引文
陈洪,朱立煌,徐吉臣,陈美玲. RAPD标记构建水稻分子连锁图.植物学报, 1995, 37(9): 677~684.
    慈晓燕,姚方印,朱常香,宋云枝,温孚江,姜明松.含Cry1Ab和Xa21基因抗病虫水稻选育研究及其田间表现.中国农业科学, 2005(38): 313~319.
    陈艳,胡军,钱韦,田颖川,赵文明,何朝族.水稻白叶枯病抗性基因Xa-min(t)的抗谱鉴定及其分子标记的筛选.自然科学进展, 2003, 13(9): 1001~1004.
    邓其明,周宇爝,蒋昭雪,万映秀,赵斌,杨莉,李平.白叶枯病抗性基因Xa21、Xa4和Xa23的聚合及其效应分析.作物学报, 2005, 31(9): 1241~1246.
    方进,翟文学,王文明,李素文,朱立煌.转基因水稻T-DNA侧翼序列的扩增与分析.遗传学报, 2001, 28(4): 345~351.
    方宣钧,吴为人,唐纪良.作物DNA标记辅助育种.北京:科学出版社, 2001. pp7~21.
    方中达,许志刚,过崇俭,殷尚智,伍尚忠,徐羡明,章琦.中国水稻白叶枯病菌致病型的研究.植物病理学报, 1990(20): 81~88.
    冯明姬.水稻香味基因fgr和抗稻瘿蚊基因Gm6的精细定位.华南农业大学硕士学位论文, 2006.
    高东迎,郭士伟,何冰,张彦,焦德茂.玉米PEPC基因和抗白叶枯病基因Xa25的水稻聚合育种.江苏农业学报, 2006, 22(1): 5~9.
    过崇俭.水稻白叶枯病.见:中国农业科学院职务保护所主编,中国农作物病虫害(第二版).北京,中国农业出版社,1995,14~24.
    黄大昉,林敏.农业微生物基因工程.科学出版社,北京, 2001.
    贾士荣.转基因植物食品中标记基因的安全性评价.中国农业科学, 1997, 30(2): 1~15.
    贾士荣,郭三堆,安道昌.转基因棉花.北京:科学出版社, 2001, pp19~23.
    李宣铿.杂交稻三系资源及组合抗白叶枯病研究.杂交水稻, 1993 (2): 35~36.
    马伯军,王文明,邹军煌,刘国庆,方进,朱立煌,翟文学.利用测交和PCR快速选择Xa21转基因杂交稻恢复系的纯合系.科学通报, 2000, 45(14): 1516~1520.
    倪大虎,易成新,李莉,汪秀峰,王文相,杨剑波.利用分子标记辅助选择聚合水稻基因Xa21和Pi9(t).分子植物育种, 2005, 3(3): 329~334.
    倪大虎,易成新,杨剑波,汪秀峰,张毅,章琦,王春连,赵开军,王文相,李莉.利用分子标记辅助选择聚合Pi9(t)和Xa23基因.分子植物育种, 2007, 5(4): 491~496.
    潘海军,王春莲,赵开军,章琦,樊颖伦,周少川,朱立煌.水稻抗白叶枯病基因Xa23的PCR分子标记及辅助选择.作物学报, 2003, 29(4): 501~507.
    秦钢,李杨瑞,李道远,梁海福,莫海玲,于松保,唐梅,郑希.水稻白叶枯病抗性基因Xa4、Xa23聚合及分子标记检测.分子植物育种, 2007, 5(5): 625~630.
    沙学延,朱立宏.中国水稻抗白叶枯病品种的抗源分析.南京农业大学学报, 1993, 16(4): 6~12.
    舒庆尧,叶恭银,崔海瑞,项友斌,高明尉.转基因水稻“克螟稻”选育.浙江农业大学学报, 1998, 24(6): 579~580.
    苏金,吴瑞.水稻中转基因表达的“位置效应”初报.农业生物技术学报, 1999, 7(4): 311~315.
    谭光轩,任翔,翁清妹,时振英,祝莉莉,何光存.药用野生稻转育后代一个抗白叶枯病新基因的定位.遗传学报, 2004, 31(7): 724~729.
    谭震波,章琦,朱立煌,王春连.水稻抗白叶枯病基因Xa-14在分子标记连锁图上的定位. 遗传, 1998, 20(6): 30~33.
    唐柞舜,李良材,田文忠.基因枪法转基因水稻后代农艺性状的表现.中国农业科学, 2001, 34(6): 581~586.
    王兴春,杨长登,李西明,马良勇.分子标记辅助选择与花药培养相结合快速聚合水稻白叶枯病抗性基因.中国水稻科学, 2004, 18(1): 7~10.
    吴家道,杨剑波,许传万,李莉,向太和,倪大虎,汪秀峰,贾士荣,唐益雄,张世平, Fauquet C.M..水稻抗白叶枯病基因Xa21转基因水稻及其杂交稻研究.作物学报, 2001(27): 29~34.
    徐建龙,林贻滋,翁锦屏,赵新立.水稻白叶枯病抗性基因的聚合及其遗传效应.作物学报, 1996, 22(2): 129~134.
    熊卫,余功新,林兴华,张端品,谢岳峰.五个云南稻种抗白叶枯病基因分析.华中农业大学学报, 1994, 13(2): 99~110.
    徐雍皋,徐敬友.农业植物病理学.江苏科学技术出版社. 1995.
    姚方印,李广贤,杨磊,李中华,翟红梅,杨海旭.农杆菌介导将白叶枯病抗性基因Xa21导入水稻获得转基因植株.西北植物学报, 2002(22): 1038~1043.
    杨文才,章琦,王春莲.六个籼稻品种对水稻白叶枯病的抗性遗传研究.华中农业大学学报, 1994, 13(3): 234~240.
    叶恭银,胡萃,舒庆尧.转基因抗虫水稻的转育及其合理持续利用. 1998.见:程家安,周
    伟军编.跨世纪农业发展与研究.北京:中国环境科学出版社, pp406~414.
    翟文学,李晓兵,田文忠,周永力,潘学彪,曹守云,赵显峰,赵彬,章琦,朱立煌.由农杆菌介导将白叶枯病抗性基因Xa21转入我国的5个水稻品种.中国科学, 2000, 30(2): 200~206.
    张红生,翟虎渠,陆志强,朱立宏,韩亮.水稻不同抗白叶枯病基因抗谱鉴定及在杂种Fl代的表达.南京农业大学学报, 1997, 20(2): 12~16.
    张良佑,吴洪基,萧整玉等.转基因水稻的抗虫性初探.华南农业大学学报,1998, 19(2):4~7.
    章琦.我国水稻白叶枯病抗性遗传的评价和利用.中国农业科学, 1991, 24(30): 26~36.
    章琦.我国水稻抗白叶枯病基因的利用及策略.植物病理学报,1995(22): 241~246.
    章琦.水稻白叶枯病抗性基因鉴定进展及其利用.中国水稻科学, 2005, 19(5): 453~459.
    章琦,施爱农等.水稻白叶枯病抗性遗传研究Ⅱ,水稻品种对五个菌系的抗性遗传分析. 中国水稻科学, 1990, 4(1): 1~8.
    章琦,施爱农,王春连,白建法,杨文才. 9个水稻品种对白叶枯病(Xanthomonas oryzae pv. oryzae)的抗性遗传研究.作物学报,1994(20): 84~92.
    章琦,王春莲,林世成.水稻抗病育种研究Ⅲ.我国主栽品种对若干白叶枯病菌系的抗性遗传分析.中国农业科学, 1988(21): 41~48.
    赵彬,王文明,郑先武,王春莲,马伯军,薛庆中,朱立煌,翟文学.水稻白叶枯病广谱抗性基因Xa21导入两用核不育系培矮64S.生物工程学报, 2000, 16(2): 137~141.
    郑康乐,庄杰云,王汉荣.基因聚合提高了水稻对白叶枯病的抗性.遗传, 1998(4): 4~6.
    周盛梅,汪晓玲,华志华,薛锐,王慧中,黄纯农,黄大年.用基因枪法转bar基因以培育抗除草剂的直播稻.杭州师范学院学报, 2001(18): 20~25.
    周毓珍,陆作相,吕兴泉,丁伦友,孙荣才.杂交稻恢复系白叶枯病(Xanthomonas oryzae pv. oryzae)抗性改良研究.南京农业大学学报, 1993, 16(4): 1~5.
    朱常香,胡全安,温孚江,郑成超,张杰.用两个抗虫基因分别转化水稻及抗虫株系的获得.农业生物技术学报, 1999(7): 259~266.
    朱常香,潘春欣,温孚江.利用PIG基因枪进行水稻基因转化的研究. 1996(27): 123~128.
    朱生伟,黄国存,孙敬三.外源DNA直接导入受体植物的研究进展.植物学通报2000, 17(1): 11~16.
    Adang M.J., firoozabady E. and Klein J.. Application of a Bacillus thuringiensis crystal protein for insect control. In: Alan R L ed. Molecular srtategies for crop protein. New York, 1987, pp345~353.
    Balzergue S., Dubreucq B., Chauvin S., Le Clainche L., Le Boulaire F., de Rose R., Samson F., Biaudet V., Lecharny A., Cruaud C., Weissenbaeh J., Caboche M. and Lepiniec L.. Improved PCR-walking for large-scale isolation of plant T-DNA borders. Biotechniques, 2001(30): 496~498, 502, 504.
    Barton K.A., Wllitely H.R. and Yang N.S.. Bacillus thuringiensis delta-endotoxin expressed in transgenic Nicotiana tabaeum provides resistance to lepidopteran insects. Plant Physiol, 1987(85): 1103~1109.
    Bates S.L., Zhao J.Z., Roush R.T. and Shelton A.M.. Insect resistance management in GM crops: past, present and future. Nat Biotechnol, 2005(23): 57~62.
    Bauer L.S.. Resistance: a threat to the insecticidal crystal proteins of Bacillus thuringiensis. Fla Entomol, 1995(78): 414~443.
    Borines L., Redona E., Porter B., White F., Vera Cruz C. and Leung H.. Molecular markers for detecting bacterial blight resistance genes in maintainer lines of rice hybrids. In:Khush G.S., Brar D.S., Hardy B.. Advance in Rice Genetics. Supplement to Rice GeneticsⅣ. Proceedings of the Fourth International Rice Genetics Symposium, 22-37 October 2000, IRRI, Los Banos, Philippines, 2001, pp245~247.
    Bostein D., White R.L. and Skolnick M.. Construction of genetic linkage map in man using restriction fragment length polymorphism. Amer J Hun Genet, 1993, 31(3): 314~318.
    Buttery R.G., Ling C.L., Bienvenido O.J. and Turnbaugh J.G.. Cooked rice aroma and 2-acetyl-1-pyrroline. Agric Food Chem. 1983, 31(4): 823~826.
    Cao J., Tang J.D., Strizhov N., Shelton A.M. and Earle E.D.. Transgenic broccoli with high levels of Bacillus thuringiensis Cry1C protein control diamondback moth larvae resistant to Cry1A or Cry1C. Mol Breed, 1999(5): 131~141.
    Casas A.M., Kononowiez A.K., Zehr U.B., Tomes D.T., Axtell J.D., Butler L.G., Bressan R.A. and Hasegawa P.M.. Transgenic sorghum plants via mocroprojectile bombardment. Proc Natl Acad Sci USA, 1993(90): 11212~11216.
    Causse M.A. and Fulon T.M.. Saturated molecular mapping of rice genome based on an interspecific backcross population. Genetics, 1994, 138(4): 1251~1274.
    Chattopadhyay A., Bhatnagar N.B. and Bhatnagar R.. Bacterial insecticidal toxins. Crit Rev microbiol, 2004(30): 33~54.
    Chen H.L., Wang S.P. and Zhang Q.F.. New gene for bacterial blight resistance in rice located on chromosome 12 identifid from Minghui 63, an elite restorer line. Phytopathology, 2002(92): 750~754.
    Chen S., Lin X.H., Xu C.G. and Zhang Q.F.. Improvement of bacterial blight resistance of ‘Minghui 63’, an elite restorer line of htbrid rice, by molecular marker-assisted selection. Crop Science, 2000, 40(1):239~244.
    Chen X.J., Curtiss A., Alcantara E. and Dean D.H.. Mutations in domainⅠof Bacillus thuringiensis delta-endotoxin Cry1Ab reduce the irreversible bingding of toxin to manduca sexta brush border membrane vesicles. J Biol Chem, 1995(270): 6412~6419.
    Cheng X.Y., Sardana R., Kaplan H. and Altosaar I.. Agrobacterium-transformed rice plants expressing synthetic cry1A(b) and cry1A(c) genes are highly toxic to yellow stem borer and striped stem borer. Proc Natl Acad Sci USA, 1998(95): 2767~2772.
    Chong S., Adang M.J., Lynch R.E., Anderson W.F., Wang A.M., Cardineau G. and Ozias-Akins P.. Expression of a Bacillus thuringiensis cry1Ac gene in transgenic peanut plants and its efficacy against lesser cornstalk borer. Transgenic Research, 1997(6): 169~176.
    Chu X.H., Fu B.Y., Yang H., Xu C.G., Li Z.K., Sanchez A., Park Y.J., Bennetzen J.L., Zhang Q.F. and Wang S.P.. Targeting xa13, a recessive gene for bacterial blight resistance in rice. Theor Appl Genet, 2006(112): 455~461.
    Datta K., Baisakh N., Maung Thet K., Tu J. and Datta S.K.. Pyramiding transgenes for multiple resistance in rice against bacterial blight, yellow stem borer and sheath blight. Theor Appl Genet, 2002(106): 1~8.
    Datta K., Vasquez A., Tu J., Torrizo L., Alam M.F., Oliva N., Abrigo E., Khush G.S. and Datta S.K.. Constitutive and tissue-specific differential expression of the cry1A(b) gene in transgenic rice plants conferring resistance to rice insect pest. Theor Appl Genet, 1998(97): 20~30.
    Datta S.K., Datta K., Soltanifar N., Donn G. and Potrykus I.. Herbicide-resistant indica rice plants from IRRI breeding line IR72 after PEG-meiated transformation of protoplasts. Plnat Mol Bio, 1992(20): 619~629.
    De Block M., Debrouwer D. and Tenning T.. Transformation of Brassica napus and Brassica oleracea using Agrobacterium tumefaciens and the expression of the bar neo genes in the transgenic plants. Plant Physiol, 1989(91): 694~701.
    Ezuka A. and Horino O.. Classfication of rice varieties and Xanthomonas oryzae strains on the basis of their differential interactions. Bull Tokai-Kinki natl Afric Stn, 1974(27): 1~29.
    Ferre J., Real M.D., Rie J.V., Jansens S. and Peferoen M.. Resistance to the Bacillus thuringiensis bioinsecticide in a field population of Plutella xylostella is due to a change in a midgut membrane receptor. Proc Natl Acad Sci USA, 1991(88): 119~123.
    Ferre J. and Van Rie J.. Biochemistry and genetics of insect resistance to Bacillus thuringiensis. Annu Rev Entomaol, 2002(47): 501~533.
    Fischhoff D.A., Bowdish K.S. and Perlak F.J.. Insect tolerant transgenic tomato plants. Bio Technol, 1987(5): 807~813.
    Fujimoto H., Itoh K. and Yamomoto M.. Insect resistance rice genetated by introduction of a modified delta-endotoxin gene of Bacillus thuringiensis. Bio Technol, 1993(11): 1151~1155.
    Gao D.Y., Xu Z.G., Chen Z.Y., Sun L.H., Sun Q.M., Lu F., Hu B.S. and Liu Y.F.. Identification of a new gene for resistance to bacterial blight in a somaclonal mutant HX-3 (indica). Rice Genet Newsl, 2001(18): 66~68.
    Garczynski S.F., Crim J.W. and Adang M.J.. Idengtification fo putative insect brush border membrane-binding molecules specific to Bacillus thuringiensis delta-endotoxin by protein blot analysis. Appl Environ Microbiol, 1991(57): 2816~2820.
    Ghareyazie B., Alinia F., Menguito C.A., Rubia L.G., de Palma J.M., Liwanag E.A. and Cohen M.B.. Enhanced resistance to two stem borers in an aromatic rice containing asynthetic cry1(A)b gene. Mol Breed, 1997(3): 401~414.
    Gould F., Anderson A., Reynolds A., Bumgarner L. and Moar W.. Selection and genetic analysis of a Helinthis virescens (Lepidoptera: Noctuidae) strain with high levels of resistance to Bacillus thuringiensis toxins. J Econom Entomol, 1995(88): 1545~1559.
    Gu K., Tian D., Yang F., Wu L., Sreekala C., Wang D., Wang G.L. and Yin Z.. High-resolution genetic mapping of Xa27(t), a new bacterial blight resistance gene in rice, Oryza sativa L. Theor Appl Genet, 2004, 108(5): 800~807.
    Gu K., Yang B., Tian D., Wu L., Wang D., Sreekala C., Yang F., Chu Z., Wang G.L., White F.F. and Yin Z.. R gene expression induced by a type-Ⅲeffector triggers disease resistance in rice. Nature, 2005(435): 1122~1125.
    Guiford P.. Microsatellites in Malus×domestica (apple): abundance, polymorphism and cultivar identification. Thero Appl Genet, 1997994): 249~254.
    Hama H., Suzuki K. and Tanaka H.. Inheritance and stability of resistance to Bacillus thuringiensis formulations of the diamondback moth, Plutella xylostella (Linnaeus) (Lepidoptera: Yponomeutidae). Appl Entomol Zool, 1992(27): 355~362.
    Hamada H., Marianne G.P. and Kakunaga T.. A novel repeated element with Z-DNA-forming protential is widely found in evolutionarily diverse eukaryotic genomes. Proc Natl Acad Sci USA, 1982(79): 6465~6469.
    Hannay C.L.. Crystalline inclusions in aerobic spore-forming bacteria. Nature, 1953(172): 1004.
    Hannay C.L. and Fitz-James P.. The protein crystals of Bacillus thuringieisis Berliner. Can J Microbiol, 1955(1): 694~709.
    Heale S.M. and Petes T.D. The stabilization of repetitive tracts of DNA by variant repeats requires a functional DNA mismatch repair system. Cell, 1995(83): 539~545.
    Heckel D.G.. The complex genetic basis of resistance to Bacillus thuringiensis toxin in insects. Biocontrol Sci Technol, 1994(4): 405~417.
    Hiei Y.. Efficient transformation of riee (Orazae sativa L) mediated by Agrobacterium and sequence analysis of the boundaries of theT-DNA. The Plant Journal, 1994, 6(2): 271~282.
    Hilder V.A., Gatehouse A.M.R., Sheerman S.E., Barker R.F. and Boulter D.. A novel mechanism of insect resistance engineered into tobacco. Nature, 1987(330): 160~163.
    Hittalmanni S., Parco A., Mew T.V., Zeigler R.S. and Huang N.. Fine mapping and DNA marker-assisted pyramiding of the three major genes for blast resistance in rice. Theor Appl Genet, 2000(100): 1121~1128.
    Hofmann C., Vanderbruggen H., Hofte H., Van Rie J., Jansens S. and Van Mellaert H.. Specificity of Bacillus thutingiensis delta-endotoxins is correlated with the presence ofhigh-affinity binding sites in the brush border membrane of target insect midguts. Proc Natl Acad Sci USA, 1998(85): 7844~7848.
    Huang N., Angeles E.R., Domingo J., Magpantay G., Singh S., Zhang G., Kumaravadivel N., Bennett J. and Khush G.S.. Pyramiding of bacterial blight resistance genes in rice: marker-assisted selection using RFLP and PCR. Theor Appl Genet, 1997(95): 313~320.
    IRRI. Annual program report. IRRI, Los Banos. 1995. pp19~54.
    Iyer A.S. and McCouch S.R.. The rice bacterial blight resistance gene xa5 encodes a novel form of disease resistance. Mol Plant Microbe Interact, 2004, 17(12): 1348~1354.
    James C.. Global status of transgenic crops in 1997. ISAAA Brief No 5, 1997. Ithaca, NY, ISAAA.
    James C.. Global status of commercialized biotech/GM crops: 2007. ISAAA Brief 37-2007: Executive Summary. Ithaca, NY, ISAAA.
    Jefferson R.A., Kavanagh T.A. and Bevan M.W.. GUS fusions:β-glucuronidase as a sensitive and versatile gene fusion marker in higher plants. EMBO J, 1987(6): 3901~3907.
    Jena K.K., Jeung J.U. and Shin M.S.. Pyramiding specific BB resistance genes for improvement of temperate Japanic rice in South Korea. Abstracts of the 1st International Conference on Bacterial Blight of Rice. March 17-19, 2004, Tsukuba, Japan: 16.
    Jiang J., Linscombe S.D., Wang J. and Oard J.H.. High efficiency transformation of U.S. rice lines from mature seedderived calli and segregation of glufosinate resistance under field conditions. Crop Sci, 2000(40): 1729~1741.
    Kain W.C., Zhao J.Z., Janmaat A.F., Myers J., Shelton A.M. and Wang P.. Inheritance of resistance to Bacillus thuringiensis Cry1Ac toxin in a greenhouse-derived strain of cabbage looper (Lepidoptera: Noctuidae). J Ecom Entomol, 2004(97): 2073~2078.
    Kaji R. and Ogawa T.. Identification of the located chromosome of the resistance gene Xa7 to bacterial leaf blight in rice. Jpn J Breeding, 1995, 45(suppl 1): 79.
    Kauffman H.E., Reddy A.P.K., Hsieh S.P.Y. and Merca S.D.. An improved technique for evaluating resistance of rice varieties to Xanthomonas oryzae. Plant Dis Rep, 1973(57): 737~741.
    Khush G.S. and Angeles E.R.. A new gene for resistance to race 6 of bacterial blight in rice, Oryza sativa L.Rice Genet Newsl, 1999(16): 92~93.
    Khush G.S., Bacalangco E. and Ogawa T.. A new gene for resistance to bacterial blight from O. longistaminata. Rice Genet Newsl, 1990(7): 121~122.
    Khush G.S., Mackill D.J. and Sidhu G.S.. Breeding rice for resistance to bacterial blight, In Bacterial Blight of Rice, First Edition. IRRI. 1989, pp207~218.
    Kohli A., Fu X., Twyman R. and Christou P.. Transgene integration, organization and expression. Rice Genetics News, 1999(16): 135~138.
    Kolesnik T., Szeverenyi L., Bachmann D., Kumar C.S., Jiang S.Y., Ramamoorthy R., Cai M.N., Ma Z.G., Sundaresan V. and Ramachandran S.. Establishing an efficient Ac/Ds tagging system in rice: large~scale analysis of Ds flanking sequences. The Plant Journal, 2004(37): 301~314.
    Koziel M.G., Beland G.L., Bowman C., Carozzi N.B., Crenshaw R., Crossland L., Dawson J., Desai N., Hill M., Kadwell S., Launis K., Lewis K., Maddox D., McPherson K., Meghji M.R., Merlin E., Rhodes R., Warren G.W., Wright M. and Evola S.V.. Field performance of elite transgenic maize plants expressing an insecticidal protein derived from Bacillus thuringiensis. Bio Tech, 1993(11): 194~200.
    Lander E.S., Green P., Abrahamson J., Barlow A., Daly M.J., Lincoln S.E. and Newburg L.. Mapmaker: an interactive computer package for construction primary genetic linkage maps of experimental and natural populations. Genomics, 1987(1): 174~181.
    Laursen C.M., Krzyzek R.A., Flick C.E., Anderson P.C., and Spencer T.M.. Production of fertile transgenic maize by electroporation of suspension culture cells. Plant Mol Biol, 1994(24): 51~61.
    Leach J.E., Roberts P.D., Guo A. and Barton-Willis P.. Multiplication of Xanthomonas campestris pv. oryzae in rice leaves. In: IRRI eds, Bacterial blight of rice. IRRI, Manila, 1989, pp43~53.
    Lee K.S., Rasabandith S., Angeles E.R. and Khush G.S.. Inheritance of resistance to bacterial blight in 21 cultivars of rice. Phytopathology, 2003, 93(2): 147~152.
    Leuhrsen K.R. and Walbot V.. Insertion of Mul elements in the first intron of the Adh1-s gene of maize results in novel RNA processing events. The Plant Cell, 1990(2): 1225~1238.
    Liang Y., Patel S.S. and Dean D.H.. Irreversible bingding kinetics of Bacillus thuringiensis Cry1A delta-endotoxins to gypsy moth brush border membrane vesicles is directly correlated to toxicity. J Biol Chem, 1995(270): 24719~24724.
    Li Z.K., Luo L.J., Mei H.W., Paterson A.H., Zhao X.H., Zhong D.B., Wang Y.P., Yu X.Q., Zhu L., Tabien R., Stansel J.W. and Ying C.S.. A“defeated”rice resistance gene acts as a QTL against a virulent strain of Xanthomonas oryzae pv. oryzae. Mol Gen Genet, 1999(261): 58~63.
    Li Z.K., Sanchez A., Angeles E., Singh S., Domingo J., Huang N. and Khush G.S.. Are the dominant and recessive plant disease resistance genes similar? A case study of rice R gene and Xanthomonas oryzae pv. oryzae races. Genetics, 2001(159): 757~765.
    Lin X.H., Zhang D.P., Xie Y.F., Gao H.P. and Zhang Q.. Identifying and mapping a new gene for bacterial blight resistance in rice based on RFLP markers. Phytopathology, 1996(86) : 1156~1159.
    Liu Y.G., Mitsukawa N., Oosumi T. and Whittier R.F.. Efficient isolation and mapping of Arobidopsis thaliana T-DNA insert junctions by thermal asymmetric interlaced PCR. Plant J, 1995a(8): 457~463.
    Liu Y.G. and Whittier R.F.. Thermal asymmetric interlaced PCR: automatable amplification and sequencing of insert and fragments from P1 and YAC clones for chromosome walking. Genomics, 1995b(25): 674~681.
    Lorieux M., Petrov M., Huang N., Guiderdoni E. and Ghesquière A.. 1996. Aroma in rice: genetic analysis of quantitative trait. Theor Appl Genet, 1996(93): 1145~1151.
    Lynch P.T., Jones J, Blackhall N.W., Davey M.R., Power J.B., Cocking E.C., Nelson M.R., Bigelow D.M., Orum T.V., Orth C.E. and Schuh W.. The phenotypic characterization of R2 generation transgenic rice plants under field and glasshouse conditions. Euphytica, 1995, 85(1-3): 395~401.
    Mahatheeranont S., Keawsa-ard S. and Dumri K.. Qualification of the rice aroma compound, 2-Acetyl-1-pyrroline, in uncooked Khao Dawk Mali 105 brown rice. Agric Food Chem, 2001(49): 773~779.
    Martin P.A.W. and Travers T.S.. Worldwide abundance and distribution of BT isolates. Appl Environ Microbiol, 1989(55): 2437~2442.
    McCouch S.R., Abenes M.L., Angeles R., Khush G.S. and Tanksley S.D.. Molecular tagging of a resistance gene, xa5, for resistance to bacterial blight of rice. Rice Genet Newsl, 1991(8): 143~145.
    McCouch S.R., Kochert G., Yu Z.H., Wang Z.Y., Khush G.S., Coffman W.R. and Tanksley S.D.. Molecular mapping of rice chromosomes. Theor Appl Genet, 1988, 76(6): 815~829.
    McGaughey W.H.. Insect resistance to the biological insecticide Bacillus thuringiensis. Science, 1985(229): 193~195.
    McGaughey W.H and Whlton M.E.. Managing insect resistance to Bacillus thuringiensis toxins. Science, 1992(58): 1451~1455.
    McPherson K., Meghji M.R., Merlin E., Rhodes R., Warren G.W., Wright M. and Evola S.V.. Field performance of elite transgenic maize plants expressing an insecticidal protein derived from Bacillus thuringiensis. Bio Tech, 1993(11): 194~200.
    Mew T.W.. Current status and future prospects of research on bacterial blight of rice. Annu Rev Phytopathol, 1987(25): 359~382.
    Mew T.W.. Characterization of Resistance in Rice to Bacterial Blight. Ann. Phytopath. Soc. Mew T.W. Focus on bacterial blight of rice, Plant disease, 1993, 77(1): 5~12.
    Moore S.S., Sargeant L.L. and King T.G.. Conservation of dinucleotide microsatellites among mammalian genomes allows use of heterologous PCR primer pairs in closely related species. Genomics, 1991(10): 654~660.
    Nakai H., Kuwahara M. and Saito M. Studies of an induced mutant resistant to multiple races of bacterial leaf blight. Rice Genet Newsl, 1988(5): 101~103.
    Nayak P., Basu D., Das S., Basu A., Ghosh D., Ramakrishnan N.A., Ghosh M. and Sen S.K.. Transgenic elite indica rice plants expressing cry1A(c) endotoxin of Bacillus thuringiensis are resistant against yellow stem borer(scirpophaga incertulas). Proc Natl Acad Sci USA, 1997(94): 2111~2116.
    Noda T. and Ohuchi A.. A new pathogenic race of Xanthomonas campestris pv. oryzae and inheritance of resistance of differential rice variety, Tetep to it. Ann Phytopathol Soc Jpn, 1989(55): 201~207.
    Oard J.H., Linscombe S.D., Braverman M.P., Jodari F., Blouin D.C., Leech M., Kohli A., Vain P., Cooley J.C. and Christou P.. Development, field evaluation, and agronomic performance of transgenic herbicide resistant rice. Mol Breeding, 1996(2): 359~368.
    Ogawa T., Busto G.A., Tabien R.E., Romero G.O., Endo N. and Khush G.S.. Grouping of rice cultivars based on reaction pattern to Philippine races of bacterial blight pathogen (Xanthomonas oryzae pv. oryzae). Japan J Breed, 1991, 41(1): 109~119.
    Ogawa T.. Methods and strategy for monitoring race distribution and identification of resistance genes to bacterial leaf blight (Xanthomonas compestrispv. oryzae) in rice. JARQ, 1993, 27(2): 71~80.
    Ogawa T. and Khush G.S.. Major genes for resistance to bacterial blight in rice. Proceediongs of the international workshop on bacterial blight of rice. IRRI. Los Banos, Phillippines, 1989, pp177~192.
    Ogawa T., Lin L., Tabien R.E. and Khush G.S.. A new recessive gene for resistance to bacterial blight of rice. Rice Genet Newsl, 1987(4): 98~100.
    Ogawa T., Morinaka T., Fuiii K. and Kiruma T.. Inheritance of resistance of rice varieties Kogyoku and Java14 to bacterial grouopⅤof Xanthomonas oryzae. Ann Phytopath Soc Jpn, 1978(44): 137~141.
    Ogawa T. and Yamamoto T.. Resistance gene of rice cultivar, Asominori to bacterial blight of rice. Jpn J Breeding, 1989(39)(suppl 1): 196~197.
    Olufowote J.O., Xu Y.B., Chen X.L., Park W.D., Beachell H.M., Dilday R.H., Goto M. and McCouch S.R.. Comparative evalution of within-cultivar variation of rice (Oryzae sativa L) using microsatellite and RFLP markers. Genome, 1997(40): 370~378.
    Ou S.H.. Rice disease. Kew: Commomwealth Mycological Institute, 1972, pp368.
    Ou S.H.. Rice disease. Commomwealth Agricultural Bereaux. England, 1985, pp380. Panaud O., Chen X. and McCouch S.R.. Frequency of microsatellites sequences in rice (Oryza sativa L.). Genome, 1995, 38(6): 1170~1176.
    Peng J., Kononowicz H. and Hodges T.K.. Transgenic indica rice plants. Theo Appl Genet, 1992, 83(6-7): 855~863.
    Perlak F.J., Deaton R.W., Armstrong T.A., Fuchs R.L., Sims S.R., Greenplate J.T. andFischhoff D.A.. Insect resistance cotton plants. Bio Technol, 1990(8): 939~943.
    Perlak F.J., Fuchs R.L., Dean D.A., McPherson S.L. and Fischhoff D.A.. Modification of the coding sequence enhances plant expression of insect control protein genes. Proc Natl Acad Sci USA, 1991(88): 3324~3328.
    Perucho M., Hanahan D., Lipsich L. and Wigler M.. Isolation of the chicken thymidine kinase gene by plasmid rescue. Nature, 1980(285): 207~210.
    Petpisit V., Khush G.S. and Kauffman H.E.. Inheritance of resistance to bacterial blight in rice. Crop Sci, 1977(17): 551~554.
    Pingali P.L. and Roger P.A.. Impact of pesticides on farmer health and the rice environment. Manila, International Rice Research Institute, 1995.
    Powell W., Morgante M., Andre C., Hannfey M., Vogel J., Tingey S. and Rafalski A.. The comparison of RFLP, RAPD, AFLP and SSR(microsatellite) markers for germplasm analysis. Mol Breed, 1996(2): 225~238.
    Rajamohan F., Alcantara E., Lee M.K., Chen X.J., Curtiss A. and Dean D.H.. Single amino acid changes in domainⅡof Bacillus thuringiensis Cry1Ab delta-endotoxin affect irreversible binding to Manduca sexta midgut membrane vesicles. J Bacteriol, 1995(177): 2276~2282.
    Ramsey M.D. and Moffett M.L.. Bacterial blight in North Queensland, Australia. In: IRRI eds, Bacterial blight of rice. IRRI. Manila, 1989, pp219.
    Rashild H., Yokoi S., Toriyam K. and Hinata K.. Transgenic plant production mediated by Agrobacterium in Indica rice. Plant Cell Rep, 1996(15): 727~730.
    Rathmore K.S., Chowdhury V.K. and Hodges T.K.. Use of bar as a selectable marker gene and for the production of herbicide-resistant rice plants from protoplasts.Plant Mol Biol, 1993(21): 871~884.
    Reckhaus P.M.. Occurrence of bacterial blight of rice in Niger, West Africa. Plant Dis, 1983(67): 1039.
    Reddy A.P.K.. Bacterial blight crop less assessment and disease management. In: IRRI eds, Bacterial blight of rice. IRRI. Manila, 1989.
    Reddy V.D. and Reddy G.M.. Genetic and biochemical basis of scent in rice (Oryza sativa L.). Theor Appl Genet, 1987, 73(5): 699~700.
    Riazuddin S.. Transformation of indica rice with Bt pesticidal genes. In: Rice GeneticsⅢ: Proceeding of the Third International Rice Genetics Symposium, October 16-20, 1995, Manila, Philippines. IRRI, 1996, pp730~734.
    Ronald P.C. and Tanksley D.. Genetic and physical mapping of the bacterial blight resistance gene Xa21. Rice Genet Newsl, 1991(8): 142~143.
    Ronald P.C., Albano B. and Tabien R., Abenes L., Wu K., McCouch S. and Tanksley S.D..Genetic and physical analysis of the rice bacterial blight resistance locus, Xa21. Mol Gen Genet, 1992(236): 113~120.
    Roush R.T.. Two-toxin strategies for management of insect resistant transgenic crops: Can pyramiding succeed where pesticide mixtures have not? Philos Trans R Soc London Ser B, 1998(353): 1777~1786.
    Sanchez A.C., Brar D.S., Huang N., Li Z.K. and Khush G.S.. Sequence tagged site marker-assisted selection for three bacterial blight resistance genes in rice. Crop Sci, 2000(40): 792~797.
    Sayyed A.H., Haward R., Herrero S., Ferre J. and Wright D.J.. Genetics and biochemical approach for characterization of resistance to Bacillus thuringiensis toxin Cry1Ac in a field population of the diamondback moth, Plutella xylostella. Appl Environ Microbiol, 2000(66): 1509~1516.
    Schnepf H.E. and Whiteley H.R.. Cloning and expression of the Bacillus thuringiensis. Crystal protein gene in Escherichia coli. Proc Natl Acad Sci USA, 1981, 78(5): 2893~2897.
    Sessions A., Burke E., Presting G., Aux G., McEllver J., Patton D., Dietrich B., Ho P., Bacwaden J., Ko C., Clarke J.D., Cotton D., Bullis D., Snell J., Miguel T., Hutchison D., Kimmerly B., Mitzel T., Katagiri F., Glazebrook J, Law M. and Goff S.A.. A high-throughput Arabidopsis reverse genetics system. Plant Cell, 2002(14): 2985~2994.
    Sharon D., Adato A., Mhameed S., Lavi U., Hillel J., Gomolk M., Epplen C. and Epplen J.T.. DNA fingerprints in plants using simple-sequence repeat and minisatellite probes. Hortscience, 1995, 30(1): 109~112.
    Shelton A.M., Tang J.D., Roush R.T., Metz T.D. and Earle E.D..Field tests on managing resistance to Bt-engineered plants. Nat Biotechnol, 2000(18): 339~342.
    Shelton A.M., Zhao J.Z. and Roush R.T.. Economic, ecologic, food, safety and social consequence of the deployment of Bt transgenic plants. Annu Rev Entomol, 2002(47): 845~881.
    Shu Q.Y., Ye Q.Y., Cui H.T., Cheng X.Y., Xiang Y.B., Wu D.X., Gao M.W., Xia Y.W., Hu C., Sardana R. and Altosaar I.. Transgenic rice plants with a synthetic cry1A(b) gene from Bacillus thuringiensis were highly resistant to eight lepidopteran rice pest species. Mol Breed, 2000(6): 433~439.
    Sidhu G.S., Khush G.S. and Mew T W. Genetic analysis of bacterial blight resistance in seventy-four cultivars of rice, Oryzae sativa L.Theor Appl Genet, 1978(53): 105~111.
    Singh G.P., Srivastara M.K., Singh R.V. and Singh R.M.. Variation in quantitative and qualitative losses caused by bacterial blight on rice varieties. Indian Phytopathol, 1977(30): 180~185.
    Singh R.K., Singh U.S. and Khush G.S.. Indigenous aromatic rice of India: Present scenarioand needs. Agricultural Situation in India, 1997(8): 491~496.
    Singh S., Sidhu J.S., Huang N., Vikal Y., Li Z., Brar D.S., Dhaliwal H. and Khush G.S.. Pyramiding three bacterial blight resistance genes (xa-5, xa-13 and Xa21) using marker-assisted selection into indica rice cultivar PR106. Theor Appl Genet, 2001(102): 1011~1015.
    Song W.Y., Wang G.L., Chen L.L., Kim H.S., Pi L.Y., Holsten T., Gardener J., Wang B., Zhai W.X., Zhu L.H., Fauquet C. and Ronald P.. A receptor kinase-like protein edcoded by the rice disease resistance gene, Xa21. Science, 1995(270): 1804~1806.
    Sood B.C. and Siddiq E.A.. A rapid technique for scent determination in rice. Indian J Genet Plant Breed, 1978(38): 268~271.
    Stahly D.P., Dingman D.W., Bulla L.A. and Aronson A.I.. Possible origin and function of the parasporal crystals in Bacillus thutingiensis. 1978, 84(3): 581~588.
    Strizhov N., Keller M., Mathur J., Koncz-Kalman Z., Bosch D., Prudovsky E., Schell, J. Sneh B., Koncz C. and Zilberstein A.. A synthetic cry1C gene, encoding a Bacillus thuringiensisδ-endotoxin, confers Spodoptera resistance in alfalfa and tobacco. Proc Natl Acad Sci USA, 1996(13): 15012~15017.
    Sun X., Cao Y., Yang Z., Xu C., Li X., Wang S. and Zhang Q.. Xa26, a gene conferring resistance to Xanthomonas oryzae pv. oryzae in rice, encodes an LRR receptor kinase-like. Plant J, 2004(37): 517~527.
    Sun X., Yang Z., Wang S. and Zhang Q.. Identification of a 47 kb DNA fragment containing Xa4, a locus for bacterial blight resistance in rice. Theor Appl Genet, 2003(106): 683~687.
    Swing J.M.. Reclassification of the causal agents of bacterial blight (Xoo) and bacterial leaf streak of rice. Nom Rev J Bacterial, 1990(40): 309~311.
    Tabashnik B.E., Finson N., Groeters F.R., Moar W.J., Johnson M.W., Luo K. and Adang M.J.. Reversal of resistance to Bacillus thuringiensis in Plutella xylostella. Proc Natl Acad Sci USA, 1994(91): 4120~4124.
    Tabashnik B.E.. Evolution of resistance to Bacillus thuringiensis. Annu rev Entomol, 1994(39): 47~79.
    Tabien R.E., Abalos M.C. and Bustaman M.. Multilines for bacterial blight control in rice. Abstract, 1st International Conference on Bacterial Blight of Rice. March 17-19, 2004, Tsukuba, Japan: 51.
    Tagami Y. and Mizukami T.. Historical review of the researches on bacterial blight of rice caused by Xanthomonas oryzae (Uyeda et Ishiyama) Dowson. Spec Rep Plant Dis Ins Pest Forcasting Serv, 1962(10): 112.
    Tanksley S.D., Young N.D., Paterson A.H. and Bonierbale M.W.. RFLP mapping in plant breeding: new tools for old sciences. Biotechnology, 1989(7): 257~264.
    Tautz D.. Hypervariability of simple sequences as a general source for polymorphic DNA markers. Nucleic Acids Res, 1989(17): 6463~6471.
    Taura S., Ogawa T., Yoshimur A., Ileda R. and Omura T.. Identification of a recessive resistance gene in induced mutant line XM5 of rice to rice bacterial blight. Jpn J Breeding, 1991, 41(3): 427~432.
    Taura S., Ogawa T., Yoshimura A., Ileda R. and Iwata N.. Identification of a recessive gene to rice bacterial blight of mutant line XM6, Oryza sativa L. Jpn J Breeding, 1992, 42(1): 7~13.
    Temnykh S., Park W.D., Ayres N., Cartinhour S., Hauck N., Lipovich L., Cho Y.G., Ishii T. and McCouch S.R.. Mapping and genome organization of microsatellite sequences in rice (Oryza satica L.). Thero Appl Genet, 2000(100): 697~712.
    Tsuzuki E., Tanaka K. and Shida S.. Studies on the characteristics of scented riceⅥ. Varietal differences in fatty acid composition of brown rice. Bull Fac Agric Miyazaki Univ, 1979(26): 443~449.
    Tu J., Ona I., Zhang Q., Mew T.W., Khush G.S. and Datta S.K.. Transgenic rice varity“IR72”with Xa21 is resistant to bacterial blight. Theor Appl Genet, 1998(97): 31~36.
    Tu J., Zhang G., Datta K., Xu C., He Y., Zhang Q., Khush G.S. and Datta S.K.. Field performance of transgenic elite commercial hybrid rice expressing Bacillus thuringiensisδ-endotoxin. Nature Biotechnol, 2000(18): 1101~1104.
    Vaeck M., Reynaerts A. and Hofte H.. Transgenic plants protected from insect attack. Nature, 1987(328): 33~37.
    Van Rie J., Jansens S., Hofte H., Degheele D. and Van Mellaert H.. Specificity of Bacillus thuringiensis delt-endotoxins. Importance of specific receptors on the brush border membrane of the mid-gut of target insects. Eur J Biochem, 1989(186): 239~247.
    Van Rie J., McGaughey W.H., Johnson D.E., Barnett B.D. and Van Mellaert H.. Mechanism of insect resistance to the microbial insecticide Bacillus thuringiensis. Science, 1990(247): 72~74.
    Vasil V., Castillo A.M., Fromm M.E. and Vasil I.K.. Herbicide resistant fertile transgenic wheat plants obtained by microprojectile bombardment of regenerable embryonic callus. Biotechnology, 1993(10): 667~674.
    Vera Cruz C.M. and Mew T.W.. Phenotypic analysis of the rice of Xanthomonas campestris pv. oryzae. Phytopathology, 1986(76): 11~17.
    Wan Y. and Lemaux P.G.. Generation of large numbers of independently transformed fertile barley plants.Plant Physiol, 1994(104): 37~48.
    Wang G.L., Ruan D.L., Song W.Y., Sideris S., Chen L.L., Pi L.Y., Zhang S.P., Zhang Z., Fauquet C., Gsut B.S., Whalen M.C. and Ronald P.. Xa21D encodes a receptor-like molecular with a leucine-rich repeat domain that determines race-specific recognitionand is subject to adaptive evolution. Plant Cell, 1998(10): 765~779.
    Wang G.L., Song W.Y., Ruan D.L., Sideris S. and Ronald P.. The cloned gene, Xa21, confers resistance to multiple Xanthomonas oryzae pv. oryzae isolates in transgenic plants. Mol Plant Microbe Interact, 1996(9): 850~855.
    Wang G.W., He Y.Q., Xu C.G. and Zhang Q.F.. Identification and confirmation of three neutral alleles conferring wide compatibility in inter-subspecific hybrids of rice (Oryza sativa L.) using near-isogenic lines. Theor Appl Genet, 2005(111): 702~710.
    Wei Y.P., Yao F.Y., Zhu C.X., Jiang M.S., Li G.X., Song Y.Z. and Wen F.J.. Breeding of transgenic rice restorer line for multiple resistance against bacterial blight, striped stem borer and herbicide. Euphytica, 2007, online first.
    Welson J. and Mc Clelland M.. Fingerprinting genomes using PCR with arbitrary primers. Nucl Acids Res, 1990, 18(24): 7213~7218.
    White P.T.. Rice: The essential harvest. Natl Geogr, 1994(185): 48~79. Williams J.G.K., Kubelik A.R., Livak K.J., Rafalski J.A. and Tingey S.V.. DNA polymorphisms amplified by arbitrary primers are useful as genetic markers. Nucl Acids Res, 1990, 18(22): 6531~6535.
    Wolfersberger M.G.. The toxicity of two Bacillus thuringiensis delta-endotoxins to gypsy moth larve is inversely related to the affinity of binding sites on midgut brush border membranes for the toxins. Experientia, 1990(46): 475~477.
    Wu C., Fan Y., Zhang C., Oliva N. and Datta S.K.. Transgenic fertile japannica rice plants expressing a modified cry1A(b) gene resistant to yellow stem borer. Plant Cell Rep, 1997(17): 129~132.
    Wu K.S., Jone R., Danneberger L. and Scolnik P.A.. Detection of microsatellite polymorphisms without cloning. Nucl Acids Res, 1994, 22(15): 3257~3258.
    Wu K.S. and Tanksley S.D.. Abundance, polymorphism and genetic mapping of microsatellite in rice. Mol Gen Genet, 1993(241): 225~235.
    Wunn J., Kloti A., Burkhardt P.K., Bidwas G.C.G., Launis K., Iglesia V.A. and Potrykus I.. tTransgenic indica rice breeding line IR58 expressing a synthecie cry1A(b) gene from Bacillus thuringiensis provies effective insect pest control. Bio Tech, 1996, 14(2): 171~176.
    Xe S. and Muir W.M.. Multistage selection for genetic gain by orthogonal transformation. Genetics, 1991(129): 963~974.
    Yamada T., Horino O. and Samoto S.. Studies on Genetics and Breeding of Resistance to Bacterial Leaf Blight in RiceⅠ, Discovery of new Varietal Groups on The Basis of Reaction Patterns to Five Different Pathotypes of Xanthomonas oryzae (Uyeda et Ishiyama) Dowson in Japan. Ann Phytopathol Soc Japan, 1979, 45(2): 240~246.
    Yamamoto T. and Ogawa T.. Inheritance of resistance in rice cultivars, Toyonishiki, Milyang 23 and IR24 to Myanmar isolates of bacterial leaf blight pathogen. JARQ, 1990(24): 74~77.
    Yang Z.F., Sun X.L., Wang S.P. and Zhang Q.. Genetic and physical mapping of a new gene for bacterial blight resistance in rice. Theor Appl Genet, 2003(106): 1467~1472.
    Ye G.Y., Tu J., Hu C., Datta K. and Datta S.K.. Transgenic IR72 with fused Bt gene cry1Ab/cry1Ac from Bacillus thuringiensis is resistant against four lepidopteran species under field conditions. Plant Biotechnol, 2001(18): 125~133.
    Yoshimura A., Ideta O., Takebayashi S., Takebayashi S., Matasumoto T., Yoshimura S., Ogawa T. and Iwata N.. Integration of RFLP and conventional maps in rice.Ⅱ. The location of gl-Ⅰand xa5. Jpn J Breeding, 1992b, 42(suppl2): 124~125.
    Yoshimura A., Lei J.X., Matsumoto T., Tsunematsu H. and Yoshimura S.. Analysis and pyramiding of bacterial blight resistance genes in rice by using DNA markers. In: Khush G.S. ed., Rice geneticsⅢ, Prodedings of the Third International Rice Genetics Symposium. IRRI. Manila, Philippines, 1996a, pp577~581.
    Yoshimura A., Mew T.W., Khush G.S. and Omura T.. Inheritance of resistance to bacterial blight in rice cultivar Cas209. Phytopathology, 1983, 73(10): 1409~1412.
    Yoshimura S., Yamanouchi U., Katayose Y., Toki S., Wang Z.K., Kono I., Kurata N., Yano M., Iwata N. and Sasaki T.. Expression of Xa1, a bacterial blight resistance gene in rice, is induced by bacterial inoculation. Proc natl Acad Sci USA, 1998(95): 1663~1668.
    Yoshimura S., Yamanouchi U., Kurata N., Nagamura Y., Sasaki T., Minobe Y. and Iwata N.. Identification of a YAC clone carrying the Xa1 allele, a bacterial blight resistance gene in rice. Theor Appl Genet, 1996b(93): 117~122.
    Yoshimura S., Yoshimura A., Iwata N., McCouch S.R., Abenes M.L., Baraoidan M.R., Mew T.W. and Nelson R.J.. Tagging and combining bacterial blight resistance genes in rice using RAPD and RFLP markers. Mol Breeding, 1995(1): 375~387.
    Yoshimura S., Yoshimura A., Saito A., Kishimoto N., Kawase M., Yano M., Nakagahra M., Ogawa T. and Iwata N.. RFLP analysis of introgressed chromosomal segments in three near-isogenic line of rice for bacterial blight resistance genes, Xa-1, Xa-3, and Xa-4. Jpn J Genet, 1992a(67): 29~37.
    Zambryski P.C., Joos H., Genetello C., Leemans J., Van M. and Schell J.. Ti plasmid vector for the introduction of DNA into plant cell without alteration of their normal regeneration capacity. EMBO J, 1983(2): 2143~2150.
    Zhang G., Angeles E.R., Abenes M.L.P., Khush G.S. and Huang N.. RAPD and RFLP mapping of the bacterial blight resistance gene xa-13 in rice. Theor Appl Genet, 1996(93): 65~70.
    Zhang Q., Lin S.C., Zhao B.Y., Wang C.L., Yang W.C., Zhou Y.L., Li D.Y., Chen C.B. andZhu L.H.. Identification and tagging a new gene for resistance to bacterial blight (Xanthomonas oryzae pv. oryzae) from O. rufipogon. Rice Genet Newsl, 1998(15): 138~142.
    Zhang Q., Wang C.L., Zhao K.J., Yang W.C., Qiao F., Zhou Y.L., Jiang Q.X. and Liu G.C.. Development of near-iso-genic line CBB23 with new resistance gene to bacterial blight in rice and its application. Chinese J Rice Sci, 2002, 16(3): 206~210.
    Zhang S., Song W.Y., Chen L.L., Ruan D.L., Taylor N., Ronald P., Beachy R. and Fauquet C.. Transgenic elite Indica rice varieties, resistant to Xanthomonas oryzae pv. oryzae. Molecular breeding, 1998(4): 551~558.
    Zheng K.L., Huang N., Bennett J. and Khush G.S.. PCR-based marker-assisted selection in rice breeding. 1995, IRRI discussion paper, series No.12.
    Zhou P.H., Tan Y.F., He Y.Q., Xu C.G. and Zhang Q.F.. Simultaneous improvement for four traits of Zhenshan97, an elite parent of hybrid rice, by molecular marker-assisted selection. Theor Appl Genet, 2003 (106): 326~331.
    Zoschke A.. Yield losses in tropical rice as influenced by the composition of weed flora and the timing of its elimination. In: B.T. Grayson M.B. Green & L.G. Copping (Eds), Pest Management in Rice, 1990: 300~313. New York, Elsevier Applied Science.

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