水稻三系恢复系航1号的白叶枯病和螟虫抗性改良研究
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摘要
水稻白叶枯病是世界水稻三大病害之一,利用水稻自身的抗病基因培育抗病品种,是白叶枯病防治的必由之路。虫害也是导致水稻减产的重要因素。水稻自身抗螟虫的基因资源匮乏,通过培育转Bt基因水稻,可探究抗虫育种的新途径。航1号是我国生产上应用面积较大的三系恢复系,其配组的超级杂交稻组合特优航1号和Ⅱ优航1号,具有优质高产的特性。为进一步提高恢复系航1号对白叶枯病的抗性、同时增加其对螟虫如稻纵卷叶螟的抗性,木研究采用常规育种方法并结合分子标记辅助选择技术先将Xa7基因、Xa21基因和cry1C*基因从供体亲木华恢20与T1C-19分别导入航1号中,再通过复交聚合双基因(Xa7+cry1C*或Xa21+cry1C*)和三基因(Xa7+Xa21+cry1C*),经田间人工接种白叶枯病菌株ZHE173(Ⅳ型)和GD1358(Ⅴ型)对育成的改良株系进行白叶枯病抗性鉴定,在自然发虫条件下对育成株系进行螟虫抗性鉴定,并对其及其所配组合,按照中华人民共和国农业部标准进行稻米品质测定。最后采用NCⅡ设计选择9个航1号的改良株系分别与5个不育系C815S、Ⅱ-32A、川农1A、金科1A和巨风A进行配组,并对各组合的8个主要农艺性状进行配合力分析。木研究主要结果如下：
1.育成了含Xa7+cry1C*或Xa21+cry1C*或Xa7+Xa21+cry1C*基因的同时抗白叶枯病和螟虫的改良株系各2个。
2.人工接种鉴定结果表明,只含Xa7或Xa21基因以及基因聚合的航1号改良株系的白叶枯病抗性明显提高。经自然条件下田间稻纵卷叶螟鉴定,含cry1C*基因的改良株系抗性也明显提高。
3.稻米品质分析结果表明,各供试材料的稻米品质等级难以超过对照品种扬两优6号,整精米率和垩白性状为各供试材料品质改良的主要限制因子。
4.杂交组合的杂种优势分析表明,有部分改良株系在农艺性状和配合力等方面基本保持了航1号的主要特征特性,同时筛选出金科1A/JY8021-1、巨风A/航1号、巨风A/JY8020-1、巨A/JY8022-1、巨风A/JY8024-1和巨风A/JY8232-1等六个强优势组合。
综合上述结果,本研究已成功地将抗白叶枯病基因Xa7、Xa21,抗螟虫基因cry1C*导入水稻恢复系航1号中,育成的改良株系和所配的组合白叶枯病和螟虫抗性明显提高,这些基因聚合株系具有良好的应用前景。
Bacterial blight of rice is one of the three most destructive diseases throughout the world. As an effective measure, the exploitation of host resistance genes can be used for the improvement of disease resistance. On the other hand, pests are the main bottleneck for rice production due to the lack of host resistance genes. Nowadays, we can develop resistant cultivars by transferring pests-resistant Bt gene. Hang 1 is an elite restorer line of three-line hybrid, which is widely used in the cultivation of hybrid rice. The super hybrids, Teyouhang 1 andⅡyouhang 1, which were mated using Hang 1 as male parent, have the characteristics of high yield and novel quality. In order to improve its resistance to both bacterial blight and stem bore such as Cnaphalocrocis medinalis. Xa7, Xa21 and cry1C* genes were transferred from Huahui 20 and TIC-19 respectively to the genetic background of Hang 1 by traditional breeding combining with MAS firstly, then the two-gene lines (Xa7+cry1C* or Xa21+cry1C*) and three-gene lines (Xa7+Xa21+cry1C*) were obtained by multiple cross. The specific Xanthomonas oryzae pv. oryzae strains, ZHE173 (Ⅳ) and GD1358 (Ⅴ) were used for identifying the improved lines' resistance to bacterial blight by artificial inoculation under the field condition. The experiment of resistance to stem borer for the improved lines was carried out under the field condition also. The quality traits of improved lines and their hybrids have been measured and analyzed in the research. The 9 improved lines of Hang 1 were hybridized with 5 sterile lines C815S,Ⅱ-32A, Chuannong 1 A, Jinke 1A and Jufeng A according to NCII design, and the combining ability for 8 main agronomic traits of the hybrids were investigated. The main result in the research is as follows:
1. We have obtained two improved lines with genetic background of Hang 1 which contain Xa7 and cry1C* genes, Xa21 and cry1C* genes and Xa7, Xa21 and cry1C* genes respectively.
2. The improved lines and their hybrids' resistance level to bacterial blight has been greatly improved. The improved lines and their hybrids showed high resistance level to Cnaphalocrocis medinalis.
3. The result of rice quality traits shows that all the meterials studied are generally inferior to CK (Yangliangyou 6) on quality traits, and chalky trait and milled rice rate are main limiting factors.
4. The result of usage of rice heterosis shows that only a part of improved lines' agronomic traits and combining ability are similar to Hang 1. And Jinke 1A/JY8020-1, Jufeng A/Hang 1, Jufeng A/JY8021-1, Jufeng A/JY8022-1, Jufeng A/JY8024-1 and Jufeng A/JY8232-1 were selected in the research.
All in all, Xa7, Xa21 and cry1C* genes were transferred into the genetic background of Hang 1 successfully. The improved lines and their hybrids's resistance levels to both bacterial blight and stem borer were increased obviously. These lines have a good prospect in future practical use.
1.育成了含Xa7+cry1C*或Xa21+cry1C*或Xa7+Xa21+cry1C*基因的同时抗白叶枯病和螟虫的改良株系各2个。
2.人工接种鉴定结果表明,只含Xa7或Xa21基因以及基因聚合的航1号改良株系的白叶枯病抗性明显提高。经自然条件下田间稻纵卷叶螟鉴定,含cry1C*基因的改良株系抗性也明显提高。
3.稻米品质分析结果表明,各供试材料的稻米品质等级难以超过对照品种扬两优6号,整精米率和垩白性状为各供试材料品质改良的主要限制因子。
4.杂交组合的杂种优势分析表明,有部分改良株系在农艺性状和配合力等方面基本保持了航1号的主要特征特性,同时筛选出金科1A/JY8021-1、巨风A/航1号、巨风A/JY8020-1、巨A/JY8022-1、巨风A/JY8024-1和巨风A/JY8232-1等六个强优势组合。
综合上述结果,本研究已成功地将抗白叶枯病基因Xa7、Xa21,抗螟虫基因cry1C*导入水稻恢复系航1号中,育成的改良株系和所配的组合白叶枯病和螟虫抗性明显提高,这些基因聚合株系具有良好的应用前景。
Bacterial blight of rice is one of the three most destructive diseases throughout the world. As an effective measure, the exploitation of host resistance genes can be used for the improvement of disease resistance. On the other hand, pests are the main bottleneck for rice production due to the lack of host resistance genes. Nowadays, we can develop resistant cultivars by transferring pests-resistant Bt gene. Hang 1 is an elite restorer line of three-line hybrid, which is widely used in the cultivation of hybrid rice. The super hybrids, Teyouhang 1 andⅡyouhang 1, which were mated using Hang 1 as male parent, have the characteristics of high yield and novel quality. In order to improve its resistance to both bacterial blight and stem bore such as Cnaphalocrocis medinalis. Xa7, Xa21 and cry1C* genes were transferred from Huahui 20 and TIC-19 respectively to the genetic background of Hang 1 by traditional breeding combining with MAS firstly, then the two-gene lines (Xa7+cry1C* or Xa21+cry1C*) and three-gene lines (Xa7+Xa21+cry1C*) were obtained by multiple cross. The specific Xanthomonas oryzae pv. oryzae strains, ZHE173 (Ⅳ) and GD1358 (Ⅴ) were used for identifying the improved lines' resistance to bacterial blight by artificial inoculation under the field condition. The experiment of resistance to stem borer for the improved lines was carried out under the field condition also. The quality traits of improved lines and their hybrids have been measured and analyzed in the research. The 9 improved lines of Hang 1 were hybridized with 5 sterile lines C815S,Ⅱ-32A, Chuannong 1 A, Jinke 1A and Jufeng A according to NCII design, and the combining ability for 8 main agronomic traits of the hybrids were investigated. The main result in the research is as follows:
1. We have obtained two improved lines with genetic background of Hang 1 which contain Xa7 and cry1C* genes, Xa21 and cry1C* genes and Xa7, Xa21 and cry1C* genes respectively.
2. The improved lines and their hybrids' resistance level to bacterial blight has been greatly improved. The improved lines and their hybrids showed high resistance level to Cnaphalocrocis medinalis.
3. The result of rice quality traits shows that all the meterials studied are generally inferior to CK (Yangliangyou 6) on quality traits, and chalky trait and milled rice rate are main limiting factors.
4. The result of usage of rice heterosis shows that only a part of improved lines' agronomic traits and combining ability are similar to Hang 1. And Jinke 1A/JY8020-1, Jufeng A/Hang 1, Jufeng A/JY8021-1, Jufeng A/JY8022-1, Jufeng A/JY8024-1 and Jufeng A/JY8232-1 were selected in the research.
All in all, Xa7, Xa21 and cry1C* genes were transferred into the genetic background of Hang 1 successfully. The improved lines and their hybrids's resistance levels to both bacterial blight and stem borer were increased obviously. These lines have a good prospect in future practical use.
引文
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59. Lynch P T, Jones J, Blackball N W, et al. The phenotypic characterization of R2 generation transgenic rice plants under field and glasshouse conditions. Euphytica,1995,85:395-491
60. Mizukami T. Studies on the ecologocal properties of Xanthomonas oryzae, the casual organism of bacterial leaf blight of rice plant. Agriculture Bulletin, Saga University,1961,13:1-85
61. Mew T W, Vera Cruz C M, Reyes R C. Pathotypes of Xanthomonas campestris pv. oryzicala in Asia. IRRI research Paper Series,1982a, No.75
62. Mew T W, Vera Cruz C M, Reyes R C. Interaction of Xanthomonas campestris pv. oryzicala and a resistance rice cultivar. Phytopathology,1982b,72(7): 786-789
63. Mew T W, Vera Cruz C M. Virulence of Xanthomonas campestris pv. oryzae in the Philippines. Phytopathology,1985,75:13-16
64. Mew T W. Current status and future prospects of research on bacterial blight of rice. Annu Rev of Phytopathology,1987,72:786-789
65. Maqbool S B, Christou P. Multiple traits of agronomic importance in transgenic indic rice plants:analysis of transgene integration patterns, expression levels and stability. Mol Breed,1999,5:471-480
66. Nayak P, Basu D, Das S, Basu A, Ghosh D, Ramakrishnan N A, Ghosh M, Sen S K. Transgenic elite Indica rice plants expressing CrylAc δ-endotoxin of Bacillus thuringiensis are resistant against yellow stem borer (Scirpophaga incertulas). Proc Natl Acad Sci USA,1997,94:2111-2116
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69. Swings J, Van den Mooter M, Mew T W et al. Reclassification of the casual agents of bacterial blight(Xanthomonas campestris pv. oryzae) and bacterial leaf streak(Xanthomonas campestris pv. oryzicala) of rice as pathovars of Xanthomonas oryzae (ex Ishiyama 1922) sp. Nov., nom. Rev. International Journal of Systematic Bacteriology,1990, p.309-311
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71. Tu J M, Zhang G A, Datta K, Xu C G, He Y Q, Zhang Q F, Khush G S, Datta S K. Field performance of transgenic elite commercial hybrid rice expressing Bacillus thuringiensis δ-endotoxin. Nat Biotechnol,2000,18:1101-1104
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53. Fujimoto H, Itoh K, Yamamoto M, Kyozuka J, Shimamoto K. Insect resistant rice generated by introduction of a modified δ-endotoxin gene of Bacillus thuringiensis. Nat Biotechnol,1993,11:1151-1155
54. Horino O, Mew T W, Khush G S et al. Resistance of Japanese and IRRI differential rice varieties to pathotypes of Xanthomonas oryzae in the Philippines and in Japan. IRRI Research Paper Series,1980,53:11
55. Ho N H, Baisakh N, Oliva N, Datta K, Frutos R, Datta S K. Translational fusion hybrid Bt genes confer resistance against yellow stem borer in transgenic elite vietnamese rice (Oryza sativa L.) cultivars. Crop Sci,2006,46:781-789
56. Khush G S, Bacalangco E, Ogawa T. A new gene for resistance to bacterial blight from O. longistaminata. RGN,1990,7:121-122
57. Kaji R, Qgawa T. Identification of the located chromosome of the resistance gene Xa7 to bacterial leaf blight in rice. Jpn. J. Breed,1995,45:79
58. Leach J F, Rhoads M L, White F F et al. A repetitive DNA sequence from Xanthomonas campestris pathovars. J. Cell Biochem,1988,12C:254
59. Lynch P T, Jones J, Blackball N W, et al. The phenotypic characterization of R2 generation transgenic rice plants under field and glasshouse conditions. Euphytica,1995,85:395-491
60. Mizukami T. Studies on the ecologocal properties of Xanthomonas oryzae, the casual organism of bacterial leaf blight of rice plant. Agriculture Bulletin, Saga University,1961,13:1-85
61. Mew T W, Vera Cruz C M, Reyes R C. Pathotypes of Xanthomonas campestris pv. oryzicala in Asia. IRRI research Paper Series,1982a, No.75
62. Mew T W, Vera Cruz C M, Reyes R C. Interaction of Xanthomonas campestris pv. oryzicala and a resistance rice cultivar. Phytopathology,1982b,72(7): 786-789
63. Mew T W, Vera Cruz C M. Virulence of Xanthomonas campestris pv. oryzae in the Philippines. Phytopathology,1985,75:13-16
64. Mew T W. Current status and future prospects of research on bacterial blight of rice. Annu Rev of Phytopathology,1987,72:786-789
65. Maqbool S B, Christou P. Multiple traits of agronomic importance in transgenic indic rice plants:analysis of transgene integration patterns, expression levels and stability. Mol Breed,1999,5:471-480
66. Nayak P, Basu D, Das S, Basu A, Ghosh D, Ramakrishnan N A, Ghosh M, Sen S K. Transgenic elite Indica rice plants expressing CrylAc δ-endotoxin of Bacillus thuringiensis are resistant against yellow stem borer (Scirpophaga incertulas). Proc Natl Acad Sci USA,1997,94:2111-2116
67. Ou S H. Bacterial Diseases. In:Rice Diseases.2nd ed. Kew, England:Commonw. Mycol. Inst.,1985,61-69
68. Ronald P C, Albano B, Tabien R, et al. Genetic and physical analysis of the rice bacterial blight resistance locus, Xa-21. Mol Gen Genet,1992,236:113-120
69. Swings J, Van den Mooter M, Mew T W et al. Reclassification of the casual agents of bacterial blight(Xanthomonas campestris pv. oryzae) and bacterial leaf streak(Xanthomonas campestris pv. oryzicala) of rice as pathovars of Xanthomonas oryzae (ex Ishiyama 1922) sp. Nov., nom. Rev. International Journal of Systematic Bacteriology,1990, p.309-311
70. Tang W, Chen H, Xu C G, Li X H, Lin Y J, Zhang Q F. Development of insect resistant transgenic indica rice with a synthetic cry1C* gene. Mol Breed,2006, 18:1-10
71. Tu J M, Zhang G A, Datta K, Xu C G, He Y Q, Zhang Q F, Khush G S, Datta S K. Field performance of transgenic elite commercial hybrid rice expressing Bacillus thuringiensis δ-endotoxin. Nat Biotechnol,2000,18:1101-1104
72. Tagami Y, Kuhara S, Kurita T et al. Epidemiological studies on the bacterial leaf blight of rice, Xanthomonas oryzae(Uyeda et Ishiyama)Dowson. I. The overwintering of the pathogen. Bulletin of the Kyushu Agricultural Experiment station,1963,9:89-122
73. Wunn J, Kloti A, Burkhardt P K, Biswas G C G, Launis K, Iglesias V A, Potrykus I. Transgenic indica rice breeding line IR58 expressing a synthetic cry1A(b) gene from Bacillus thuringiensis provides effective insect pest control. Nat Biotechnol,1996,14:171-176