水稻品种抗褐飞虱基因的定位及分子标记辅助选择
详细信息    本馆镜像全文|  推荐本文 |  |   获取CNKI官网全文
摘要
褐飞虱(Nilaparvata lugens St(?)l)又称褐稻虱,属同翅目(Homoptera)飞虱科(Delphacide),是一种世界著名的水稻单食性、迁飞性害虫。褐飞虱不仅直接刺吸稻株韧皮部或在叶鞘内产卵造成直接危害,而且还传播草状丛矮病(Grass Stunt)和齿叶矮缩病(Ragged Stunt)等病毒病。20世纪70年代以前,褐飞虱在亚洲大多数国家仅仅是水稻的一种次要害虫。但在20世纪70年代以后,随着亚洲各国水稻品种和栽培耕作措施的变化,以及大量氮肥和广谱性杀虫剂的使用,褐飞虱跃居为亚洲水稻生产的头号害虫,对亚洲各国的水稻生产造成严重危害。近30年来我国褐飞虱发生和为害具有发生面积扩大、暴发频率增加和为害程度增强等特点。
     长期以来,化学防治在减轻褐飞虱为害中发挥了重要作用。但是广谱杀虫剂在灭杀褐飞虱的同时,往往也杀死或驱赶了褐飞虱天敌,破坏了褐飞虱与天敌的生态平衡,导致褐飞虱数量大量增加;而且,有些杀虫剂,如甲基对硫磷和二嗪农等还能够刺激褐飞虱雌虫产卵。此外,化学药剂的使用不可避免地对自然环境造成污染和破坏。
     利用品种自身的抗性被认为是防治褐飞虱为害的最经济有效途径之一。然而,在育种家努力选育一个又一个抗虫品种以抵抗褐飞虱为害的同时,褐飞虱也同样通过生物型的改变一次又一次地克服了水稻品种的抗性。为了延缓褐飞虱新生物型的产生,增加抗虫品种的使用年限,科学家提出了抗虫品种轮作、不同抗虫品种合理布局以及聚合育种等策略。同时,许多研究还表明,具有中等抗性的抗虫品种或由多基因控制的抗虫品种比由单个主基因控制的抗虫品种表现出更持久的抗性。因而,不断挖掘和利用新的抗褐飞虱主基因和抗褐飞虱数量性状基因座(quantitative trait locus, OTL),选育具有持久抗虫性的水稻品种,已成为利用水稻品种自身抗性防治褐飞虱危害的首要任务。
     本研究对ASD7的抗褐飞虱主基因bph2进行了SSR定位分析及分子标记辅助选择;还利用SSR标记定位了斯里兰卡籼稻品种Rathu Heenati中的显性抗褐飞虱基因Bph3,同时分析了Col.5 Thailalld的抗褐飞虱QTL位点。有关研究结果如下:
     1.利用综合性状较好对褐飞虱敏感的粳稻恢复系C418为父本,以含有bph2基因的抗褐飞虱品种ASD7为母本构建了包含134个F_(2:3)家系的群体,利用苗期接虫鉴定法对F_(2:3)家系进行抗性鉴定,并根据F_(2:3)家系的抗虫性表现推测相应F_2单株的基因型。根据http://www.dna-res.kazusa.or.jp/9/6/05/spl-table1/table1.pdf和http://www.gramene.org/网站的Gramene数据库公布的SSR序列合成引物,进行ASD7/C418 F_2分离群体的分子标记分离分析和分子连锁图的构建。利用MAPMAKER/3.0软件对F_2作图群体进行分子标记和褐飞虱抗性之间连锁分析,将ASD7的bph2基因定位于第12染色体长臂上的两个SSR
The brown planthopper (BPH), Nilaparvata lugens (Stal), is one of the most serious insect pests of rice (Oryza sativa L.) in Asian rice growing areas. It not only causes direct damage by sucking plant sap but also transmits several viral diseases such as rice grassy stunt and rugged stunt. BPH has been called a pest of the green revolution because of the adoption pf modern agricultural practices (e.g., flooding fields, increased use of nitrogenous fertilizer, and the use of high yielding varieties). Prior to 1970, the rice BPH was not regarded as a pest of tropical rice and received only a brief mention in an illustrated field guide to rice pests in the Philippines. Since 1970s, however, this monophagous planthopper has achieved major pest status, causing massive losses in rice production throughout Asia.Protection against BPH pest insects has depended mostly on insecticides in the past. However, one common cause of insect pest outbreaks is the resurgence of insect pest population in response to inecticide application. One of the primary factors responsible for pest resurgence has long been believed to be the elimination of natural enemies as a result of inecticide application. Another implicated resurgence mechanism is the enhancement of insect fecundity by chemical stimulation, and researchers have demonstrated that certain chemicals, such as decamethrin, methyl parathion and diazinon, could stimulate reproduction of planthopper females.Outbreaks of BPH promoted the international Rice Research Institute (IRRI) to initiate a breeding program for the development of resistant varieties of rice. To slow down the evolution of BPH population, several approaches have been proposed over the years, such as rotating different cultivars, planting multiple resistant lines within the same field, and combing different resistance genes within the same cultivar. It also has been proposed that moderate and/or polygenic resistance to insect pests, including BPH, should provide more durable resistance than single major genes. Therefore, new genes and quantitive trait locus (QTL) for BPH resistance must be constantly identified and incorporated into breeding program.In this study, SSR mapping and marker-assisted selection of a brown planthopper resistance gene bph2 in ASD7. Fine mapping of Bph3 in Rathu Heenati and analysis of
    QTL for resistance to BPH in an indica rice Co1.5 Thailand were the other two main research topics in this study. The results were as folio wings:(1) In the study reported here, ASD7 harboring a BPH resistance gene bph2 was crossed to a susceptible cultivar C418, a japonica restorer line. A F2 segregating population composed of 134 plants was constructed from the cross between ASD7 and C418, and each F2 plant was selfed to obtain F2:3 line. BPH resistance was evaluated using 134 F2:3 lines, and the genotype of each F2 plant was inferred from the phenotype of corresponding F2.3 line. SSR assay and linkage analysis were carried out to detect bph2. As a result, the resistant gene bph2 in ASD7 was successfully mapped between RM7102 and RM463 on the long arm of chromosome 12, with distances of 7.6cM and 7.2 cM, respectively. Meanwhile, both phenotypic selection and marker assisted selection (MAS) were conducted in the BC1F1 and BC2F1 populations. Selection efficiencies of RM7102 and RM463 were determined to be 89.9% and 91.2% respectively. It would be very beneficial for BPH resistance improvement by using MAS of this gene.(2) A Sri Lankan indica rice (Otyza sativa L.) cultivar Rathu Heenati harboring a brown planthopper (BPH) (Nilaparvata lugens Stal) resistance gene Bph3, showed resistanqe to all the four biotypes of BPH. But Bph3 has not been mapped with molecular markers to date. In this study, we constructed a linkage map to determine the locus for BPH resistance gene, using an F2 population from a cross between 'Rathu Heenati' and a susceptible cultivar '02428'. Insect resistance was evaluated using 156 F23 lines and the genotype of each F2 plant was inferred from the phenotype of corresponding F2:3 lines. The e
引文
1. Alam S N, and M B Cohen. Detection and analysis of QTLs for resistance to the brown planthopper in a double-haploid rice population. Theor. Appl. Genet. 1998b, 97: 1370-1379.
    2. Alam S N, and M B Cohen. Durability of brown planthopper, Nilaparvata lugens, resistance in rice variety IR64 in greenhouse selection studies. Ent. Exp. Appl. 1998a, 89: 71~78
    3. Athwal D S, M D Pathak, E H Bacalangco, and C D Pura. Genetics of resistance to brown planthoppers and green leaf hoppers in Oryza sativa L. Crop Sci. 1971, 11: 747-750.
    4. Athwal D S and E H Pathak. Genetics of resistance to rice insect. In: Rice Breeding. IRRI. 1972, p375-368
    5. Besson E, G Dellamonica, J Chopin, K R Markham, M Kim, H Koh, and H Fuka. C-Glycosylflavones from rice plant involved in planthopper feeding. Phytochemistry. 1985, 24: 1061-1064.
    6. Bonman J M, G S Khush and R J Nelson. Breeding rice for resistance to pests. Ann. Rev. of Phytopathol. 1992. 30: 507-528
    7. Bosque-Perez N A, and I W Buddenlhagen. The development of host-plant resistance to insect pests: outlook for the tropics. Pp235~249. In Menken SsBJ, Visser JH, Harrewijn P. ed. Proc 8th Int Symp Insect-Plant Relationships. Dordrecht: Kluwer Academic Publishers. 1992.
    8. Bui Van Ich. The occurrence and migration of the brown planthopper in Vietnam. In: Proceedings of the international Seminar on migration and dispersal insects. September 25-28, 1991, Tsukuba, Japan. P183-204
    9. Chatterjee P B. Occurrence of brown planthopper on rice in West Bengal, India. International Rice Research Newsletter. 1978, 3(2): 12
    10. Chelliah S, and A Subramanian. A note on the chemical control of the brown planthopper, Nilaparvata lugens (Stal). of rice. Annamalai University Agricultural Research Annual 1972/1973. 1974, 4/5: 213-216.
    11. Chelliah S, and E A Heinrichs. Factors affecting insecticide-induced resurgence of the brown planthopper, Nilaparvata lugens on rice. Environ. Entomol. 1980, 9: 773-777.
    12. Chelliah S, and E A Heinrichs. Identification of insecticides that induce BPH resurgence when applied as foliar spray. International Rice Research Newsletter. 1979, 4(1): 14-15
    13. Chen X, S Temnykh, Y Xu, Y G Cho, and S R McCouch. Development of a microsatellite framework map providing genome-wide coverage in rice (Oryza sativa L.). Theor. Appl. Genet. 1997. 97: 370-380
    14. Churchill G A, R W Doerge. Empirical threshold values for quantitative trait mapping. Genetics 1994, 138: 963-971
    15. Claridge M F, and J D Hokkander. The biotype concept and its application to insect pests of agriculture. Crop Prot. 1983, 2: 85~95
    16. Claridge M F, and J D Hollander. The "biotypes" of the rice brown planthopper, Nilaparvata lugens. Ent. Exp. Appl. 1980, 27: 23-30.
    17. Claridge M F, J D Hokkander, and D Haslam. The significance of morphometric and fecundity differences between the "biotypes" of planthopper, Nilaparvata lugens. Ent. Exp. Appl. 1984, 39: 107-114.
    18. Cohen M B, S N Alam, E B Medina, and C C Bernal. Brown planthopper, Nilaparvata lugens, resistance in rice cultivar IR64: mechanism and role in successful N. Lugens management in central Luzon. Philippones. Ent. Exp. Appl. 1997, 85: 221~229.
    19. Cook A G, and R F Denno. Planthopper/plant interactions: Feeding behavior, Plant nutrition, plant defense and host plant specialiazation. Pp114-138. In Robert F. Denno and T. John perfect, ed. Planthoppers: their ecology & management. New York: Chapman & Hall, 1994
    20. Croft B A, and A W A Brown. Response of arthropod natural enemies to insecticides Ann. Rev. Entomol. 1975, 20: 285-336
    21. Das N M, and M J Thomas. Effect of water level in rice fields on the population build up of the brown planthopper, Nilaparvata lugens, and on the incidence of hopper burn. Agricultural Research Journal of Kerala. 1977, 15 (1): 104-105
    22. Das N M, K V Mammen, and S P Christudas. Occurrence of Nilaparvata lugens (Delphacidae: Homopera) as a serious pest of paddy in Kerala, India. Agricultural Research Journal of Kerala. 1972, 10 (1): 191-192.
    23. Dellaporta S L, T Wood, and T B Hicks., 1983. A plant DNA mini preparation: version Ⅱ. Plant Mol Bioi Rep. 1: 19-21
    24. Demayo C C, R C Saxena, and A A Barrion. Allozyme variation in local populations of brown planthopper, Nilaparvata lugens (Stal) in the Philippines. Philipp. Ent. 1990, 8: 737~748
    25. Duan X, X Li, and Q Xue. Transgenic rice plants harboring an introduced potato proteinase inhibitor Ⅱ gene are insect resistant. Nature Biothchnol. 1996, 14: 494-498.
    26. Dyck V A, and B Thomas. The brown planthopper problem. P3-17. In Brown planthopper: threat to rice production in Asia. International Rice Research Institute, Los Banos, Philippines. 1979.
    27. Gallagher K D, P E Kenmore, and K Sogawa. Judicial use of insecticides deter planthopper outbtreaks and extend the life of resistant varieties in southeast Asian Rice. Pp599~614. In Robert F. Denno, T. John perfect, ed. Planthoppers: their ecology & management. New York: Chapman & Hall, 1994.
    28. Hatchett J H, and R L Gallum. Genetics of the ability of the Hessian fly, Mayetiola destructor, to survive on weat having different genes for resistance. Ann. Entomol Soc. Am. 1970, 63: 1400-1407.
    29. Heinrichs E A, F G Medrano, and H R Rapusas. Genetic evaluation for insect resistance in rice International Rice Research Institution. Los Banos, Philippines. 1985.
    30. Heinrichs E A, W H Reissig, S Valencia, and S Chelliah. Rates and effect o fresurgence-inducing insecticides on population of Nilaparvata lugens (Homoptera: Delphacidae) and its predators. Envrion. Entomol. 1982a, 11: 1269-1273.
    31. Heinrichs E A. Perspectives and directions for the continued development of insect-resistant rice varieties. Agric. Ecosystems Environ. 1986, 18: 9~36.
    32. Heinrichs E A. Impact of insecticides on the resistance and resurgence of rice planhoppers. Pp571~599. In Robert E Denno, T. John perfect, ed. Planthoppers: their ecology & management. New York: Chapman & Hall. 1994.
    33. Hirabayashi H, and T Ogawa. Identification and utilization of DNA markers linked to genes for resistance to brown planthopper (BPH) in rice. Recent. Adv. Breed. Sci. 1999, 41: 71-74 (in Japanese).
    34. Hirabayashi H, and T Ogawa. RFLP Mapping of Bph-1 (Brown planthopper resistance gene) in rice. Breed. Sci 1995, 45: 369-371.
    35. Ho V. C. Brown planthopper management in south Vietnam. Intercountry forcasting system and management for brown planthopper in east asia. May 19-21, 1999. Suwon, Korea. 121-128
    36. Hollander J D, and P K Pathak. The genetics of the "biotypes" of the rice brown planthopper, Nilaparvata lugens. Ent. Exp. Appl. 1981, 29: 76~86
    37. Huang N, A Pareo, and T Mew. RFLP mapping of Bph-lisozymes, RAPD and QTLs for grain shape, brown planthopper resistance in a DH population rice population. Mol. Breed. 1997, 3: 105-113.
    38. Huang Z, G He, L Shu, X Li, and Q Zhang. Identification and mapping of two brown planthopper resistance genes in rice. Theor. Appl. Genet. 2001, 102: 929-934
    39. Huynh N V. Brown planthopper and white-backed planthopper infestations in the Mekong Delta (Vietnam). The Rice Entomology Newsletter. 1975, 2: 4
    40. Ikeda R, and C Kaneda. Genetic analysis of resistance to brown planthopper, Nilaparvata lugens (Stal), in rice. Japan. J. Breed. 1981, 31(3): 279-285.
    41. Ikeda R, and C Kaneda. Trisomic analysis of resistance to brown planthopper, Nilaparvata lugens Stal, in rice. Japan. J. Breed. 1983, 33: 40-44.
    42. Ikeda R, and D A Vaughau. The distribution of resistance genes to the brown planthopper in rice germplasm. RGN. 1991, 8: 125.
    43. Ikeda R. Studies of the inheritance of resistance to the rice brown planthopper (Nilaparvata lugens stal) and the breeding of resistance rice cultivar. Bull. Natl. Agric. Res. Cent. 1985, 3: 1-54 (in Japanese)
    44. IRRI. A brown planthopper population that feed on IR36. Ann. Rep. For 1981, 1982, 56~58
    45. IRRI. Standard evaluation Systems for rice. IRRI, Manila, Philippines. 1988
    46. Ishii T, D S Brar, and D S Multani. Molecular tagging of genes for brown planthopper resistance and earliness introgressed from Oryza australiensis into cultivated rice, O. Sativa. Genome, 1994, 37: 217~221.
    47. Jeon Y H, S N Ahn, H C Choi, T R Hahn, and H P Moon. Identification of a RAPD marker linked to a brown planthopper resistance gene in rice. Euphytica. 1999, 107: 23-28.
    48. Jongsma M R, P L Bakker, and J Peters. Adaptation of spodoptera exigua larvae to plant proteirtase inhibitors by induction of gut proteinase activity insensitive to inhibition. PNAS. 1995, 92: 8041-8045
    49. Kabir M A, and G S Kush. Genetic analysis of resistance to brown planthopper in rice (Oryza sativa L.). Plant Breed 1988, 100: 54-58.
    50. Kabir M A, and G S Kush. Genetic analysis of resistance to brown planthopper in rice (Oryza sativa L.). Plant Breed 1988, 100: 54-58.
    51. Kaneda C, H Nemoto, R Ikeda and M Yokoo. A Kobayashi. Breeding of rice Norin-PL3, a new germplasm with brown planthopper resistance. Bull. Natl. Agric. Res. Cent. 1985, 5: 93-103 (in Japanese)
    52. Kaneda C, H Nemoto, R Ikeda. M Yokoo, and A Kobayashi. Ikehashi H. Takita T. Breeding of rice Norin-PL4, a new germplasm with brown planthopper resistance gene 'bph2'. Bull. Natl. Agric. Res. Cent. 1986, 6: 19-32 (in Japanese)
    53. Kaneda C, K Ito and R Ikeda. Screening of rice cultivars for resistance to the brown planthopper, Nilaparvata lugens (Stal). by three biotypes. Jpn. J. Breed. 1981, 31 (2): 141-151
    54. Kaneda C, R Ikeda, and A Kobayashi. 'Kanto PL4' and 'Kanto PL5' New parental lines of japonica rice resistant to the brown planthopper (Nilaparvata lugens). Japan J. Breed. 1980, 30 (Suppl. 2): 100-101 (in Japanese)
    55. Kaneda C, R Ikeda, and A Kobayashi. New parental lines of rice, 'Kanto PL1' and 'Kanto PL2' resistant to the brown planthopper. Japan J. Breed 1979, 29 (Suppl. 1): 74-75 (in Japanese)
    56. Kawaguchi M, K Murata, T Ishii, S Takumi and N Mori. Assignment of a Brown Planthopper (Nilaparvata lugens Stal) Resistance Gene bph4 to the Rice Chromosome 6. Breed. Sci. 2001, 51: 13-18.
    57. Kenmore P E, F O Carino, C A Perez, V A Dyck, and A P Gutierrez. Population regulation of the rice brown planthopper (Nilaparvata lugens Stal) within rice fields in the Philippines. J Plant Prot. Tropics.1984, 1: 19-37
    58. Kessler A, and I T Baldwin. Defensive function of herbivore-induced plant volatile emission in nature. Science. 2001, 291: 2141~2144
    59. Khuong Q V. Ecological features and integrated management of brown planthopper in the Red river Delta. Intercountry forecasting system and management for brown planthopper in East Asia. May 19-21, 1999, Suwon, Korea, P37-52
    60. Khush G S, A N M Rezaul Karim, and E R Angeles. Genetics of resistance of rice cultivar AR10550 to Bangladesh brown planthopper biotype. J. Genet. 1985, 64: 121-125.
    61. Khush G S, and D S Brar. Genetics of resistance to insects in crop plants. Advances in Agronomy, Academic Press, Inc. 1991. 45: 224-228.
    62. Khush G S, Disease and insect resistance in rice. Advances in Agronomy, New York: Academic Press. 1977, 29: 265-341
    63. Khush G S. Breeding rice for resistance to insects, Prot. Ecol. 1984, 7: 147-165
    64. Khush G S., 1979. Genetics of and breeding resistance to the brown planthopper. In: Brown Planthopper: Threat to Rice Production in Asia. IRRI, Los Banos, Philippines. p321-332
    65. Kisimoto R. Development, behaviour, population dynamics and control of the brown planthopper, Nilaparvata lugens Stal. Rev. Plant Prot. Res. 1981, 14: 26-58.
    66. Kogan M and E E Ortman. Antixenosis-a new term proposed to replace Painter's 'nonpreference' modality of resistance. Bull. Entomol. Soc. Am. 1978, 24: 175-176.
    67. Kuno E. Ecology of the brown planthopper in temperate regions. P 45-60. In Brown planthopper: threat to rice production in Asia. International Rice Research Institute, Los Banos, Philippines. 1979.
    68. Lakshiminarayana A and G S Khush. New genes for resistance to the brown planthopper in rice. Crop Sci. 1977, 17: 96-100.
    69. Lander E S, P Green, J Abrahamson, M J Barlow, M J Daly, S E Lincoln and L Newburg. An interactive computer for constructing primary genetics linkage maps of experimental and natural populations, Genomics. 1987. (1): 174-181
    70. Lee S I, S H Lee, J C Koo, H J Chun, C O Lim, J H Mun, Y H Song, and M J Cho. Soybean Kunitz trypsin inhibitor (SKTI) confers resistance to the brown planthoper (Nilaparvata lugens Stal) in transgenic rice. Molecular Breeding. 1999, 5: 1-9.
    71. Lincoln SE, Daly MJ, Lander ES, et al. Mapping genes controlling quantitative traits using MAPMAKER/QTL version 1.1: A tutorial and reference manual. A Whitehead Institute for biomedical research technical report third edition. Cambridge, Mass, 1993
    72. Ling. K. C, E. R. Tiongco, and V. M Aguiero Rice ragged stunt a new virus disease. Plant. Dis. Rep. 1978, 62, 701-705
    73. Maliepaard C, N Bas, H S Van, S Heusden, J Kos, G Pet, R Verkerk, R Vrielink, P Zabel, and L Newburg. Mapping of QTLs for glandular trichome densities and Trialeurodes vaporariorum (greenhouse whitefly) resistance in an F2 from Lycopersicon esculentum × Lycopersicon hirsutum f. Glubratum. Heredity. 1995, 75: 425~433.
    74. Maqbool S B, S Riazuddin, N T Loc, A M R Gatehous, J A Hatehous, and P Christou. Expression of multiple insecticidal genes confers broad resistance against a range of different rice pests. Molecular Breeding. 2001, 7: 85-93.
    75. Martinez C R, and G S Khush. Sources and Inheritance of Resistance to Brown Planthopper in Some Breedigng Lines of Rice. Crop Science. 1974, 14:264~267.
    76. McCouch. S.R., L. Teytelman, Y. Xu, et al. Development and Mapping of 2240 New SSR Markers for Rice (Oryza sativa L.) (Supplement). DNA Research. 2002. 9: 257 - 270
    77. Multani D S, K K Jena, and D S Brar. Development of monosomic alien addition lines and introgression of genes from Oryza australiensis Domin. to cultivated rice, O. sativata L . Theor. Appli. Genet. 1994,88:102-109.
    78. Murata K, M Fujiwara, C Kaneda, S Takumi, N Mori and C Nakamura. RFLP mapping of a brown planthopper (Nilaparvata lugens Stal) resistance gene bph2 of indica rice introgressed into a japonica breeding line 'Norin-PL4'. Genes Genet. Syst. 1998a, 73: 359-364.
    79. Murata K, M Fujiwara, H Murai, S Takumi, N Mori and C Nakamura. Bph9, a dominant brown planthopper resistance gene, is located on the long arm of chromosome 12. Rice Genetics Newsletter. 2000, 17: 84 - 86
    80. Murata K, M Fujiwara, Nakamura C, N Mori, and C Kaneda. Mapping of brown planthopper resistance genes bph2 and Bph9 in rice. Journal of Crop Science and Breeding. 1998b, 43:4 -7. (in Japanese)
    81. Murata K. Molecular linkage analysis of brown planthopper resistance genes in rice, Doctoral Dissertation, The graduate School of Science and technology Kobe University, 1998 p64-78
    82. Nagata T, and T Hayakawa. Activity of aconiti acid and oxalic acid on brown planthopper, Nilaparvata lugens (Stal), and green rice leafhopper, Nephoterrix cincticeps (Uhler). Jpn, J. Appl. Entomol. Zool. 1998,42: 115-121
    83. Nemoto H, M Yokoo, C Kaneda, E Shimura, R Ikeda, and A Kobayashi. Breeding of rice Norin-PL7, a new germplasm with the brown planthopper resistance gene 'bph4'. Bull. Natl. Agric. Res. Cent. 1988, 9: 93-105 (in Japanese)
    84. Nemoto H, M Yokoo, Kaneda C, E Shimura, R Ikeda, and A Kobayashi. Breeding of rice Norin-PL7, a new germplasm with the brown planthopper resistance gene 'bph3'. Bull. Natl. Agric. Res. Cent. 1989b, 15: 47-57 (in Japanese)
    85. Nemoto H, R Ikeda, and C Kaneda. New genes for resistance to brown planthopper, Nilaparvata lugens (Stal), in rice]. Japan. J. Breed. 1989a, 39:23-28.
    86. Ooi P A C, and K L Heong. Operation of a brown planthopper surveillance system in the Tanjung Karang Irrigation Scheme in Malaysia. Crop Prot. 1988, 7:273-278.
    87. Painter R H. Biological strains of Hessian fly. J. Econ. Entomol. 1930, 23: 322-326.
    88. Painter R H. Insect resistance in crop plants. Macmillan, New York. 1951.
    89. Pathak M D, and G S Khush. Studies of varietal resistance in rice to the brown planthopper at the international rice research institute. Pp285- 301. In Brown Planthopper: Threat to Rice Production in Asia. IRRI, Los Banos, Philippines. 1979.
    90. Pathak M D, and R C Saxena. Breeding approaches in rice. Pp 421-455. In F G Maxwell and P E Jenniings, ed. Breeding Plants Resistant to Insects. Wiley, New York. 1980.
    91. Powell K S, A M R Gatehouse, and V A Hilder. et al. Different antimetabolic effects of related lectins towards nymphal stages of Nilaparvata lugens. Entomol. Exp. Appl. 1995, 75:61-65
    92. Powell K S, J Spence, and M Bharathi. et al. Immunohistochemical and developmental studies to elucidate the mechanism of action of the snow drop lectin on the rice brown planthopper, Nilaparvata lugens (Stal). J. Insect Physiol. 1998,44: 529-539
    93. Pusztai A, S W B Ewen, and G Grant. et al. Lections and also bacteria modify the glycosylation of gut surface receptors in the rat. Glycoconjugate J. 1995,12: 22-35
    94. Rao K V, K S Rathore, T K Hodges, X Fu, E Stoger, D Sudhakar, S William, P Christou, M Bharathi, D Bown, K S Powell, J Spemce, A M R Gatehouse, and J A Gatehouse. Expression of snowdrop lectin (GNA) in transgenic plants confers resistance to rice brown planthopper. The Plant Journal. 1998,15 (4): 469-477.
    95. Reissig W H, E A Heinrichs, and S L Valencia. Insecticide induced resurgence of the brown planthopper, Nilaparvata lugens on rice varieties with different levels of resistance. Environ. Entomol. 1982,11:165-168.
    96. Rivera. C.T, S.H. Ou and T.T. Lida Grassy stunt disease of rice and its transmission by Nilaparvata lugens (Stal). Plant. Dis. Rep. 1966, 50:453-456
    97. Roderick G K. Genetics of host plant adaptation in Dephacid planthoppers. Pp551~570. In Robert F. Denno and T. John perfect, eds. Planthoppers:their ecology & management. Chapman & Hall, New York. 1994
    98. Sakai T, and K Sogawa. Effects of nutrient compounds on sucking response of the brown planthopper. Nilaparvata lugens(Homoptera: Delphacidae). Appl. Entomol. Zool. 1976,11:82-88
    99. Sanguinetti C J, N E Dias, and A J G Simpson. Rapid silver staining and recover of PCR products separated on polyacrylamide gels. Biotechniques. 1994. 17: 915-919
    100. Saxena R C, and A A Barrion. Biotypes of the brown planthopper, Nilaparvata lugens (Stal) and strategies in development of host plant resistance. Insect Sci. Applic. 1985, 6(3): 271-289.
    101. Saxena R C, and Z R Khan. Factors affecting resistance of rice varieties to planthopper and leafhopper pests. Agri. Zool. Rev. 1989,3: 97 - 132.
    102. Saxena R C, C C Demayo, and A A Barrion. Allozyme variation among biotypes of the brown planthopper, Nilaparvata lugens in the Philippines. Biochem. Gen. 1991,314:115-123
    103. Saxena R C, M V Velasco, and A A Barrion. Morphological variation between brown planthopper biotypes on leersia hexandra and rice in the Philippines. Int. Rice Res. Newsl. 1983, 8 (3): 3.
    104. Sekido S, and K Sogawa. Effects of salicylic acid on probing and oviposition of the rice plant-and leafhoppers (Homoptera: Delphacidae and Deltocephalinae). Appl. Entomol. Zool. 1976, 11: 75~81
    105. Shepard B M, Z R Khan, M D Pathak, and E A Heinrichs. Management of insect pests of rice in Asia: P255-278. In D. Pimentea, ed. CRC handbook of Pest Management in Agriculture. Second edition. CRC Press. Boca Raton, FL. 1990
    106. Shigematsu Y, N Murofushi, K Ito, C Kaneda. S Kawabe, and N Takahashi. Sterol and asparagines in the rice plant, endogenous factors related to resistance against the brown planthopper, Nilaparvata lugens. Agric. Biol. Chem. 1982, 46: 2877-2896.
    107. Shufran K A, and M E Whalon. Genetics analysis of brown planthopper biotypes using random amplified DNA-polymerase chain reaction (RAPD-PCR). Insect Sci. Appl. 1995, 16(1): 27~33
    108. Sidhu G S and G S Khush. Linkage relationships of some genes for disease and insect resistance and semidwarf stature in rice. Euphytica. 1979, 28: 233-237
    109. Sidhu G S, and G S Khush. Genetic analysis of brown planthopper resistance in twenty varieties of rice, Oryza sativa L. Theor. Appl. Genet. 1978, 53: 199-203.
    110. Sogawa K, and M D pathak. Mechanism of brown planthopper resistance in Mudgo vatiety of rice (Hemiptera Delphacidae). Appl. Entomol. Zool. 1970, 5: 145~158
    111. Sogawa K. Overseas immigration of rice planthopper into Japan and associated meteorological systems. In. Proceedings of China-Japan Joint Workshop on "Migration and Management of Insect Pest of Rice in Monsoon Asia", China National Rice Research Institute (ed.). 1997. P13-35. November 27-29, Hangzhou, P. R. China.
    112. Sogawa K. The rice brown planthopper: Feeding physiology and host plant interactions. Annu. Rev. Entomol. 1982, 27: 49~73
    113. Su C. C., X. N. Cheng, H. Q. Zhai, J. M. Wan Detection and analysis of QTL for resistance to brown planthopper, Nilaparvata lugens (Stal), in rice (Oryza sativs L.), using backcross inbred lines. Acta. Genetica. Sinica. 2002, 29 (4): 332~338 (in Chinese)
    114. Su C. C., J. Wan, H. Q. Zhai, C. M. Wang, L. H. Sun, H. Yasui, A. Yoshimura A new locus for resistance to brown planthopper in DV85, an indica rice (Oryza. Sativa L.). Plant. Breeding. 2005, 124: 93-95
    115. Sudhakar D, X Fu, E Stoger, S Williams, M Barathi, J Gatehouse, and P Christou. Expression and immunolocalization of the snowdrop lectin insecticidal protein GNA, in transgenic rice plants. Transgenic Res. 1998, 7(4): 371-378
    116. Tanaka K. Quantitative genetic analysis of biotypes of the brown planthopper Nilaparvata lugens: heritability of virulence to resistance rice varieties. Ent. Exp. Appl. 1999, 90: 278~287.
    117. Tang K X, X F Sun, J H Yao, H X Qi, and X G Lu. Studis on Resistance to Brown Planthopper Transgenic Rice Pure Lines. Journal of Fudan University (Natural Sciences). 2000, 39(4): 436-440.
    118. Temnykh, S., W. D. Park, N. Ayres, S. Cartinhour, N. Hauck, L. Lipovich, Y. G. Cho, T. Ishii and S. R. McCouch Mapping and genome organization of microsatellite sequence in rice (Oryza sativa L.). Theor. Appl. Genet. 2000. 100: 697-712
    119. Thomas B. Studies on the varietal resistance to the brown planthopper (Nilaparvata lugens) in Kerala. The Rice Entomology Newsletter. 1979, 4: 10-11.
    120. Tooyama T, T Yamamoto, T Tsuji. Chromosomal location of the brown planthopper resistance gene Bph-1 revealed by RFLP mapping. Breed Sci. 1995. 45(Suppl. 2): 171. (in Japanese).
    121. Turlings T C, J H Loughrin, P J McCall. How caterpillar-damaged plants protect themselves by attracting parasitic wasps. PNAS. 1995, 92: 4169~4174
    122. Wang, S., C. Basten and Z.-B. Zeng. Window QTL Cartographer, V2.0. North Carolina State University. Raleigh, NC. 2003
    123. Xu D, Q Xue, D McElroy. et al. Constitutive expression of a cowpea trypsin inhibitor gene, Cpti, transgenic rice plants confers resistance to two major rice insect pests. Mol. Breed. 1996, 2: 167-173
    124. Xu X F, H W Mei, L J Luo, X N Cheng, and Z K Li. RFLP-facilitated investigation of the quantitative resistance of rice to brown planthopper (Nilaparvata lugens). Theor Appl. Genet. 2002, 104: 248-253
    125. Yoshihara T, K Sogawa, M D Pathak, B Juliana and S Sakamara. Soluble silicic acid as a sucking inhibitory substance on rice against the brown planthopper (Hemiptera Delphacidae)). Entomol Exp. Appl. 1970, 26: 314-322
    126. Zheng K L, Huang N, John B, et al. PCR basedmarker assistedselection in rice breeding. IRRI discussion paper series No12. 1995, Manila, Philippines
    127.程遐年 等.稻褐飞虱迁飞规律的研究.昆虫学报.1979,22(1):1-21
    128.程遐年,浦茂华,陈钰英.褐飞虱防治理论研究与进展,南京:江苏科学技术出版社,1993
    129.程遐年,吴进才,马飞.编著.褐飞虱研究与防治.北京:中国农业出版社,2003.
    130.丁锦华,张富满.东北飞虱志同翅目飞虱科.北京:中国农业科技出版社.1994,83-85
    131.葛钟麟,丁锦华,田立新等.中国经济昆虫志同翅目飞虱科.北京:科学出版社,1984,138
    132.胡伯海,姜瑞中.农作物病虫长期运动规律与预测.北京:中国农业出版社.1997,7-12
    133.胡国文,唐启义,马巨法等.中国褐飞虱的分布和为害.昆虫知识.1997,34(1):50-51
    134.蒋志农 主编,云南稻作.昆明:云南科技出版社.1995,88-89
    135.李容柏,秦学毅,韦素美,黄凤宽,李青,罗善昱.普通野生稻抗源94-42-5-1对稻褐飞虱的抗性评价及其遗传研究.中国水稻科学,2002,16(2):115-118
    136.刘国庆,颜辉煌,傅强,钱前,张志涛,翟文学,朱立煌.栽培稻的紧穗野生稻抗褐飞虱主效基因的遗传定位.科学通报.2001,46(9):738-742.
    137.吕仲贤,俞哓平,陶林勇,巫国瑞,陈建明,郑许松,徐红星.水稻新品种(系)对褐飞虱抗性的评价.中国农业科学,2002,35(2):225~229.
    138.王布哪,黄臻,舒理慧,任翔,李香花,何光存.两个来源于野生稻的抗褐飞虱新基因的分子标记定位.科学通报.2001,46(1):46~49.
    139.王桂荣,樊叶杨,庄杰云,郑康乐,张志涛.稻褐飞虱的遗传变异性分析.昆虫学报,2001,44(1):123~126
    140.巫国瑞,俞晓平,陶林勇.褐飞虱和白背飞虱灾害的长期预测.中国农业科学.1997,30(4):25~29
    141.阳体冰.粳稻抗褐飞虱育种中亲本选配问题.江苏农业科学,1982,(12) 1-5
    142.应存山 主编.中国稻种资源.北京:中国农业科技出版社.1993
    143.中华人民共和国国家统计局和中华人民共和国民政部.中国灾情报告.北京:中国统计出版社,1995,119-120