分类号 S511 学 号 2008201068 博 士 学 位 论 文 两个水稻品种抗灰飞虱的遗传分析 LE QUANG TUYEN 黎光泉 指导教师 万建民 教授 专业名称 作物遗传育种 研究方向 分子遗传育种 答辩日期 二○一一年六月 Genetic analysis for the resistance to small brown planthopper (Laodelphax striatellus Fallén) in two rice varieties By Le Quang Tuyen A Dissertation Submitted to Nanjing Agricultural University In Partial Fulfillment of the Requirements for the Doctoral Degree Supervised by Professor Wan Jianmin College of Agronomy Nanjing Agricultural University Nanjing 210095, P R China 原创性声明 本人郑重声明：所呈交的学位论文、是本人在导师的指导下、独立进行研究工作所取 得的成果。除文中已经注明引用的内容外、本论文不包含任何其他个人或集体已经发表或撰 写过的作品成果。对本文的研究做出重要贡献的个人和集体、均已在文中以明确方式标明。 本人完全意识到本声明的法律结果由本人承担。 学位论文作者（需亲笔）签名： 年 月 日 学位论文版权使用授权书 本学位论文作者完全了解学校有关保留、使用学位论文的规定、同意学校保留并向国家 有关部门或机构送交论文的复印件和电子版、允许论文被查阅和借阅。本人授权南京农业大 学可以将本学位论文的全部或部分内容编入有关数据库进行检索、可以采用影印、缩印或扫 描等复制手段保存和汇编学位论文。 保密□、在 年解密后适用本授权书。本学位论文属于不保密□。 （请在以上方框内打“√”） 学位论文作者（需亲笔）签名： 年 月 日 导师（需亲笔）签名： 年 月 日 目 录 目 录 摘 要 .I ABSTRACT III 英文缩略表 V 第一章 文献综述 第一节 稻飞虱的生物学特性及其危害 1 灰飞虱 Laodelphax striatellus (Fallén) 1.1 灰飞虱的形态特征 1.2 灰飞虱的生活习惯特性 .2 1.3 灰飞虱的分布、发生及危害 1.3.1 灰飞虱对水稻直接取食造成损失 .3 1.3.2 灰飞虱对水稻条纹病毒病的传毒机制及其危害 .4 1.3.3 灰飞虱对水稻黑条矮缩病病毒的传毒特性及其危害 1.4 灰飞虱的遗传多样性研究 褐飞虱 Nilaparvata lugens (Stål) 2.1 褐飞虱的生物学特性 2.2 褐飞虱在世界的分布和危害 白背飞虱 Sogatalla furcifera (Horváth) 11 3.1 白背飞虱的生物学特性 11 3.2 白背飞虱的分布及其危害 .11 第二节 水稻抗稻飞虱遗传基础研究 .12 水稻抗褐飞虱遗传基础研究 12 水稻品种对白背飞虱抗性的遗传分析 14 水稻抗灰飞虱遗传基础研究 15 第三节 植物抗虫的三种类型 .16 植物对害虫的抗生性机制 .16 植物对害虫的趋避性机制 .16 植物的耐虫性机制 17 第四节 本研究的目的及意义 .19 第二章 水稻品种 Rathu Heenati 抗灰飞虱基因的分子定位 21 材料与方法 .22 i 两个水稻品种抗灰飞虱的遗传分析 1.1 材料 .22 1.2 灰飞虱的饲养与繁殖 .23 1.3 抗灰飞虱鉴定 23 1.3.1 苗期集团接种法 （Standard Seedling-box Screening Test） 23 1.3.2 趋避性检测 .25 1.3.3 抗生性检测 .25 1.4 DNA 样品制备 26 1.5 SSR 标记分析 26 1.5.1 试剂 .26 1.5.2 PCR 反应体系 27 1.5.3 PCR 反应程序 27 1.5.4 SSR 标记的 PCR 产物检测 27 1.6 连锁图谱的构建 27 1.7 QTLs 定位 28 1.8 数据分析 28 结果与分析 .29 2.1 亲本 Rathu Heenati 和 02428 表现 29 2.1.1 亲本和对照品种的抗虫性表现 29 2.1.2 SSR 分子标记在基因组的分布频率 30 2.2 利用 02428/Rathu Heenati F2 群体检测灰飞虱抗性 QTL 30 2.2.1 F2 群体分子遗传连锁图谱的构建 30 2.2.2 苗期集团鉴定检测 F2 群体中抗灰飞虱 QTL。 32 2.2.3 F2 群体中灰飞虱趋避性相关 QTL 检测 35 2.2.4 抗生性 QTL 定位 36 2.3 利用 02428/Rathu Heenati//02428 BC1 群体分析水稻抗灰飞虱 QTL 38 2.3.1 BC1 分子遗传连锁图谱的构建 .38 2.3.2 苗期集团鉴定检测抗灰飞虱 QTL .40 2.3.3 BC1 群体趋避性相关 QTL 检测 42 2.3.4 BC1 群体灰飞虱抗生性相关 QTLs 的检测 43 讨论 43 3.1 水稻第 12 染色体存在一个抗灰飞虱主效 QTL 43 3.2 水稻第 12 染色体 RM519-RM3331 之间存在稻飞虱基因簇该区间与吸汁类 害虫抗性密切相关 .45 第三章 利用昌恢 891/02428 F2 群体检测灰飞虱抗性 QTL 47 ii 目 录 材料与方法 .48 1.1 植物材料 48 1.2 灰飞虱的饲养与繁殖 48 1.3 试验方法 48 结果与分析 .49 2.1 苗期集团鉴定法检测抗灰飞虱 QTLs .49 2.1.1 亲本昌恢 891 和 02428 抗性表现 .49 2.1.2 149 个 02428/昌恢 891 F2：3 家系对灰飞虱抗性表现 .49 2.1.3 抗灰飞虱 QTL 定位 50 2.2 抗生性测验及 QTL 定位 51 2.3 趋避性抗性相关 QTL 的检测 .52 讨论 53 第四章 全文结论 55 参考文献 .57 在读期间发表论文 .73 附 录 75 致 谢 77 iii 摘 要 两个水稻品种抗灰飞虱的遗传分析 摘 要 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 Laodelphax striatellus Fallén几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几几几几几几几几几几几几几(QTL)几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几 Rathu Heenati 几几几 891 几几几几几几几几几几几几几几几几几几几几 1几几几几几几 Rathu Heenati 几几几几几几几几 02428 几几几几几几几几几 162 几几几几 02428/Rathu Heenati F2 几 150 几几几几 02428/Rathu Heenati//02428 BC1 几几几几几几几 162 F2几3 几几几 150 几 BC1F2 几几几几几几几几几几几几几几几几几几几几几几几几几 Windows QTL Cartographer 2.5 几几几几几几几几 F2 几几几几几几 几几几几 QTLs几qSBPH2-b1, qSBPH5b 几 qSBPH6-b几几几几几几 2几5 几 几几几几几几几几几几几几几 30.75%几几 BC1 几几几几几几 几几 QTL几qSBPH2-b2几几几几几 几几几几几几几几几几几几 6.61%几几几几几几几几几几 F2 几 几几几几几 几 QTL几qSBPH8-c, qSBPH9-c 几 qSBPH12-c几几LOD 几几 2.49几2.59 几 2.56几几几几几几几几几 21.87%几几 BC1 几几几几几几几 几 QTL几qSBPH5-c几几LOD 几几 2.34几几几几几 7.21%几几几几几几几几几几几几几 F2 几几几几几几几几 几 QTL几qSBPH1-a） qSBPH2-a）qSBPH4-a）qSBPH12-a2 几几几几几 qSBPH12-a1几几几几几几几 1几2几4 几 12 几几几几LOD 几几几几 4.45几2.40几3.77几2.57 几 18.99几几几几几 7.6%几4.2%几9.8%几 4.47%几 37.87%几几几几几几几几 几 QTLs 几几几几几几几 Rathu Heenati几几几几几几几 qSBPH12-a1 几 qSBPH4-a 几 BC1 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 几几几几几 QTLs几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 2几几几几几几几几几几几 891 几几几几几 02428 几几几几几几 149 几几几几 F2 几几几几几几几 几几几 149 几 F2:3 几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几几 I 两个水稻品种抗灰飞虱的遗传分析 几 几几几几几几几几 几几几几几 QTL 几几 qSBPH6-a 几 qSBPH7-a几几 LOD 几几 2.61 几 2.91几几几几几 8.75%几 13.01%几几几几几几几几几几几几 891几几几 几 几几几几几几几几 几 几几几 QTL几qSBPH1-c 几 qSBPH2-c几LOD 几几几几 2.40 几 4.11几几几几几 15.03%几 15.13%几几几几几几几几几几几几几几几 QTL几几几几几几几几几 891 几几几几几几几几几几几几几几 几几几几几几几 关键词:几几(Oryza sativa L.)几几几几几QTL几几几几几几几几几几几几 II ABSTRACT Genetic analysis for the resistance to small brown planthopper (Laodelphax striatellus Fallén) in two rice varieties ABSTRACT Rice (Oryza sativa L.) is the staple food for more than 50% of the world’s population，although insect pests are the major biotic constraints to production of this crop The Small brown planthoper，Laodelphax striatellus Fallén (Homoptera: Delphacide)， is one of the most destructive and wide spread insect pests found throughout the temperate rice-growing regions such as China，Japan and Korea The population of Laodelphax striatellus Fallén has increased steadily due to alteration of the cropping system and infested South East China The dults and nymphs of Laodelphax striatellus Fallén suck rice sap causing yellowing of leaves，wilting and eventually death resulting in yield loss.The pest also transmits viral diseases such as Rice stripe virus，and rice black streaked dwarf virus causing further yield loss Chemicals have mostly been used to control Laodelphax striatellus Fallén，although it has developed resistance to most of them Use of these chemicals has also resulted into death of natural enemies and pollution of the environment leading to the resurgence of the pest Development of resistant varieties is the most effective and economical way in controlling this pest The feeding behavoiur of the pest was evaluated on two rice varieties bearing resistance genes derived from three basic resistance mechanisms : antibiosis，antixenosis and tolerance by identifying the resistance genes in rice varieties with the help of the QTL A set of F2 and BC1 population derived from the cross between 02428 and Rathu Heenati were used to investigate the small brown planthopper resistance loci Using the F2 population，three QTLs for antixenosis resistance were located on chromosome 2，5 and 6，respectively accounting for 30.75% of the phenotypic variance The QTL on chromosome was also identified in BC1 population Three QTLs for antibiosis against the pest were detected on chromosome 8，9 and 12，respectively in F2 population qSBPH5-c explaining 7.21% of phenotypic variance for antibiosis against the insect was identified on chromosome using BC1 population A major QTL qSBPH12-a1 explained about 40% of phenotypic variance and a minor QTL (qSBPH4-a) were detected by SSST method using both the F2 and BC1 population The QTLs identified in present study will be useful for III 两个水稻品种抗灰飞虱的遗传分析 mark assisted selection for SBPH resistance in rice The Japonica rice 02428 and the indica rice Changhui891，was used to detect quantitative trait loci (QTLs) for the resistance to SBPH Modified seedling screening test (MSST)，along with antixenosis test and antibiosis test were applied to evaluate the resistance response of the two parents and 149 plant F2 to the insect and composite interval mapping (CIM) was used for QTL analysis When the resistance was measured by MSST method，two QTLs conferring resistance to Small brown planthopper were mapped on chromosome and chromosome namely qSBPH6-a and qSBPH7-a，with log of odds (LOD) scores 2.61 and 2.91，respectively; and two QTLs explained 8.75% and 13.1% of the phenotypic variance in this population，respectively Two QTLs，namely qSBPH1-c and qSBPH2-c, expressing antibiosis to SBPH were mapped on chromosomes and 2， respectively，explaining 30.16% of the total phenotypic variance Keywords: Rice; SBPH ( Laodelphax striatellus Fallén); Resistance; Quantitative trait 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Molecular Breeding，2003，12:133-143 Yang H X，Ren X，Weng Q，Zhu L，He G Molecular mapping and genetic analysis of a rice brown plant-hopper (Nilaparvata lugens Stål) resistance gene Hereditas， 2002，136:39-43 Zhang H M，Cheng J P， Lei J L，et al Sequence analysis shows that a dwarfing disease on rice， wheat and maize in China is caused by rice black-streaked dwarf virus[J] Eur J Plant Pathol， 2001，107 :563 - 567 71 在读期间发表论文 在读期间发表论文 Yuqiang Liu，Jun He，Ling Jiang，HanWu，Yuehua Xiao，Yanlin Liu， Guangquan Li，Yueqiang Du，Chenyang Liu，JianminWan Nitric oxide production is associated with response to brown planthopper infestation in rice Journal of Plant Physiology 168 (2011) 739–745 王宝祥、江玲、陈亮明、卢百关、王琦、黎光泉、樊继伟、程遐年、翟虎渠、徐 大勇、万建民。水稻黑条矮缩病抗性资源的筛选和抗性 QTL 的定位。作物学报、 2010、 36（8）：1258-1264 已整理待发表的论文 Le Quang Tuyen，Yuqiang Liu，Ling Jiang，Baoxiang Wang，Qi Wang，Than Thi Thu Hanh，Jianmin Wan Identification of quantitative trait loci associated with small brown planthopper (Laodelphax striatellus Fallén) resistance in rice (Oryza sativa L.) 黎光泉、刘裕强、江玲、刘晨阳、王宝祥、王琦、申氏秋幸、万建民。水稻品种 昌恢 891 抗灰飞虱相关 QTL 的检测 73 附 录 附 录 附录 1： MS salt Stock (Murashige Skoog、 1962) NH4NO3 :1650 mg/L H3BO3 : 6.2 mg/L FeSO4.7H2O : 27.8 mg/L CuSO4.5H2O : 0.025 mg/L KI :0.83 mg/L MgSO4.7H2O : 370 mg/L KNO3 : 1900 mg/L ZnSO4.7H2O: 8.6 mg/L MnSO4.4H2O : 22.3 mg/L CoCl2.6H2O : 0.025 mg/L Na2MoO4.2H2O : 0.25 mg/L CaCl2.2H2O: 440 mg/L KH2PO4 : 170 mg/L Na2EDTA : 37.3 mg/L 附录 2： Indel 标记序列 编码 正义引物 反义引物 Code Forward primer(5’-3’) Reverse prime(5’-3’) L1 CAAGTGGATGCGGGAAAT GGGCTGGCGTGTAGGATA L2 TTTCGGACGGAGGGAGTA TCGGGATCTAACACTGTAATGC L3 GAAAAGCTAACTTACATGA AGATATGCATCATCAGTGGAAGCTT 75 致 谢 致 谢 本论文是在导师万建民教授的悉心指导下完成的、从论文的选题、课题的实施及 论文写作等每个环节上均得到了导师大量的指导和帮助。值此论文完成之际我谨向 导师表示最诚挚的谢意！ 感谢南京农业大学水稻所江玲教授、杨世湖教授、程遐年教授、张文伟副教授、 赵志刚老师、陈赛华老师、王益华老师对我学习、实验、生活等多方面的关心！ 特别感谢刘裕强老师对我学习和实验多方面的帮助及在论文撰写时的悉心指导！ 特别感谢抗性组:王宝祥、张迎信、何俊、王琦、吴寒、刘晨阳、杜越强、宋兆 强、刘艳玲、胡金龙、康海燕在实验过程中给与的帮助！ 感谢南京农业大学水稻所研究生陈海梅、谢坤、龙起樟、边建民、郑蕾娜、汪勇、 边小峰、任玉龙、韩小花、周峰、卢丙越、李晶晶、赵宝泉、陈红、吕佳、刘峰、 彭城、李林芳、钟铮铮、张云辉、刘凯、万宇、肖月华、金锡明、朱成强、周振铃、 石彦荣、陈明江、吴薇、黄洁雪、杨春艳、李江涛、纪晓卿、陈苗苗、吴盛阳、张 俊杰、金检生、孙林鹤、阮氏如簪、郑明、沈雨民、骆卫峰、陈丽萍、余晓文、张 禹舜、张龙、刘凯、吴涛、赵绍路、张正尧、陈韦韦、张世利、陈益琳、禹阳、陈 新刚、冯志明、孔维一、宋攀、王超龙、王建 、王云龙、王立伟、韦琮、许扬、张 超、李晓慧、林秋云、牛梅、王新华、沈贝贝、周丽、马婧、梁婷婷等同学在实验 和学习中的关心和帮助！ 感谢南京农业大学国际教育学院多年来对我的帮助！ 特别感谢越南政府和人民给予我最好的留学条件！ 特别感谢我的父亲、母亲、哥哥、弟弟及同事对我无微不至的爱和全面的帮助！ 我要特别感谢我的夫人申氏秋幸和女儿黎玉英、她们的鼓励和支持使我又走上了 求学的道路、她们的无私奉献和默默关爱使我完成了学业！ 最后、谨向参加本论文评阅、答辩的各位专家致以衷心的感谢！ 南京农业大学、2011 年 月 Le Quang Tuyen 77 作者简历 作者简历 姓 名：LE QUANG TUYEN- 越南留学生 中国名字：黎光泉 出生年月：1971 年 月 籍 贯：越南兴安省 学习经历： 1998 年 月：越南河内农业大学农学院、农学学士毕业 2007 年 月：越南胡志明农林大学、农学硕士毕业 2008 年 10 月至今：中国南京农业大学水稻研究所、攻读作物遗传育种学专业博士 工作经历： 1998 年 月- 2006 年 10 月：越南甘蔗和糖科学院研究员 2006 年 10 月至今：越南南部农科院研究员 79 [...]... 扩增片段长度多态性 BIL backcross inbred lines 回交重组自交系 BPH brown planthopper 褐飞虱 CIM composite interval mapping 复合区间作图 DH doubled haploid 双单倍体 DIBA dot-immunobinding assay 斑点免疫法 EST expressed sequence tags 表达序列标签 HR highly resistance 高抗 HS highly susceptible 高感 Indel insert/delete 插入缺失 MR moderately resistance 中抗 MSST modified seedbox screening test 改进的苗期筛选法 QTL quantitative trait locus 数量性状位点 R resistance 抗 RAPD... 2-2 苗期集团接种示意图 Fig 2-2 The pattern of standard seedling-box screening test 表 2-1 灰飞虱对水稻苗期的致害性及其评价标准 Table 2-1 Evaluation criteria for resistance to SBPH at rice seedling stage 抗性级别 抗性水平 Resistance scale Resistance level 未受害 (No visible damage) 0 免疫 (I) 受害级轻微 (Very slightly damage) 1 高抗 (HR) 3 抗虫 (R) 5 中抗 (MR) 7 感虫 (S) 9 高感 (HS) 危害症状（Damage symptom） 第一、二片叶都分发黄 (Partial yellowing of the first and second... 质。稻飞虱俗名火蠓、响虫、火旋、稻虱子等、常见的有三种：褐飞虱、白背飞虱 和灰飞虱。三种飞虱以褐飞虱 (Nilaparvata lugens Stål)为主、白背飞虱（Sogatella furcifera Horvath）次之、灰飞虱（Laodelphax striatellus Fallén）更次之。 灰飞虱 (Laodelphax striatellus) 褐飞虱 (Nilaparvata lugens ) 白背飞虱（Sogatella furcifera） Small brown planthopper Brown planthopper Whitebacked planthopper 图 1-1： 稻飞虱 Figure 1-1: Rice planthopper 一般年份、这类害虫的危害、可使水稻减产 1 成以上、大发生年可减产 2-3 成。 稻飞虱以刺吸式口器刺入稻株组织吸取汁液、造成各种不规则的白色或褐色条斑、 并以产卵器刺伤叶鞘、嫩茎和叶中脉等组织、产卵于其中、使稻株枯黄或倒伏、严... 从南京市土桥镇郊区稻田采集灰飞虱、在小麦上饲养 4 代、然后转入种有武育粳 3 号的培养皿和塑料盒中饲养、每个培养皿或塑料盒转移一头待产灰飞虱繁殖后代。 采用斑点免疫法(dot- immunobinding assay、 DIBA) 和 PCR 扩增法对每个培养皿或塑 料盒进行带毒检测、无毒灰飞虱用于接虫鉴定水稻的抗虫性（图 2-1）。饲养于南京 农业大学温室内。 水稻繁殖 小麦繁殖 图 2-1 灰飞虱室内繁殖体系 Fig 2- 1 The propagation of small brown planthopper in house 1.3 抗灰飞虱鉴定 1.3.1 苗期集团接种法 （Standard Seedling-box Screening Test） 采用标准苗期集团接种法 (Heinrichs 等、1985)对两个亲本 02428 和 Rathu Heenati、 162 个 02428/Rathu Heenati F2：3... polymorphic DNA 快速扩增多态性 DNA RBSDV rice black-streaked dwarf virus 水稻黑条矮缩病 RFLP restriction fragment length polymorphism 限制性片段长度多态性 RIL recombinant inbred lines 重组自交系 RSV rice stripe virus 水稻条纹病毒 S susceptible 感 SBPH small brown planthopper 灰飞虱 SSR simple sequence repeat 简单序列重复 SSST standard seedbox screening test 标准苗期筛选法 STS sequence-tagged site 标记序列位点 WBPH whitebacked planthopper 白背飞虱 V 第一章 文献综述 第一章... 叶片明显发黄、部分苗矮化 (Pronounced yellowing, and some seedlings slight stunning) 稻苗严重矮化或枯死 (Seedlings showing signs of severe stunning or wilting) 90% 稻苗枯死 (90% seedling dead) Note: I- immune; HR- highly resistant; R- resistant; MR- moderately resistant; S- susceptible; HR- highly susceptible 当感虫对照武育粳 3 号死苗率达 90%时、参照 IRRI (1996)和 Duan 等 (2008、 2009) 制定的抗虫性评价标准（Standard Seedling-box Screening Test, SSST）、 对亲 本和各家系的每个单株进行目测评级（表... (Ikeda 和 Vaughau, 1991) Table 1-1 Originations and distribution of four BPH Biotypes (Ikeda and Vaughau, 1991) 生物型 地理分布 来源 Biotypes Distributions Originations 生物型 1 东亚和东南亚 野生型（产生于东亚和东南亚） biotype 1 East and Southeast Asia Wild type (wildly distributed in East and Southeast Asia 生物型 2 东亚和东南亚 在菲律宾由取食带有 Bph1 水稻品种后演变而来 biotype 2 East and Southeast Asia Originated in Philippines after widescale cultivation of... after widescale cultivation of varities with Bph1 生物型 3 东亚和东南亚 在日本和菲律宾的实验室中产生 biotype 3 East and Southeast Asia Produced in laboratories in both Japan and the Philippines 生物型 4 南亚 南亚 9 两个水稻品种抗灰飞虱的遗传分析 biotype 4 South Asia South Asia 表 1-2 越南红河三角洲稻区褐飞虱发生概况 Table 1-2 Situation of Laodelphax striatellus Fallén in Vietnam red river delta 发生 代次 高峰期（月/旬、月/日） 水稻生育期 迁飞的可能性 1 3/中 到 3/下 早稻分蘖期 无 2 4/中 到 4/下 早稻孕穗期... 带毒灰飞虱刺吸植物、植株染病后终身带毒、并可经卵传给下一代（Hibino, 1989；Wu 等、2001）、且能在体内增殖。带毒的灰飞虱通过刺吸健康的寄主植物、 将病毒传播给健康植物、一般地说、传播能力不断下降、但那些不能将病毒传到植 株上的昆虫仍可以 75%一 100%的比例经卵传毒至后代(Shinkai、1962)。带毒灰飞虱 可将病毒传递 23 代、经卵传递率为 90%(Kisimoto, 1967；Washio, 1968a, 1968b)。灰 飞虱传播水稻条纹病毒的过程如下： 灰飞虱经卵传毒示意图： 带毒♀X 带毒 or 无毒♂ 卵（后代）带毒 无毒♀X 带毒 or 无毒♂ 卵（后代）无毒 4 第一章 文献综述 图 1-2 灰飞虱繁殖及传播 RSV 危害的世代示意图(张迎信、2010) Fig 1-2 The life circle and transmitting stripe virus of small brown planthopper. .. 除水稻外、还有很多植物也是灰飞虱的寄主植物、如下：小麦（Triticum aestivum L.）、大麦（Hordeum vulgare L.）、游草（Leersia japonica Makino）、看麦 娘（Alopecurus aequalis Sobol.）、双穗雀稗（Paspalum distichum L.）、千金子 （Leptochloa chinensis L.) Nees）、蟋蟀草（Eleusine indica L.）、白茅（Imperata cylindrica Beauv）、节缕草（Zoysia japonica Steud）、日本看麦娘（Alopecurus japonica Steud.）、竹叶茅（Microstegium nudum(trin.) A.Camns）等（朱绍先等、 1984）。灰飞虱取食不同寄主植物、其发育历期、成活率、繁殖率差异较大（乔慧 等、2009）。寄主植物的可溶性糖含量与氨基酸总量都能影响灰飞虱的繁殖率（张