水稻（Oryza sativa L.）作为全球主要粮食作物之一，其安全生产对维护粮食安全具有重要意义。然而，水稻生产常年遭受各类病虫害危害，严重威胁水稻的产量和品质。其中，灰飞虱[Laodelphax striatellus Fallén (Jomoptera:Delphacidae), small brown planthopper, SBPH]是水稻的主要害虫之一，其寄主广泛，除水稻外，还可以危害小麦和玉米等多种禾谷类作物。灰飞虱通过口针吸食水稻汁液，使植株枯黄，甚至枯死；此外，灰飞虱还是水稻条纹病毒（rice stripe virus, RSV）、水稻黑条矮缩病毒（rice black-streaked dwarf virus, RBSDV）、玉米粗缩病（maize rough dwarf virus, MRDV）和小麦丛矮病（wheat rosette stunt virus, WRSV）等作物主要病毒病的传播介体。因此，培育和推广抗性品种被认为是更为经济、高效且环境友好的防治策略。然而，目前为止尚未有抗灰飞虱基因克隆的报道，作物抗灰飞虱的分子机制还知之甚少。

为发掘水稻抗灰飞虱新抗源和新基因，本研究完成部分水稻3K资源的灰飞虱抗性鉴定。其中一份来自东南亚的地方品种W41129，其高抗灰飞虱。通过对该群体进行分析，在水稻第6号染色体上定位到一个主效抗灰飞虱位点，并将其命名为qSBPH6。本研究在此基础上，对qSBPH6进行精细定位，发现qSBPH6定位区间包含两个抗性位点（后续将两个位点分别命名为SBPH1和SBPH2），且两个位点的灰飞虱抗性具有累加作用。本研究进一步完成了其中SBPH1的图位克隆和功能分析。

此外，开展了水稻转录因子OsWRKY36抗灰飞虱机制研究。发现OsWRKY36抑制OsPAL6介导的木质素生物合成途径，进而削弱水稻细胞壁的物理防御屏障，OsWRKY36负调控水稻对灰飞虱的抗性。OsWRKY36的分子机制研究为水稻抗性品种改良提供重要的基因资源和理论基础。具体研究结果如下：

1.籼粳亚种间灰飞虱抗性存在分化

为发掘水稻灰飞虱新抗源，搜集了水稻3K资源中338份品种，其中包括193份籼稻、94份粳稻和51份Aus/boro稻。利用苗期集团接种法完成338份水稻资源的抗性鉴定，结果发现死苗率超过80%的品种有186份，低于10%的品种有41份，其中20份品种的死苗率为0。上述高抗灰飞虱资源的筛选为后续抗灰飞虱基因的发掘和克隆奠定基础。此外，分析发现Aus/boro稻和粳稻品种的平均幼苗死亡率分别达75.1%和90.43%，而籼稻品种的平均幼苗死亡率仅为55.61%。上述结果表明，籼粳亚种间对灰飞虱的抗性存在显著分化。

2.抗灰飞虱基因SBPH1的精细定位及候选基因分析

W41129对灰飞虱具有较强的排驱性和抗生性。此外，扫描电镜观察发现，SYN叶鞘厚壁组织较为疏松，而W41129叶鞘厚壁组织更为致密，表明厚壁组织的结构差异可能影响W41129和SYN对灰飞虱的抗性。

本研究以SYN为轮回亲本，构建了BC₁F₂和BC₂F₂群体，利用上述群体将该基因定位900 kb区间内，进一步精细定位分析发现该区间可能存在两个抗灰飞虱位点（SBPH1和SBPH2），且两个位点对灰飞虱的抗性存在加性效应。利用分子标记将两个位点进行分离，并构建了SBPH1的次级分离群体，最终将SBPH1精细定位在64 kb的区间内，该区间包含6个ORFs。结合定位区间双亲的基因组比较、基因功能预测和灰飞虱诱导表达分析等，将区间内的一个果胶甲基酯酶基因19（pectin methylesterase 19, OsPME19）确定为SBPH1的候选基因。通过构建OsPME19-KO和OsPME19-OX转基因家系，结果证实OsPME19负调控水稻灰飞虱抗性。

3.OsPME19的生物学功能分析

灰飞虱取食诱导表达分析表明，OsPME19在感虫品种SYN中的表达受灰飞虱取食诱导，且显著高于抗虫品种。组织表达分析发现，OsPME19在幼苗地上部分表达水平较高。原位杂交结果显示OsPME19在水稻叶鞘厚壁组织细胞中高表达。亚细胞定位和质壁分离结果发现，抗感等位基因编码蛋白OsPME19^W41129和OsPME19^SYN均定位于细胞壁。果胶甲基酯酶总活性结果表明OsPME19-OX植株的果胶甲基酯酶活性显著高于野生型，而OsPME19-KO活性显著低于野生型。此外，体外重组蛋白表达和酶活性测定表明，OsPME19^SYN和OsPME19^W41129的酶活没有显著差异。上述结果表明，抗感等位基因OsPME19^SYN和OsPME19^W41129编码区差异并不影响其蛋白的功能，其灰飞虱抗性水平的差异可能是表达水平的差异导致。

植株叶鞘果胶甲酯化免疫荧光标记分析的结果表明，OsPME19参与水稻细胞壁果胶去甲酯化。进一步利用扫描电镜观察叶鞘横切面结构，表明OsPME19可能通过调控果胶甲酯化水平影响厚壁组织的结构。

甲醇的含量测定结果表明OsPME19-OX甲醇含量显著高于野生型，而OsPME19-KO植株甲醇含量显著低于野生型。此外，与对照相比，灰飞虱明显倾向于取食甲醇预处理的水稻幼苗。结果表明，SYN比W41129更吸引灰飞虱取食是由于甲醇引起的。

4.OsERF47调控OsPME19表达的机制解析

为解析抗感品种OsPME19表达差异的原因，对比分析了抗感等位基因OsPME19^SYN和OsPME19^W41129启动子序列，发现与OsPME19^W41129相比，OsPME19^SYN的启动子中多了5个ERF基序。酵母单杂交（Y1H）发现一个AP2/ERF类转录因子OsERF47特异性与OsPME19^SYN启动子结合，而不与OsPME19^W41129启动子结合。启动子截短实验发现，OsERF47仅与最靠近OsPME19^SYN转录起始位点（TSS）的ERF基序（ERF#5）结合。双分子荧光素酶分析（LUC）进一步证实OsERF47可以显著促进OsPME19^SYN的表达，而对OsPME19^W41129的转录激活作用不显著。此外，与野生型相比，OsERF47-KO中OsPME19的表达水平显著降低，且甲基酯化的果胶显著增加，灰飞虱抗性显著增强。上述结果表明，OsERF47直接结合OsPME19^SYN启动子中的ERF基序，增强OsPME19^SYN的表达，从而负调控水稻灰飞虱抗性。

5.OsPME19启动子变异与籼粳亚种间灰飞虱抗性分化的相关性分析

利用3K数据库对OsPME19启动子ERF#5基序中的SNP变异进行分析，结果发现，99.5%的粳稻品种携带感虫类型（Haplotype_C），而67.7%的籼稻携带抗虫类型（Haplotype_T）。同时，携带Haplotype_C类型水稻资源的平均死苗率为83.75%，而Haplotype_T的平均死苗率仅约为56.47%。表明OsPME19启动子ERF#5基序中的SNP变异与籼粳亚种间灰飞虱抗性分化高度相关。此外，我们还分析了Haplotype_C和Haplotype_T在145份不同类型和来源的野生稻中的分布，结果发现145份野生稻全部携带感虫类型Haplotype_C。上述结果表明，Haplotype_T可能是在籼稻和粳稻两个亚种分化后在籼稻中产生并保留下来。

6.OsWRKY36对灰飞虱抗性机制研究

通过对OsWRKY36相关转基因家系进行灰飞虱表型鉴定，结果表明，OsWRKY36负调控水稻对灰飞虱的抗性。启动子结合实验表明，OsWRKY36特异性结合OsPAL6启动子区域的W-box元件，并抑制其转录水平。遗传互补实验证实，在Bphs1-D背景下异源表达OsPAL6可有效逆转其感虫表型。生化检测发现，Bphs1-D的木质素含量和厚壁组织细胞层数显著下降，而OsPAL6-OX/Bphs1-D和OsPAL6-OX的木质素含量和厚壁组织细胞层数显著升高。综上，OsWRKY36通过抑制OsPAL6的转录，抑制木质素合成途径，从而降低水稻的抗虫性。

综上所述，本论文完成了水稻抗灰飞虱基因SBPH1的图位克隆和功能分析，即通过影响果胶甲酯化修饰和甲醇释放介导水稻灰飞虱抗性的分子机制。相关研究不仅揭示了一条作物抗虫的新途径，为抗灰飞虱水稻品种培育提供基因资源，同时也为其他作物抗灰飞虱研究提供借鉴。此外，OsWRKY36的鉴定和功能分析为阐明水稻对灰飞虱抗性的分子调节网络奠定基础。

Rice (Oryza sativa L.), as one of the world's most important staple crops, plays a vital role in global food security. However, rice production faces persistent threats from various pests and diseases that significantly compromise both yield and grain quality. Among these, the small brown planthopper (Laodelphax striatellus Fallén, Hemiptera:Delphacidae; SBPH) is a major pest with a broad host range that includes wheat, maize and other cereal crops. SBPH damages rice plants by sucking phloem sap through its stylet, leading to yellowing and even plant death. More critically, it serves as a vector for several devastating viral diseases, including rice stripe virus (RSV), rice black-streaked dwarf virus (RBSDV), maize rough dwarf virus (MRDV), and wheat rosette stunt virus (WRSV). Nevertheless, no SBPH resistance gene has been cloned to date, and the molecular mechanisms underlying SBPH resistance remain largely unknown.

To identify novel resistance sources and genes against SBPH, we conducted resistance screening of a subset of the rice 3K germplasm collection. A landrace from Southeast Asia, W41129, exhibited strong resistance to SBPH. Through linkage mapping, we identified a major SBPH resistance locus, qSBPH6, on chromosome 6. Subsequent fine-mapping revealed that qSBPH6 contains two resistance loci (designated SBPH1 and SBPH2) with additive effects on SBPH resistance. This study focused on the map-based cloning and functional characterization of SBPH1.

In addition, this study found that OsWRKY36 negatively regulates rice resistance to SBPH by inhibiting OSPAL6-mediated lignin biosynthesis pathway, thereby weakening the physical defense barrier of rice cell wall. The molecular machinery study of OsWRKY36 provides important genetic resources and theoretical basis for the improvement of rice resistant varieties. The specific research outcomes are summarized as follows:

1. Differentiation of resistance to the small brown planthopper exists between indica and japonica subspecies.

To explore new sources of resistance to the SBPH in rice, 338 rice 3K resources were collected, including 193 indica rice varieties, 94 japonica rice varieties, and 51 Aus/boro rice varieties. The resistance identification of 338 rice resources was completed using the seedling-stage group inoculation method. The results showed that there were 186 varieties with a seedling mortality rate exceeding 80%, 41 varieties with a rate lower than 10%, and 20 varieties with a seedling mortality rate of 0. The screening of the above highly resistant SBPH resources laid the foundation for the subsequent discovery and cloning of genes resistant to the SBPH. In addition, the analysis found that the average seedling mortality rates of Aus/boro rice and japonica rice varieties were 75.1% and 90.43% respectively, while the average seedling mortality rate of indica rice varieties was only 55.61%. The above results indicate that there is a significant differentiation in the resistance to the SBPH between indica and japonica subspecies.

2. Fine mapping of the SBPH1 gene resistant to the SBPH and analysis of candidate genes.

W41129 had strong repellent and antibiologic activity against SBPH. In addition, scanning electron microscopy showed that the sachyma of SYN sheath was loose, while that of W41129 was denser, indicating that the structural difference of sachyma may affect the resistance of W41129 and SYN to SBPH.

In this study, using SYN as the recurrent parent, BC₁F₂ and BC₂F₂ populations were constructed. The gene was mapped within a 900 kb interval using the above populations. Further fine mapping analysis found that there may be two SBPH resistance loci (SBPH1 and SBPH2) in this interval, and the two loci have an additive effect on the resistance to the SBPH. Subsequently, the two loci were separated using molecular markers, and a secondary segregation population of SBPH1 was constructed. Finally, SBPH1 was finely mapped within a 64 kb interval, which contains 6 ORFs. Combining the genomic comparison of the two parents in the mapping interval, gene function prediction, and the induced expression analysis by the SBPH, a pectin methylesterase gene 19 (OsPME19) within the interval was identified as a candidate gene for SBPH1. Transgenic families of OSPME19-KO and OSPME19-OX showed that OsPME19 negatively regulated SBPH resistance.

3. Biological function analysis of OsPME19.

The induced expression analysis by the feeding of the SBPH showed that the expression of OsPME19 in the insect-susceptible variety SYN was induced by the feeding of the SBPH and was significantly higher than that in the insect-resistant variety. The tissue expression analysis found that the expression level of OsPME19 was higher in the above-ground parts of the seedlings. In situ hybridization results showed that OsPME19 was highly expressed in the sclerenchyma cells of the leaf sheath of rice. Subcellular localization and plasmolysis results found that the proteins encoded by the resistant and susceptible alleles, OsPME19^W41129 and OsPME19^SYN, were both localized in the cell wall. The results of total pectin methylesterase activity showed that the pectin methylesterase activity of OsPME19-OX plants was significantly higher than that of the wild type, while the activity of OsPME19-KO was significantly lower than that of the wild type. In addition, the in vitro recombinant protein expression and enzyme activity measurement further confirmed that there was no significant difference in the enzyme activity between OsPME19^SYN and OsPME19^W41129. The above results indicate that the differences in the coding regions of the resistant and susceptible alleles OsPME19^SYN and OsPME19^W41129 do not affect the function of their proteins, and the differences in the resistance level to the SBPH may be caused by the differences in the expression level.

The immunofluorescence labeling analysis of the pectin methyl-esterification of the leaf sheath showed that OsPME19 is involved in the demethyl-esterification of pectin in rice. Further, the structure of the cross-section of the leaf sheath was observed using scanning electron microscopy. Compared with the wild type, the sclerenchyma of the leaf sheath of rice in OsPME19-OX was looser than that of the wild type, while the sclerenchyma of OsPME19-KO was denser, indicating that OsPME19 may affect the structure of the sclerenchyma by regulating the level of pectin methyl-esterification.

The measurement results of the methanol content showed that the methanol content in OsPME19-OX was significantly higher than that in the wild type, while the methanol content in OsPME19-KO plants was significantly lower than that in the wild type. Therefore, the results showed that compared with the control, the SBPH obviously preferred to feed on the rice seedlings pretreated with methanol. The fact that SYN is more attractive to SBPH than W41129 is due to methanol.

4. Analysis of the mechanism by which OsERF47 regulates the expression of OsPME19.

To analyze the reasons for the expression differences of OsPME19 between the resistant and susceptible varieties, the promoter sequences of the resistant and susceptible alleles OsPME19^SYN and OsPME19^W41129 were compared and analyzed. It was found that compared with OsPME19^W41129, the promoter of OsPME19^SYN had 5 more ERF motifs. Yeast one-hybrid (Y1H) found that an AP2/ERF transcription factor OsERF47 specifically bound to the promoter of OsPME19^SYN, but not to the promoter of OsPME19^W41129. The promoter truncation experiment found that OsERF47 only bound to the ERF motif (ERF#5) closest to the transcription start site (TSS) of OsPME19^SYN. Bimolecular luciferase analysis (LUC) further confirmed that OsERF47 could significantly promote the expression of OsPME19^SYN, but had no significant transcriptional activation effect on OsPME19^W41129. In addition, compared with the wild type, the expression level of OsPME19 in the knockout mutant oserf47 was significantly reduced, the methyl-esterified pectin was significantly increased, and the resistance to the SBPH was significantly enhanced. The above results indicate that OsERF47 directly binds to the ERF motif in the promoter of OsPME19^SYN, enhances the expression of OsPME19^SYN and thus negatively regulates the resistance of rice to SBPH.

5. Correlation analysis between the variation of the OsPME19 promoter and the differentiation of resistance to the SBPH between indica and japonica subspecies.

The SNP variation in the ERF#5 motif of the OsPME19 promoter was analyzed using the 3K database. The results showed that 99.5% of japonica rice varieties carried the insect-susceptible type (Haplotype_C), while 67.7% of indica rice varieties carried the insect-resistant type (Haplotype_T). Further, the results showed that the average seedling mortality rate of rice resources carrying the Haplotype_C type was 83.75%, while that of Haplotype_T was only about 56.47%. This indicates that the SNP variation in the ERF#5 motif of the OsPME19 promoter is highly correlated with the differentiation of resistance to the SBPH between indica and japonica subspecies. In addition, we also analyzed the distribution of Haplotype_C and Haplotype_T in 145 wild rice varieties of different types and sources. The results showed that all 145 wild rice varieties carried the insect-susceptible type Haplotype_C. The above results indicate that Haplotype_T may have been generated and retained in indica rice after the differentiation of the two subspecies, indica and japonica rice.

6. Study on the resistance mechanism of OsWRKY36 to SBPH.

Through the phenotypic identification of the SBPH in the transgenic lines related to OsWRKY36, the results showed that OsWRKY36 negatively regulates the resistance of rice to the SBPH. The promoter binding experiment showed that OsWRKY36 specifically binds to the W-box element in the promoter region of OsPAL6 and inhibits its transcription level. The genetic complementation experiment confirmed that the heterologous expression of OsPAL6 in the Bphs1-D background can effectively reverse its insect-susceptible phenotype. Biochemical detection found that the lignin content and the number of sclerenchyma cell layers in Bphs1-D significantly decreased, while those in OsPAL6-OX/Bphs1-D and OsPAL6-OX significantly increased. In summary, OsWRKY36 inhibits the transcription of OsPAL6 and the lignin synthesis pathway, thereby reducing the insect resistance of rice.

In conclusion, this thesis completed the map-based cloning and functional analysis of a gene SBPH1 resistant to the SBPH in rice, that is, the molecular mechanism by which it mediates the resistance of rice to SBPH by affecting the pectin methyl-esterification modification and the release of methanol. The related research not only reveals a new pathway for crop insect resistance, provides gene resources for the breeding of rice varieties resistant to the SBPH, but also provides a reference for the research on the resistance of other crops to the SBPH. In addition, the identification and functional analysis of OsWRKY36 lay the foundation for clarifying the molecular regulatory network of the resistance of rice to the SBPH.

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