水稻抗病调控机制研究进展

doi:10.11983/CBB25011

摘要/Abstract

摘要： 水稻(Oryza sativa)是世界上最重要的粮食作物之一, 其产量影响全球粮食安全。然而, 水稻的各种病害对粮食安全构成极大威胁。其中, 稻瘟病、白叶枯病和纹枯病是影响全球水稻生产的3种主要病害, 培育具有广谱抗病性的水稻品种迫在眉睫。近年来, 水稻抗病调控机制研究取得了突破性进展。该文从水稻自身免疫反应以及抗病相关基因功能等方面综述了水稻抗病调控机制, 提出今后需解决的问题, 以期为水稻广谱抗病育种提供参考。

关键词: 水稻, 抗病调控机制, 植物免疫响应

Abstract: Rice (Oryza sativa) is one of the most vital food crops globally, and its yield plays a crucial role in ensuring food security. However, various diseases affecting rice pose significant threats to this security. Among these, rice blast, bacterial blight, and sheath blight are the three predominant diseases impacting global rice production. Consequently, there is an urgent need to breed and cultivate rice varieties with broad-spectrum disease resistance. In recent years, substantial advancements have been made in understanding the regulatory mechanisms underlying disease resistance in rice. This paper reviews these mechanisms from multiple perspectives, including the plant’s intrinsic immune responses and the functional dynamics of resistance genes. Furthermore, it highlights pressing issues that require immediate attention to facilitate broad-spectrum disease-resistant breeding efforts for rice.

Key words: rice, disease resistance regulation mechanism, plant immune response

江亚楠, 徐雨青, 魏毅铤, 陈钧, 张容菀, 赵蓓蓓, 林宇翔, 饶玉春. 水稻抗病调控机制研究进展. 植物学报, 2025, 60(5): 734-748.

Jiang Yanan, Xu Yuqing, Wei Yiting, Chen Jun, Zhang Rongwan, Zhao Beibei, Lin Yuxiang, Rao Yuchun. Research Progress on the Regulatory Mechanism of Rice Disease Resistance. Chinese Bulletin of Botany, 2025, 60(5): 734-748.

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB25011

https://www.chinbullbotany.com/CN/Y2025/V60/I5/734

图/表 4

参考文献 124

[1]	Andaya CB, Ronald PC (2003). A catalytically impaired mutant of the rice Xa21 receptor kinase confers partial resistance to Xanthomonas oryzae pvoryzae. Physiol Mol Plant Pathol 62, 203-208.
[2]	Ashikawa I, Hayashi N, Yamane H, Kanamori H, Wu JZ, Matsumoto T, Ono K, Yano M (2008). Two adjacent nucleotide-binding site-leucine-rich repeat class genes are required to confer Pikm-specific rice blast resistance. Genetics 180, 2267-2276. DOI PMID
[3]	Boutrot F, Zipfel C (2017). Function, discovery, and exploitation of plant pattern recognition receptors for broad- spectrum disease resistance. Annu Rev Phytopathol 55, 257-286. DOI PMID
[4]	Bryan GT, Wu KS, Farrall L, Jia YL, Hershey HP, McAdams SA, Faulk KN, Donaldson GK, Tarchini R, Valent B (2000). A single amino acid difference distinguishes resistant and susceptible alleles of the rice blast resistance gene Pi-ta. Plant Cell 12, 2033-2046. DOI PMID
[5]	Cao WL, Zhang HM, Zhou Y, Zhao JH, Lu SB, Wang XQ, Chen XJ, Yuan LM, Guan HY, Wang GD, Shen WX, De Vleesschauwer D, Li ZQ, Shi XP, Gu JF, Guo M, Feng ZM, Chen ZX, Zhang YF, Pan XB, Liu WD, Liang GH, Yan CJ, Hu KM, Liu QQ, Zuo SM (2022). Suppressing chlorophyll degradation by silencing OsNYC3 improves rice resistance to Rhizoctonia solani, the causal agent of sheath blight. Plant Biotechnol J 20, 335-349.
[6]	Cao YL, Duan L, Li HJ, Sun XL, Zhao Y, Xu CG, Li XH, Wang SP (2007). Functional analysis of Xa3/Xa26 family members in rice resistance to Xanthomonas oryzae pv.oryzae. Theor Appl Genet 115, 887-895.
[7]	Césari S, Kanzaki H, Fujiwara T, Bernoux M, Chalvon V, Kawano Y, Shimamoto K, Dodds P, Terauchi R, Kroj T (2014). The NB-LRR proteins RGA4 and RGA5 interact functionally and physically to confer disease resistance. EMBO J 33, 1941-1959. DOI PMID
[8]	Chen J, Peng P, Tian JS, He YG, Zhang LP, Liu ZX, Yin DD, Zhang ZH (2015). Pike, a rice blast resistance allele consisting of two adjacent NBS-LRR genes, was identified as a novel allele at the Pik locus. Mol Breed 35, 117.
[9]	Chen XF, Liu PC, Mei L, He XL, Chen L, Liu H, Shen SR, Ji ZD, Zheng XX, Zhang YC, Gao ZY, Zeng DL, Qian Q, Ma BJ (2021). Xa7, a new executor R gene that confers durable and broad-spectrum resistance to bacterial blight disease in rice. Plant Commun 2, 100143.
[10]	Cheng XY, Zhou GH, Chen W, Tan L, Long QS, Cui FS, Tan L, Zou GX, Tan Y (2024). Current status of molecular rice breeding for durable and broad-spectrum resistance to major diseases and insect pests. Theor Appl Genet 137, 219. DOI PMID
[11]	Chu J, Xu H, Dong H, Xuan YH (2021). Loose plant architecture 1-interacting kinesin-like protein KLP promotes rice resistance to sheath blight disease. Rice (NY) 14, 60.
[12]	Chu ZH, Yuan M, Yao JL, Ge XJ, Yuan B, Xu CG, Li XH, Fu BY, Li ZK, Bennetzen JL, Zhang QF, Wang SP (2006). Promoter mutations of an essential gene for pollen development result in disease resistance in rice. Genes Dev 20, 1250-1255.
[13]	Das A, Soubam D, Singh PK, Thakur S, Singh NK, Sharma TR (2012). A novel blast resistance gene, Pi54rh cloned from wild species of rice, Oryza rhizomatis confers broad spectrum resistance to Magnaporthe oryzae. Funct Integr Genom 12, 215-228.
[14]	De Vleesschauwer D, Van Buyten E, Satoh K, Balidion J, Mauleon R, Choi IR, Vera-Cruz C, Kikuchi S, Höfte M (2012). Brassinosteroids antagonize gibberellin- and salicylate-mediated root immunity in rice. Plant Physiol 158, 1833-1846. DOI PMID
[15]	Deng YW, Zhai KR, Xie Z, Yang DY, Zhu XD, Liu JZ, Wang X, Qin P, Yang YZ, Zhang GM, Li Q, Zhang JF, Wu SQ, Milazzo J, Mao BZ, Wang ET, Xie HA, Tharreau D, He ZH (2017). Epigenetic regulation of antagonistic receptors confers rice blast resistance with yield balance. Science 355, 962-965. DOI PMID
[16]	Devanna NB, Vijayan J, Sharma TR (2014). The blast resistance gene Pi54 of cloned from Oryza officinalis interacts with Avr-Pi54 through its novel non-LRR domains. PLoS One 9, 104840.
[17]	Dong JF, Zhou L, Feng AQ, Zhang SH, Fu H, Chen L, Zhao JL, Yang TF, Yang W, Ma YM, Wang J, Zhu XY, Liu Q, Liu B (2021). The OsOXO2, OsOXO3 and OsOXO4 positively regulate panicle blast resistance in rice. Rice (NY) 14, 51.
[18]	Fan J, Quan WL, Li GB, Hu XH, Wang Q, Wang H, Li XP, Luo XT, Feng Q, Hu ZJ, Feng H, Pu M, Zhao JQ, Huang YY, Li Y, Zhang Y, Wang WM (2020). circRNAs are involved in the rice-Magnaporthe oryzae interaction. Plant Physiol 182, 272-286.
[19]	Feng T, Zhang ZY, Gao P, Feng ZM, Zuo SM, Ouyang SQ (2023). Suppression of rice Osa-miR444.2 improves the resistance to sheath blight in rice mediating through the phytohormone pathway. Int J Mol Sci 24, 3653.
[20]	Figueroa-Macías JP, García YC, Núñez M, Díaz K, Olea AF, Espinoza L (2021). Plant growth-defense trade-offs: molecular processes leading to physiological changes. Int J Mol Sci 22, 693.
[21]	Fukuoka S, Saka N, Koga H, Ono K, Shimizu T, Ebana K, Hayashi N, Takahashi A, Hirochika H, Okuno K, Yano M (2009). Loss of function of a proline-containing protein confers durable disease resistance in rice. Science 325, 998-1001. DOI PMID
[22]	Fukuoka S, Yamamoto SI, Mizobuchi R, Yamanouchi U, Ono K, Kitazawa N, Yasuda N, Fujita Y, Nguyen TTT, Koizumi S, Sugimoto K, Matsumoto T, Yano M (2014). Multiple functional polymorphisms in a single disease resistance gene in rice enhance durable resistance to blast. Sci Rep 4, 4550.
[23]	Gao Y, Xiang XJ, Zhang YX, Cao YR, Wang BF, Zhang Y, Wang C, Jiang M, Duan WJ, Chen DB, Zhan XD, Cheng SH, Liu QE, Cao LY (2022). Disruption of OsPHD1, encoding a UDP-glucose epimerase, causes JA accumulation and enhanced bacterial blight resistance in rice. Int J Mol Sci 23, 751.
[24]	Gao ZQ, Liu QE, Zhang YX, Chen DB, Zhan XD, Deng CW, Cheng SH, Cao LY (2020). OsCUL3a-associated molecular switches have functions in cell metabolism, cell death, and disease resistance. J Agric Food Chem 68, 5471-5482.
[25]	Gu KY, Yang B, Tian DS, Wu LF, Wang DJ, Sreekala C, Yang F, Chu ZQ, Wang GL, White FF, Yin ZC (2005). R gene expression induced by a type-III effector triggers disease resistance in rice. Nature 435, 1122-1125.
[26]	Guo F, Huang YZ, Qi PP, Lian GW, Hu XM, Han N, Wang JH, Zhu MY, Qian Q, Bian HW (2021). Functional analysis of auxin receptor OsTIR1/OsAFB family members in rice grain yield, tillering, plant height, root system, germination, and auxinic herbicide resistance. New Phytol 229, 2676-2692. DOI PMID
[27]	Hayashi K, Fujita Y, Ashizawa T, Suzuki F, Nagamura Y, Hayano-Saito Y (2016). Serotonin attenuates biotic stress and leads to lesion browning caused by a hypersensitive response to Magnaporthe oryzae penetration in rice. Plant J 85, 46-56.
[28]	Hayashi N, Inoue H, Kato T, Funao T, Shirota M, Shimizu T, Kanamori H, Yamane H, Hayano-Saito Y, Matsumoto T, Yano M, Takatsuji H (2010). Durable panicle blast-resistance gene Pb1 encodes an atypical CC-NBS- LRR protein and was generated by acquiring a promoter through local genome duplication. Plant J 64, 498-510.
[29]	He M, Yin JJ, Feng ZM, Zhu XB, Zhao JH, Zuo SM, Chen XW (2020). Methods for evaluation of rice resistance to blast and sheath blight diseases. Chin Bull Bot 55, 577-587. (in Chinese)
	贺闽, 尹俊杰, 冯志明, 朱孝波, 赵剑华, 左示敏, 陈学伟 (2020). 水稻稻瘟病和纹枯病抗性鉴定方法. 植物学报 55, 577-587.
[30]	Hu KM, Cao JB, Zhang J, Xia F, Ke YG, Zhang HT, Xie WY, Liu HB, Cui Y, Cao YL, Sun XL, Xiao JH, Li XH, Zhang QL, Wang SP (2017). Improvement of multiple agronomic traits by a disease resistance gene via cell wall reinforcement. Nat Plants 3, 17009. DOI PMID
[31]	Hu XH, Shen S, Wu JL, Liu J, Wang H, He JX, Yao ZL, Bai YF, Zhang X, Zhu Y, Li GB, Zhao JH, You XM, Xu J, Ji YP, Li DQ, Pu M, Zhao ZX, Zhou SX, Zhang JW, Huang YY, Li Y, Ning YS, Lu YL, Huang F, Wang WM, Fan J (2023). A natural allele of proteasome maturation factor improves rice resistance to multiple pathogens. Nat Plants 9, 228-237.
[32]	Hua LX, Wu JZ, Chen CX, Wu WH, He XY, Lin F, Wang L, Ashikawa I, Matsumoto T, Wang L, Pan QH (2012). The isolation of Pi1, an allele at the Pik locus which confers broad spectrum resistance to rice blast. Theor Appl Genet 125, 1047-1055.
[33]	Hutin M, Sabot F, Ghesquière A, Koebnik R, Szurek B (2015). A knowledge-based molecular screen uncovers a broad-spectrum OsSWEET14 resistance allele to bacterial blight from wild rice. Plant J 84, 694-703.
[34]	Im JH, Choi C, Park SR, Hwang DJ (2022). The OsWRKY6 transcriptional cascade functions in basal defense and Xa1-mediated defense of rice against Xanthomonas oryzae pv. oryzae. Planta 255, 47.
[35]	Inukai T, Nagashima S, Kato M (2019). Pid3-I1 is a race- specific partial-resistance allele at the Pid3 blast resistance locus in rice. Theor Appl Genet 132, 395-404.
[36]	Iyer AS, McCouch SR (2004). The rice bacterial blight resistance gene xa5 encodes a novel form of disease resistance. Mol Plant Microbe Interact 17, 1348-1354.
[37]	Ji CH, Ji ZY, Liu B, Cheng H, Liu H, Liu SZ, Yang B, Chen GY (2020). Xa1 allelic R genes activate rice blight resistance suppressed by interfering TAL effectors. Plant Commun 1, 100087.
[38]	Jiao L, Fu SF, Zhang YL, Lu J (2016). U-box E3 ubiquitin ligases regulate stress tolerance and growth of plants. Chin Bull Bot 51, 724-735. (in Chinese)
	缴莉, 付淑芳, 张雅丽, 卢江 (2016). U-box泛素连接酶调控植物抗逆和生长发育. 植物学报 51, 724-735. DOI
[39]	Jones JDG, Staskawicz BJ, Dangl JL (2024). The plant immune system: from discovery to deployment. Cell 187, 2095-2116. DOI PMID
[40]	Kim P, Xue CY, Song HD, Gao Y, Feng L, Li YH, Xuan YH (2021). Tissue-specific activation of DOF11 promotes rice resistance to sheath blight disease and increases grain weight via activation of SWEET14. Plant Biotechnol J 19, 409-411.
[41]	Lee SK, Song MY, Seo YS, Kim HK, Ko S, Cao PJ, Suh JP, Yi G, Roh JH, Lee S, An G, Hahn TR, Wang GL, Ronald P, Jeon JS (2009). Rice Pi5-mediated resistance to Magnaporthe oryzae requires the presence of two coiled-coil-nucleotide-binding-leucine-rich repeat genes. Genetics 181, 1627-1638.
[42]	Li DY, Zhou J, Zheng C, Zheng ES, Liang WF, Tan XJ, Xu RM, Yan CQ, Yang Y, Yi KK, Liu XL, Chen JP, Wang XM (2022). OsTGAL1 suppresses the resistance of rice to bacterial blight disease by regulating the expression of salicylic acid glucosyltransferase OsSGT1. Plant Cell Environ 45, 1584-1602.
[43]	Li N, Wei ST, Chen J, Yang FF, Kong LG, Chen CX, Ding XH, Chu ZH (2018). OsASR2 regulates the expression of a defence-related gene, Os2H16, by targeting the GT-1 cis-element. Plant Biotechnol J 16, 771-783.
[44]	Li WT, Chern M, Yin JJ, Wang J, Chen XW (2019). Recent advances in broad-spectrum resistance to the rice blast disease. Curr Opin Plant Biol 50, 114-120. DOI PMID
[45]	Li XC, Liao YY, Leung DWM, Wang HY, Chen BL, Peng XX, Liu EE (2015). Divergent biochemical and enzymatic properties of oxalate oxidase isoforms encoded by four similar genes in rice. Phytochemistry 118, 216-223. DOI PMID
[46]	Lin F, Chen S, Que ZQ, Wang L, Liu XQ, Pan QH (2007). The blast resistance gene Pi37 encodes a nucleotide binding site-leucine-rich repeat protein and is a member of a resistance gene cluster on rice chromosome 1. Genetics 177, 1871-1880.
[47]	Liu JP, Nie B, Yu BL, Xu FY, Zhang Q, Wang Y, Xu WF (2023). Rice ubiquitin-conjugating enzyme OsUbc13 negatively regulates immunity against pathogens by enhancing the activity of OsSnRK1a. Plant Biotechnol J 21, 1590-1610.
[48]	Liu QS, Yuan M, Zhou Y, Li XH, Xiao JH, Wang SP (2011). A paralog of the MtN3/saliva family recessively confers race-specific resistance to Xanthomonas oryzae in rice. Plant Cell Environ 34, 1958-1969.
[49]	Liu X, Song LL, Zhang H, Lin YJ, Shen XL, Guo JY, Su ML, Shi GS, Wang ZH, Lu GD (2021a). Rice ubiquitin-conjugating enzyme OsUBC26 is essential for immunity to the blast fungus Magnaporthe oryzae. Mol Plant Pathol 22, 1613-1623.
[50]	Liu XQ, Zhang D, Zhang XM, Wang CT, Liu XQ, Tan YP, Wu YH (2012). Study on the interaction between methyl jasmonate and the coiled-coil domain of rice blast resistance protein Pi36 by spectroscopic methods. Spectrochim Acta A Mol Biomol Spectrosc 88, 72-76. DOI PMID
[51]	Liu XY, Zhang ZG (2022). A double-edged sword: reactive oxygen species (ROS) during the rice blast fungus and host interaction. FEBS J 289, 5505-5515.
[52]	Liu Y, Liu B, Zhu XY, Yang JY, Bordeos A, Wang GL, Leach JE, Leung H (2013). Fine-mapping and molecular marker development for Pi56(t), a NBS-LRR gene conferring broad-spectrum resistance to Magnaporthe oryzae in rice. Theor Appl Genet 126, 985-998. DOI PMID
[53]	Liu ZQ, Zhu YJ, Shi HB, Qiu JH, Ding XH, Kou YJ (2021b). Recent progress in rice broad-spectrum disease resistance. Int J Mol Sci 22, 11658.
[54]	Lu X, He Y, Guo JQ, Wang Y, Yan Q, Xiong Q, Shi H, Hou QQ, Yin JJ, An YB, Chen YD, Yang CS, Mao Y, Zhu XB, Tang YY, Liu JL, Bi Y, Song L, Wang L, Yang YH, He M, Li WT, Chen XW, Wang J (2024). Dynamics of epitranscriptomes uncover translational reprogramming directed by ac4C in rice during pathogen infection. Nat Plants 10, 1548-1561.
[55]	Lu Y, Tsuda K (2021). Intimate association of PRR- and NLR-mediated signaling in plant immunity. Mol Plant Microbe Interact 34, 3-14.
[56]	Ma J, Lei CL, Xu XT, Hao K, Wang JL, Cheng ZJ, Ma XD, Ma J, Zhou KN, Zhang X, Guo XP, Wu FQ, Lin QB, Wang CM, Zhai HQ, Wang HY, Wan JM (2015). Pi64, encoding a novel CC-NBS-LRR protein, confers resistance to leaf and neck blast in rice. Mol Plant Microbe Interact 28, 558-568.
[57]	Ma JN, Morel JB, Riemann M, Nick P (2022). Jasmonic acid contributes to rice resistance against Magnaporthe oryzae. BMC Plant Biol 22, 601.
[58]	Ma L, Fang Y, Xiao SQ, Zhou C, Jin ZL, Ye WL, Rao YC (2018). QTL exploration of bacterial leaf streak and their gene expression in rice. Chin Bull Bot 53, 468-476. (in Chinese)
	马路, 方媛, 肖飒清, 周纯, 金哲伦, 叶雯澜, 饶玉春 (2018). 水稻条斑病抗性QTL的挖掘及相关基因的表达. 植物学报 53, 468-476. DOI
[59]	Mohan Babu R, Sajeena A, Vijaya Samundeeswari A, Sreedhar A, Vidhyasekaran P, Seetharaman K, Reddy MS (2003). Induction of systemic resistance to Xanthomonas oryzae pv. oryzae by salicylic acid in Oryza sativa (L.). J Plant Dis Prot 110, 419-431.
[60]	Moon H, Jeong AR, Kwon OK, Park CJ (2022). Oryza- specific orphan protein triggers enhanced resistance to Xanthomonas oryzae pv. oryzae in rice. Front Plant Sci 13, 859375.
[61]	Nabi Z, Manzoor S, Nabi SU, Wani TA, Gulzar H, Farooq M, Arya VM, Baloch FS, Vlădulescu C, Popescu SM, Mansoor S (2024). Pattern-triggered immunity and effector-triggered immunity: crosstalk and cooperation of PRR and NLR-mediated plant defense pathways during host-pathogen interactions. Physiol Mol Biol Plants 30, 587-604.
[62]	Nahar K, Kyndt T, Hause B, Höfte M, Gheysen G (2013). Brassinosteroids suppress rice defense against root-knot nematodes through antagonism with the jasmonate pathway. Mol Plant Microbe Interact 26, 106-115.
[63]	Ngou BPM, Jones JDG, Ding PT (2022). Plant immune networks. Trends Plant Sci 27, 255-273.
[64]	Okuyama Y, Kanzaki H, Abe A, Yoshida K, Tamiru M, Saitoh H, Fujibe T, Matsumura H, Shenton M, Galam DC, Undan J, Ito A, Sone T, Terauchi R (2011). A multifaceted genomics approach allows the isolation of the rice Pia-blast resistance gene consisting of two adjacent NBS-LRR protein genes. Plant J 66, 467-479.
[65]	Panthapulakkal Narayanan S, Lung SC, Liao P, Lo C, Chye ML (2020). The overexpression of OsACBP5 protects transgenic rice against necrotrophic, hemibiotrophic and biotrophic pathogens. Sci Rep 10, 14918. DOI PMID
[66]	Park SC, Kim IR, Kim JY, Lee Y, Yoo SH, Jung JH, Cheong GW, Lee SY, Jang MK, Lee JR (2019). Functional characterization of a rice thioredoxin protein OsTrxm and its cysteine mutant variant with antifungal activity. Antioxidants (Basel) 8, 598.
[67]	Peng XQ, Wang ML (2022). Research advances on resistance genes to bacterial blight disease in rice. Plant Physiol J 58, 472-482. (in Chinese)
	彭小群, 王梦龙 (2022). 水稻白叶枯病抗性基因研究进展. 植物生理学报 58, 472-482.
[68]	Peng YJ, Yang JF, Li X, Zhang YL (2021). Salicylic acid: biosynthesis and signaling. Annu Rev Plant Biol 72, 761-791. DOI PMID
[69]	Phukan UJ, Jeena GS, Shukla RK (2016). WRKY transcription factors: molecular regulation and stress responses in plants. Front Plant Sci 7, 760. DOI PMID
[70]	Rao YC, Luo YL, Ye YH, Xu JM (2025). Research progress on hormones and regulation of rice leaf senescence. J. Zhejiang Normal Univ (Nat Sci) 48, 1-11. (in Chinese)
	饶玉春, 罗怡琳, 叶语涵, 徐江民 (2025). 激素与水稻叶片衰老调控的研究进展. 浙江师范大学学报(自然科学版) 48, 1-11.
[71]	Rao YC, Wu RC, Liu FY, Dai RH (2024). On research progress of RNAi application in prevention and control of rice pests and diseases. J Zhejiang Normal Univ (Nat Sci) 47, 361-369. (in Chinese)
	饶玉春, 吴日成, 刘富远, 戴若惠 (2024). RNAi在水稻病虫害防控中的应用研究进展. 浙江师范大学学报(自然科学版) 47, 361-369.
[72]	Ren ZY, Tang BZ, Xing JJ, Liu CY, Cai X, Hendy A, Kamran M, Liu H, Zheng L, Huang JB, Chen XL (2022). MTA1-mediated RNA m⁶A modification regulates autophagy and is required for infection of the rice blast fungus. New Phytol 235, 247-262.
[73]	Sahu PK, Sao R, Choudhary DK, Thada A, Kumar V, Mondal S, Das BK, Jankuloski L, Sharma D (2022). Advancement in the breeding, biotechnological and genomic tools towards development of durable genetic resistance against the rice blast disease. Plants 11, 2386.
[74]	Shao J, Zhang ZJ, Shi Y, Jiang WQ, Siddique F, Chen LY, Liu GY, Zhu JK, Luo XF, Liu YQ, An JX, Yang CJ, Cui ZN (2024). Application and mechanism of cryptolepine and neocryptolepine derivatives as T3SS inhibitors for control of bacterial leaf blight on rice. J Agric Food Chem 72, 6988-6997.
[75]	Sharma TR, Rai AK, Gupta SK, Singh NK (2010). Broad-spectrum blast resistance gene Pi-k^h cloned from rice line tetep designated as Pi54. J Plant Biochem Biotechnol 19, 87-89.
[76]	Shen XL, Liu HB, Yuan B, Li XH, Xu CG, Wang SP (2011). OsEDR1 negatively regulates rice bacterial resistance via activation of ethylene biosynthesis. Plant Cell Environ 34, 179-191.
[77]	Shi H, Yin JJ, Zhao ZJ, Yu H, Yi H, Xu L, Tong HM, He M, Zhu XB, Lu X, Xiong Q, Li WT, Tang YY, Hou QQ, Song L, Wang L, Chen XQ, Sun CH, Li T, Fan J, Li Y, Qin P, Wang WM, Li SG, Chen XW, Li JY, Wang J (2024). Fine-tuning of IPA1 transactivation activity by E3 ligase IPI7-mediated non-proteolytic K29-ubiquitination during Magnaporthe oryzae infection. Nat Commun 15, 7608.
[78]	Shinde H, Dudhate A, Kadam US, Hong JC (2023). RNA methylation in plants: an overview. Front Plant Sci 14, 1132959.
[79]	Su J, Wang WJ, Han JL, Chen S, Wang CY, Zeng LX, Feng AQ, Yang JY, Zhou B, Zhu XY (2015). Functional divergence of duplicated genes results in a novel blast resistance gene Pi50 at the Pi2/9 locus. Theor Appl Genet 128, 2213-2225.
[80]	Suzuki G, Fukuda M, Lucob-Agustin N, Inukai Y, Gomi K (2022). The mutation of rice MEDIATOR25, OsMED25, induces rice bacterial blight resistance through altering jasmonate- and auxin-signaling. Plants (Basel) 11, 1601.
[81]	Takagi H, Uemura A, Yaegashi H, Tamiru M, Abe A, Mitsuoka C, Utsushi H, Natsume S, Kanzaki H, Matsumura H, Saitoh H, Yoshida K, Cano LM, Kamoun S, Terauchi R (2013). progeny bulk combined with de novo assembly of gap regions identifies the rice blast resistance gene Pii New Phytol 200, 276-283.
[82]	Tian DS, Wang JX, Zeng X, Gu KY, Qiu CX, Yang XB, Zhou ZY, Goh M, Luo YC, Murata-Hori M, White FF, Yin ZC (2014). The rice TAL effector-dependent resistance protein XA10 triggers cell death and calcium depletion in the endoplasmic reticulum. Plant Cell 26, 497-515.
[83]	Uji Y, Kashihara K, Kiyama H, Mochizuki S, Akimitsu K, Gomi K (2019). Jasmonic acid-induced VQ-motif-containing protein OsVQ13 influences the OsWRKY45 signaling pathway and grain size by associating with OsMPK6 in rice. Int J Mol Sci 20, 2917.
[84]	Vidhyasekaran P, Ponmalar TR, Samiyappan R, Velazhahan R, Vimala R, Ramanathan A, Paranidharan V, Muthukrishnan S (1997). Host-specific toxin production by Rhizoctonia solani, the rice sheath blight pathogen. Phytopathology 87, 1258-1263. DOI PMID
[85]	Wang AJ, Ma L, Shu XY, Jiang YQ, Liang J, Zheng AP (2022a). Rice (Oryza sativa L.) cytochrome P450 protein 716A subfamily CYP716A16 regulates disease resistance. BMC Genomics 23, 343.
[86]	Wang AJ, Shu XY, Jing X, Jiao CZ, Chen L, Zhang JF, Ma L, Jiang YQ, Yamamoto N, Li SC, Deng QM, Wang SQ, Zhu J, Liang YY, Zou T, Liu HN, Wang LX, Huang YB, Li P, Zheng AP (2021a). Identification of rice (Oryza sativa L.) genes involved in sheath blight resistance via a genome-wide association study. Plant Biotechnol J 19, 1553-1566.
[87]	Wang CL, Zhang XP, Fan YL, Gao Y, Zhu QL, Zheng CK, Qin TF, Li YQ, Che JY, Zhang MW, Yang B, Liu YG, Zhao KJ (2015a). XA23 is an executor R protein and confers broad-spectrum disease resistance in rice. Mol Plant 8, 290-302.
[88]	Wang G, Chen X, Yu CZ, Shi XB, Lan WX, Gao CF, Yang J, Dai HL, Zhang XW, Zhang HL, Zhao BY, Xie Q, Yu N, He ZH, Zhang Y, Wang ET (2024). Release of a ubiquitin brake activates OsCERK1-triggered immunity in rice. Nature 629, 1158-1164.
[89]	Wang H, Li Y, Chern M, Zhu Y, Zhang LL, Lu JH, Li XP, Dang WQ, Ma XC, Yang ZR, Yao SZ, Zhao ZX, Fan J, Huang YY, Zhang JW, Pu M, Wang J, He M, Li WT, Chen XW, Wu XJ, Li SG, Li P, Li Y, Ronald PC, Wang WM (2021b). Suppression of rice miR168 improves yield, flowering time and immunity. Nat Plants 7, 129-136.
[90]	Wang J, Qu BY, Dou SJ, Li LY, Yin DD, Pang ZQ, Zhou ZZ, Tian MM, Liu GZ, Xie Q, Tang DZ, Chen XW, Zhu LH (2015b). The E3 ligase OsPUB15 interacts with the receptor-like kinase PID2 and regulates plant cell death and innate immunity. BMC Plant Biol 15, 49.
[91]	Wang L, Xu XK, Lin F, Pan QH (2009). Characterization of rice blast resistance genes in the Pik cluster and fine mapping of the Pik-p locus. Phytopathology 99, 900-905.
[92]	Wang LH, Chen J, Zhao YQ, Wang SP, Yuan M (2022b). OsMAPK6 phosphorylates a zinc finger protein OsLIC to promote downstream OsWRKY30 for rice resistance to bacterial blight and leaf streak. J Integr Plant Biol 64, 1116-1130.
[93]	Wang Y, Yue JL, Yang N, Zheng C, Zheng YN, Wu X, Yang J, Zhang HW, Liu LJ, Ning YS, Bhadauria V, Zhao WS, Xie Q, Peng YL, Chen Q (2023). An ERAD- related ubiquitin-conjugating enzyme boosts broad-spectrum disease resistance and yield in rice. Nat Food 4, 774-787. DOI PMID
[94]	Xie YJ, Wang YP, Yu XZ, Lin YL, Zhu YS, Chen JW, Xie HG, Zhang QQ, Wang LN, Wei YD, Xiao YJ, Cai QH, Zheng YM, Wang M, Xie HA, Zhang JF (2022). SH3P2, an SH3 domain-containing protein that interacts with both Pib and AvrPib, suppresses effector-triggered, Pib-mediated immunity in rice. Mol Plant 15, 1931-1946.
[95]	Xie Z, Yan BX, Shou JY, Tang J, Wang X, Zhai KR, Liu JY, Li Q, Luo MZ, Deng YW, He ZH (2019). A nucleotide-binding site-leucine-rich repeat receptor pair confers broad-spectrum disease resistance through physical association in rice. Philos Trans Royal Soc B Biol Sci 374, 20180308.
[96]	Xing JX, Zhang DY, Yin FY, Zhong QF, Wang B, Xiao SQ, Ke X, Wang LX, Zhang Y, Zhao CM, Lu YD, Chen L, Cheng ZQ, Chen LJ (2021). Identification and fine- mapping of a new bacterial blight resistance gene, Xa47(t), in G252, an introgression line of Yuanjiang common wild rice (Oryza rufipogon). Plant Dis 105, 4106-4112.
[97]	Xu DJ, Lin QX, Li ZK, Zhuang XQ, Ling Y, Lai ML, Chen XT, Lu GD (2024). OsOPR10 positively regulates rice blast and bacterial blight resistance. Chin J Rice Sci 38, 364-374. (in Chinese)
	许丹洁, 林巧霞, 李正康, 庄小倩, 凌宇, 赖美玲, 陈晓婷, 鲁国东 (2024). OsOPR10正调控水稻对稻瘟病和白叶枯病的抗性. 中国水稻科学 38, 364-374. DOI
[98]	Xu X, Hayashi N, Wang CT, Fukuoka S, Kawasaki S, Takatsuji H, Jiang CJ (2014a). Rice blast resistance gene Pikahei-1(t), a member of a resistance gene cluster on chromosome 4, encodes a nucleotide-binding site and leucine-rich repeat protein. Mol Breed 34, 691-700.
[99]	Xu X, Lv QM, Shang JJ, Pang ZQ, Zhou ZZ, Wang J, Jiang GH, Tao Y, Xu Q, Li XB, Zhao XF, Li SG, Xu JC, Zhu LH (2014b). Excavation of Pid3 orthologs with differential resistance spectra to Magnaporthe oryzae in rice resource. PLoS One 9, 93275.
[100]	Yan YQ, Wang H, Bi Y, Song FM (2024). Rice E3 ubiquitin ligases: from key modulators of host immunity to potential breeding applications. Plant Commun 5, 101128.
[101]	Yang C, Li W, Cao JD, Meng FW, Yu YQ, Huang JK, Jiang L, Liu MX, Zhang ZG, Chen XW, Miyamoto K, Yamane H, Zhang JS, Chen SY, Liu J (2017). Activation of ethylene signaling pathways enhances disease resistance by regulating ROS and phytoalexin production in rice. Plant J 89, 338-353.
[102]	Yang S, Fu YW, Zhang Y, Yuan DP, Li S, Kumar V, Mei Q, Xuan YH (2023a). Rhizoctonia solani transcriptional activator interacts with rice WRKY53 and grassy tiller 1 to activate SWEET transporters for nutrition. J Adv Res 50, 1-12.
[103]	Yang Y, Li J, Li H, Ding Y, Wu W, Qin R, Ni J, Xu R, Wei P, Yang J (2023b). OsGSTU5 and OsGSTU37 encoding glutathione reductases are required for cadmium tolerance in rice. Int J Environ Sci Technol 20, 10253-10260.
[104]	Yang Y, Zhou YH, Sun J, Liang WF, Chen XY, Wang XM, Zhou J, Yu CL, Wang JM, Wu SL, Yao XM, Zhou YJ, Zhu J, Yan CQ, Zheng BS, Chen JP (2022). Research progress on cloning and function of Xa genes against rice bacterial blight. Front Plant Sci 13, 847199.
[105]	Yang YX, Ahammed GJ, Wu CJ, Fan SY, Zhou YH (2015). Crosstalk among jasmonate, salicylate and ethylene signaling pathways in plant disease and immune responses. Curr Protein Pept Sci 16, 450-461.
[106]	Yin X, Zou BH, Hong XX, Gao MJ, Yang WB, Zhong XB, He Y, Kuai P, Lou YG, Huang JR, Hua J, He ZH (2018). Rice copine genes OsBON1 and OsBON3 function as suppressors of broad-spectrum disease resistance. Plant Biotechnol J 16, 1476-1487.
[107]	Yoshimura S, Yamanouchi U, Katayose Y, Toki S, Wang ZX, Kono I, Kurata N, Yano M, Iwata N, Sasaki T (1998). Expression of Xa1, a bacterial blight-resistance gene in rice, is induced by bacterial inoculation. Proc Natl Acad Sci USA 95, 1663-1668. DOI PMID
[108]	Younas MU, Qasim M, Ahmad I, Feng ZM, Iqbal R, Abdelbacki AMM, Rajput N, Jiang XH, Rao B, Zuo SM (2024). Allelic variation in rice blast resistance: a pathway to sustainable disease management. Mol Biol Rep 51, 935. DOI PMID
[109]	Yu SB, Ali J, Zhang CP, Li ZK, Zhang QF (2020). Genomic breeding of green super rice varieties and their deployment in Asia and Africa. Theor Appl Genet 133, 1427-1442. DOI PMID
[110]	Yu XQ, Niu HQ, Liu C, Wang HL, Yin WL, Xia XL (2024). PTI-ETI synergistic signal mechanisms in plant immunity. Plant Biotechnol J 22, 2113-2128.
[111]	Yu Y, Ma L, Wang XY, Zhao Z, Wang W, Fan YX, Liu KQ, Jiang TT, Xiong ZW, Song QS, Li CQ, Wang PT, Ma WJ, Xu HN, Wang XY, Zhao ZJ, Wang JF, Zhang HS, Bao YM (2022). Genome-wide association study identifies a rice panicle blast resistance gene, Pb2, encoding NLR protein. Int J Mol Sci 23, 5668.
[112]	Yu ZH, Mackill DJ, Bonman JM, Tanksley SD (1991). Tagging genes for blast resistance in rice via linkage to RFLP markers. Theor Appl Genet 81, 471-476. DOI PMID
[113]	Yuan DP, Yang S, Feng L, Chu J, Dong H, Sun J, Chen H, Li Z, Yamamoto N, Zheng AP, Li S, Yoon HC, Chen JS, Ma DR, Xuan YH (2023). Red-light receptor phytochrome B inhibits BZR1-NAC028-CAD8B signaling to negatively regulate rice resistance to sheath blight. Plant Cell Environ 46, 1249-1263.
[114]	Yuan DP, Zhang C, Wang ZY, Zhu XF, Xuan YH (2018). RAVL1 activates brassinosteroids and ethylene signaling to modulate response to sheath blight disease in rice. Phytopathology 108, 1104-1113.
[115]	Zhai C, Lin F, Dong ZQ, He XY, Yuan B, Zeng XS, Wang L, Pan QH (2011). The isolation and characterization of Pik, a rice blast resistance gene which emerged after rice domestication. New Phytol 189, 321-334.
[116]	Zhai C, Zhang Y, Yao N, Lin F, Liu Z, Dong ZQ, Wang L, Pan QH (2014). Function and interaction of the coupled genes responsible for Pik-h encoded rice blast resistance. PLoS One 9, 98067.
[117]	Zhai KR, Deng YW, Liang D, Tang J, Liu J, Yan BX, Yin X, Lin H, Chen FD, Yang DY, Xie Z, Liu JY, Li Q, Zhang L, He ZH (2019). RRM transcription factors interact with NLRs and regulate broad-spectrum blast resistance in rice. Mol Cell 74, 996-1009. DOI PMID
[118]	Zhang DD, Tian CJ, Yin KQ, Wang WY, Qiu JL (2019). Postinvasive bacterial resistance conferred by open stomata in rice. Mol Plant Microbe Interact 32, 255-266.
[119]	Zhao HJ, Wang XY, Jia YL, Minkenberg B, Wheatley M, Fan JB, Jia MH, Famoso A, Edwards JD, Wamishe Y, Valent B, Wang GL, Yang YN (2018). The rice blast resistance gene Ptr encodes an atypical protein required for broad-spectrum disease resistance. Nat Commun 9, 2039.
[120]	Zhao YD, Zhong XH, Xu GJ, Zhu XY, Shi YL, Liu MH, Wang RY, Kang HX, You XM, Ning YS, Wang GL, Wang XL (2024). The F-box protein OsFBX156 positively regulates rice defence against the blast fungus Magnaporthe oryzae by mediating ubiquitination-dependent degradation of OsHSP71.1. Mol Plant Pathol 25, 13459.
[121]	Zhou XG, Liao HC, Chern M, Yin JJ, Chen YF, Wang JP, Zhu XB, Chen ZX, Yuan C, Zhao W, Wang J, Li WT, He M, Ma BT, Wang JC, Qin P, Chen WL, Wang YP, Liu JL, Qian YW, Wang WM, Wu XJ, Li P, Zhu LH, Li SG, Ronald PC, Chen XW (2018). Loss of function of a rice TPR-domain RNA-binding protein confers broad-spectrum disease resistance. Proc Natl Acad Sci USA 115, 3174-3179. DOI PMID
[122]	Zhou YL, Shen XL, Zhou LS, Lin QX, Wang ZL, Chen J, Feng HJ, Zhang ZW, Chen XT, Lu GD (2022). OsLOX10 positively regulates defense responses of rice to rice blast and bacterial blight. Chin J Rice Sci 36, 348-356. (in Chinese)
	周永林, 申小磊, 周立帅, 林巧霞, 王朝露, 陈静, 冯慧捷, 张振文, 陈晓婷, 鲁国东 (2022). OsLOX10正调控水稻对稻瘟病和白叶枯病的抗性. 中国水稻科学 36, 348-356. DOI
[123]	Zhou ZZ, Pang ZQ, Zhao SL, Zhang LL, Lv QM, Yin DD, Li DY, Liu X, Zhao XF, Li XB, Wang WM, Zhu LH (2019). Importance of OsRac1 and RAI1 in signaling of nucleotide-binding site leucine-rich repeat protein-mediated resistance to rice blast disease. New Phytol 223, 828-838.
[124]	Zou T, Li GW, Liu MM, Liu R, Yang SY, Wang K, Lu LH, Ye QY, Liu JX, Liang J, Deng QM, Wang SQ, Zhu J, Liang YY, Liu HN, Yu XM, Sun CH, Li P, Li SC (2023). A ubiquitin-specific protease functions in regulating cell death and immune responses in rice. Plant Cell Environ 46, 1312-1326.

基因名称	RAP号	编码蛋白类型	基因功能	参考文献
Pit	Os01g0149500	NBS-LRR	编码受体蛋白, 识别病原物	Hayashi et al., 2016
Pi64	Os01g0781200	NBS-LRR	编码受体蛋白, 识别病原物	Ma et al., 2015
Pish	Os01g0782100	NBS-LRR	编码受体蛋白, 识别病原物	Cheng et al., 2024
Pi35	Os01g0782100	NBS-LRR	Pish的等位基因	Fukuoka et al., 2014
Pi37	Os01g0781700	NBS-LRR	编码受体蛋白, 识别病原物	Lin et al., 2007
Pib	Os02g0818500	NBS-LRR	编码受体蛋白, 识别病原物	Xie et al., 2022
pi21	Os04g0401000	含脯氨酸蛋白	编码富含脯氨酸的蛋白, 加快免疫反应	Fukuoka et al., 2009
Pi63/Pikahei-1(t)	-	NBS-LRR	4号染色体抗病基因簇组成部分	Xu et al., 2014a
Pi50	Os06g0286700	NBS-LRR	6号染色体抗病基因簇组成部分	Su et al., 2015
Pi2	Os06g0286700	NBS-LRR	6号染色体抗病基因簇组成部分	Xie et al., 2019
Pi9	Os06g0286700	NBS-LRR	6号染色体抗病基因簇组成部分	Su et al., 2015
Piz-t	Os06g0286700	NBS-LRR	调控蛋白同源二聚化活性	Yu et al., 1991
Pigm	Os06g0286700	NBS-LRR	调控蛋白同源二聚化活性	Zhai et al., 2019
Pid3	Os06g0330100	NBS-LRR	调控防御反应调节相关的宿主因子活性	Zhou et al., 2019
Pi25	Os06g0330100	NBS-LRR	调控防御反应调节相关的宿主因子活性	Xu et al., 2014b
Pid3-I1	Os06g0330100	NBS-LRR	调控防御反应调节相关的宿主因子活性	Inukai et al., 2019
Pid2	Os06g0494100	受体蛋白激酶	编码植物凝集素类受体激酶	Wang et al., 2015b
Pi36	Os08g0150150	NBS-LRR	编码受体蛋白, 识别病原物	Liu et al., 2012
Pii	Os09g0327600	NBS-LRR	编码受体蛋白, 识别病原物	Takagi et al., 2013
Pi5	Os09g0327600	NBS-LRR	编码Pi5-1和Pi5-2蛋白, 蛋白互作调控免疫反应	Lee et al., 2009
Pi56	Os09g0328951	NBS-LRR	编码受体蛋白, 识别病原物	Liu et al., 2013
Pia	Os11g0225100	NBS-LRR	编码受体蛋白, 识别病原物	Okuyama et al., 2011
PiCO39	Os11g0225300	NBS-LRR	编码受体蛋白, 识别病原物	Césari et al., 2014
Pb1	Os11g0598500	NBS-LRR	编码受体蛋白, 识别病原物	Hayashi et al., 2010
Pik-h	Os11g0639100	NBS-LRR	编码Pikh-1和Pikh-2蛋白, 蛋白互作调控免疫反应	Zhai et al., 2014
Pi54rh	Os11g0639100	NBS-LRR	编码受体蛋白, 影响防御信号转导	Das et al., 2012
Pi54/Pi-kh	Os11g0639100	NBS-LRR	编码受体蛋白, 影响防御信号转导	Sharma et al., 2010
Pi54of	Os11g0639100	NBS-LRR	编码受体蛋白, 影响防御信号转导	Devanna et al., 2014
Pb2	Os11g0682400	NBS-LRR	编码受体蛋白, 识别病原物	Yu et al., 2022
Pi-k	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Zhai et al., 2011
Pik-m	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Ashikawa et al., 2008
Pik-p	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Wang et al., 2009
Pike	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Chen et al., 2015
Pi1	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Hua et al., 2012
Pita	Os12g0281300	NBS-LRR	编码受体蛋白, 识别病原物	Bryan et al., 2000
Ptr	Os12g0285100	ARM重复序列的抗性蛋白	推测编码E3泛素连接酶	Zhao et al., 2018

基因名称	RAP号	编码蛋白类型	基因功能	参考文献
Pit	Os01g0149500	NBS-LRR	编码受体蛋白, 识别病原物	Hayashi et al., 2016
Pi64	Os01g0781200	NBS-LRR	编码受体蛋白, 识别病原物	Ma et al., 2015
Pish	Os01g0782100	NBS-LRR	编码受体蛋白, 识别病原物	Cheng et al., 2024
Pi35	Os01g0782100	NBS-LRR	Pish的等位基因	Fukuoka et al., 2014
Pi37	Os01g0781700	NBS-LRR	编码受体蛋白, 识别病原物	Lin et al., 2007
Pib	Os02g0818500	NBS-LRR	编码受体蛋白, 识别病原物	Xie et al., 2022
pi21	Os04g0401000	含脯氨酸蛋白	编码富含脯氨酸的蛋白, 加快免疫反应	Fukuoka et al., 2009
Pi63/Pikahei-1(t)	-	NBS-LRR	4号染色体抗病基因簇组成部分	Xu et al., 2014a
Pi50	Os06g0286700	NBS-LRR	6号染色体抗病基因簇组成部分	Su et al., 2015
Pi2	Os06g0286700	NBS-LRR	6号染色体抗病基因簇组成部分	Xie et al., 2019
Pi9	Os06g0286700	NBS-LRR	6号染色体抗病基因簇组成部分	Su et al., 2015
Piz-t	Os06g0286700	NBS-LRR	调控蛋白同源二聚化活性	Yu et al., 1991
Pigm	Os06g0286700	NBS-LRR	调控蛋白同源二聚化活性	Zhai et al., 2019
Pid3	Os06g0330100	NBS-LRR	调控防御反应调节相关的宿主因子活性	Zhou et al., 2019
Pi25	Os06g0330100	NBS-LRR	调控防御反应调节相关的宿主因子活性	Xu et al., 2014b
Pid3-I1	Os06g0330100	NBS-LRR	调控防御反应调节相关的宿主因子活性	Inukai et al., 2019
Pid2	Os06g0494100	受体蛋白激酶	编码植物凝集素类受体激酶	Wang et al., 2015b
Pi36	Os08g0150150	NBS-LRR	编码受体蛋白, 识别病原物	Liu et al., 2012
Pii	Os09g0327600	NBS-LRR	编码受体蛋白, 识别病原物	Takagi et al., 2013
Pi5	Os09g0327600	NBS-LRR	编码Pi5-1和Pi5-2蛋白, 蛋白互作调控免疫反应	Lee et al., 2009
Pi56	Os09g0328951	NBS-LRR	编码受体蛋白, 识别病原物	Liu et al., 2013
Pia	Os11g0225100	NBS-LRR	编码受体蛋白, 识别病原物	Okuyama et al., 2011
PiCO39	Os11g0225300	NBS-LRR	编码受体蛋白, 识别病原物	Césari et al., 2014
Pb1	Os11g0598500	NBS-LRR	编码受体蛋白, 识别病原物	Hayashi et al., 2010
Pik-h	Os11g0639100	NBS-LRR	编码Pikh-1和Pikh-2蛋白, 蛋白互作调控免疫反应	Zhai et al., 2014
Pi54rh	Os11g0639100	NBS-LRR	编码受体蛋白, 影响防御信号转导	Das et al., 2012
Pi54/Pi-kh	Os11g0639100	NBS-LRR	编码受体蛋白, 影响防御信号转导	Sharma et al., 2010
Pi54of	Os11g0639100	NBS-LRR	编码受体蛋白, 影响防御信号转导	Devanna et al., 2014
Pb2	Os11g0682400	NBS-LRR	编码受体蛋白, 识别病原物	Yu et al., 2022
Pi-k	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Zhai et al., 2011
Pik-m	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Ashikawa et al., 2008
Pik-p	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Wang et al., 2009
Pike	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Chen et al., 2015
Pi1	Os11g0689100	NBS-LRR	11号染色体抗病基因簇组成部分	Hua et al., 2012
Pita	Os12g0281300	NBS-LRR	编码受体蛋白, 识别病原物	Bryan et al., 2000
Ptr	Os12g0285100	ARM重复序列的抗性蛋白	推测编码E3泛素连接酶	Zhao et al., 2018

基因名称	RAP号	编码蛋白类型	基因功能	参考文献
Xa1	Os04g0622600	NLR类	NBS-LRR家族; 诱导细胞凋亡	Yoshimura et al., 1998
Xa2	Os04g0622600	NLR类	Xa2、Xa14、Xa31(t)、Xa45(t)都是Xa1的等位基因, 能被TALE激活抗性(效应子触发的抗性)并被TALE所抑制(效应子触发的感病性)	Ji et al., 2020
Xa14	Os04g0622600	NLR类
Xa31(t)	Os04g0622600	NLR类
Xa45(t)	Os04g0622600	NLR类
xa5	Os05g0107700	其它	编码真核生物转录因子IIA伽马亚基	Iyer and McCouch, 2004
Xa7	Os06g0280200	Executor类	受AvrXa7诱导表达, 激活超敏反应	Chen et al., 2021
Xa27	Os06g0599600	Executor类	受AvrXa27诱导表达, 激活防御反应	Gu et al., 2005
xa13	Os08g0535200	SWEET类	调控铜离子在水稻体内的重新分布	Chu et al., 2006
Xa10	Os11g0137600	Executor类	受AvrXa10诱导表达, 诱导细胞程序性死亡	Tian et al., 2014
xa41(t)	Os11g0508600	SWEET类	启动子与TALE蛋白的结合出现差错	Hutin et al., 2015
Xa21	Os11g0569733	激酶类	编码类受体蛋白激酶	Andaya and Ronald, 2003
Xa23	Os11g0586701	Executor类	受AvrXa23诱导表达, 激活超敏反应	Wang et al., 2015a
Xa47(t)	Os11g0688832	NLR类	编码NLR类蛋白, 受病原菌诱导表达	Xing et al., 2021
Xa3/Xa26	Os11g0694850	激酶类	编码LRR受体激酶, 识别病原物	Cao et al., 2007
Xa4	-	激酶类	编码细胞壁相关激酶	Hu et al., 2017
xa25	Os12g0476200	SWEET类	编码MtN3/saliva家族成员	Liu et al., 2011

基因名称	RAP号	编码蛋白类型	基因功能	参考文献
Xa1	Os04g0622600	NLR类	NBS-LRR家族; 诱导细胞凋亡	Yoshimura et al., 1998
Xa2	Os04g0622600	NLR类	Xa2、Xa14、Xa31(t)、Xa45(t)都是Xa1的等位基因, 能被TALE激活抗性(效应子触发的抗性)并被TALE所抑制(效应子触发的感病性)	Ji et al., 2020
Xa14	Os04g0622600	NLR类
Xa31(t)	Os04g0622600	NLR类
Xa45(t)	Os04g0622600	NLR类
xa5	Os05g0107700	其它	编码真核生物转录因子IIA伽马亚基	Iyer and McCouch, 2004
Xa7	Os06g0280200	Executor类	受AvrXa7诱导表达, 激活超敏反应	Chen et al., 2021
Xa27	Os06g0599600	Executor类	受AvrXa27诱导表达, 激活防御反应	Gu et al., 2005
xa13	Os08g0535200	SWEET类	调控铜离子在水稻体内的重新分布	Chu et al., 2006
Xa10	Os11g0137600	Executor类	受AvrXa10诱导表达, 诱导细胞程序性死亡	Tian et al., 2014
xa41(t)	Os11g0508600	SWEET类	启动子与TALE蛋白的结合出现差错	Hutin et al., 2015
Xa21	Os11g0569733	激酶类	编码类受体蛋白激酶	Andaya and Ronald, 2003
Xa23	Os11g0586701	Executor类	受AvrXa23诱导表达, 激活超敏反应	Wang et al., 2015a
Xa47(t)	Os11g0688832	NLR类	编码NLR类蛋白, 受病原菌诱导表达	Xing et al., 2021
Xa3/Xa26	Os11g0694850	激酶类	编码LRR受体激酶, 识别病原物	Cao et al., 2007
Xa4	-	激酶类	编码细胞壁相关激酶	Hu et al., 2017
xa25	Os12g0476200	SWEET类	编码MtN3/saliva家族成员	Liu et al., 2011

基因	RAP号	基因功能	参考文献
OsRLCK5	Os01g0114100	与丝氨酸羟甲基转移酶互作调控活性氧(ROS)水平	Wang et al., 2021a
OsPAL06	Os02g0627100	编码苯丙氨酸脱氨酶	Zhou et al., 2018
OsBON1	Os02g0521300	调控茉莉酸(JA)与水杨酸(SA)介导通路	Yin et al., 2018
OsPhyB	Os03g0309200	调控红光/远红光受体活性	Yuan et al., 2023
OsACBP5	Os03g0243600	调控JA与SA介导通路	Panthapulakkal Narayanan et al., 2020
OsUMP1	Os03g0583700	调控过氧化氢分解与蛋白分解代谢	Hu et al., 2023
Osoxo4	Os03g0694000	调控JA、脱落酸(ABA)与SA信号通路	Dong et al., 2021
OsGLP3-4	Os03g0693800	调控JA、ABA与SA信号通路	Li et al., 2015
RAVL1	Os04g0581400	参与油菜素内酯信号途径和合成途径基因表达	Yuan et al., 2018
OsWRKY45	Os05g0322900	SA信号通路介导的中心调节因子	Uji et al., 2019
OsWRKY53	Os05g0343400	正调控水稻油菜素内酯信号	Yang et al., 2023a
OsNYC3	Os06g0354700	α/β折叠水解酶家族蛋白, 调控氧化还原酶活性	Cao et al., 2022
OsBZR1	Os07g0580500	调控DNA转录因子活性	Yuan et al., 2023
CYP716A16	Os07g0518100	调控JA与SA介导通路和ROS水平	Wang et al., 2022a
OsWRKY30	Os08g0499300	促进内源JA积累和病程相关蛋白(PR)基因表达	Wang et al., 2022b
OsGSTU5	Os09g0367700	增强抗氧化酶和谷胱甘肽还原酶活性	Yang et al., 2023b
KLP	Os10g0512800	调控转录辅助激活因子活性	Chu et al., 2021
OsSWEET14	Os11g0508600	调控对糖的非特异性运输	Kim et al., 2021
OsAFB3	Os11g0515500	调控生长素受体活性	Guo et al., 2021
OsASR2	Os11g0167800	靶向GT-1顺式作用元件, 激活防御相关基因表达	Li et al., 2018
OsTrxm	Os12g0188700	硫氧还蛋白互作调控ROS清除	Park et al., 2019