中国水稻主产区白叶枯病菌致病型分析及近等基因系鉴别寄主的构建

doi:10.3864/j.issn.0578-1752.2022.21.007

中国农业科学 ›› 2022, Vol. 55 ›› Issue (21): 4175-4195.doi: 10.3864/j.issn.0578-1752.2022.21.007

中国水稻主产区白叶枯病菌致病型分析及近等基因系鉴别寄主的构建

冯爱卿(),汪聪颖,张梅英,陈炳,封金奇,陈凯玲,汪文娟,杨健源,苏菁,曾列先,陈深,朱小源()

广东省农业科学院植物保护研究所/广东省植物保护新技术重点实验室，广州 510640

收稿日期:2022-05-10 接受日期:2022-06-03 出版日期:2022-11-01 发布日期:2022-11-09
通讯作者: 朱小源
作者简介:冯爱卿，E-mail：fengaq@gdppri.com。
基金资助:
国家水稻产业技术体系(CARS-01-35);国家重点研发计划(2016YFD0300707);广东省现代农业产业技术体系专项资金(2021KJ105);广东省农业科学院学科团队建设项目(201613TD);广东省农业科学院学科团队建设项目(202116TD);广东省自然科学基金面上项目(2021A1515012497);广东省农业科学院协同创新中心项目(XT202211)

Pathotype Analysis of Xanthomonas oryzae pv. oryzae in Main Rice Producing Regions of China and Establishment of Differential Hosts of Near-Isogenic Lines

FENG AiQing(),WANG CongYing,ZHANG MeiYing,CHEN Bing,FENG JinQi,CHEN KaiLing,WANG WenJuan,YANG JianYuan,SU Jing,ZENG LieXian,CHEN Shen,ZHU XiaoYuan()

Plant Protection Research Institute, Guangdong Academy of Agricultural Sciences/Guangdong Provincial Key Laboratory of High Technology for Plant Protection, Guangzhou 510640

Received:2022-05-10 Accepted:2022-06-03 Online:2022-11-01 Published:2022-11-09
Contact: XiaoYuan ZHU

摘要/Abstract

摘要：

【目的】分析中国不同稻区白叶枯病菌（Xanthomonas oryzae pv. oryzae）致病型，建立近等基因系鉴别寄主，为白叶枯病菌群体结构田间实时准确监测、抗性品种应用以及抗病育种提供科学依据。【方法】利用中国鉴别寄主、IR24以及15个抗白叶枯病近等基因系等共21个鉴别寄主，采用人工剪叶接种方法，对2018—2021年采自广东、广西、海南、浙江、湖南、辽宁、云南共7个省（自治区）的954个单菌落分离菌株进行致病型测定，探明白叶枯病菌致病型种类、分布及毒性分化;基于测试菌株与15个近等基因系及IR24的抗感互作，应用主成分因子分析法，开展近等基因系与病菌互作的变量因子分析，构建白叶枯病菌致病型近等基因系鉴别寄主;基于抗病基因与测试菌株的抗感反应，分析抗病基因聚合效应。【结果】954个测试菌株在中国鉴别寄主上鉴定出11个致病型，包括SRRRR（I）、SSRRR（Ⅱ）、SSSRR（Ⅲ）、SSSSR（Ⅳ）、SSRRS（V）、SRSRR（Ⅵ）、SSSSS（Ⅸ）、SSSRS（新型1）、SRSRS（新型2）、SRSSS（新型3）以及SSRSS（新型4），占测试菌株的比率分别为11.53％、4.82％、7.34％、6.18％、7.23％、1.05%、59.96％、1.57%、0.10%、0.10%、0.10%。Ⅸ型菌作为致病性最广的强毒菌系已上升为华南和长江中下游湖南和浙江稻区的优势致病型，西南稻区的云南以Ⅳ型菌为主，东北稻区的辽宁以I型菌为主。15个水稻抗白叶枯病近等基因系对954个菌株的抗感性分析结果表明，测试的15个近等基因系可分为5种类型，第Ⅰ类为高感基因系，包括IRBB1、IRBB2、IRBB10、IRBB11、IRBB4;第Ⅱ类为中感基因系，包括IRBB3、IRBB203、IRBB14;第Ⅲ类为中抗基因系，包括IRBB8、IRBB13;第Ⅳ类为抗病基因系，有IRBB21;第Ⅴ类为高抗基因系，包括IRBB5、IRBB7、CBB23、GDBB23;测试菌株中，出现可侵染抗病基因xa5的有42个、Xa7的有34个、Xa23的有31个。对以白叶枯病近等基因系为主的16个品种（系）与954个菌株组成的抗感互作变量数据矩阵进行因子分析，以解释总变量>85.0%为界，提取出8个主成分因子，组建了以近等基因系为主的10个品种(系)组成白叶枯病菌近等基因系鉴别寄主，按其对变量方差贡献大小，这些寄主分别为IRBB10（Xa10）、IRBB4（Xa4）、GDBB23（Xa23）、IRBB5（xa5）、IRBB7（Xa7）、IRBB21（Xa21）、IR24（Xa18）、IRBB13（xa13）、IRBB3（Xa3）、金刚30;新鉴别寄主可将954个测试菌株划分为55个致病型，对测试稻区的白叶枯病菌菌株表现出较好的鉴别力。基因聚合联合抗性分析表明，不同抗病基因聚合对病菌的抗性频率有一定的提升，不同抗病基因对测试菌株的抗性具有一定的互补性。【结论】监测稻区的白叶枯病菌系趋向多样化，毒性分化明显，强毒菌系Ⅸ型菌在部分稻区已上升为优势致病型，侵染xa5、Xa7及Xa23等广谱抗性基因的菌株有上升趋势;抗病基因聚合可拓宽品种对病原菌系的抗性谱，是培育广谱抗性品种的有效途径;近等基因系鉴别寄主的建立与应用可为白叶枯病发生流行的精准监测以及田间实时预警提供技术支撑。

关键词: 水稻白叶枯病菌, 致病型, 近等基因系, 鉴别寄主, 抗性

Abstract:

【Objective】The pathotypes of Xanthomonas oryzae pv. oryzae (Xoo) in diverse rice regions in China were analyzed, and the differential hosts of near-isogenic lines (NILs) were established to provide a scientific basis for accurate field monitoring of Xoo population, application of resistant varieties and breeding of resistant varieties against rice bacterial blight.【Method】To explore virulence diversity and distribution of pathotypes of Xoo, the pathotypes of 954 single colony strains collected from Guangdong, Guangxi, Hainan, Zhejiang, Hunan, Liaoning and Yunnan provinces (Autonomous Region) from 2018 to 2021 were identified by artificial leaf-cutting inoculation on 21 differential hosts including Chinese differential hosts, IR24 and 15 bacterial blight NILs. Based on the resistant and susceptible interactions among the tested Xoo strains, 15 NILs and IR24, the principal component analysis (PCA) was used to analyze the variable factors of which help abstraction of the candidate differentials. Based on the resistant and susceptible interactions between resistance genes and tested strains, the pyramiding effect of resistance genes was analyzed.【Result】The tested 954 Xoo strains were divided into 11 pathotypes including SRRRR (I), SSRRR (Ⅱ), SSSRR (Ⅲ), SSSSR (Ⅳ), SSRRS (Ⅴ), SRSRR (Ⅵ), SSSSS (Ⅸ), SSSRS (new pathotype 1), SRSRS (new pathotype 2), SRSSS (new pathotype 3) and SSRSS (new pathotype 4) based on Chinese differential hosts. The percentage of each pathotype mentioned above was 11.53%, 4.82%, 7.34%, 6.18%, 7.23%, 1.05%, 59.96%, 1.57%, 0.10%, 0.10% and 0.10%, respectively. Pathotype Ⅸ, with broad pathogenicity and more virulence, had become the predominant race in the South China rice region and the Yangtze River rice region (Hunan, Zhejiang), and the predominant pathotypes in Yunnan (Southwest China) and Liaoning (Northeast China) were pathotype Ⅳ and pathotype I, respectively. The resistance and susceptibility of 15 rice NILs to 954 Xoo strains were analyzed. The results showed that the 15 NILs could be divided into five types, highly susceptible lines with IRBB1, IRBB2, IRBB10, IRBB11, IRBB4, the moderately susceptible lines with IRBB3, IRBB203, IRBB14, moderately resistant lines with IRBB8, IRBB13, resistant lines with IRBB21, highly resistant lines with IRBB5, IRBB7, CBB23, GDBB23. Among the tested Xoo strains, 42 strains could infect xa5, 34 strains could infect Xa7, and 31 strains could infect Xa23. Factors were extracted from the interaction variable data matrix among 954 strains and 16 varieties (15 NILs and its recurrent parent IR24). With the total explained variable >85.0% as the boundary, eight principal components were extracted, and ten varieties (lines) mainly with NILs were constructed as differential hosts. Accord to their contribution to interaction variances, these differential hosts were IRBB10 (Xa10), IRBB4 (Xa4), GDBB23 (Xa23), IRBB5 (xa5), IRBB7 (Xa7), IRBB21 (Xa21), IR24 (Xa18), IRBB13 (xa13), IRBB3 (Xa3), Jingang30. With new differential hosts, 954 tested Xoo strains could be divided into 55 pathotypes, the newly developed differential hosts showed good discrimination ability for monitoring the virulence of Xoo in the tested rice regions. The results of gene pyramiding combined resistance analysis showed that the resistance frequency of different resistance genes pyramiding to Xoo was increased, and different resistance genes had diverse complementarity to the resistance of the test strains. 【Conclusion】The Xoo strains in the monitoring rice regions tended to be diversified, and the virulence differentiation was obvious. Pathotype Ⅸ of the virulent strain had become the prevailing races in some rice production regions, and the number of the strains compactible with broad-spectrum resistance genes xa5, Xa7 and Xa23 showed an increasing trend. Resistance gene polymerization can broaden the resistance spectrum of varieties to pathogen lines, which is an effective way to breed broad-spectrum resistant varieties. The establishment and application of NILs differential host could provide technical support for precise monitoring of the occurrence and early warning of epidemic of bacterial blight disease in the field.

Key words: Xanthomonas oryzae pv. oryzae (Xoo), pathotype, near-isogenic line (NIL), differential host, resistance

冯爱卿,汪聪颖,张梅英,陈炳,封金奇,陈凯玲,汪文娟,杨健源,苏菁,曾列先,陈深,朱小源. 中国水稻主产区白叶枯病菌致病型分析及近等基因系鉴别寄主的构建[J]. 中国农业科学, 2022, 55(21): 4175-4195.

FENG AiQing,WANG CongYing,ZHANG MeiYing,CHEN Bing,FENG JinQi,CHEN KaiLing,WANG WenJuan,YANG JianYuan,SU Jing,ZENG LieXian,CHEN Shen,ZHU XiaoYuan. Pathotype Analysis of Xanthomonas oryzae pv. oryzae in Main Rice Producing Regions of China and Establishment of Differential Hosts of Near-Isogenic Lines[J]. Scientia Agricultura Sinica, 2022, 55(21): 4175-4195.

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https://www.chinaagrisci.com/CN/Y2022/V55/I21/4175

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测试菌株在新鉴别寄主上的致病反应"

序号 Serial number	致病型 <BOLD>P</BOLD>athotype	寄主反应Host response										致病谱Pathogenicity spectrum (%)	各致病型的菌株数 Number of strains of each pathotype	各致病型发生频率 Percentage of each pathotype (%)
序号 Serial number	致病型 <BOLD>P</BOLD>athotype	GDBB23	IRBB7	IRBB5	IRBB21	IRBB13	IRBB3	IRBB4	IRBB10	IR24	CBBD1	致病谱Pathogenicity spectrum (%)	各致病型的菌株数 Number of strains of each pathotype	各致病型发生频率 Percentage of each pathotype (%)
R1	RRRRRRRRRS	R	R	R	R	R	R	R	R	R	S	9.09	36	3.77
R2	RRRRRRRRSS	R	R	R	R	R	R	R	R	S	S	18.18	59	6.18
R3	RRRRRRRSRS	R	R	R	R	R	R	R	S	R	S	18.18	4	0.42
R4	RRRRRRSRRS	R	R	R	R	R	R	S	R	R	S	18.18	9	0.94
R5	RRRRSRRRRS	R	R	R	R	S	R	R	R	R	S	18.18	1	0.10
R6	SRRRRRRRRS	S	R	R	R	R	R	R	R	R	S	18.18	1	0.10
R7	RRRRRRRSSS	R	R	R	R	R	R	R	S	S	S	27.27	15	1.57
R8	RRRRRRSRSS	R	R	R	R	R	R	S	R	S	S	27.27	10	1.05
R9	RRRRRRSSRS	R	R	R	R	R	R	S	S	R	S	27.27	3	0.31
R10	RRRRRSRRSS	R	R	R	R	R	S	R	R	S	S	27.27	6	0.63
R11	RRRRRSRSRS	R	R	R	R	R	S	R	S	R	S	27.27	1	0.10
R12	RRRRRSSRRS	R	R	R	R	R	S	S	R	R	S	27.27	2	0.21
R13	RRRRSRRRSS	R	R	R	R	S	R	R	R	S	S	27.27	2	0.21
R14	SRRRRRRRSS	S	R	R	R	R	R	R	R	S	S	27.27	2	0.21
R15	RRRRRRSSSS	R	R	R	R	R	R	S	S	S	S	36.36	10	1.05
R16	RRRRRSRSSS	R	R	R	R	R	S	R	S	S	S	36.36	22	2.31
R17	RRRRRSSRSS	R	R	R	R	R	S	S	R	S	S	36.36	3	0.31
R18	RRRRRSSSRS	R	R	R	R	R	S	S	S	R	S	36.36	2	0.21
R19	RRRRSRRSSS	R	R	R	R	S	R	R	S	S	S	36.36	5	0.52
R20	RRRRSRSRSS	R	R	R	R	S	R	S	R	S	S	36.36	3	0.31
R21	RRRRSRSSRS	R	R	R	R	S	R	S	S	R	S	36.36	2	0.21
R22	RRRRRSSSSS	R	R	R	R	R	S	S	S	S	S	45.45	90	9.43
R23	RRRRSRSSSS	R	R	R	R	S	R	S	S	S	S	45.45	31	3.25
R24	RRRRSSRSSS	R	R	R	R	S	S	R	S	S	S	45.45	5	0.52
R25	RRRRSSSRSS	R	R	R	R	S	S	S	R	S	S	45.45	2	0.21
R26	RRRRSSSSRS	R	R	R	R	S	S	S	S	R	S	45.45	1	0.10
R27	RRRSRRSSSS	R	R	R	S	R	R	S	S	S	S	45.45	3	0.31
R28	RRRSRSRSSS	R	R	R	S	R	S	R	S	S	S	45.45	2	0.21
R29	RSSRRSRRSS	R	S	S	R	R	S	R	R	S	S	45.45	1	0.10
R30	SRRRRSRSSS	S	R	R	R	R	S	R	S	S	S	45.45	1	0.10
R31	SRSRRRRSSS	S	R	S	R	R	R	R	S	S	S	45.45	1	0.10
R32	RRRRSSSSSS	R	R	R	R	S	S	S	S	S	S	54.55	370	38.78
R33	RRRSRSSSSS	R	R	R	S	R	S	S	S	S	S	54.55	18	1.89
R34	RRRSSRSSSS	R	R	R	S	S	R	S	S	S	S	54.55	8	0.84
R35	RRRSSSRSSS	R	R	R	S	S	S	R	S	S	S	54.55	3	0.31
序号 Serial number	致病型 <BOLD>P</BOLD>athotype	寄主反应Host response										致病谱Pathogenicity spectrum (%)	各致病型的菌株数 Number of strains of each pathotype	各致病型发生频率 Percentage of each pathotype (%)
序号 Serial number	致病型 <BOLD>P</BOLD>athotype	GDBB23	IRBB7	IRBB5	IRBB21	IRBB13	IRBB3	IRBB4	IRBB10	IR24	CBBD1	致病谱Pathogenicity spectrum (%)	各致病型的菌株数 Number of strains of each pathotype	各致病型发生频率 Percentage of each pathotype (%)
R36	RRSRRSSSSS	R	R	S	R	R	S	S	S	S	S	54.55	2	0.21
R37	RSRRRSSSSS	R	S	R	R	R	S	S	S	S	S	54.55	4	0.42
R38	RSSRRRSSSS	R	S	S	R	R	R	S	S	S	S	54.55	1	0.10
R39	RSSSRSRRSS	R	S	S	S	R	S	R	R	S	S	54.55	1	0.10
R40	SRRRRSSSSS	S	R	R	R	R	S	S	S	S	S	54.55	2	0.21
R41	SRRRSRSSSS	S	R	R	R	S	R	S	S	S	S	54.55	1	0.10
R42	SRRRSSRSSS	S	R	R	R	S	S	R	S	S	S	54.55	2	0.21
R43	SSSRRRRSSS	S	S	S	R	R	R	R	S	S	S	54.55	1	0.10
R44	RRRSSSSSSS	R	R	R	S	S	S	S	S	S	S	63.64	148	15.51
R45	RRSRSSSSSS	R	R	S	R	S	S	S	S	S	S	63.64	10	1.05
R46	RSRRSSSSSS	R	S	R	R	S	S	S	S	S	S	63.64	2	0.21
R47	RSRSRSSSSS	R	S	R	S	R	S	S	S	S	S	63.64	1	0.10
R48	RSSRRSSSSS	R	S	S	R	R	S	S	S	S	S	63.64	19	1.99
R49	SRRRSSSSSS	S	R	R	R	S	S	S	S	S	S	63.64	12	1.26
R50	SRRSSSRSSS	S	R	R	S	S	S	R	S	S	S	63.64	2	0.21
R51	SSSRRRSSSS	S	S	S	R	R	R	S	S	S	S	63.64	1	0.10
R52	RRSSSSSSSS	R	R	S	S	S	S	S	S	S	S	72.73	4	0.42
R53	RSRSSSSSSS	R	S	R	S	S	S	S	S	S	S	72.73	3	0.31
R54	SRRSSSSSSS	S	R	R	S	S	S	S	S	S	S	72.73	3	0.31
R55	SRSRSSSSSS	S	R	S	R	S	S	S	S	S	S	72.73	1	0.10

表11

表12

图4

表13

参考文献 48

[1]	方中达. 水稻白叶枯病. 南京: 江苏人民出版社, 1963: 1-2.
	FANG Z D. Rice Bacterial Blight. Nanjing: Jiangsu People’s Publishing Press, 1963: 1-2. (in Chinese)
[2]	章琦. 水稻白叶枯病抗性的遗传及改良. 北京: 科学出版社, 2007: 2-4.
	ZHANG Q. Genetics and Improvement of Resistance to Bacterial Blight in Rice. Beijing: Science Press, 2007: 2-4. (in Chinese)
[3]	LEACH J E, WHITE F F. Bacterial avirulence genes. Annual Review of Phytopathology, 1996, 34: 153-179.
[4]	陈功友, 徐正银, 杨阳阳, 邹丽芳, 朱勃. 我国水稻白叶枯病菌致病型划分和水稻抗病育种中应注意的问题. 上海交通大学学报(农业科学版), 2019, 37(1): 67-73.
	CHEN G Y, XU Z Y, YANG Y Y, ZOU L F, ZHU B. Classification of pathotypes of Chinese Xanthomonas oryzae pv. oryzae and resistance breeding strategies for bacterial blight. Journal of Shanghai Jiaotong University (Agricultural Science), 2019, 37(1): 67-73. (in Chinese)
[5]	堀野修, 季伯衡. 水稻白叶枯病菌的小种分化与水平抗性鉴定法. 农业译文, 1991(2): 1-7.
	HORINO O, JI B H. Race differentiation of Xanthomonas oryzae pv. oryzae and horizontal resistance identification. Agricultural Translation, 1991(2): 1-7. (in Chinese)
[6]	NODA T, HORINO O, OHUCHI A. A survey of geographical distribution of pathogenic races of Xanthomonas campestris pv. oryzae from Japan. Proceedings of the Association for Plant Protection of Hokuriku, 1987, 35: 7-13. (in Japanese with English summary)
[7]	NODA T, OHUCHI A. A new pathogenic race of Xanthomonas campestris pv. oryzae and inheritance of resistance of differential rice variety, Te-tep to it. Japanese Journal of Phytopathology, 1989, 55(2): 201-207.
[8]	MEW T W. Current status and future prospects of research on bacterial blight of rice. Annual Review of Phytopathology, 1987, 25: 359-382.
[9]	伍尚忠, 徐羡明, 刘景梅, 苗东华, 维拉·克鲁茨. 华南及菲律宾稻白叶枯病病原菌株致病性比较研究. 植物病理学报, 1985, 15(2): 65-72.
	WU S Z, XU X M, LIU J M, MEW T W, VERA CRUZ C M. Comparison of virulence of Xanthomonas campestris pv. oryzae in south China and in the Philippines. Acta Phytopathologica Sinica, 1985, 15(2): 65-72. (in Chinese)
[10]	方中达, 许志刚, 过崇俭, 殷尚智, 伍尚忠, 徐羡明, 章琦. 中国水稻白叶枯病菌致病型的研究. 植物病理学报, 1990, 20(2): 81-88.
	FANG Z D, XU Z G, GUO C J, YIN S Z, WU S Z, XU X M, ZHANG Q. Studies on pathotypes of Xanthomonas campestris pv. oryzae in China. Acta Phytopathologica Sinica, 1990, 20(2): 81-88. (in Chinese)
[11]	王春连, 章琦, 周永力, 赵炳宇. 我国长江以南地区水稻白叶枯病原菌遗传多样性分析. 中国水稻科学, 2001, 15(2): 131-136.
	WANG C L, ZHANG Q, ZHOU Y L, ZHAO B Y. Genetic diversity of pathogen Xanthomonas oryzae pv. oryzae from southern regions of Yangtze River in China. Chinese Journal of Rice Science, 2001, 15(2): 131-136. (in Chinese)
[12]	曾列先, 朱小源, 杨健源, 伍圣远, 陈珍, 陈深. 广东水稻白叶枯病菌新致病型的发现及致病性测定. 广东农业科学, 2005(2): 58-59.
	ZENG L X, ZHU X Y, YANG J Y, WU S Y, CHEN Z, CHEN S. A new pathotype of Xanthomonas oryzae pv. oryzae was found and tested for pathogenicity in Guangdong. Guangdong Agricultural Sciences, 2005(2): 58-59. (in Chinese)
[13]	陈深, 汪聪颖, 苏菁, 冯爱卿, 朱小源, 曾列先. 华南水稻白叶枯病菌致病性分化检测与分析. 植物保护学报, 2017, 44(2): 217-222.
	CHEN S, WANG C Y, SU J, FENG A Q, ZHU X Y, ZENG L X. Differential detection and analysis of pathotypes and differentiation against Xanthomonas oryzae pv. oryzae in southern China. Journal of Plant Protection, 2017, 44(2): 217-222. (in Chinese)
[14]	OGAWA T, KHUSH G S. Major genes for resistance to bacterial blight in rice//Bacterial Blight of Rice. Internationa Rice Research Institute, 1989.
[15]	OGAWA T, YAMAMOTO T, KHUSH G S, MEW T W. Breeding of near-isogenic lines of rice with single genes for resistance to bacterial blight pathogen (Xanthomonas campestris pv. oryzae). Japanese Journal of Breeding, 1991, 41(3): 523-529.
[16]	章琦, 杨文才, 施爱农, 王春莲, 阙更生, 赵炳宇, 邢全党. 3个粳稻抗白叶枯病近等基因系的构建. 作物学报, 1998, 24(6): 799-804.
	ZHANG Q, YANG W C, SHI A N, WANG C L, QUE G S, ZHAO B Y, XING Q D. Breeding of three near-isogenic japonica rice lines with major genes for resistance to bacterial-blight. Acta Agronomica Sinica, 1998, 24(6): 799-804. (in Chinese)
[17]	章琦, 王春连, 赵开军, 杨文才, 乔枫, 周永力, 江祺祥, 刘古春. 携有抗白叶枯病新基因Xa23水稻近等基因系的构建及应用. 中国水稻科学, 2002, 16(3): 206-210.
	ZHANG Q, WANG C L, ZHAO K J, YANG W C, QIAO F, ZHOU Y L, JIANG Q X, LIU G C. Development of near-isogenic line CBB23 with a new resistance gene to bacterial blight in rice and its application. Chinese Journal of Rice Science, 2002, 16(3): 206-210. (in Chinese)
[18]	JIANG N, YAN J, LIANG Y, SHI Y, HE Z, WU Y, ZENG Q, LIU X, PENG J. Resistance genes and their interactions with bacterial blight/leaf streak pathogens (Xanthomonas oryzae) in rice (Oryza sativa L.)—An updated review. Rice, 2020, 13(1): 3.
[19]	NEELAM K, MAHAJAN R, GUPTA V, BHATIA D, GILL B K, KOMAL R, LORE J S, MANGAT G S, SINGH K. High-resolution genetic mapping of a novel bacterial glight resistance gene xa-45(t) identified from Oryza glaberrima and transferred to Oryza sativa. Theoretical and Applied Genetics, 2020, 133(3): 689-705.
[20]	许志刚, 孙启明, 刘凤权, 陈志谊, 胡白石, 郭亚辉, 刘永峰, 刘红霞. 水稻白叶枯病菌小种分化的监测. 中国水稻科学, 2004, 18(5): 469-472.
	XU Z G, SUN Q M, LIU F Q, CHEN Z Y, HU B S, GUO Y H, LIU Y F, LIU H X. Race monitoring of rice bacterial blight (Xanthomonas oryzae pv. oryzae) in China. Chinese Journal of Rice Science, 2004, 18(5): 469-472. (in Chinese)
[21]	杨万风, 刘红霞, 胡白石, 许志刚, 刘凤权. 中国水稻白叶枯病菌毒性变异研究. 植物病理学报, 2006, 36(3): 244-248.
	YANG W F, LIU H X, HU B S, XU Z G, LIU F Q. Virulence variation of Xanthomonas oryzae pv. oryzae on rice near-isogenic lines in China. Acta Phytopathologica Sinica, 2006, 36(3): 244-248. (in Chinese)
[22]	夏立琼, 李明容, 谢仕猛, 翟文学, 夏志辉. 海南水稻白叶枯病菌优势生理小种的分离及致病力分析. 分子植物育种, 2016, 14(5): 1336-1340.
	XIA L Q, LI M R, XIE S M, ZHAI W X, XIA Z H. Isolation and pathogenicity analysis of the predominant race of Xanthomonas oryzae pv. oryzae in Hainan. Molecular Plant Breeding, 2016, 14(5): 1336-1340. (in Chinese)
[23]	KOGEETHAVANI R, FATIN N A, SUZIANTI I V, ERWAN S S. Characterization of pathogenic variability of Xanthomonas oryzae pv. oryzae isolates causing bacterial leaf blight disease in Malaysian rice granaries. Australasian Plant Pathology, 2021, 50: 293-298.
[24]	余建英, 何旭宏. 数据统计分析与SPSS应用. 北京: 人民邮电出版社, 2003.
	YU J Y, HE X H. Data Statistical Analysis and SPSS Application. Beijing: People’s Posts and Telecommunications Press, 2003. (in Chinese)
[25]	周俊飞, 章山, 孙婧, 张伟, 高利芬. 水稻抗白叶枯病近等基因系的抗性鉴定揭示基因聚合的多向性效应. 华北农学报, 2020, 35(6): 67-73.
	ZHOU J F, ZHANG S, SUN J, ZHANG W, GAO L F. Resistance identification of near-isogenic lines to bacterial blight in rice reveals the multidirectional effect of gene pyramiding. Acta Agriculture Boreali-Sinica, 2020, 35(6): 67-73. (in Chinese)
[26]	成太辉, 陈深, 杨健源, 朱小源, 伍圣远, 洪启金, 曾列先. 水稻抗白叶枯病V型菌xa5基因利用现状及前景. 广东农业科学, 2020, 47(1): 92-97.
	CHENG T H, CHEN S, YANG J Y, ZHU X Y, WU S Y, HONG Q J, ZENG L X. Dtilization situation and prospect of gene xa5against pathotype V of rice bacterial blight. Guangdong Agricultural Sciences, 2020, 47(1): 92-97. (in Chinese)
[27]	倪大虎, 易成新, 杨剑波, 汪秀峰, 张毅, 章琦, 王春连, 赵开军, 王文相, 李莉. 利用分子标记辅助选择聚合Pi9(t)和Xa23基因. 分子植物育种, 2007, 5(4): 491-496.
	NI D H, YI C X, YANG J B, WANG X F, ZHANG Y, ZHANG Q, WANG C L, ZHAO K J, WANG W X, LI L. Pyramiding Pi9(t) and Xa23 genes by molecular marker-assisted selection. Molecular Plant Breeding, 2007, 5(4): 491-496. (in Chinese)
[28]	ABDUL FIYAZ R, SHIVANI D, CHAITHANYA K, MOUNIKA K, CHIRANJEEVI M, LAHA G S, VIRAKTAMATH B C, SUBBA RAO L V, SUNDARAM R M. Genetic improvement of rice for bacterial blight resistance: Present status and future prospects. Rice Science, 2022, 29(2): 118-132.
[29]	陈小林, 颜群, 高利军, 韦善富, 李道远, 高汉亮. 广西水稻白叶枯病菌致病型的初步鉴定. 南方农业学报, 2015, 46(2): 236-240.
	CHEN X L, YAN Q, GAO L J, WEI S F, LI D Y, GAO H L. Preliminary identification of pathotype of Xanthomonas oryzae pv. oryzae in Guangxi. Journal of Southern Agriculture, 2015, 46(2): 236-240. (in Chinese)
[30]	袁斌, 刘友梅, 黄薇, 张舒, 吕亮, 常向前, 杨小林. 湖北省水稻白叶枯病菌致病型分化检测与分析. 湖北农业科学, 2018, 57(24): 100-103.
	YUAN B, LIU Y M, HUANG W, ZHANG S, LÜ L, CHANG X Q, YANG X L. Differential detection and analysis of pathotypes of Xanthomonas oryzae pv. oryzae in Hubei Province. Hubei Agricultural Sciences, 2018, 57(24): 100-103. (in Chinese)
[31]	阎园园, 陈华, 姜波. 基于群智能算法分类模型的番茄病害识别. 江苏农业科学, 2020, 48(1): 219-224.
	YAN Y Y, CHEN H, JIANG B. Tomato disease recognition based on swarm intelligence classification model. Jiangsu Agricultural Science, 2020, 48(1): 219-224. (in Chinese)
[32]	徐笑锋, 肖英杰, 章学来, 徐亚伟. 基于PCA-相对熵模型的海上中转引航平台选址研究. 安全与环境学报, 2021, 21(6): 2438-2443.
	XU X F, XIAO Y J, ZHANG X L, XU Y W. Research on site selection of maritime transit pilot platform based on PCA-relative entropy model. Journal of Safety and Environment, 2021, 21(6): 2438-2443. (in Chinese)
[33]	吕开云, 鞠厦轶, 龚循强, 鲁铁定. 基于PCA和IGG权函数的人脸图像鲁棒线性回归分类方法. 电子测量技术, 2021, 44(21): 152-157.
	LÜ K Y, JU X Y, GONG X Q, LU T D. Face recognition using robust linear regression classification based on PCA and IGG weight function. Electronic Measurement Technology, 2021, 44(21): 152-157. (in Chinese)
[34]	郭金玉, 刘玉超, 李元. 加权局部近邻标准化PCA的工业过程故障检测. 沈阳化工大学学报, 2021, 35(3): 265-274.
	GUO J Y, LIU Y C, LI Y. Fault detection of industrial process based on weighed local neighborhood standardization PCA. Journal of Shenyang University of Chemical Technology, 2021, 35(3): 265-274. (in Chinese)
[35]	REICH D, PRICE A L, PATTERSON N. Principal component analysis of genetic data. Nature Genetics, 2008, 40(5): 491-492.
[36]	HUANG X, WEI X, SANG T, ZHAO Q, FENG Q, ZHAO Y, LI C, ZHU C, LU T, ZHANG Z, et al. Genome-wide association studies of 14 agronomic traits in rice landraces. Nature Genetics, 2010, 42(11): 961-967.
[37]	ZHAO K, TUNG C W, EIZENGA G C, WRIGHT M H, ALI M L, PRICE A H, NORTON G J, ISLAM M R, REYNOLDS A, MEZEY J, MCCLUNG A M, BUSTAMANTE C D, MCCOUCH S R. Genome-wide association mapping reveals a rich genetic architecture of complex traits in Oryza sativa. Nature Communications, 2011, 2: 467.
[38]	JONNALAGADDA S, SRINIVASAN R. Principal components analysis based methodology to identify differentially expressed genes in time-course microarray data. BMC Bioinformatics, 2008, 9: 267.
[39]	周丽娜, 于海业, 张蕾, 任顺, 隋媛媛, 于连军. 基于叶绿素荧光光谱分析的稻瘟病害预测模型. 光谱学与光谱分析, 2014, 34(4): 1003-1006.
	ZHOU L N, YU H Y, ZHANG L, REN S, SUI Y Y, YU L J. Rice blast prediction model based on analysis of chlorophyll fluorescence spectrum. Spectroscopy and Spectral Analysis, 2014, 34(4): 1003-1006. (in Chinese)
[40]	WANG C Y, CHEN S, FENG A Q, SU J, WANG W J, FENG J Q, CHEN B, ZHANG M Y, YANG J Y, ZENG L X, ZHU X Y. Xa7, a small orphan gene harboring promoter trap for AvrXa7, leads to the durable resistance to Xanthomonas oryzae pv. oryzae. Rice, 2021, 14(1): 48.
[41]	CHEN X F, LIU P C, MEI L, HE X L, CHEN L, LIU H, SHEN S R, JI Z D, ZHENG X X, ZHANG Y C, GAO Z Y, ZENG D L, QIAN Q, MA B J. Xa7, a new executor R gene that confers durable and broad-spectrum resistance to bacterial blight disease in rice. Plant Communications, 2021, 2(3): 100143.
[42]	LUO D, HUGUET-TAPIA J C, RABORN R T, WHITE F F, YANG B. The Xa7 resistance gene guards the susceptibility gene SWEET14 of rice against exploitation by bacterial blight pathogen. Plant Communications, 2021, 2(3): 100164.
[43]	JI C H, JI Z Y, LIU B, CHENG H, LIU H, LIU S Z, YANG B, CHEN G Y. Xa1 allelic R genes activate rice blight resistance suppressed by interfering TAL effectors. Plant Communications, 2020, 1(4): 100087.
[44]	ZHANG B M, ZHANG H T, LI F, OUYANG Y D, YUAN M, LI X H, XIAO J H, WANG S P. Multiple alleles encoding atypical NLRs with unique central tandem repeats in rice confer resistance to Xanthomonas oryzae pv. oryzae. Plant Communications, 2020, 1(4): 100088.
[45]	PRADHAN S K, BARIK S R, SAHOO A, MOHAPATRA S, NAYAK D K, MAHENDER A, MEHER J, ANANDAN A, PANDIT E. Population structure, genetic diversity and molecular marker-trait association analysis for high temperature stress tolerance in rice. PLoS ONE, 2016, 11(8): e0160027.
[46]	BALIYAN N, MALIK R, RANI R, MEHTA K, VASHISTH U, DHILLON S, BOORA K S. Integrating marker-assisted background analysis with foreground selection for pyramiding bacterial blight resistance genes into Basmati rice. Comptes Rendus Biologies, 2018, 341(1): 1-8.
[47]	HSU Y C, CHIU C H, YAP R, TSENG Y C, WU Y P. Pyramiding bacterial blight resistance genes in Tainung82 for broad-spectrum resistance using marker-assisted selection. International Journal of Molecular Sciences, 2020, 21(4): 1281.
[48]	WANG Y, PRUITT R N, NÜRNBERGER T, WANG Y C. Evasion of plant immunity by microbial pathogens. Nature Reviews Microbiology, 2022, 20(8): 449-464.

年份Year	生态区Ecological region	采集地Origin	菌株数Number of strains
2018	华南稻区South China rice region	广东Guangdong	89
		广西Guangxi	16
		海南Hainan	13
	长江中下游稻区 Middle and lower reaches of the Yangtze River rice region	湖南Hunan	0
		浙江Zhejiang	47
	西南稻区Southwest China rice region	云南Yunnan	10
	东北稻区Northeast China rice region	辽宁Liaoning	18
2019	华南稻区South China rice region	广东Guangdong	185
		广西Guangxi	45
		海南Hainan	0
	长江中下游稻区 Middle and lower reaches of the Yangtze River rice region	湖南Hunan	10
		浙江Zhejiang	43
	西南稻区Southwest China rice region	云南Yunnan	0
	东北稻区Northeast China rice region	辽宁Liaoning	5
2020	华南稻区South China rice region	广东Guangdong	140
		广西Guangxi	7
		海南Hainan	1
	长江中下游稻区 Middle and lower reaches of the Yangtze River rice region	湖南Hunan	4
		浙江Zhejiang	23
	西南稻区Southwest China rice region	云南Yunnan	1
	东北稻区Northeast China rice region	辽宁Liaoning	3
2021	华南稻区South China rice region	广东Guangdong	36
		广西Guangxi	65
		海南Hainan	0
	长江中下游稻区 Middle and lower reaches of the Yangtze River rice region	湖南Hunan	0
		浙江Zhejiang	193
	西南稻区Southwest China rice region	云南Yunnan	0
	东北稻区Northeast China rice region	辽宁Liaoning	0

年份Year	菌株数Number of strains	各致病型发生频率Percentage of each pathotype (%)
年份Year	菌株数Number of strains	Ⅰ	Ⅱ	Ⅲ	Ⅳ	Ⅴ	Ⅵ	Ⅸ	新型1 New pathotype 1	新型2 New pathotype 2	新型3 New pathotype 3	新型4 New pathotype 4
2018	193	15.03	3.11	7.25	7.77	7.77	0.52	57.51	0.52	0	0.52	0
2019	288	11.46	9.03	10.42	6.60	11.81	0	50.69	0	0	0	0
2020	179	17.32	6.70	7.26	6.70	6.15	0	53.07	2.23	0	0	0.56
2021	294	5.78	0.68	4.42	4.42	3.06	3.06	74.83	3.40	0.34	0	0

年份Year	菌株数Number of strains	各致病型发生频率Percentage of each pathotype (%)
年份Year	菌株数Number of strains	Ⅰ	Ⅱ	Ⅲ	Ⅳ	Ⅴ	Ⅵ	Ⅸ	新型1 New pathotype 1	新型2 New pathotype 2	新型3 New pathotype 3	新型4 New pathotype 4
1999	30	16.70	20.00	20.00	26.70	16.70	0	0	0	0	0	0
2000	43	18.60	20.90	16.30	23.30	20.90	0	0	0	0	0	0
2004	53	17.00	22.60	11.30	24.50	18.90	0	5.70	0	0	0	0
2014	150	14.00	15.30	7.30	19.30	27.40	0	16.67	0	0	0	0
2015	68	44.12	7.35	0.00	2.94	20.59	0	25.00	0	0	0	0
2016	50	16.00	10.00	6.00	22.00	26.00	0	20.00	0	0	0	0
2017	60	16.67	11.67	8.33	18.34	23.34	0	21.67	0	0	0	0
2018	118	9.32	4.24	5.08	5.08	12.71	0.85	61.02	0.85	0	0.85	0
2019	230	10.43	10.43	10.87	6.09	14.78	0	47.39	0	0	0	0
2020	148	15.54	6.76	8.11	7.43	6.76	0	52.70	2.03	0	0	0.68
2021	101	5.94	0.99	3.96	1.98	8.91	0.99	72.28	4.95	0	0	0

分类 Category	单基因系 Monogenic line	基因 Gene	平均抗性频率 Average resistance frequency (%)
第Ⅰ类Class I	IRBB1、IRBB2、IRBB10、IRBB11、IRBB4	Xa1、Xa2、Xa10、Xa11、Xa4	15.32
第Ⅱ类Class Ⅱ	IRBB3、IRBB203、IRBB14	Xa3、Xa14	23.24
第Ⅲ类Class Ⅲ	IRBB8、IRBB13	Xa8、xa13	36.06
第Ⅳ类Class Ⅳ	IRBB21	Xa21	79.45
第Ⅴ类Class Ⅴ	IRBB5、IRBB7、CBB23、GDBB23	xa5、Xa7、Xa23	96.41

来源Origin	菌号Number of bacteria	IRBB5	IRBB7	IRBB21	CBB23
广东广州Guangzhou, Guangdong	2848	R	R	R	S
	2850	R	R	R	S
	2851	R	R	R	S
	2852	R	R	R	S
	2853	R	R	R	S
	2854	R	R	R	S
广东四会Sihui, Guangdong	1387	S	R	R	S
	1392	R	R	R	S
	1394	R	R	R	S
	1389	R	R	R	S
	1390	R	R	R	S
	1391	R	R	R	S
广东新兴Xinxing, Guangdong	2614	R	R	R	S
广东珠海Zhuhai, Guangdong	2749	R	R	R	S
广西靖西Jingxi, Guangxi	2821	S	S	R	S
广西靖西Jingxi, Guangxi	2824	S	R	R	S
湖南浏阳Liuyang, Hunan	1823	R	R	S	S
浙江常山Changshan, Zhejiang	595	R	R	R	S
	890	R	R	R	S
	891	R	R	R	S
浙江忂州Quzhou, Zhejiang	579	R	R	S	S
	855	R	R	S	S
	856	R	R	S	S
	859	R	R	R	S
	1252	R	R	S	S
	1254	R	R	R	S
	1792	R	R	S	S
浙江台州Taizhou, Zhejiang	2420	R	R	R	S
浙江台州Taizhou, Zhejiang	2491	R	R	R	S
浙江温岭Wenling, Zhejiang	2573	R	R	R	S
浙江仙居 Xianju, Zhejiang	2581	S	S	R	S

中国水稻主产区白叶枯病菌致病型分析及近等基因系鉴别寄主的构建

Pathotype Analysis of Xanthomonas oryzae pv. oryzae in Main Rice Producing Regions of China and Establishment of Differential Hosts of Near-Isogenic Lines

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致病型 Pathotype	鉴别寄主反应Phynotypes of differential host to the strains					各致病型的菌株数Number of strains of each pathotype	各致病型发生频率Percentage of each pathotype (%)
致病型 Pathotype	CBBD1	CBBD2	CBBD3	CBBD4	CBBD5	各致病型的菌株数Number of strains of each pathotype	各致病型发生频率Percentage of each pathotype (%)
Ⅰ	S	R	R	R	R	110	11.53
Ⅱ	S	S	R	R	R	46	4.82
Ⅲ	S	S	S	R	R	70	7.34
Ⅳ	S	S	S	S	R	59	6.18
Ⅴ	S	S	R	R	S	69	7.23
Ⅵ	S	R	S	R	R	10	1.05
Ⅸ	S	S	S	S	S	572	59.96
新型1 New pathotype 1	S	S	S	R	S	15	1.57
新型2 New pathotype 2	S	R	S	R	S	1	0.10
新型3 New pathotype 3	S	R	S	S	S	1	0.10
新型4 New pathotype 4	S	S	R	S	S	1	0.10
合计Total						954	100.00

近等基因系 Near-isogenic line	基因 Gene	抗性频率 Resistance frequency (%)
IRBB1	Xa1	15.62
IRBB2	Xa2	15.20
IRBB3	Xa3	21.91
IRBB203	Xa3	25.26
IRBB4	Xa4	18.13
IRBB5	xa5	95.60
IRBB7	Xa7	96.44
IRBB8	Xa8	37.21
IRBB10	Xa10	14.47
IRBB11	Xa11	13.21
IRBB13	xa13	34.91
IRBB14	Xa14	22.54
IRBB21	Xa21	79.45
CBB23	Xa23	96.75
GDBB23	Xa23	96.86

中国鉴别寄主的致病型 Pathotypes identified by Chinese differential hosts	菌株数 Number of strains	近等基因系鉴定的致病型个数 Number of pathotypes identified by near-isogenic lines
Ⅰ	110	35
Ⅱ	46	37
Ⅲ	70	44
Ⅳ	59	33
Ⅴ	69	21
Ⅵ	10	6
Ⅸ	572	51
新型 New pathotype	18	9

成分因子Component	因子提取Extraction of factors			旋转后因子的提取Extraction of factors after rotation
成分因子Component	特征值 Eigenvalues	变异量 Amount of variation (%)	累计变异量 Cumulative variance (%)	特征值 Eigenvalues	变异量 Amount of variation (%)	累计变异量 Cumulative variance (%)
1	6.572	41.076	41.076	5.078	31.739	31.739
2	2.034	12.713	53.789	1.996	12.477	44.217
3	1.725	10.779	64.569	1.653	10.328	54.545
4	0.948	5.927	70.495	1.489	9.306	63.851
5	0.769	4.806	75.301	1.017	6.358	70.209
6	0.648	4.049	79.350	0.961	6.003	76.212
7	0.572	3.578	82.929	0.936	5.849	82.061
8	0.462	2.886	85.815	0.601	3.753	85.815
9	0.425	2.655	88.470
10	0.396	2.477	90.947
11	0.365	2.281	93.228
12	0.346	2.163	95.391
13	0.289	1.807	97.198
14	0.255	1.596	98.794
15	0.176	1.101	99.896
16	1.67E-02	0.104	100

近等基因系<BOLD>N</BOLD>ear-isogenic line	成分因子Component
近等基因系<BOLD>N</BOLD>ear-isogenic line	1	2	3	4	5	6	7	8
IRBB1 (Xa1)	0.851	5.67E-02	6.68E-02	-9.91E-02	6.70E-02	-6.84E-02	9.87E-02	-8.39E-02
IRBB2 (Xa2)	0.859	3.50E-02	8.06E-02	-8.52E-02	5.44E-02	-0.134	2.94E-02	-8.06E-02
IRBB3 (Xa3)	0.779	-7.25E-03	6.24E-02	-6.42E-02	6.87E-02	-0.124	1.42E-02	0.337
IRBB203 (Xa3)	0.703	5.54E-02	6.03E-02	8.40E-02	-0.191	-2.99E-02	-0.59	0.238
IRBB4 (Xa4)	0.798	-9.38E-02	4.57E-02	-3.55E-02	-0.19	-0.162	0.124	9.91E-02
IRBB5 (xa5)	5.83E-02	0.344	0.789	0.134	-0.115	0.174	0.155	7.14E-02
IRBB7 (Xa7)	4.85E-02	0.292	0.828	0.158	-7.21E-02	6.90E-02	-6.10E-03	-5.27E-02
IRBB8 (Xa8)	0.673	-3.92E-02	-0.247	0.185	-0.334	0.366	1.63E-02	-0.229
IRBB10 (Xa10)	0.864	1.45E-02	2.67E-02	-0.111	6.76E-02	-8.65E-02	0.177	-4.02E-02
IRBB11 (Xa11)	0.837	-5.39E-02	3.58E-02	-0.163	0.174	-0.104	0.161	2.07E-02
IRBB13 (xa13)	0.661	-8.83E-02	-0.353	0.115	-0.265	0.351	0.201	0.202
IRBB14 (Xa14)	0.763	1.03E-02	3.52E-02	5.75E-02	-0.134	-0.198	-0.194	-0.402
IRBB21 (Xa21)	0.303	-6.27E-02	-0.145	0.866	0.339	-0.116	5.00E-02	2.31E-02
CBB23 (Xa23)	1.95E-02	0.947	-0.302	-5.64E-03	1.16E-02	-3.72E-02	1.27E-02	8.76E-03
GDBB23 (Xa23)	1.54E-02	0.948	-0.299	-1.61E-02	8.89E-03	-3.56E-02	1.24E-02	7.44E-03
IR24 (Xa18)	0.596	4.11E-02	5.85E-02	-0.188	0.562	0.465	-0.188	-5.40E-02

分类 Category	鉴别寄主数 <BOLD>N</BOLD>umber of differential hosts	鉴别寄主名称 Name of differential hosts	鉴定出的致病型个数 Number of pathotypes identified
本研究新鉴别寄主 New differential hosts in present study	10	CBBD1、IR24、IRBB10、IRBB4、IRBB3、IRBB13、IRBB21、IRBB5、IRBB7、GDBB23	55
中国鉴别寄主Chinese differential hosts	5	CBBD1、CBBD2、CBBD3、CBBD4、CBBD5	11
杨万风等建立的鉴别寄主 Differential hosts developed by YANG WanFeng et al.^[21]	6	IRBB5、IRBB13、IRBB3、IRBB14、IRBB2、IR24	33
陈功友等建立的鉴别寄主 Differential hosts developed by CHEN GongYou et al.^[4]	8	IR24、IRBB3、IRBB4、IRBB5、IRBB7、IRBB13、IRBB21、CBB23	45
许志刚等建立的鉴别寄主 Differential hosts developed by XU ZhiGang et al.^[20]	6	CBBD1、IRBB14、IRBB3、IRBB4、CBBD4、IRBB5	23
夏立琼等建立的鉴别寄主 Differential hosts developed by XIA LiQiong et al.^[22]	7	IRBB3、IRBB4、IRBB5、IRBB13、IRBB21、CBB23、IR24	39

基因 Gene	抗性频率 <BOLD>R</BOLD>esistance frequency (%)	聚合的基因 Polymerized gene	基因聚合后抗性频率 Resistance frequency after gene pyramiding (%)	提升的抗性频率 Elevated resistance frequency (%)	聚合的基因 Polymerized gene	基因聚合后抗性频率 Resistance frequency after gene pyramiding (%)	提升的抗性频率 Elevated resistance frequency (%)
Xa11	13.21	Xa11+Xa21	79.56	0.11	xa5+Xa7	97.48	1.04
Xa10	14.47	Xa10+Xa21	79.56	0.11	xa5+Xa23	99.58	2.83
Xa2	15.20	Xa2+Xa21	79.56	0.11	Xa7+Xa23	99.79	3.04
Xa1	15.62	Xa1+Xa21	79.77	0.32	xa5+Xa21	99.48	3.88
Xa4	18.13	Xa4+Xa21	80.29	0.84	Xa4+Xa10+Xa21	80.29	0.84
Xa3	21.91	Xa14+Xa21	80.29	0.84	Xa10+xa13+Xa21	82.08	2.63
Xa14	22.54	Xa3+Xa21	81.24	1.79	Xa10+Xa8+Xa21	82.49	3.04
xa13	34.91	Xa8+Xa21	82.39	2.94	Xa4+xa13+Xa21	82.60	3.15
Xa8	37.21	xa13+Xa21	82.08	2.63	Xa14+xa13+Xa21	82.70	3.25
Xa21	79.45	Xa10+xa13	35.74	0.83	Xa3+xa13+Xa21	82.91	3.46
xa5	95.60	Xa11+xa13	35.74	0.83	Xa4+Xa8+Xa21	82.91	3.46
Xa7	96.44	Xa1+xa13	36.79	1.88	Xa3+Xa8+Xa21	83.23	3.78
Xa23	96.75	Xa2+xa13	36.90	1.99	Xa8+xa13+Xa21	83.33	3.88
		Xa4+xa13	37.00	2.09	xa5+xa13+Xa21	99.58	3.98
		Xa14+xa13	40.99	6.08	xa5+Xa4+Xa21	99.58	3.98
		Xa3+xa13	42.77	7.86	Xa4+Xa10+xa13	37.53	2.62
		Xa8+xa13	45.39	8.18	Xa4+Xa8+xa13	46.44	9.23
		Xa10+Xa8	37.74	0.53	Xa14+Xa8+xa13	48.43	11.22
		Xa2+Xa8	38.16	0.95	Xa3+Xa8+xa13	48.74	11.53
		Xa11+Xa8	38.16	0.95	Xa4+xa5+Xa7	97.80	1.36
		Xa1+Xa8	38.16	0.95	Xa4+xa5+xa13+Xa21	99.58	3.98
		Xa4+Xa8	39.62	2.41	Xa10+Xa4+Xa8+xa13	46.44	9.23
		Xa14+Xa8	41.51	4.30	Xa3+Xa4+Xa8+xa13	49.58	12.37
		Xa3+Xa8	43.50	6.29	Xa3+Xa4+Xa8+xa13+Xa21	84.49	5.04
		xa5+Xa4	96.02	0.42	Xa4+xa5+Xa7+xa13+Xa21	100.00	3.56

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