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

Abstract

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

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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Figures/Tables 17

Table 1

Fig. 1

Table 2

Table 3

Fig. 2

Table 4

Table 5

Fig. 3

Table 6

Table 7

Table 8

Table 9

Table 10

Table 11

Reaction of new differential hosts to Xoo strains"

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

Table 11

Table 12

Fig. 4

Table 13

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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

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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