江苏近年育成粳稻新品种/系的稻瘟病抗性基因及穗颈瘟抗性分析

doi:10.16819/j.1001-7216.2020.0208

中国水稻科学 ›› 2020, Vol. 34 ›› Issue (5): 413-424.DOI: 10.16819/j.1001-7216.2020.0208

江苏近年育成粳稻新品种/系的稻瘟病抗性基因及穗颈瘟抗性分析

王小秋¹, 杜海波¹, 陈夕军², 李明友¹, 王嘉楠¹, 许志文², 冯志明^1,², 陈宗祥^1,², 左示敏^1,^2,^*()

¹植物功能基因组学教育部重点实验室/江苏省作物基因组学和分子育种重点实验室/扬州大学农学院,江苏扬州 225009
²扬州大学江苏省粮食作物现代产业技术协同创新中心/江苏省作物遗传生理重点实验室,江苏扬州 225009

收稿日期:2020-02-20 修回日期:2020-03-30 出版日期:2020-09-10 发布日期:2020-09-10
通讯作者: 左示敏
基金资助:
江苏省重点研发计划资助项目(BE2019339);江苏省农业重大新品种创制项目(PZCZ201703);扬州大学研究生国际学术交流专项基金资助项目(201701023)

Analysis of Blast Resistance Genes and Neck Blast Resistance of japonica Rice Varieties/Lines Recently Developed in Jiangsu Province

Xiaoqiu WANG¹, Haibo DU¹, Xijun CHEN², Mingyou LI¹, Jianan WANG¹, Zhiwen XU², Zhiming FENG^1,², Zongxiang CHEN^1,², Shimin ZUO^1,^2,^*()

¹Key Laboratory of Plant Functional Genomics of the Ministry of Education/ Jiangsu Key Laboratory of Crop Genomics and Molecular Breeding, College of Agriculture, Yangzhou University, Yangzhou 225009, China
²Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops/ Jiangsu Key Laboratory of Crop Genetics and Physiology, Yangzhou University, Yangzhou 225009, China

Received:2020-02-20 Revised:2020-03-30 Online:2020-09-10 Published:2020-09-10
Contact: Shimin ZUO

摘要/Abstract

摘要：

【目的】明确水稻品种携带的抗稻瘟病基育种应用价值,是利用抗病基因培育抗病品种控制病害流行的重要前期工作。【方法】利用功能标记分析了14个抗稻瘟病基因在江苏近年育成的195个粳稻新品种/系中的分布情况,并对其中158个品种和17份携带抗稻瘟病基因Pigm的高世代回交株系进行穗颈瘟接种鉴定。【结果】大多数品种携带2~5个抗病基因,但所有品种均不含有Pigm基因;Pib、Pita和Pikh基因在供试品种中的分布频率较高,均在45%以上,其余基因均在30%以内;Pid3、Pid2、Pia、Pb1在新育成品种中的出现频率明显高于审定品种。测试品种对穗颈瘟的抗性主要与3个基因显著相关,贡献率由高至低依次为Pia、Pi3/5/i和Pita;其中,Pia或Pi3/5/i与Pita间的聚合效应均显著高于各基因单独存在时的抗病效应,且以Pia和Pita间的聚合效应最强,携带该基因组合的所有品种对穗颈瘟均表现抗至高抗水平抗性。回交导入Pigm基因的所有17份株系对穗颈瘟的抗性均显著高于各自轮回亲本,且均达到了抗病以上水平。【结论】抗病基因Pigm及基因组合“Pia+Pita”在江苏粳稻抗穗颈瘟育种中具有重要的育种应用价值。

关键词: 粳稻, 穗颈瘟, 抗病基因, 基因分布, 育种价值

Abstract:

【Objective】Evaluating the breeding potential of the known blast resistance (R) genes harbored in rice varieties is one of prerequisites for developing rice blast resistant varieties and R genes-based control of the disease epidemic. 【Method】A total of 195 new japonica varieties/lines from Jiangsu Province were genotyped using functional markers corresponding to 14 rice blast R gene loci. In order to identify gene or gene combination with breeding potential against neck-blast, 158 japonica rice varieties were then artificially inoculated with blast strains as well as 17 advanced backcrossing lines containing Pigm gene in two different genetic backgrounds at booting stage. 【Result】Most varieties carried 2 to 5 R genes, and none of new varieties contained Pigm; the distribution frequencies of Pib, Pita and Pikh were relatively high and all exceeded 45%, the rest 10 genes were under 30%; Pid3, Pid2, Pia and Pb1 had apparently higher frequency in lines from regional tests than that in authorized and released varieties. Three genes with effects ordered as Pia > Pi3/5/i > Pita significantly contributed to neck-blast resistance, and significantly positive interaction effects were found between Pia and Pita, and between Pi3/5/i and Pita. Combination effect of Pia+Pita was significantly higher than that of Pi3/5/i+Pita, and the varieties harboring Pia+Pita reached resistant or highly resistant levels. The resistance of 17 advanced backcrossing lines carrying Pigm was all significantly improved as compared with corresponding controls, and all reached resistant level or even higher. 【Conclusion】 The results suggested that Pigm and the combination of Pia+Pita have critically important breeding potential in japonica rice breeding against neck blast in Jiangsu Province.

Key words: japonica rice, neck blast, resistance gene, gene distribution, breeding value

中图分类号:

S435.111.4⁺1

王小秋, 杜海波, 陈夕军, 李明友, 王嘉楠, 许志文, 冯志明, 陈宗祥, 左示敏. 江苏近年育成粳稻新品种/系的稻瘟病抗性基因及穗颈瘟抗性分析[J]. 中国水稻科学, 2020, 34(5): 413-424.

Xiaoqiu WANG, Haibo DU, Xijun CHEN, Mingyou LI, Jianan WANG, Zhiwen XU, Zhiming FENG, Zongxiang CHEN, Shimin ZUO. Analysis of Blast Resistance Genes and Neck Blast Resistance of japonica Rice Varieties/Lines Recently Developed in Jiangsu Province[J]. Chinese Journal OF Rice Science, 2020, 34(5): 413-424.

图/表 11

表1 本研究用于各抗病基因检测的阴阳性对照

Table 1 Information of controls used in detection of resistance genes to blast disease in present study.

基因	染色体	阳性对照	阴性对照	基因	染色体	阳性对照	阴性对照
Gene	Chr.	Control 1	Control 2	Gene	Chr.	Control 1	Control 2
Pit	1	IRBLt-K59	丽江新团黑谷LTH	Pia	11	IRBLa-A	丽江新团黑谷LTH
Pib	2	IRBLb-B	丽江新团黑谷LTH	Pik	11	IRBLk-Ka	丽江新团黑谷LTH
Pigm	6	谷梅4号Gumei 4	丽江新团黑谷LTH	Pikm	11	IRBLkm-Ts	日本晴Nipponbare
Pid3	6	地谷Digu	丽江新团黑谷LTH	Pikh	11	IRBLkh-K3	丽江新团黑谷LTH
Pi25	6	地谷Digu	丽江新团黑谷LTH	Pi1	11	IRBL1-CL	日本晴Nipponbare
Pid2	6	地谷Digu	丽江新团黑谷LTH	Pb1	11	镇稻18 Zhendao 18	日本晴Nipponbare
Pid2	6	地谷Digu	丽江新团黑谷LTH	Pita	12	IRBLta-K1	丽江新团黑谷LTH
Pi3/5/i	9	IRBL5-M	丽江新团黑谷LTH

表2 本研究用于各抗病基因检测的功能标记引物信息

Table 2 Information of molecular markers used in detection of resistant genes to blast disease in present study.

基因 Gene	染色体 Chr.	标记 Marker	引物序列(5'-3') Primer sequences(5'-3')	片段大小 Fragment size / bp	抗/感病功能 Function	限制性内切酶 Enzyme	参考文献 Reference
Pit	1	tRn1	F: GTTGGAGCTACGGTTGTTCAG	733	抗 R		[20]
		tRn2	F: CCAAGGGATTAGGTCCTAGTG	733	抗 R
		tN11	R: CAGCGATGGTATGAGCACAA	530	感 S
Pib	2	Pibdom	F: GAACAATGCCCAAACTTGAGA	365	抗 R		[21]
		Pibdom	R: GGGTCCACATGTCAGTGAGC	365	抗 R
		Lys145	F: TCGGTGCCTCGGTAGTCAGT	803	感 S
		Lys145	R: GGGAAGCGGATCCTAGGTCT	803	感 S
Pigm	6	GMR-3	F: AGTTCTACTTACGGAGGAGC	146+98	抗 R		[22]
		GMR-3	R: AGAATTATGATAAAGAGAAAGGAA	146+98	抗 R
		Actin1-1	F: ACAAAGTTTTCAACCGGCCTA	146	感 S
		Actin1-1	R: CTGGTACCCTCATCAGGCATC	146	感 S
Pid3	6	Pid3-dCAPS	F: TACTACTCATGGAAGCTAGTTCTC	751	抗 R	BamH I	[23]
Pid3	6	Pid3-dCAPS	R: ACGTCACAAATCATTCGCTC	751	抗 R	BamH I	[23]
Pi25	6	CAP3	F: CCTCACGTTTCTACGTCTTG	490	抗 R	Nde I	[24]
Pi25	6	CAP3	R: CACACCATTTCTGATGAACC	490	抗 R	Nde I	[24]
Pid2	6	D2	F: TTGGCTATCATAGGCGTCC	1057	抗 R	MIu I	[25]
Pid2	6	D2	R: ATTTGAAGGCGTTTGCGTAGA	1057	抗 R	MIu I	[25]
Pi3/5/i	9	M-Pi5	F: ATAGATCATGCGCCCTCTTG	206	抗 R		[26]
Pi3/5/i	9	M-Pi5	R: ATAGATCATGCGCCCTCTTG	307	感 S		[26]
Pia	11	Pia	F: CTTTTGAGCTTGATTGGTCTGC	116	抗 R		[27]
Pia	11	Pia	R: CTATTGCACCAGAGGGACCAG	125	感 S		[27]
Pik	11	RGA4	F: GGAAAGCTGATATGTTGTCG	1650 -	抗 R
Pik	11	RGA4	R:ACTCGGAGTCGGAGAGTCAG	1650 -	感 S
Pikh	11	FM143	F: CCCAACATTGGTAGTAGTGC	285	抗 R		[28]
Pikh	11	FM143	R: TCCTTCATACGCAAVCCAATCT	401	感 S		[28]
Pikm	11	DKm1	F: TGAGCTCAAGGCAAGAGTTGAGGA	174	抗 R		[29]
		DKm1	R: TGTTCCAGCAACTCGATGAG	213	感 S
		DKm2	F: CAGTAGCTGTGTCTCAGAACTATG	290	抗 R
		DKm2	R: AAGGTACCTCTTTTCGGCCAG	332	感 S
Pi1	11	M-Pi1	F: GTGCTGCTGTGGCTAGTTTG	460 -	抗 R		[30]
Pi1	11	M-Pi1	R: AGTCCCCGCTCAATTTTTCT	460 -	感 S		[30]
Pb1	11	Pb1	F: ATCAACGCTACCTTCCC	159 -	抗 R		[31]
Pb1	11	Pb1	R: GTGCCATCACAATTTCTTC	159 -	感 S		[31]
Pita	12	Pita-R	F: AGCAGGTTATAAGCTAGGCC	1042	抗 R		[32]
		Pita-R	R: CTACCAACAAGTTCATCAAA	1042	抗 R
		Pita-S	F: AGCAGGTTATAAGCTAGCTAT	1042	感 S
		Pita-S	R: CTACCAACAAGTTCATCAAA	1042	感 S

表2 本研究用于各抗病基因检测的功能标记引物信息

Table 2 Information of molecular markers used in detection of resistant genes to blast disease in present study.

基因 Gene	染色体 Chr.	标记 Marker	引物序列(5'-3') Primer sequences(5'-3')	片段大小 Fragment size / bp	抗/感病功能 Function	限制性内切酶 Enzyme	参考文献 Reference
Pit	1	tRn1	F: GTTGGAGCTACGGTTGTTCAG	733	抗 R		[20]
		tRn2	F: CCAAGGGATTAGGTCCTAGTG	733	抗 R
		tN11	R: CAGCGATGGTATGAGCACAA	530	感 S
Pib	2	Pibdom	F: GAACAATGCCCAAACTTGAGA	365	抗 R		[21]
		Pibdom	R: GGGTCCACATGTCAGTGAGC	365	抗 R
		Lys145	F: TCGGTGCCTCGGTAGTCAGT	803	感 S
		Lys145	R: GGGAAGCGGATCCTAGGTCT	803	感 S
Pigm	6	GMR-3	F: AGTTCTACTTACGGAGGAGC	146+98	抗 R		[22]
		GMR-3	R: AGAATTATGATAAAGAGAAAGGAA	146+98	抗 R
		Actin1-1	F: ACAAAGTTTTCAACCGGCCTA	146	感 S
		Actin1-1	R: CTGGTACCCTCATCAGGCATC	146	感 S
Pid3	6	Pid3-dCAPS	F: TACTACTCATGGAAGCTAGTTCTC	751	抗 R	BamH I	[23]
Pid3	6	Pid3-dCAPS	R: ACGTCACAAATCATTCGCTC	751	抗 R	BamH I	[23]
Pi25	6	CAP3	F: CCTCACGTTTCTACGTCTTG	490	抗 R	Nde I	[24]
Pi25	6	CAP3	R: CACACCATTTCTGATGAACC	490	抗 R	Nde I	[24]
Pid2	6	D2	F: TTGGCTATCATAGGCGTCC	1057	抗 R	MIu I	[25]
Pid2	6	D2	R: ATTTGAAGGCGTTTGCGTAGA	1057	抗 R	MIu I	[25]
Pi3/5/i	9	M-Pi5	F: ATAGATCATGCGCCCTCTTG	206	抗 R		[26]
Pi3/5/i	9	M-Pi5	R: ATAGATCATGCGCCCTCTTG	307	感 S		[26]
Pia	11	Pia	F: CTTTTGAGCTTGATTGGTCTGC	116	抗 R		[27]
Pia	11	Pia	R: CTATTGCACCAGAGGGACCAG	125	感 S		[27]
Pik	11	RGA4	F: GGAAAGCTGATATGTTGTCG	1650 -	抗 R
Pik	11	RGA4	R:ACTCGGAGTCGGAGAGTCAG	1650 -	感 S
Pikh	11	FM143	F: CCCAACATTGGTAGTAGTGC	285	抗 R		[28]
Pikh	11	FM143	R: TCCTTCATACGCAAVCCAATCT	401	感 S		[28]
Pikm	11	DKm1	F: TGAGCTCAAGGCAAGAGTTGAGGA	174	抗 R		[29]
		DKm1	R: TGTTCCAGCAACTCGATGAG	213	感 S
		DKm2	F: CAGTAGCTGTGTCTCAGAACTATG	290	抗 R
		DKm2	R: AAGGTACCTCTTTTCGGCCAG	332	感 S
Pi1	11	M-Pi1	F: GTGCTGCTGTGGCTAGTTTG	460 -	抗 R		[30]
Pi1	11	M-Pi1	R: AGTCCCCGCTCAATTTTTCT	460 -	感 S		[30]
Pb1	11	Pb1	F: ATCAACGCTACCTTCCC	159 -	抗 R		[31]
Pb1	11	Pb1	R: GTGCCATCACAATTTCTTC	159 -	感 S		[31]
Pita	12	Pita-R	F: AGCAGGTTATAAGCTAGGCC	1042	抗 R		[32]
		Pita-R	R: CTACCAACAAGTTCATCAAA	1042	抗 R
		Pita-S	F: AGCAGGTTATAAGCTAGCTAT	1042	感 S
		Pita-S	R: CTACCAACAAGTTCATCAAA	1042	感 S

图1 各抗病基因功能标记在阳性、阴性等品种/系上的扩增验证图中各基因标记检测图谱中,泳道从左至右依次为分子量标记、阳性对照、阴性对照及随机选择的10个江苏粳稻品种;在分子量标记上,除了Pid2和Pia检测时使用DL2000、Pit检测时使用的20 bp DNA ladder外,其余基因检测中采用的标记均是DL10000;各基因采用的阳性和阴性对照具体名称见表1。

Fig. 1. Validation of the molecular markers in determining the corresponding resistance gene using known positive and negative varieties or lines as well as 10 japonica varieties randomly selected from Jiangsu Province. In the picture, the lanes from left to right are DNA Markers, positive control, negative control and 10 random japonica varieties from Jiangsu Province; About the DNA markers, expect Pid2 and Pia were DL2000, Pit was 20 bp DNA ladder, others was DL10000; The information of positive and negative control is listed in Table 1.

图2 各抗病基因在江苏粳稻品种中的分布频率（A）及品种携带不同抗病基因数分布（B）

Fig. 2. Distribution frequency of resistance genes in japonica rice from Jiangsu Province(A) and the distribution of gene number carried by varieties(B).

图3 抗病基因在江苏不同生态型粳稻品种中的分布频率

Fig. 3. Distribution of resistance genes in different types of japonica rice.

图4 各抗病基因在2019年前审定及2019年区试品种/系间的频率分布比较

Fig. 4. Comparison of resistance gene distribution frequency between varieties authorized before 2019 and regional test varieties in 2019.

图5 供试品种稻瘟病病级分布

Fig. 5. Distribution of disease grade in varieties tested. EMMLJ, Early- and medium-maturity late japonica rice; LMMJ, Late-maturity medium japonica rice; MMMJ, Medium-maturity medium japonica rice.

图6 品种携带抗病基因数与病级间的关系分析

Fig. 6. Analysis of the resistance gene number carried by each variety and disease grade.

图7 各抗病基因对穗颈瘟抗性的贡献逻辑回归统计参数 A、B图分别为单基因逐一逻辑回归和多基因逻辑回归的统计参数;回归分析中的基因回归系数达到统计分析中的显著或极显著水平的,其P值分别以蓝色字体标注在相应的基因位置;图中黑点为优势比值,黑点左右黑线分别为各优势比的95%置信区间。

Fig. 7. Logistic regression statistical analysis about the contribution of each resistance genes to neck blast resistance. A and B mean single gene one by one logistic regression statistical parameter chart and multigene logistic regression statistical parameter chart, respectively; In regressive analysis, if the regression coefficient reached significant or extremely significant level, then there will be a P value around the specific gene; The dark dots mean dominance ration, the dark lines mean 95% confidence interval of each dominance ration.

图8 三个对穗颈瘟抗性有显著贡献的基因及组合间的品种病级差异比较-表示不带有相应的抗病等位基因;柱状图上的数值表示携带相应基因或组合的品种平均病级;不同的小写字母表示彼此间的病级差异达到5%水平上的统计显著性差异。

Fig. 8. Contribute significantly of three genes to the resistance of neck blast resistance and the comparison of disease grade differences among them and their combinations. - means the absence of the specific resistance gene; The value above each bar means the average disease grade of each gene or gene combination; The lowercase letter means significance difference at 5% level between two disease grades.

图9 携带Pigm高世代回交株系的穗颈瘟抗性表现

Fig. 9. Resistance performance of advanced backcrossing lines with Pigm and the corresponding recurrent parents.

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江苏近年育成粳稻新品种/系的稻瘟病抗性基因及穗颈瘟抗性分析

Analysis of Blast Resistance Genes and Neck Blast Resistance of japonica Rice Varieties/Lines Recently Developed in Jiangsu Province

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