持有Pi9基因的水稻单基因系IRBL9-W对稻瘟病菌苗期和成株期抗性差异

doi:10.16819/j.1001-7216.2021.0717

中国水稻科学 ›› 2021, Vol. 35 ›› Issue (3): 303-310.DOI: 10.16819/j.1001-7216.2021.0717

• 研究报告 • 上一篇

持有Pi9基因的水稻单基因系IRBL9-W对稻瘟病菌苗期和成株期抗性差异

刘树芳^1,²^,^#, 董丽英^1,²^,^#, 李迅东^1,², 周伍民^1,³, 杨勤忠^1,^2,^*()

¹云南省农业科学院农业环境资源研究所/云南省农业跨境有害生物绿色防控重点实验室,昆明 650205
²农业农村部昆明作物有害生物科学观测实验站,昆明 650205
³云南大学农学院,昆明 650091

收稿日期:2020-07-24 修回日期:2020-11-24 出版日期:2021-05-10 发布日期:2021-05-10
通讯作者: 杨勤忠
作者简介:
^#共同第一作者;
基金资助:
云南省基础研究计划重点资助项目(202001AS070006);科技部国家重点研发专项(2016YFD0100101);国家自然科学基金资助项目(31860524);云南省农业科学院院基金资助项目(YJZ201803)

Different Reactions of Rice Monogenic Line IRBL9-W Harboring Pi9 Gene to Magnaporthe oryzae Containing AvrPi9 During Seedling and Adult-plant Stages

Shufang LIU^1,²^,^#, Liying DONG^1,²^,^#, Xundong LI^1,², Wumin ZHOU^1,³, Qinzhong YANG^1,^2,^*()

¹Agricultural Environment Resources Research Institute, Yunnan Academy of Agricultural Sciences/Key Laboratory of Green Prevention and Control of Agricultural Transboundary Pests of Yunnan Province, Kunming 650205, China
²Scientific Observing and Experimental Station of Crop Pests in Kunming, Ministry of Agriculture and Rural Affairs, Kunming 650205, China
³College of Agriculture, Yunnan University, Kunming 650091, China

Received:2020-07-24 Revised:2020-11-24 Online:2021-05-10 Published:2021-05-10
Contact: Qinzhong YANG
About author:
^#These authors contributed equally to this work;

摘要/Abstract

摘要：

【目的】Pi9是一个广谱稻瘟病抗性基因,田间病圃监测发现持有Pi9的水稻单基因系IRBL9-W在苗期高抗稻瘟病,但却感穗瘟。探明水稻单基因系IRBL9-W在苗期抗病而孕穗末期感染穗颈瘟的原因,为Pi9基因在水稻抗病育种中的有效利用提供参考。【方法】利用从IRBL9-W穗颈瘟病斑上分离的8个单孢菌株以及实验室保存的单孢菌株Y363,在温室分别对单基因系IRBL9-W苗期和孕穗末期进行接种鉴定;并利用病原菌AvrPi9基因的特异引物对9个单孢菌株进行PCR扩增及产物测序;提取水稻单基因系IRBL9-W苗期叶片和抽穗期穗部的总RNA,通过半定量RT-PCR以及实时qRT-PCR分析Pi9基因在苗期和穗期的表达。【结果】在温室人工喷雾接种条件下,IRBL9-W在苗期对从穗颈瘟上分离的8个单孢及对照菌株Y363均表现为抗病;随机选取的2个从IRBL9-W穗颈瘟病样分离的单孢菌株(YX2-7-1和YX2-15-1)及对照菌株Y363对孕穗末期IRBL9-W注射接种,接种的植株表现出典型的穗颈瘟症状;AvrPi9的等位基因分析结果表明,与AvrPi9相比,Y363中的等位基因与AvrPi9完全相同,而从IRBL9-W穗瘟分离的8个单孢菌株中编码区与AvrPi9基因完全相同,但在编码起始位置上游-264 bp处缺失16 bp的一段序列。由于IRBL9-W苗期对这些菌株均表现抗病,推测这段序列的缺失并不影响AvrPi9基因的功能;实时qRT-PCR分析结果表明,Pi9基因在穗部的表达量为苗期叶片表达量的47.3%。【结论】在水稻单基因系IRBL9-W中,与苗期叶片中Pi9基因的表达量相比,Pi9基因在穗部表达量的明显降低可能是造成IRBL9-W穗期感稻瘟病的原因。

关键词: 水稻, 稻瘟病菌, Pi9, 苗期, 成株期

Abstract:

【Objective】 The Pi9 gene confers broad-spectrum resistance against Magnaporthe oryzae. Rice monogenic line IRBL9-W carrying single blast resistance gene Pi9 is highly resistant to leaf blast during the seedling stage, but it is susceptible to neck blast at adult-plant stage in blast nursery. It is important to analyze the mechanism behind the reversion of resistance to Magnaporthe oryzae, which will provide important information for effective utilization of Pi9 in disease-resistant rice breeding program.【Method】Eight single spore strains of M. oryzae isolated from neck blast samples of diseased IRBL9-W, and one isolate Y363 were artificially inoculated by spraying at seedling stage to pathotype their pathogenicity on 24 monogenic lines including IRBL9-W. Two out of eight strains and control strain Y363 were further selected and inoculated on IRBL9-W at late-booting stage by injecting to evaluate their pathogenicity. The alleles of M. oryzae AvrPi9 gene cognate to rice blast resistance gene Pi9 were amplified and sequenced with AvrPi9-specific primers in nine strains. Total RNA of the rice monogenic line IRBL9-W in seedling and late-booting stage was extracted from young leaf and panicle, respectively. The expression analysis of Pi9 gene was performed by semi-quantitative RT-PCR and real-time qRT-PCR. 【Result】 The monogenic line IRBL9-W was completely resistant to the nine M. oryzae strains at the seedling stage exposed to the artificial spraying. However, IRBL9-W was susceptible to panicle blast when inoculated with both two randomly selected strains (YX2-7-1 and YX2-15-1) isolated from panicle blast lesions of IRBL9-W, and the control strain Y363 by injecting inoculation at the late-booting stage. Compared with AvrPi9, allelic sequence amplified from Y363 was identical to cloned AvrPi9, and the coding region of the other eight strains isolated from IRBL9-W panicle blast was identical to AvrPi9 gene, except for a 16-bp deletion at -264 bp upstream of the coding starting position in the eight strains. Given high resistance of IRBL9-W to these strains at the seedling stage, we contended that this 16-bp deletion did not affect the function of AvrPi9 gene; qRT-PCR results showed that the relative expression level of Pi9 gene in panicle was only 47.3% as compared to that in seedling leaves. 【Conclusion】Compared with the relative expression level of Pi9 gene at the seedling stage in IRBL9-W, the significant decrease of Pi9 gene expression level in panicle of IRBL9-W could be the cause of susceptibility to M. oryzae at late-booting stage.

Key words: Oryza sativa L., Magnaporthe oryzae, Pi9, seedling stage, adult-plant stage

刘树芳, 董丽英, 李迅东, 周伍民, 杨勤忠. 持有Pi9基因的水稻单基因系IRBL9-W对稻瘟病菌苗期和成株期抗性差异[J]. 中国水稻科学, 2021, 35(3): 303-310.

Shufang LIU, Liying DONG, Xundong LI, Wumin ZHOU, Qinzhong YANG. Different Reactions of Rice Monogenic Line IRBL9-W Harboring Pi9 Gene to Magnaporthe oryzae Containing AvrPi9 During Seedling and Adult-plant Stages[J]. Chinese Journal OF Rice Science, 2021, 35(3): 303-310.

图/表 5

表1 稻瘟病菌单孢菌株在单基因系上的致病性测定

Table 1 Pathotyping of Magnaporthe oryzae strains on monogenic lines.

单基因系 Monogenic line	抗性基因 R gene	稻瘟病菌株 Magnaporthe oryzae strains
单基因系 Monogenic line	抗性基因 R gene	YX2-2-1	YX2-3-2	YX2-5-1	YX2-6-1	YX2-10-1	YX2-14-1	YX2-7-1	YX2-15-1	Y363
IRBLA-A	Pia	R	R	R	R	R	R	S	S	S
IRBLI-F5	Pii	R	S	R	S	S	R	S	S	S
IRBLKS-F5	Piks	S	S	S	S	S	S	S	S	S
KRBLK KA	Pik	S	S	S	S	S	S	S	S	S
KRBLKP-K60	Pikp	S	S	S	S	S	S	S	S	R
KRBLKH-K3	Pikh	R	R	R	R	R	R	R	S	R
IRBLZ5-CA	Pi2	S	S	S	S	S	S	S	S	S
IRBLZ FU	Piz	S	S	S	S	S	S	S	S	S
IRBLZT-T	Pizt	R	R	R	R	R	R	R	R	R
IRBLTA-K1	Pita	S	S	S	S	S	S	S	S	S
IRBLB-B	Pib	S	S	S	S	S	S	S	S	S
IRBLT-K59	Pit	R	R	R	R	R	R	S	S	S
IRBLSH-B	Pish	S	S	S	S	S	S	S	S	R
IRBL1-CL	Pi1	R	R	R	R	R	R	R	S	S
IRBL3-CP4	Pi3	S	S	S	S	S	S	S	S	S
IRBL5-M	Pi5	R	R	R	R	R	R	R	R	R
IRBL7-M	Pi7	S	S	S	S	R	S	S	S	R
IRBL12-M	Pi12	S	S	S	S	S	S	S	S	R
IRBL19-A	Pi19	S	S	S	S	R	S	S	S	S
IRBLKM-TS	Pikm	R	R	R	R	R	R	R	S	S
IRBL20-IR24	Pi20	R	R	R	R	R	R	R	R	R
IRBLTA2-PI	Pita2	S	S	S	S	S	S	R	S	R
IRBL11-ZH	Pi11	S	S	S	S	S	S	S	S	S
IRBL9-W	Pi9	R	R	R	R	R	R	R	R	R
LTH	-	S	S	S	S	S	S	S	S	S

图2 不同菌株中AvrPi9的扩增分析 M–DNA分子量标记; 条带1~9分别指稻瘟病菌菌株YX2-2-1, YX2-3-2, YX2-5-1, YX2-6-1, YX2-10-1, YX2-14-1, YX2-7-1, YX2-15-1和Y363。

Fig. 2. Amplification of AvrPi9 from different M. oryzae strains. M, DNA molecular marker; DL2000; Lane 1 to Lane 9 refer to YX2-2-1, YX2-3-2, YX2-5-1, YX2-6-1, YX2-10-1, YX2-14-1, YX2-7-1, YX2-15-1 and Y363, respectively.

表2 孕穗末期利用3个稻瘟病菌菌株接种IRBL9-W后的穗瘟调查结果

Table 2 Investigating results of panicle blast of IRBL9-W inoculated with three M. oryzae strains at late-booting stage.

稻瘟病菌株 M. oryzae strain	发病级别 Disease scores
稻瘟病菌株 M. oryzae strain	0	1	3	5	7	9
YX2-7-1	0^a	0	0	1	1	13
YX2-15-1	0	0	0	0	1	13
Y363	0	0	0	0	1	14
空白对照CK^b	15	0	0	0	0	0

图1 单基因系IRBL9-W孕穗末期及苗期接种的症状 A~C–IRBL9-W接种菌株YX2-7-1、YX2-15-1和Y363在孕穗末期的症状; D~F–IRBL9-W（左）和丽江新团黑谷（右）苗期接种菌株YX2-7-1、YX2-15-1和Y363后的症状。

Fig. 1. Symptoms of monogenic line IRBL9-W inoculated with M. oryzae fungus at late-booting stage and seedling stage. A–C, Symptoms of monogenic line IRBL9-W inoculated with M. oryze strains YX2-7-1, YX2-15-1 and Y363 at late booting stage, respectively. D–F: Symptoms of monogenic line IRBL9-W (left) and Lijiangxintuanheigu (right) inoculated with M. oryzae strains YX2-7-1, YX2-15-1 and Y363 at seedling stage, respectively.

图3 Pi9基因的表达分析 A–Pi9基因的半定量分析。从成株期穗部(P)及苗期叶片(L)制备的cDNA用作PCR模板进行扩增,Actin1 用作内参基因;B–实时荧光定量RT-PCR比较分析Pi9基因在苗期和穗期的表达量。

Fig. 3. Expression level of Pi9 gene. A, Expression analysis of Pi9 gene by semi-quantitative RT-PCR method. The cDNA prepared from panicles (P) of adult plants and seedling leaves (L) of IRBL9-W carrying Pi9 gene was used as template for PCR amplification, with Actin 1 as reference gene; B, Comparative analysis of relative expression level of Pi9 gene in panicle and seedling leaves of IRBL9-W by real-time quantitative RT-PCR.

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持有Pi9基因的水稻单基因系IRBL9-W对稻瘟病菌苗期和成株期抗性差异

Different Reactions of Rice Monogenic Line IRBL9-W Harboring Pi9 Gene to Magnaporthe oryzae Containing AvrPi9 During Seedling and Adult-plant Stages

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[1]	郭展, 张运波. 水稻对干旱胁迫的生理生化响应及分子调控研究进展[J]. 中国水稻科学, 2024, 38(4): 335-349.
[2]	韦还和, 马唯一, 左博源, 汪璐璐, 朱旺, 耿孝宇, 张翔, 孟天瑶, 陈英龙, 高平磊, 许轲, 霍中洋, 戴其根. 盐、干旱及其复合胁迫对水稻产量和品质形成影响的研究进展[J]. 中国水稻科学, 2024, 38(4): 350-363.
[3]	许丹洁, 林巧霞, 李正康, 庄小倩, 凌宇, 赖美玲, 陈晓婷, 鲁国东. OsOPR10正调控水稻对稻瘟病和白叶枯病的抗性[J]. 中国水稻科学, 2024, 38(4): 364-374.
[4]	候小琴, 王莹, 余贝, 符卫蒙, 奉保华, 沈煜潮, 谢杭军, 王焕然, 许用强, 武志海, 王建军, 陶龙兴, 符冠富. 黄腐酸钾提高水稻秧苗耐盐性的作用途径分析[J]. 中国水稻科学, 2024, 38(4): 409-421.
[5]	胡继杰, 胡志华, 张均华, 曹小闯, 金千瑜, 章志远, 朱练峰. 根际饱和溶解氧对水稻分蘖期光合及生长特性的影响[J]. 中国水稻科学, 2024, 38(4): 437-446.
[6]	刘福祥, 甄浩洋, 彭焕, 郑刘春, 彭德良, 文艳华. 广东省水稻孢囊线虫病调查与鉴定[J]. 中国水稻科学, 2024, 38(4): 456-461.
[7]	陈浩田, 秦缘, 钟笑涵, 林晨语, 秦竞航, 杨建昌, 张伟杨. 水稻根系和土壤性状与稻田甲烷排放关系的研究进展[J]. 中国水稻科学, 2024, 38(3): 233-245.
[8]	缪军, 冉金晖, 徐梦彬, 卜柳冰, 王平, 梁国华, 周勇. 过量表达异三聚体G蛋白γ亚基基因RGG2提高水稻抗旱性[J]. 中国水稻科学, 2024, 38(3): 246-255.
[9]	尹潇潇, 张芷菡, 颜绣莲, 廖蓉, 杨思葭, 郭岱铭, 樊晶, 赵志学, 王文明. 多个稻曲病菌效应因子的信号肽验证和表达分析[J]. 中国水稻科学, 2024, 38(3): 256-265.
[10]	朱裕敬, 桂金鑫, 龚成云, 罗新阳, 石居斌, 张海清, 贺记外. 全基因组关联分析定位水稻分蘖角度QTL[J]. 中国水稻科学, 2024, 38(3): 266-276.
[11]	魏倩倩, 汪玉磊, 孔海民, 徐青山, 颜玉莲, 潘林, 迟春欣, 孔亚丽, 田文昊, 朱练峰, 曹小闯, 张均华, 朱春权. 信号分子硫化氢参与硫肥缓解铝对水稻生长抑制作用的机制[J]. 中国水稻科学, 2024, 38(3): 290-302.
[12]	周甜, 吴少华, 康建宏, 吴宏亮, 杨生龙, 王星强, 李昱, 黄玉峰. 不同种植模式对水稻籽粒淀粉含量及淀粉关键酶活性的影响[J]. 中国水稻科学, 2024, 38(3): 303-315.
[13]	关雅琪, 鄂志国, 王磊, 申红芳. 影响中国水稻生产环节外包发展因素的实证研究：基于群体效应视角[J]. 中国水稻科学, 2024, 38(3): 324-334.
[14]	许用强, 姜宁, 奉保华, 肖晶晶, 陶龙兴, 符冠富. 水稻开花期高温热害响应机理及其调控技术研究进展[J]. 中国水稻科学, 2024, 38(2): 111-126.
[15]	吕海涛, 李建忠, 鲁艳辉, 徐红星, 郑许松, 吕仲贤. 稻田福寿螺的发生、危害及其防控技术研究进展[J]. 中国水稻科学, 2024, 38(2): 127-139.