水稻类病斑突变体lmm7的鉴定与基因定位

doi:10.16819/j.1001-7216.2022.210711

中国水稻科学 ›› 2022, Vol. 36 ›› Issue (4): 357-366.DOI: 10.16819/j.1001-7216.2022.210711

水稻类病斑突变体lmm7的鉴定与基因定位

孙志广¹, 代慧敏², 陈庭木¹, 李景芳¹, 迟铭¹, 周振玲¹, 刘艳¹, 刘金波¹, 徐波¹, 邢运高¹, 杨波¹, 李健¹, 卢百关¹, 方兆伟¹, 王宝祥¹^,^*(), 徐大勇¹^,^*()

¹连云港市农业科学院/江苏省现代作物生产协同创新中心, 江苏连云港 222006
²连云港市生产力促进中心, 江苏连云港 222006

收稿日期:2021-07-30 修回日期:2021-08-30 出版日期:2022-07-10 发布日期:2022-07-12
通讯作者: 王宝祥,徐大勇
基金资助:
现代农业技术体系建设专项资金资助项目(CARS-01-61);江苏省农业重大品种创制计划资助项目(PZCZ201704);江苏省自然科学基金面上项目(BK20201214);苏北科技专项(LYG-SZ201930);苏北科技专项(LYG-SZ202040);连云港市财政专项资金资助项目(QNJJ1902);连云港市财政专项资金资助项目(QNJJ2001)

Phenotypic Identification and Gene Mapping of a Lesion Mimic Mutant lmm7 in Rice

SUN Zhiguang¹, DAI Huimin², CHEN Tingmu¹, LI Jingfang¹, CHI Ming¹, ZHOU Zhenling¹, LIU Yan¹, LIU Jinbo¹, XU Bo¹, XING Yungao¹, YANG Bo¹, LI Jian¹, LU Baiguan¹, FANG Zhaowei¹, WANG Baoxiang¹^,^*(), XU Dayong¹^,^*()

¹Lianyungang Academy of Agricultural Sciences/Jiangsu Collaborative Innovation Center for Modern Corp Production, Lianyungang 222006, China
²Lianyungang Productivity Promotion Center, Lianyungang 222006, China

Received:2021-07-30 Revised:2021-08-30 Online:2022-07-10 Published:2022-07-12
Contact: WANG Baoxiang, XU Dayong

摘要/Abstract

摘要：

【目的】对水稻类病斑突变体的研究有助于解析其与植物生长和防御反应的关系。【方法】本研究在粳稻品系FI135胚培养过程中获得了1个类病斑突变体lmm7(lesion mimic mutant 7)。通过对其进行系统的表型鉴定、农艺性状考查、超微结构观察、生理学特性分析，阐明LMM7基因对植物生长的调控。通过病原菌抗性鉴定，明确lmm7对植物防御反应的影响。利用9311B与突变体lmm7杂交所得F₂群体对该突变体进行了遗传分析和基因精细定位。【结果】该突变体苗期表型正常，分蘖初期，植株基部叶片从叶尖开始不断出现褐色斑点，并向整株扩散，且斑点数目随植株生长不断增加。与野生型相比，突变体的株高、穗长、有效穗数、每穗粒数、结实率及剑叶长宽都显著降低，但籽粒性状和抽穗期没有显著性差异。遮光处理表明，突变体lmm7的表型受到光照诱导，抽穗期突变体lmm7叶肉细胞严重失绿，光合色素含量显著降低。组织化学分析表明，突变体病斑处的H₂O₂含量显著升高。透射电镜观察结果表明，突变体lmm7叶肉细胞的叶绿体数目减少，叶绿体类囊体片层结构严重受损，细胞器肿胀解体，并出现大量嗜锇小体，同时病斑内部和周围区域积累了大量的ROS。抗性鉴定结果显示突变体lmm7稻瘟病抗性水平显著高于野生型。遗传分析表明lmm7的突变表型受单个隐性基因控制。利用图位克隆的方法，目的基因被定位于水稻第7染色体短臂两InDel标记7B35和7B43之间，区间范围约260 kb。测序结果表明该区间内候选基因LOC_Os07g0203700第2891位碱基T发生了单碱基缺失，导致后续移码突变及翻译提前终止。【结论】lmm7与spl5互为等位基因，其突变抑制了植株的生长，同时增强了对稻瘟病的抗性。

关键词: 水稻, 类病斑突变体, 表型鉴定, 基因定位

Abstract:

【Objective】The study of lesion mimic mutants (LMMs) is helpful to understand the interaction between growth and defense response in plants.【Method】We herein reported a novel lesion mimic mutant 7 (lmm7) derived from a japonica rice variety FI135 by embryo culture. For deciphering the effect of lmm7 on plant growth, phenotype and agronomic traits were compared between the wild type (WT) and lmm7 plants. The ultrastructure of mesophyll cells and histochemical characterization was done in WT and lmm7 plants. Evaluation of WT and lmm7 was observed for resistance to pathogens was employed to explore the underlying effect of lmm7 on plant defense response. An F₂ population derived from the cross between lmm7 mutant and 9311B was used for genetic analysis and gene mapping.【Result】The seedlings of lmm7 were normal at the seedling stage, but the basal leaves began to exhibit brown spots at the initial tillering stage, which then spread to the whole plant, and the number of mosaic spots increased along with the development of lmm7 plants. Compared with the wild type, agronomic traits including plant height, panicle length, effective panicle number, grain number per panicle, and seed setting rate were significantly decreased in the mutant. However, no significant difference in seed traits and heading date was observed between WT and lmm7 plants. The shading assay indicated that lesion formation in lmm7 was induced by light. The mesophyll cells of lmm7 mutant were seriously chlorotic, and the photosynthetic pigment contents of the mutant were significantly decreased at the heading stage. Histochemical analysis showed that a large amount of hydrogen peroxide (H₂O₂) accumulated at the lesion sites. The observation with a transmission electronic microscope (TEM) demonstrated that the number of chloroplasts in mesophyll cells of lmm7 plants was decreased, the lamellar structure of chloroplasts was seriously damaged, and osmiophilic granules were obviously increased in the mutant. Moreover, the levels of reactive oxygen species (ROS) were increased in the internal and surrounding regions of the lesions. The lmm7 plants displayed higher resistance to rice blast compared to the wild type. Genetic analysis revealed that in lmm7 the mutation was controlled by a single recessive gene. Using a map-based cloning strategy, lmm7 was narrowed down to a 260 kb region between InDel markers 7B35 and 7B43 on the short arm of chromosome 7 in rice. Sequencing analysis revealed that a deletion of T base at 2891 nucleotide position occurred in the DNA sequence of LOC_Os07g0203700, resulting in a frame-shift mutation and a premature stop codon.【Conclusion】The results reveal the target gene is allelic to spl5, and the mutation of lmm7 enhances resistance to rice blast, while it curbs the growth and development of lmm7 plants.

Key words: Oryza sativa L., lesion mimic mutants, phenotypic identification, gene mapping

孙志广, 代慧敏, 陈庭木, 李景芳, 迟铭, 周振玲, 刘艳, 刘金波, 徐波, 邢运高, 杨波, 李健, 卢百关, 方兆伟, 王宝祥, 徐大勇. 水稻类病斑突变体lmm7的鉴定与基因定位[J]. 中国水稻科学, 2022, 36(4): 357-366.

SUN Zhiguang, DAI Huimin, CHEN Tingmu, LI Jingfang, CHI Ming, ZHOU Zhenling, LIU Yan, LIU Jinbo, XU Bo, XING Yungao, YANG Bo, LI Jian, LU Baiguan, FANG Zhaowei, WANG Baoxiang, XU Dayong. Phenotypic Identification and Gene Mapping of a Lesion Mimic Mutant lmm7 in Rice[J]. Chinese Journal OF Rice Science, 2022, 36(4): 357-366.

图/表 7

图1 野生型（WT）和突变体lmm7的表型鉴定 A―分蘖期野生型（WT）和突变体（lmm7）植株表型，红色箭头指示出现类病斑叶片;B―分蘖期野生型（WT）和突变体（lmm7）基部叶片;C―大田条件下，野生型（WT）和突变体（lmm7）成熟期植株;D―野生型（WT）和突变体（lmm7）成熟种子大小的比较;E~F―植株主茎穗和节间长度的比较。在A中，比例尺为5 cm;在B中比例尺为2 cm;在C和E中，比例尺为10 cm;在D中，比例尺为10 mm。

Fig. 1. Phenotypic identification of WT plants and lmm7. A, Plants of wild type (WT) and lmm7 mutant at the tillering stage, red arrows indicate lesion mimic leaves; B, Basal leaves of wild type (WT) and lmm7 at the tillering stage; C, Plants of wild type (WT) and lmm7 at the mature period in field conditions; D, Comparisons of the size of mature seeds from WT and lmm7 plants; E-F: Panicles and internodes of main culms. P, Panicle; I to VI indicate corresponding internodes from top to bottom. Bars =5 cm in A; 2 cm in B; 10 cm in C, E; 10 mm in D.

表1 野生型（WT）和lmm7突变体的农艺性状比较

Table 1. Agronomic traits comparison between WT plants and lmm7.

性状 Trait	野生型Wild type	突变体 lmm7	性状 Trait	野生型Wild type	突变体 lmm7
抽穗期 Heading date /d	95.3±0.6	95.7±0.6	千粒重 1000-grain weight/g	24.6±0.4	23.8±0.5
株高 Plant height /cm	102.1±2.6	80.8±1.8^**	剑叶长 Flag leaf length/cm	24.5±1.4	18.8±1.2^**
穗长 Panicle length /cm	17.4±0.8	13.0±1.2^**	剑叶宽 Flag leaf width/cm	1.8±0.3	1.6±0.2^**
有效穗数 Effective panicle	9.2±1.3	6.5±0.7^*	粒长 Grain length/mm	7.6±0.4	7.5±0.6
每穗粒数 Grains per panicle	142.3±6.5	115.4±7.6^**	粒宽 Grain width /mm	4.4±0.2	4.3±0.1
结实率 Seed setting rate /%	90.6±2.5	82.7±4.3^*	粒厚 Grain thickness /mm	2.1±0.2	2.1±0.2

图2 野生型（WT）和突变体lmm7抽穗期光合色素含量 A―抽穗期野生型(WT)和突变体lmm7的剑叶光合色素含量;B―倒2叶光合色素含量;C―倒3叶光合色素含量。*在0.05水平上差异显著;**在0.01水平上差异显著。

Fig. 2. Photosynthetic pigments contents of the wild type and the lmm7 mutant at heading stage. A, Photosynthetic pigment contents of flag leaf of the wild type and lmm7 mutant at the heading stage; B, Photosynthetic pigment contents of the second leaf from top; C, Photosynthetic pigment contents of the third leaf from top; *,** significantly different at 0.05 and 0.01 levels, respectively.

图3 成熟期野生型（WT）和突变体lmm7的叶肉细胞超微结构 A―成熟期WT剑叶叶片;B~D―WT剑叶细胞超微结构;E―成熟期突变体lmm7剑叶叶片;F~H―突变体lmm7剑叶细胞超微结构;Chl―叶绿体;Os―嗜锇小体;TM―类囊体膜。标尺为20 μm (B, F), 6 μm (C, G), 3 μm (D, H)。

Fig. 3. Ultrastructure of mesophyll cells in the wild type (WT) and the lmm7 mutant at mature period. A, Flag leaf of WT at mature period; B-D, Ultrastructure of the blade of WT; E, Flag leaf of lmm7 mutant at mature period; F-G, Ultrastructure of the blade of lmm7 mutant; Chl, Chloroplast; Os, Osmiophilic granule; TM, Thylakoid membranes. Bars: 20 μm (B, F), 6 μm (C, G), 3 μm (D, H).

图4 野生型和突变体lmm7的遮光处理及组织化学分析 A―遮光处理7 d后野生型(WT)和突变体lmm7的叶片;B―野生型(WT)和突变体lmm7叶片的台盼蓝染色观察;C―野生型(WT)和突变体lmm7叶片的DAB染色观察。在A、B、C中，比例尺均为1 cm。

Fig. 4. Shading treatment and histochemical analysis of the wild type and lmm7 mutant. A, Effects of shading on the wild type(WT) and the lmm7 mutant leaves 7 days after shading; Leaves of WT and the lmm7 mutant stained by trypan blue(B) and DAB(C). Bars =1 cm in A, B and C.

图5 野生型(WT)和突变体lmm7人工接种鉴定条件下水稻黑条矮缩病和稻瘟病抗性评价 A―人工接种鉴定条件下，野生型（WT）和突变体lmm7的水稻黑条矮缩病平均发病率;B―人工接种鉴定条件下，野生型（WT）和突变体lmm7的稻瘟病病情指数。*P < 0.05(t检验)。

Fig. 5. Resistance evaluation of wild type (WT) and lmm7 mutant to rice black-streaked dwarf virus disease (RBSDVD) and rice blast under artificial inoculation condition. A, Disease incidence of wild type (WT) and lmm7 mutant to rice black-streaked dwarf virus disease (RBSDVD) by the artificial inoculation. B, Disease index of wild type (WT) and lmm7 to rice blast by the artificial inoculation. *P < 0.05 by Student’s t-test.

图6 lmm7基因在水稻第7染色体上的分子定位

Fig. 6. Molecular mapping of lmm7 on chromosome 7.

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水稻类病斑突变体lmm7的鉴定与基因定位

Phenotypic Identification and Gene Mapping of a Lesion Mimic Mutant lmm7 in Rice

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