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作物学报 ›› 2024, Vol. 50 ›› Issue (4): 857-870.doi: 10.3724/SP.J.1006.2024.32010

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水稻类病变突变体lms1的表型鉴定与抗病分子机制分析

余瑶1(), 王紫瑶1, 周思睿1, 刘鹏程1, 叶亚峰2, 马伯军1, 刘斌美2,*(), 陈析丰1,*()   

  1. 1浙江师范大学生命科学学院, 浙江金华 321004
    2中国科学院合肥物质科学研究院, 安徽合肥 230031
  • 收稿日期:2023-03-27 接受日期:2023-10-23 出版日期:2024-04-12 网络出版日期:2023-11-23
  • 通讯作者: * 刘斌美, E-mail: liubm@ipp.ac.cnl; 陈析丰, E-mail: xfchen@zjnu.cn
  • 作者简介:E-mail: 2315318441@qq.com
  • 基金资助:
    国家自然科学基金项目(32071987);国家自然科学基金项目(32101697);浙江省自然科学基金项目(LQ22C130005);浙江省自然科学基金项目(LZ23C130004)

Phenotypic identification and disease resistance mechanism analysis of rice lesion mutant lms1

YU Yao1(), WANG Zi-Yao1, ZHOU Si-Rui1, LIU Peng-Cheng1, YE Ya-Feng2, MA Bo-Jun1, LIU Bin-Mei2,*(), CHEN Xi-Feng1,*()   

  1. 1College of Life Science, Zhejiang Normal University, Jinhua 321004, Zhejiang, China
    2Hefei Institutes of Physical Science, Chinese Academy of Science, Hefei 230031, Anhui, China
  • Received:2023-03-27 Accepted:2023-10-23 Published:2024-04-12 Published online:2023-11-23
  • Contact: * E-mail: liubm@ipp.ac.cnl; E-mail: xfchen@zjnu.cn
  • Supported by:
    National Natural Science Foundation of China(32071987);National Natural Science Foundation of China(32101697);Natural Science Foundation of Zhejiang Province(LQ22C130005);Natural Science Foundation of Zhejiang Province(LZ23C130004)

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摘要：

类病变突变体是研究植物细胞死亡和抗病分子机制的重要遗传材料。通过辐射诱变粳稻品种武运粳7号, 获得一个少见的抗病性下降的类病变突变体lms1 (lesion mimic and disease susceptible mutant 1)。与野生型相比, 该突变体叶片自发出现红褐色斑点, 其株高、穗长、每穗粒数以及单株产量降低, 但千粒重增加; 对水稻白叶枯病的抗性显著下降, 组织染色表明突变体的叶片中存在明显的细胞死亡以及活性氧的过量积累。遗传分析表明, lms1突变体的表型受单隐性核基因控制, 利用图位克隆技术将lms1基因精细定位于水稻9号染色体Indel7和Indel8两个分子标记间, 物理距离为62 kb。定位区间内候选基因的PCR扩增测序结果表明, 其中一个编码泛素羧基末端水解酶的OsLMP1 (Lesion Mimic Phenotype 1)基因第1个外显子中插入了一段长度为654 bp的序列, 导致其蛋白翻译提前终止。利用蛋白组学技术分析了lms1突变体与野生型对照的蛋白积累水平, 共鉴定到19个差异蛋白(7个上调、12个下调), 主要参与氧化还原、叶绿素合成、光合作用等代谢途径。上述结果为进一步研究OsLMP1基因的功能及其调控细胞程序性死亡与抗病性的分子机制提供借鉴。

关键词: 水稻, 类病变, lms1, 基因定位, 蛋白组学分析

Abstract:

Lesion mimic mutants are important genetic materials for studying molecular mechanisms of plant cell death and disease resistance. Through radiation mutagenesis of a japonica rice cultivar ‘Wuyunjing 7’, a rare lesion mimic and disease-susceptible mutant lms1 was obtained. Compared to the wild type, the leaves of this mutant spontaneously appeared reddish-brown spots, and its plant height, panicle length, number of grains per panicle, and yield per plant decreased, but the weight of 1000-grain increased. In addition, the resistance of lms1 to rice bacterial blight decreased significantly, and tissue staining showed significant cell death and excessive accumulation of reactive oxygen species in the mutant leaves. Genetic analysis showed that the phenotype of the lms1 mutant was controlled by a single recessive nuclear gene, and the lms1 gene was finely located between two molecular markers, Indel7 and Indel8, on chromosome 9 of rice with a physical distance of 62 kb. PCR amplification and sequencing of candidate genes in the localization interval showed that a 654 bp sequence was inserted into the first exon of the OsLMP1 (Lesion Mimic Phenotype 1) gene, which encodes a ubiquitin carboxyl-terminal hydrolase, resulting in premature termination of protein translation. The protein compositions of lms1 mutants and WT controls were analyzed by a proteomics technology, which identified a total of 19 differentially accumulated proteins (7 upregulated and 12 down-regulated), mainly involved in redox, chlorophyll synthesis, photosynthesis, and other metabolic pathways. The above results provide a reference for further research on the function of the OsLMP1 gene and its molecular mechanism of regulating programmed cell death and disease resistance.

Key words: rice, lesion mimic, lms1, gene mapping, proteomic analysis

表1

用于基因精细定位的分子标记及引物序列"

名称
Name		 正向引物序列
Former primer (5′-3′)		 反向引物序列
Reverse primer (5′-3′)
Indel1 ATGCAAATGACCTCGCTTCC GCCTGCCACTGACTCTGTTC
Indel2 TACAGGGAGATAAGAGGAAG CAAGTCGGAGGAACGAACAA
Indel3 CTTACCTCCGTCTCCAAA GTCATATTAACAGTCCCATT
Indel4 GCTTCACCCGCTTCTTCAC CAGGTTCTCGGCCCAATC
Indel5 AGAGGGTTGAGGGAGGCTGA TTCCATGCGGTCCATAAGTG
Indel6 ATGGCTGGAGGAGTGAATGA CTCGTGAGCGGATTATAGTTG
Indel7 GTATTTCCCGATGACTTCC ACATTTATTGATTGCTCCC
Indel8 AGGCTTAGTGATGGAGTAGAA CATGATGCCATTTGATTTGA

表2

用于目标基因扩增和鉴定的PCR引物"

名称
Name		 正向引物序列
Former primer (5′-3′)		 反向引物序列
Reverse primer (5′-3′)
OsLMP1-1 GGGGATTTCTGTTGGTGTAGG GTTGCATCATCAATAGGGTTTT
OsLMP1-2 GTCATGTGCTTCTGTGCCTAA CATGGACAGTGGGACCTCTTA
OsLMP1-3 AGTATGCCTTCCCATCTGTA AGGCCAACTGGACTCAATCAAA
lms1-Chr.9 AGCAGCAGCAGAAGAAGAAA AGCCCTCTGATTACACTACCAT
lms1-Chr.3 CAGTGATGAACACCTGAGAAC GCCGCAGTAGTTAGATGTTGT

图1

水稻 lms1突变体的类病变表型与抗病性鉴定 A: 苗期植株; B: 成熟期植株; C: 旗叶; D: 叶绿素含量测定; E: 染色前的叶子; F~H: 分别被TB、NBT和DAB染色后的叶子; I: 被Xoo菌株感染的叶片照片; J: Xoo菌株感染叶片病斑长度的统计(用刻度尺测量病斑长度, 每个类型水稻至少测量15片剑叶, 最终取平均值作为其病斑长度)。使用t检验测量显著性差异(*: P < 0.05; ** P < 0.01, 下同), 标尺为2 cm (A, C~G, I~J), 标尺为10 cm (B)。"

图2

水稻lms1突变体与WT的主要农艺性状比较 A~L: 分别为株高、旗叶长、旗叶宽、有效分蘖数、穗长、每穗粒数、结实率、粒长、粒宽、粒厚、千粒重和单株产量; M~O: WT和lms1突变体的10粒种子的长度、宽度和厚度的照片, 标尺为1 cm。"

图3

突变体lms1目标基因的精细定位与候选基因鉴定 A, B: lms1的初步定位和精细定位, “n”代表使用的F2群体中突变型单株的数量, 每个标记物下方的数字表示该标记检测到的重组体数量; C: 62 kb区域内的注释基因以及lms1候选基因的突变位点, 在第一个外显子上的第625 bp位点插入了一个654 bp的片段; D: lms1中目标基因在突变位点的测序结果及与WT的对比, 黑框里为整个插入序列的片段; E: lms1突变体染色体易位模式图, DSB, DNA双链断裂; NHEJ, 非同源末端连接; F: 利用突变位点两侧引物对WT和lms1扩增后的PCR产物电泳结果。M代表DL2000分子标记。"

图4

OsLMP1蛋白的保守结构域与系统发育树分析 A: OsLMP1及其同源蛋白的序列比对, 下画线和框里为保守结构域; B: OsLMP1蛋白的系统发育树, 由MEGA v7.0软件用最大似然法构建, bootstrap=1000。"

表3

差异表达蛋白的基本信息"

蛋白序号a
Protein IDa		 基因序号b
Gene IDb		 描述
Description		 表达量差异倍数
FC		 分数
Score		 趋势(下调/下调)
Trend (down/up)
Q8W250 LOC_Os01g01710 1-deoxy-D-xylulose 5-phosphate reductoisomerase 0 16.768 Down
Q9LGQ6 LOC_Os01g09010 Acyl transferase 9 0 11.008 Down
Q6Z2T6 LOC_Os02g51080 Geranylgeranyl diphosphate reductase 0 57.381 Down
Q75L11 LOC_Os05g02300 Histone H2A.6 0 139.17 Down
A2Y8E0 LOC_Os06g01850 Leaf-type ferredoxin-NADP+ oxidoreductase 0 5.7841 Down
Q69TY4 LOC_Os06g42000 Peroxiredoxin-2E-1 0 4.344 Down
Q53JF7 LOC_Os11g06720 Abscisic stress-ripening protein 5 0 323.31 Down
B6RGY0 LOC_Os01g37510 Peptide deformylase 0.2145 7.3262 Down
Q7XBW5 LOC_Os10g42500 Probable plastid-lipid-associated protein 3 0.2309 27.554 Down
Q71U98 LOC_Os03g27310 Histone H3.3 0.2328 203.14 Down
Q6ENC8 LOC_Os10g21358 50S ribosomal protein L23 0.2462 55.05 Down
Q53RM0 LOC_Os03g36540 Magnesium-chelatase subunit ChlI 0.3060 71.42 Down
P30298 LOC_Os06g09450 Sucrose synthase 2.0059 121.52 Up
Q9LSU0 LOC_Os01g59600 Proteasome subunit alpha type-3 2.5953 11.201 Up
Q7F1U0 LOC_Os07g48020 Peroxidase 3.3684 109.27 Up
Q84YK8 LOC_Os08g39850 Lipoxygenase 4.7947 20.507 Up
Q75IM9 LOC_Os05g03480 Isovaleryl-CoA dehydrogenase 8.1778 12.587 Up
Q6ZCR3 LOC_Os08g13440 Cupin domain containing protein 27468000 6.2905 Up
Q0JG75 LOC_Os01g71190 Photosystem II reaction center PSB28 protein 189290000 323.31 Up

图5

水稻lms1突变体与WT的蛋白组学比较分析 A: lms1突变体和WT之间的差异蛋白(DEP)的GO分析; B: DFP的KEGG富集, X轴表示富集率, Y轴表示KEGG途径, 气泡的大小表示蛋白质的数量, 颜色表示Q值的大小; C: H2O2的含量; D: POD的活性。"

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