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作物学报 ›› 2018, Vol. 44 ›› Issue (03): 332-342.doi: 10.3724/SP.J.1006.2018.00332

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水稻类病斑突变体spl34的鉴定与基因精细定位

刘宝玉(), 刘军化, 杜丹, 闫萌, 郑丽媛, 吴雪, 桑贤春, 张长伟*()   

  1. 西南大学水稻研究所 / 转基因植物与安全控制重庆市重点实验室, 重庆400716
  • 收稿日期:2017-05-22 接受日期:2017-11-21 出版日期:2018-03-12 网络出版日期:2017-12-04
  • 通讯作者: 张长伟
  • 作者简介:

    673325435@qq.com

  • 基金资助:
    本研究由国家转基因生物新品种培育重大专项(2016ZX08001002-002)资助

Identification and Gene Mapping of a Lesion Mimic Mutant spl34 in Rice (Oryza sativa L.)

Bao-Yu LIU(), Jun-Hua LIU, Dan DU, Meng YAN, Li-Yuan ZHENG, Xue WU, Xian-Chun SANG, Chang-Wei ZHANG*()   

  1. Rice Research Institute of Southwest University / Chongqing Key Laboratory of Application and Safety Control of Genetically Modified Crop, Chongqing 400716, China
  • Received:2017-05-22 Accepted:2017-11-21 Published:2018-03-12 Published online:2017-12-04
  • Contact: Chang-Wei ZHANG
  • Supported by:
    This study was supported by the National Major Project for Developing New GM Crops (2016ZX08001002-002).

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摘要：

利用化学诱变剂EMS处理籼型水稻恢复系“缙恢10号”, 从其后代中筛选到1个遗传稳定的类病斑突变体spl34。该突变体于分蘖后期在下部叶片的叶鞘上开始出现褐色的类病斑, 随后沿着中脉扩散至整个叶片, 成熟期扩散至整个植株。相比于野生型, 该突变体的株高显著变矮, 穗长显著变短, 穗粒数、结实率和千粒重极显著降低。遮光试验和组织化学分析表明, 突变体类病斑的形成受光诱导, 在类病斑形成部位发生大量过氧化氢沉积和细胞程序性死亡。荧光显微镜观察发现, 在紫外光照射下突变体产生的荧光较野生型弱。与野生型相比, 突变体spl34的H2O2和O2-含量较高, 而CAT、POD和T-SOD等保护酶的活性显著降低; 稻瘟病抗性无明显差异或略显降低。遗传分析表明, 突变体spl34的表型受1对隐性核基因控制。基因定位结果表明, 该基因定位于第4染色体的LR49和LR52两个分子标记之间, 物理距离为200 kb。测序分析发现该区间内的候选基因LOC_Os04g56480的第3449位碱基发生突变(G3449T), 导致色氨酸替换为半胱氨酸。qRT-PCR结果表明该基因在突变体内表达量降低, 而部分病程相关基因的表达量则升高。

关键词: 水稻, 类病斑突变体, spl34, 基因, 精细定位

Abstract:

A mutant spotted leaf 34 (spl34) was screened from the progeny of indica restorer line Jinhui 10 treated with ethyl methane sulfonate (EMS). Brown lesions in spl34 exhibit on the sheath of lower leaves at the late tillering stage, then spred from the midrib to entire leaf and finally throughout the whole plant at maturity stage. Compared with the wild type, the plant height, ear length, grain number per panicle, seed setting rate and thousand-grain weight as well as the activities of protective enzymes (CAT, POD, and T-SOD) were all significantly decreased while the content of reactive oxygen species (ROS) increased in spl34. The shading assay showed that the formation of lesions in spl34 was induced by light. Histochemical analysis showed that spl34 had excessive hydrogen peroxide (H2O2) deposition and programmed cell death in the position of lesions. In addition, the chlorophyll fluorescence was weaker in spl34 than in the wild type under the fluorescence microscopy. There was no significant difference in blast resistance between spl34 and the wild type. Genetic analysis suggested that the phenotype of spl34 was controlled by a single recessive nuclear gene, which was mapped between InDel markers LR49 and LR52 on chromosome 4 with an interval of 200 kb. Sequencing analysis revealed that a single base substitution (G to T) occurred at 3449 bp in the DNA sequence of LOC_Os04g56480, resulting in an amino acid change from tryptophane to cysteine. The qRT-PCR results showed that the transcriptional level of LOC_Os04g56480 was down-regulated in spl34, while that of some pathogenesis-related genes was highly up-regulated when compared with the wild type.

Key words: rice, lesion mimic mutant, spl34, gene, fine mapping

图1

分蘖期、成熟期野生型(WT)和突变体spl34的表型 A: 分蘖期野生型(WT)和突变体spl34植株; B: 成熟期野生型(WT)和突变体spl34植株; C: 成熟期野生型(WT)和突变体spl34的叶片。"

表1

野生型(WT)和spl34的主要农艺性状"

材料
Material		 株高
Plant height
(cm)		 有效穗数
Effective panicle		 穗长
Panicle length (cm)		 每穗粒数
Grain number per panicle		 结实率
Seed-setting rate (%)		 千粒重
1000-grain weight (g)
WT 107.12±3.35 8.60±1.20 25.26±1.40 180.90±12.48 78.41±4.45 25.57±0.81
Spl34 100.32±3.59* 8.80±0.82 22.34±1.22* 135.42±8.66** 53.65±3.96** 22.68±0.30**

图2

遮光对野生型和突变体spl34叶片的影响 A: 野生型遮光处理后; B: 野生型遮光处理后复光1周; C: 突变体spl34遮光后; D: 突变体spl34遮光处理后复光1周后。"

图3

野生型(WT)和突变体spl34抽穗期光合色素含量*在0.05水平上差异显著; **在0.01水平上差异显著。A~C: 抽穗期野生型(WT)和突变体spl34的倒一叶(A)、倒二叶(B)和倒三叶(C)光合色素含量。"

图4

野生型和突变体spl34叶片的自发荧光 A, B: 野生型在自然光和紫外光下叶片横切显微结构; C, D: 突变体spl34在自然光和紫外光下叶片横切显微结构; 标尺: 100 μm。白色箭头所指部位为类病斑形成部位, 黄色箭头所指部位为未形成类病斑部位。"

图5

野生型(WT)和突变体spl34的组织化学分析 A, B: 野生型(WT)和突变体spl34叶片的台盼蓝染色; C, D: 野生型(WT)和突变体spl34叶片的DAB染色。"

图6

抽穗期野生型(WT)和突变体spl34的生理指标*在0.05水平上差异显著; **在0.01水平上差异显著。"

表2

野生型(WT)与突变体spl34对稻瘟病菌的抗谱"

材料
Material		 对稻瘟病各生理群的抗病频率
Resistance frequency to each physiological group		 总群抗病频率
Resistance frequency to total
population
ZA ZB ZC ZD ZE ZG
WT 39.39±5.25 74.75±1.76 0 100 100 100 70.37±1.85
spl34 45.45±5.45 72.04±4.10 0 100 100 100 70.37±3.20

表3

野生型(WT)与突变体spl34的稻瘟病病情指标"

材料
Material		 苗瘟病情指数
Disease indexes of leaf blast
at seedling stage		 叶瘟病情指数
Disease indexes of leaf blast
at tillering stage		 穗颈瘟发病率
Disease incidence of neck blast
WT 32.00±2.23 71.43±3.31 3.94±0.61
spl34 36.56±2.01 77.14±2.22 4.19±0.52

表4

新开发的InDel标记"

标记
Marker		 正向引物
Forward primer (5°-3°)		 反向引物
Reverse primer (5°-3°)
LR7 TTCCTACACGGTCGAAGATCAT TCAAGTACCGGATTGAGTTTGAG
LR14 GAGATTATCCTGCGGTCTTAATC TTGATGATACGATGACCAAGTTC
LR25 ATATTCACATGCGCCGTTTG CTCCAACAGATCCATCTCAACC
LR31 GATCCAATGCGATGCTACTCC GCGATTAACTGGAGACATCACG
LR33 TTCAATGTGCTGCATTTAGAAAT AAAGATGGTTCAAGTCTTGAGGA
LR49 GCCAAACAGATTGGTTAGCG TTAGCGTAGCTGGCATAACATC
LR52 TTGATCTCGACTTCGATCACTAG GATGTCTGGACGTACACACACG

图7

突变体spl34的基因定位"

图8

目的基因spl34相对表达量分析"

图9

病程相关基因和稻瘟病抗性基因的表达分析病程相关基因为PAL、PR1a、PR1b、POX22.3、PR5、NPR1和POC1; 稻瘟病抗性基因为Pi-d3、Pish、Pi9、Pit、Pi5和Pia。"

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