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作物学报 ›› 2022, Vol. 48 ›› Issue (6): 1389-1400.doi: 10.3724/SP.J.1006.2022.12035

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水稻早衰突变体esl-H5的表型鉴定与基因定位

郑崇珂1(), 周冠华1, 牛淑琳1,2, 和亚男1, 孙伟1, 谢先芝1,*()   

  1. 1山东省水稻研究所 / 山东省农业科学院, 山东济南 250100
    2山东师范大学生命科学学院, 山东济南 250014
  • 收稿日期:2021-05-27 接受日期:2021-10-19 出版日期:2022-06-12 网络出版日期:2021-12-05
  • 通讯作者: 谢先芝
  • 作者简介:E-mail: zhengck1983@163.com
  • 基金资助:
    中国科学院(A类)战略性先导科技专项(XDA24030101-6)

Phenotypic characterization and gene mapping of an early senescence leaf H5(esl-H5) mutant in rice (Oryza sativa L.)

ZHENG Chong-Ke1(), ZHOU Guan-Hua1, NIU Shu-Lin1,2, HE Ya-Nan1, SUN wei1, XIE Xian-Zhi1,*()   

  1. 1Shandong Rice Research Institute / Shandong Academy of Agricultural Sciences, Jinan 250100 Shandong, China
    2College of Life Sciences, Shandong Normal University, Jinan 250014, Shandong, China
  • Received:2021-05-27 Accepted:2021-10-19 Published:2022-06-12 Published online:2021-12-05
  • Contact: XIE Xian-Zhi
  • Supported by:
    Fund：The study was supported by the Strategic Priority Research Program of Chinese Academy of Sciences(XDA24030101-6)

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摘要：

在甲基磺酸乙酯(EMS)诱变粳稻淮稻5号的突变体库中, 筛选到一个稳定遗传的叶片黄化早衰突变体esl-H5 (early senescence leaf H5)。该突变体幼苗期表型正常, 播种后50 d开始出现下部叶片黄化早衰表型。与野生型相比, esl-H5突变体抽穗期延迟, 株高、穗长、穗粒数、有效分蘖数、千粒重等均显著降低, 叶绿素含量显著减少。遗传分析表明该突变受一对隐性基因调控。分子标记定位结果显示, 该突变基因定位于1号染色体。通过MutMap分析发现编码胼胝质合成酶基因Os01g0533000的最后一个外显子内有一个碱基G变成了A, 这导致翻译提前终止。进化树分析结果显示, ESL-H5与拟南芥AtGSL7 (Glucan Synthase-Like 7)同源性最高。糖含量测定表明esl-H5突变体中可溶性糖和淀粉含量增高, 推测ESL-H5功能缺失后影响了光合产物的转运, 导致叶片中糖含量显著增高, 进而引起叶片衰老。qRT-PCR结果显示, esl-H5突变体中抗病相关基因PR1a、PR1b、PR2、PR4、PR5和PR10表达量均高于野生型, 这与突变体的白叶枯病抗性明显提高一致。上述研究结果为研究糖信号调控水稻衰老和抗病性奠定了基础。

关键词: 水稻, 早衰, 基因定位, 胼胝质合成酶

Abstract:

A stable mutant esl-H5 (early senescence leaf H5) was identified from the mutant library of japonica rice Huaidao 5 population induced by ethyl methane sulfonate (EMS) treatment. The mutant was normal at seedling stage. However, the lower leaves in esl-H5 mutant displayed premature senescence at about 50 days after sowing. Compared with the wild type (WT), the heading date of the esl-H5 mutant was delayed, while agronomical traits including plant height, panicle length, grain number per panicle, effective tiller numbers, and 1000-grain weight were significantly reduced. Moreover, chlorophyll content was also decreased in esl-H5 mutant. Genetic analysis indicated that the early senescence trait in esl-H5 mutant was controlled by a single recessive gene. ESL-H5 gene was localized on chromosome 1 using molecular marker. MutMap analysis further revealed that one nucleotide G to A replace occurred in the last exon of Os01g0533000 gene which encodes callose synthase. The G to A replace in the ESL-H5 introduced a premature stop codon. Phylogenetic analysis showed that ESL-H5 was homology with Arabidopsis AtGSL7 (Glucan Synthase-Like 7). At tillering stage, the contents of soluble sugar and starch were significantly increased in the leaves of the esl-H5 mutant compared with those of the WT. These results implied that the mutation of ESL-H5 affected the transport of photosynthetic products, resulting in premature leaf senescence phenotypes. The qRT-PCR analysis revealed that the expression levels of disease resistance-related genes PR1a, PR1b, PR2, PR4, PR5, and PR10 in esl-H5 mutant were higher than those in WT, which was consistent with the observation that esl-H5 mutant improved bacterial blight resistance. The present results lay the foundation for studying the roles of sugar signal in regulating rice senescence and disease resistance.

Key words: rice (Oryza sativa L.), early senescence, gene mapping, callose synthase

表1

qRT-PCR引物序列"

引物名称
Primer name		 正向引物序列
Forward sequence (5'-3')		 反向引物序列
Reverse sequence (5'-3')
OseEF-1α TTTCACTCTTGGTGTGAAGCAGAT GACTTCCTTCACGATTTCATCGTAA
OsESL-H5 GATTGGGCGACAGAAGTTTG CAGCCATCACAGAGACAAAAC
qPR1a TTCATCACCTGCAACTACTCG TGCATAAACACGTAGCATAGCAT
qPR1b GTGTGCGGGCACTACACG CGGCTTATAGTTGCATGTGA
qPR2 ACAACTCGGAGAAGCATCAG GAAACATAATCCTCGCCAAAGC
qPR4 TGGGACCTGAACAAAGTGAG GGATACACTTGCCACACGAG
qPR5 ATCGACGGCTACAACGTC GTGTCTTGGTGTTGTCTTCG
qPR10 CACCATCTACACCATGAAGC AGCACATCCGACTTTAGGAC
qOsWRKY23 TCCAGTTCCTCTCCCAGTTCTAA CACATTGTTCTCCTTTTCTTCCC
qOsWRKY72 CACCACAAATCACATCTACTCCG GCTGAAGGGAAGAGAGGTGAG
qOsSGR AGGGGTGGTACAACAAGCTG ATCCTGGATTTGGCAAGAAC
qOsNAP CAAGAAGCCGAACGGTTC GCTCCTTGCGGAAGATGTAG
qOsh36 GAACTACAGGAAGGGTCGGT GTTAGAGTGGAGCAGCAT
qOsl2 GCAGACAACAAATCGCCAAAT AGTATCCTGGTGCCAGTTCC
qOsl30 AACCTTTTTCTTGGAGATGATACGA TCTCCAGCAACTCTAACCAGCAT
qOsl85 TGGGCAAAGGAGTTACTGAA CTTGAACTGTAGGGGCTTGCTT
qOsl43 TGTGACAAGTGCTAATAATACATACGA ATCCTGGATTTGGCAAGAAC
qrbcS TCCGCTGAGTTTTGGCTATTT GGACTTGAGCCCTGGAAGG
qlhcA GTCTGTGGTTTGACCCGCT GAGCCGCCCGTTCTTGA
qlhcB GCTCAAGGTGAAGGAGATCAAGA ATGCGTTGTTGTTGACGGG
qrbcL CTTGGCAGCATTCCGAGTAA ACAACGGGCTCGATGTGATA
qPsaA GCGAGCAAATAAAACACCTTTC GTACCAGCTTAACGTGGGGAG
qPsbA CCCTCATTAGCAGATTCGTTTT ATGATTGTATTCCAGGCAGAGC
qpetD TAATGGTTTCTGTGCCGACG CCTAAAGTTAAGGATTTTTCAATGGG
qndhA CAAAGCGATATCCCAAAGGAAT CCGCATCGATACAACAACGTAT
qatpA GATCTCTCCAAACAGGCACAAG CGGCTCTTTCTAAAAGGCGTG

图1

esl-H5突变体表型 A: 播种10 d表型, 标尺为1 cm; B: 植株分蘖期表型, 标尺为5 cm; C: 植株灌浆期表型, 标尺为5 cm; D: 抽穗期上部三叶表型, 标尺为5 cm; E: 成熟期穗子表型, 标尺为1 cm; F和G粒长和粒宽表型, 标尺为1 cm。"

表2

esl-H5突变体与野生型农艺性状比较"

农艺性状
Agronomic trait		 esl-H5 野生型
Wild type
抽穗期 Heading date 124.20 ± 0.28** 120.80 ± 0.23
株高 Plant height (cm) 60.80 ± 0.42** 87.95 ± 0.62
分蘖数 Tiller number 5.75 ± 0.32** 11.65 ± 0.42
穗长Main panicle length (cm) 15.04 ± 0.23** 15.23 ± 0.21
每穗实粒数 Grain number per panicle 64.50 ± 2.65** 158.60 ± 4.80
一级枝梗数No. of primary branch 12.28 ± 0.25** 13.00 ± 0.37
二级枝梗数No. of secondary branch 20.08 ± 0.86** 27.00 ± 1.23
粒长 Grain length (mm) 7.82 ± 0.25 7.84 ± 0.21
粒宽Grain width (mm) 3.76 ± 0.17** 3.88 ± 0.11
千粒重 1000-grain weight (g) 20.90 ± 0.01** 27.20 ± 0.01

图2

衰老和光合作用相关基因在野生型和esl-H5突变体中的表达分析 A: 衰老相关基因在野生型和突变体中的相对表达水平; B: 光合作用相关基因在野生型和突变体中的相对表达水平。"

图3

esl-H5突变体和野生型苗期和分蘖期叶绿素含量 A: 播种40 d最新展开叶叶绿素含量; B: 播种40 d倒二叶叶绿素含量; C: 播种60 d最新展开叶叶绿素含量; D: 播种60 d倒二叶叶绿素含量。**: P < 0.01; *: P < 0.05。"

表3

3个F2分离群体的分离比"

杂交组合
Cross combinations		 F1表型 F1 phenotype F2表型 F2 phenotype
野生型
Wild type		 突变型
Mutant		 野生型
Wild type		 突变型
Mutant		 χ2 (3:1)测验
χ2 (3:1) text
WT/esl-H5 15 0 109 35 0.01
NPB/esl-H5 23 0 672 217 0.14
9311/esl-H5 19 0 105 31 0.24

表4

多态性标记信息"

标记名称
Marker name		 引物序列
Primer sequence (5'-3')		 使用的酶
Enzymes used
ID1-9F GTACGCAAGCGATCAAAC
ID1-9R AGTCCCGTCTAAAATTCACA
ID1-24F ATACTGGGTGTGCTCATTCT
ID1-24R TTTGTTTATTCTTGTCCGTTT
dSNP1F TGATGAAGGTAAGATTAGAGGATGAAGGAATTC EcoR І
dSNP1F TCAGTTAGAAATTACTCCCT
dSNP2F AGGGATCCGAATAGGAGTCCTACGGGCCTTGGGTACC Kpn І
dSNP2R GCACCGCAAACCAGCGTTGCCT
dSNP3F CCTGTTGTTTGTTGTTTCTTCCATCTAGA Xba І
dSNP3R GCACCAACCAGTAGGCATGA

图4

ESL-H5基因的定位和克隆 A: ESL-H5基因的精细定位(横线上方的标记表示精细定位中用的InDel或SNP标记, 下方的数字表示该标记下的重组交换株数); B: SNP3所在基因Os01g0533000 基因组G→A的改变; C: SNP3转化为dCAPS标记; D: SNP3转化为dCAPS标记后检测早衰表型单株; E: Os01g0533000在野生型和突变体中的相对表达量。"

图5

ESL-H5蛋白质结构预测和进化树分析 A: 野生型(ESL-H5)和突变体(esl-H5)蛋白质结构预测, Vta1、FKS1和Glucan synthase指蛋白质结构域; B: ESL-H5和esl-H5蛋白质的跨膜区预测; C: 拟南芥和水稻胼胝质合成酶进化树分析; D: ESL-H5与AtGSL7氨基酸序列比较。"

图6

ESL-H5基因表达模式分析"

图7

分蘖初期野生型和突变体叶片中淀粉和可溶性糖含量 A: 6:00可溶性糖含量; B: 18:00可溶性糖含量; C: 6:00淀粉含量; D: 18:00淀粉含量。*: P < 0.05; **: P < 0.01。"

图8

分蘖期野生型和突变体白叶枯抗性鉴定 A: 白叶枯病抗性表型; B: 病斑长度统计; C病程相关蛋白表达量分析。**: P < 0.01。"

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