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作物学报 ›› 2023, Vol. 49 ›› Issue (8): 2077-2087.doi: 10.3724/SP.J.1006.2023.23062

所属专题： 玉米：遗传育种·种质资源·分子遗传学

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

一个新的玉米细胞核雄性不育突变体ms6的鉴定与基因定位

王兴荣1(), 张彦军1, 涂奇奇2, 龚佃明2,*(), 邱法展2,*()   

  1. 1 甘肃省农业科学院作物研究所, 甘肃 兰州 730070
    2 湖北洪山实验室, 湖北 武汉 430070
  • 收稿日期:2022-09-23 接受日期:2023-02-21 出版日期:2023-08-12 网络出版日期:2023-03-10
  • 通讯作者: 龚佃明,邱法展
  • 作者简介:E-mail: wxr_0618@163.com
  • 基金资助:
    武汉市生物育种关键技术及新品种培育重大专项(2022021302024852);甘肃省农业科学院科研条件建设及成果转化项目(2021GAAS04);甘肃省省级重点人才项目(农作物种质资源保护利用与人才培养)和国家自然科学基金项目(31960405)

Identification and gene localization of a novel maize nuclear male sterility mutant ms6

WANG Xing-Rong1(), ZHANG Yan-Jun1, TU Qi-Qi2, GONG Dian-Ming2,*(), QIU Fa-Zhan2,*()   

  1. 1 Crops Research Institute, Gansu Academy of Agricultural Sciences, Lanzhou 730070, Gansu, China
    2 Hubei Hongshan Laboratory, Wuhan 430070, Hubei, China
  • Received:2022-09-23 Accepted:2023-02-21 Published:2023-08-12 Published online:2023-03-10
  • Contact: GONG Dian-Ming,QIU Fa-Zhan
  • Supported by:
    Wuhan Major Project of Key Technologies in Biological Breeding and New Variety Cultivation(2022021302024852);Research Condition Construction and Achievement Transformation Project of Gansu Academy of Agricultural Sciences(2021GAAS04);Key Talent Project of Gansu Province(Crop Germplasm Resources Protection and Utilization Talent Training). and National Natural Science Foundation of China(31960405)

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摘要：

玉米雄性不育基因的定位、克隆和功能机理研究, 不仅能够加深我们对玉米雄花生长发育的分子调控机理的认识, 而且能够有效推动雄性不育技术体系的发展及在玉米育种和种子生产中的运用。本研究以玉米自交系Mo17为野生型背景材料, 经EMS诱变获得了一个玉米雄性不育突变体, 命名为ms6 (male sterile 6)。表型鉴定结果表明, ms6突变体植株能够正常抽雄, 但雄花颖壳不能正常开裂和散粉, 花粉粒干瘪, 表现为无花粉型不育。同时, ms6与Mo17野生型(wild type, WT)在株型、穗部性状以及籽粒粒形等相关性状上无显著差异, 说明该基因突变后, 仅影响植株的育性, 而不影响其他农艺性状。细胞学观察显示, ms6不育突变体的小孢子发育晚期出现异常, 表现为绒毡层细胞提前降解, 小孢子不能进行有丝分裂并逐渐裂解。扫描电镜观察表明, ms6花药外壁皱缩, 内壁无完整的花粉粒, 无乌氏小体的存在。遗传学分析表明, ms6突变性状受1对隐性核基因控制。以ms6 × B73 F2遗传定位群体, 利用全基因组约200对多态性SSR分子标记, 结合表型与基因型连锁分析, 将ms6初定位于玉米6号染色体C6-19与C6-30两个标记之间, 进一步利用区间内10对新开发的多态性标记, 最终将ms6定位在分子标记M13~M14之间约480 kb的区间范围内。转录组测序结合qRT-PCR试验验证结果, 初步将Zm00001d035201确定为ms6的关键候选基因。Zm00001d035201基因编码一个酸性核糖体蛋白。本研究结果为ms6后续基因功能的研究打下了坚实的基础。同时ms6作为一个新的核不育突变体也为将来玉米新型核不育基因的生产应用提供了重要材料支持。

关键词: 玉米, 雄性不育, ms6, 基因定位, 候选基因鉴定

Abstract:

The localization, cloning, and functional mechanism research of male sterility genes can not only deepen our understanding of the molecular regulation mechanism of male flower growth and development, but also effectively promote the development of male sterility technology system and its application in maize breeding and seed production in maize. In this study, maize inbred line Mo17 was used as the wild-type background material, and a maize male sterile mutant, named male sterile 6 (ms6), was obtained by EMS mutagenesis. Phenotypic identification of ms6 mutant plants showed that they could perform normal tasseling, but the male flower glume could not dehiscence and powder could not disperse normally, and the pollen grains were dry, indicating pollen free sterility. Meanwhile, there were no significant differences between ms6 and Mo17 wild type (WT) in plant architecture, ear-related traits, kernel size, and other related traits, demonstrating that the ms6 gene only affected plant fertility, but did not affect other agronomic traits. Cytological observation revealed that the microspore of ms6 mutant was abnormal at the late stage of microspore development, which showed that tapetal cells were degenerated in advance, and the microspore could not undergo mitosis and gradually split. Scanning electron microscopy demonstrated that the anther epidermis of mutant ms6 were shrunk, and there was no intact pollen or ubisch on the inner surface of anther locule. Genetic analysis revealed that the ms6 phenotype was controlled by a pair of recessive nuclear genes. Combined with phenotypic and genotypic linkage analysis, ms6 was initially located between C6-19 and C6-30 on maize chromosome 6 by using the ms6 × B73 F2 genetic mapping population and about 200 pairs of polymorphic SSR markers in the whole genome, and 10 pairs of newly developed polymorphic markers in the interval were further used. Finally, ms6 was located within the interval of about 480 kb between M13-M14. Gene mapping located the mutation site into a 480 kb interval between M13 and M14. Zm00001d035201 was preliminarily identified as the key candidate gene of ms6 by RNA-Seq combined with qRT-PCR. Zm00001d035201 encoded an acid ribosomal protein. This study provides a solid foundation for further studies on the function of ms6 gene. Meanwhile, as a novel male sterile mutant, ms6 also provides the important material support for the production and application of new maize genetic sterile genes in the future.

Key words: maize, male sterile, ms6, gene mapping, identification of candidate genes

图1

野生型WT和ms6突变体的表型鉴定 A: 野生型WT和ms6突变体散粉期的植株雄穗。B: 野生型WT和ms6突变体的花药; 标尺为2 mm。C: I2-KI染色后, 野生型的花粉粒形态; D: I2-KI染色后, 突变体ms6的花粉粒形态; 标尺为0.1 mm。"

图2

ms6突变体和野生型农艺性状调查 ms6突变体和野生型各种农艺性状比较分析, 其中HKW指百粒重, EW指穗重, KRN指穗行数, KL指粒长, KW指粒重, PH指株高, EH指穗位高, SBN指雄穗分枝数。所有数据均为平均值±SD, 显著性通过t检验估计, NS代表无显著性差异。"

图3

野生型和ms6花药扫描电镜 A: 野生型花药的花粉粒; B: ms6花药干瘪的花粉粒; C和D分别为野生型和突变体花药外壁, E和F分别为野生型和突变体花药内壁。Ub: 乌氏体; 放大倍数已在图中标注。"

图4

野生型和ms6突变体不同发育时期花药的石蜡切片观察 A~F: 野生型花药细胞学切片; G~L: ms6突变体花药细胞学切片; A和G为孢原细胞时期, B和H为二分体时期, C和I为四分体时期, D和J为单核小孢子初期, E和K为单核小孢子中后期, F和L为三核细胞早期。E: 表皮; En: 内皮层; ML: 中层; T: 绒毡层; Dy: 二分体; Msp: 小孢子; Bp: 二孢花粉; Mp: 成熟花粉。标尺为50 μm。"

表1

ms6突变位点的遗传分析"

群体
Population		 总株数
Total number		 可育株
Fertile plants		 不育株
Sterile plants		 可育/不育(3:1)
Fertility/sterile (3:1)		 χ (3:1) χ20.05
ms6×B73 396 300 96 3.13 0.12 3.84
ms6×Mo17 159 116 43 2.70 0.35

表2

本研究所用的引物"

引物
Primer name		 序列
Primer sequence (5°-3°)		 引物
Primer name		 序列
Primer sequence (5°-3°)
IN34-F AGCACGGAGAAATGGAGTTG M13-F GAGGCCAAGCACAATGTATG
IN34-R AACAGAAAGCTTGGCGTCAT M13-R TTTCAGGTGGAACCAGCAAT
MY8860-F AATGGCAGGAGCACTACACC C6-11A-F CAACCAGGAGAAGCTGGCTA
MY8860-R GGTCTGCCTGATGCTCTTGT C6-11A-R GAAGAGCGCGTAGAAGATGC
M10-F GTGGAGCCCAGACACATTTT M14-F ACTGCAGCGTGTGAAAACAA
M10-R TTTGTTTATGCTTGGCCGTA M14-R GCGGGAGACTCATACCTGAA
M11-F ATGACCACTCCTGGGTTTTG M17-F GAGTTCTAAATAAGGGGGATGGA
M11-R CTGTAGCCAAGAAGCCTTGC M17-R TTGGTTTCTGTGATGCATTGA
M12-F AATTGGAGGAACCAGCTCAA C6-20-F ATTCGATCTAGGGTTTGGGTTCAG
M12-R ATTTGGAGGGCGGTTTTATC C6-20-R GATGCAGTAGCATGCTGGATGTAG

图5

ms6精细定位结果 红色椭圆代表着丝粒。"

表3

候选基因功能注释"

基因ID Gene ID 注释 Gene annotation
Zm00001d035199 WPP domain-associated protein
Zm00001d035200 Putative AMP-dependent synthetize and ligase superfamily protein
Zm00001d035201 60S acidic ribosomal protein P0
AC187088.3_FG002 Guanylate kinase/Guanosine monophosphate kinase
Zm00001d035203 Unknown
Zm00001d035204 Unknown
Zm00001d035206 ATP binding protein
Zm00001d035207 Putative uncharacterized protein
Zm00001d035208 BTB/POZ domain-containing protein
Zm00001d035211 Ca(2)-dependent phospholipid-binding protein (Copine) family

表4

ms6区间内的基因在ms6和WT花药中的表达"

基因ID
Gene ID		 表达差异Expression difference
花药发育时期
Anther development stage		 ms6 野生型
WT		 log2FC P-value 显著性
Significant
Zm00001d035199 四分体时期The tetrad stage 1024 1157 0.18 0.43 NS
双核期The binucleate stage 1069 911 -0.31 0.28 NS
Zm00001d035200 四分体时期The tetrad stage 103 140 0.45 0.38 NS
双核期The binucleate stage 135 75 -0.84 0.26 NS
Zm00001d035201 四分体时期The tetrad stage 9652 8941 -0.11 0.69 NS
双核期The binucleate stage 6690 17205 1.36 0.00018 DOWN
AC187088.3_FG002 四分体时期The tetrad stage NS NS NS NS NS
双核期The binucleate stage NS NS NS NS NS
Zm00001d035203 四分体时期The tetrad stage NS NS NS NS NS
双核期The binucleate stage NS NS NS NS NS
Zm00001d035204 四分体时期The tetrad stage NS NS NS NS NS
双核期The binucleate stage NS NS NS NS NS
Zm00001d035206 四分体时期The tetrad stage 600 943 0.65 0.0035 NS
双核期The binucleate stage 1196 1924 0.69 0.088 NS
Zm00001d035207 四分体时期The tetrad stage 736 819 0.15 0.56 NS
双核期The binucleate stage 763 729 -0.06 0.82 NS
Zm00001d035208 四分体时期The tetrad stage 827 1022 0.31 0.45 NS
双核期The binucleate stage 1184 877 -4.41 0.41 NS
Zm00001d035211 四分体时期The tetrad stage 167 134 -0.31 0.59 NS
双核期The binucleate stage 95 40 -1.06 0.16 DOWN

图6

候选基因qRT-PCR表达量分析 以Ubiquitin作为内参基因, 误差线表示n = 3的标准误。取野生型及突变体双核期花药进行表达量分析。WT: 野生型; ms6: 突变体。所有数据均为平均值 ± SD (n = 3)。**在P < 0.01水平上差异显著, NS: 不显著。"

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