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作物学报 ›› 2023, Vol. 49 ›› Issue (11): 2913-2922.doi: 10.3724/SP.J.1006.2023.33002

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

一个玉米ZmMs7复等位基因突变体的遗传分析与分子鉴定

曹枭雄1,2(), 刘伊凡1,2, 周玉强2, 王婧2, 吴宇锦2, 王红武2, 李坤2, 刘小刚2, 黄长玲2, 刘志芳2, 郭晋杰1,*(), 胡小娇2,*()   

  1. 1河北农业大学农学院 / 国家玉米改良中心河北分中心 / 华北作物改良与调控国家重点实验室, 河北保定 071001
    2中国农业科学院作物科学研究所 / 作物分子育种国家工程研究中心, 北京 100081
  • 收稿日期:2023-01-05 接受日期:2023-04-17 出版日期:2023-11-12 网络出版日期:2023-05-05
  • 通讯作者: 胡小娇, E-mail: huxiaojiao@caas.cn; 郭晋杰, E-mail: guojinjie512@163.com
  • 作者简介:E-mail: caoxx13@163.com
  • 基金资助:
    国家重点研发计划项目(2022YFD1200802);中国农业科学院科技创新工程(CAAS-ZDRW202004);作物分子育种国家工程研究中心和河北省玉米现代种业科技创新团队项目(21326319D)

Genetic analysis and molecular identification of a multiple allele mutant of ZmMs7 gene in maize

CAO Xiao-Xiong1,2(), LIU Yi-Fan1,2, ZHOU Yu-Qiang2, WANG Jing2, WU Yu-Jin2, WANG Hong-Wu2, LI Kun2, LIU Xiao-Gang2, HUANG Chang-Ling2, LIU Zhi-Fang2, GUO Jin-Jie1,*(), HU Xiao-Jiao2,*()   

  1. 1College of Agronomy, Hebei Sub-center of National Maize Improvement Center of China / State Key Laboratory of North China Crop Improvement and Regulation / Hebei Agricultural University, Baoding 071001, Hebei, China
    2Institute of Crop Sciences, Chinese Academy of Agricultural Sciences / National Engineering Research Center of Crop Molecular Breeding, Beijing 100081, China
  • Received:2023-01-05 Accepted:2023-04-17 Published:2023-11-12 Published online:2023-05-05
  • Supported by:
    National Key Research and Development Program of China(2022YFD1200802);Agricultural Science and Technology Innovation Program(CAAS-ZDRW202004);National Engineering Research Center of Crop Molecular Breeding, and the Science and Technology Innovation Team of Maize Modern Seed Industry in Hebei(21326319D)

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摘要：

我们在自然群体中发现了一个玉米雄性不育突变体(male sterile mutant), 命名为ms20s1。该突变体雄花育性彻底丧失, 花药干瘪皱缩, 没有花粉形成。细胞学分析发现, 与野生型相比, ms20s1突变体花药在S11期表现出明显的药室收缩, 绒毡层细胞肿胀, 小孢子破裂的表型, 表明ms20s1突变体绒毡层细胞程序性死亡出现异常, 且花粉败育。遗传分析表明该不育性状受单个隐性核基因控制。为克隆目标基因, 以ms20s1为母本分别与不同自交系杂交构建F2定位群体, 利用靶向测序技术(GBTS)分析群体基因型, 将基因定位于7号染色体124.95~128.47 Mb之间, 进一步精细定位将该区间缩小到0.68 Mb。生物信息学分析发现, 该区间存在一个已知基因ZmMs7。ZmMs7基因编码PHD-finger转录因子, 在绒毡层发育和花粉外壁的形成过程中发挥重要作用。等位测验分析发现ms20s1为ZmMs7基因的等位突变体。基因测序结果表明ms20s1突变体在外显子区存在多处序列变异, 与所报道的ZmMs7基因已知突变体ms7-6007和ms7gl的突变方式不同, 证明ms20s1是一个新的ZmMs7基因等位突变体。突变体ms20s1的发现与鉴定为探讨玉米核雄性不育的分子机制以及育种应用提供了新的材料。

关键词: 玉米, 雄性不育, ms20s1, 基因定位, 等位突变

Abstract:

We identified a maize male sterile mutant in the natural population, designated as ms20s1. The mutant had complete male sterility and lacked pollen grains in the withered anthers. Cytological analysis showed that, compared with the wild type, the ms20s1 mutant exhibited shrinkage locule, swollen tapetum cells, and aborted microspore at S11 stage, indicating that the ms20s1 mutant had abnormal tapetal programmed cell death and complete pollen abortion. Genetic analysis revealed that the male sterility trait was controlled by a single recessive nuclear gene. To clone the target gene, we constructed the F2 populations by crossing ms20s1 with different inbred lines and analyzed the population genotype using genotyping by target sequencing (GBTS) technology. The gene was initially mapped to the 124.95-128.47 Mb region on chromosome 7, and the interval was narrowed down to 0.68 Mb after fine mapping. Bioinformatics analysis indicated that there was one known gene ZmMs7 in this region. The ZmMs7 gene encoded a PHD-finger transcription factor that played an important role in tapetum development and pollen wall formation. Allelism test demonstrated that ms20s1 was an allelic mutant of ZmMs7 gene. Gene sequencing results showed that the ms20s1 mutant had multiple sequence variants in the exon region, which were different from the reported mutants ms7-6007 and ms7gl, confirming ms20s1 was a new allelic mutant of ZmMs7. The discovery and identification of the ms20s1 mutant provide a new material for exploring the molecular mechanism and breeding application of maize genic male sterility.

Key words: maize, male sterility, ms20s1, gene mapping, allelic mutant

表1

本研究采用的PCR引物"

引物名称
Primer name		 正向引物
Forward primer (5°-3°)		 反向引物
Reverse primer (5°-3°)		 用途
Usage
InD124 GCACTGAAGGCTTATTTCGTCG TTTTTGGTCGTTGCTGCTGAT 基因定位
Gene mapping
InD125 AGCAGGAACGAAAAGGCACT GCAAAACAGGACACGCATCA
InD127 CCCTGCTATGACGACTTTTTTTCT ACTCTTGGTTGTCCACCGTGC
InD128 CCCGCTCATTGCTCTGTTG CCAAGCGAGCAGGCACAT
InD130 GCTCTTGATTTGCGAGGTGGT GCTGTGAGTGTGATGCGTGTG
20q CTTGGACACCAAGCACTTCGTC CAGGGTGACCGTCTCGTACG 基因表达分析
Gene expression
Tubulin GTGTCCTGTCCACCCACTCTCT GGAACTCGTTCACATCAACGTTC
Ms7-P1 CAGAACAGAGCAGAGGAACCAT GGATAACCAAACGAAACACGAGCC 基因克隆
Gene cloning
Ms7-P2 GGAGAAACCGTCCAAAGGC AAACCTTCGTGGTAATCTTTGAC
Ms7-P3 GTCTTGGACACCAAGCACTTCGT TGCCATGGCGGCTATAGGAGTT 基因克隆/CAPS鉴定
Gene cloning/CAPS

图1

野生型和突变体ms20s1的表型比较 A: 散粉期野生型(WT)与突变体ms20s1植株的形态比较, 标尺为20 cm; B: 散粉期野生型(WT)与突变体ms20s1的雄穗比较, 标尺为20 cm; C、D: 野生型(C)与突变体ms20s1植株(D)的花药比较, 标尺为1 mm; E、F: 野生型(E)与突变体ms20s1 (F)被I2-KI染色后的花粉粒, 标尺为200 μm。"

表2

野生型和突变体重要农艺性状比较"

农艺性状
Agronomic trait		 野生型
WT		 突变体
ms20s1
株高Plant height (cm) 188.13±3.88 178.83±3.53*
穗位高Ear height (cm) 77.20±6.04 76.50±4.89
雄穗主轴长Total tassel length (cm) 41.01±2.29 38.48±2.21*
雄穗分枝数Tassel branch number 16.00±1.73 19.00±2.00*
穗上1叶长Leaf length-1 (cm) 88.33±3.94 90.65±3.27
穗上2叶长Leaf length-2 (cm) 82.26±3.03 84.32±4.25
穗上3叶长Leaf length-3 (cm) 75.17±3.20 74.75±3.80
穗上1叶宽Leaf width-1 (cm) 8.03±0.41 7.70±0.43
穗上2叶宽 Leaf width-2 (cm) 7.61±0.43 7.43±0.35
穗上3叶宽 Leaf width-3 (cm) 7.22±0.41 6.93±0.47
穗上1叶夹角Leaf angle-1 (°) 30.74±1.54 30.18±1.37
穗上2叶夹角Leaf angle-2 (°) 31.19±1.47 31.27±1.52
穗上3叶夹角Leaf angle-3 (°) 32.13±2.63 32.98±3.43

图2

WT和ms20s1突变体玉米花药(S8a~S11期)的横切面分析 E: 表皮层; En: 内皮层; ML: 中间层; Ta: 绒毡层; Msp: 小孢子; CMsp: 收缩的小孢子; Dy: 二分体; Tds: 四分体。标尺为50 μm。"

表3

F2群体表型分离的卡方检测"

群体
Population		 总株数
Total number of plants		 可育植株
Number of fertile plants		 不育植株
Number of male sterile plants		 期望比
Expected rate		 χ2
F2 (ms20s1×B73) 379 285 94 3:1 0.0009
F2 (ms20s1×Mo17) 181 139 42 3:1 0.2228
F2 (ms20s1×C7-2) 990 769 221 3:1 3.6418

图3

SNP-index全基因组频率分布图"

图4

ms20s1的精细定位和ZmMs7基因的结构示意图 A: ms20s1的精细定位。Marker: 标记名称; N: 群体大小; Recombinant: 重组单株数; B: ZmMs7基因的结构示意图。灰色框代表5°非翻译区; 黑色框代表外显子区, 横线代表内含子区。"

图5

ms7-6007的杂合体与ms20s1突变体等位测验 A: 从左到右分别为ms20s1突变体、杂交后代中的可育植株、杂交后代中的不育植株、ms7-6007突变体的雄穗, 标尺为10 cm; B: 从左到右分别为ms20s1突变体、杂交后代中的可育植株、杂交后代中的不育植株、ms7-6007突变体的花药, 标尺为1 mm; C: 从左到右分别为ms20s1突变体、杂交后代中的可育植株、杂交后代中的不育植株、ms7-6007突变体被I2-KI染色后的花粉粒, 标尺为100 μm。"

表4

ms20s1与ms7-6007突变体等位测验"

父母本基因型
Parental genotype		 表型统计Phenotypic statistics 总株数
Number of the total plants		 χ2(1:1)
可育植株Fertile plants 不育植株Sterile plants
ms20s1×+/ms7-6007 96 100 196 0.0459
ms7-6007×+/ms20s1 151 145 296 0.0844

图6

基因型鉴定 A: CAPS标记PCR产物; B: SnaB I酶切产物, AA、Aa、aa分别代表纯合野生型、杂合野生型、纯合突变体。"

图7

ZmMs7基因在不同组织中的表达量变化"

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