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作物学报 ›› 2020, Vol. 46 ›› Issue (9): 1359-1367.doi: 10.3724/SP.J.1006.2020.03005

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

玉米籽粒突变体dek48的表型鉴定与基因定位

石慧敏1(), 蒋成功1,2, 王红武1, 马庆2, 李坤1, 刘志芳1, 吴宇锦1, 李树强1, 胡小娇1,*(), 黄长玲1,*()   

  1. 1 中国农业科学院作物科学研究所作物分子育种国家工程实验室, 北京 100081
    2 安徽农业大学生命科学学院 / 作物抗逆育种与减灾国家地方联合工程实验室, 安徽合肥 230036
  • 收稿日期:2020-01-20 接受日期:2020-04-15 出版日期:2020-09-12 网络出版日期:2020-04-26
  • 通讯作者: 胡小娇,黄长玲
  • 作者简介:E-mail: 3041601854@qq.com
  • 基金资助:
    本研究由国家自然科学基金项目(31500984);中国农业科学院科技创新工程;中央级公益性科研院所基本科研业务费专项资助(Y2019CG15)

Phenotype identification and gene mapping of defective kernel 48 mutant (dek48) in maize

SHI Hui-Min1(), JIANG Cheng-Gong1,2, WANG Hong-Wu1, MA Qing2, LI Kun1, LIU Zhi-Fang1, WU Yu-Jin1, LI Shu-Qiang1, HU Xiao-Jiao1,*(), HUANG Chang-Ling1,*()   

  1. 1 Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, National Engineer Laboratory of Crop Molecular Breeding, Beijing 100081, China
    2 National Engineering Laboratory of Crop Stress Resistance Breeding / School of Life Sciences, Anhui Agricultural University, Hefei 230036, Anhui, China
  • Received:2020-01-20 Accepted:2020-04-15 Published:2020-09-12 Published online:2020-04-26
  • Contact: Xiao-Jiao HU,Chang-Ling HUANG
  • Supported by:
    National Natural Science Foundation of China(31500984);Agricultural Science and Technology Innovation Program at CAAS;Central Public-interest Scientific Institution Basal Research Fund(Y2019CG15)

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摘要：

籽粒是玉米的主要营养储存器官, 也是禾本科植物种子发育研究的模式器官。本研究对玉米自交系郑58进行甲基磺酸乙酯(EMS)诱变, 获得一个稳定遗传的籽粒缺陷突变体, 命名为defective kernel 48 (dek48)。该突变体籽粒皱缩扁小, 百粒重显著降低, 胚和胚乳发育严重缺陷, 不能成苗。在玉米授粉后12 d即可观察到明显的发育缺陷, 表明该突变发生在籽粒发育的早期阶段。扫描电镜观察发现dek48与野生型相比淀粉粒显著变小。石蜡切片显微观察发现dek48淀粉胚乳填充不饱满, 糊粉层细胞发育不规则。遗传学分析表明, 该突变性状受隐性单基因控制。进一步构建F2遗传定位群体, 将该突变体基因精细定位于3号染色体7.39 Mb~7.52 Mb之间。生物信息学分析发现该区间内有6个开放阅读框, 暂未发现与籽粒发育有关的已知基因, 后续将通过测序和基因表达分析进一步确定候选基因。

关键词: 玉米, 籽粒突变, dek48, 基因定位

Abstract:

Maize kernel is not only a main nutrient storage organ, but also a model organ for seed development research of gramineous plants. In this study, a stable defective kernel mutant 48 (dek48) was identified from a library of mutants of the maize inbred line Zheng58 treated with ethylmethane sulfonate (EMS). Compared with wild type, the hundred-kernel weight of the dek48 was decreased greatly due to shrunken appearance and small flat size. Moreover, the dek48 was incapable of growing into a plantlet owning to the severely defective embryo and endosperm. The obvious defective development of the mutant can be observed at 12 days after pollination (DAP), indicating that the mutation occurred at an early stage of kernel development. Microscopic observation by scanning electron microscopy (SEM) revealed that starch granule of the dek48 was significantly smaller than wild type (WT). The observation of the paraffin section demonstrated that the starch granule of dek48 endosperm was partially filled and the aleurone layer cells developed irregularly. Genetic analysis based on kernel form indicated that the mutant trait was controlled by a single recessive gene. Based on genetic F2 population mapping, the gene of the mutant was located between 7.39 Mb-7.52 Mb on chromosome 3. The bioinformation analysis indicated that there were six new open reading frames (ORFs) and unknow genes related to kernel development in this region. Furthermore, candidate gene will be identified through sequencing and gene expression analysis in the future.

Key words: maize, defective kernel mutant, dek48, gene mapping

表1

基因定位引物序列"

引物
Primer		 正向序列
Forward sequence (5′-3′)		 反向序列
Reverse sequence (5′-3′)
In6.38 TTGGCATCTTGTGGTTTTGTG CGAACGGAGTGAAAGTGATAGAAG
In7.29 GCTCAAGAAAGCAATCGGGTAA GTGATGAAGCCAGGATTCGCT
In7.39 TTTCTGGCTCTTTATTTGTGCTG CAAGCGTACATACACCTGTCAGA
In8.29 ATGCAGTTAAGTTGACCTGAATTG TATAGCTAGGAAACCAAGGCGT
In8.53 GCTTTTTCCACCACTGCGAC ATACTCGGGGGCTGCCTTAC
In9.06 GGAAAACACTAAAACACATAACCCT CTAAGGCGTCACTTTGCGG

图1

玉米dek48突变体表型 A: 成熟M4代果穗; B: 野生型籽粒纵切面; C: dek48突变籽粒纵切面; D: 以B73为母本的F2果穗; E: 以Mo17为母本的F2果穗; F: C7-2为母本的F2果穗。"

图2

正常籽粒和突变籽粒的百粒重 **分别表示在0.01水平差异显著。"

图3

WT和dek48籽粒成分测定分析 A: 野生型和dek48的淀粉含量; B: 野生型和dek48的蛋白和油分含量。*, **分别表示野生型与突变体在0.05和0.01水平差异显著。"

图4

WT和dek48胚乳的扫描电镜观察 A: WT扫描电镜观察, bar = 10 μm; B: dek48扫描电镜观察, bar = 10 μm; SG: 淀粉粒; MP: 基质蛋白。"

图5

授粉后不同时期胚和胚乳观察分析 A: 突变型籽粒(dek48)在12、16、20、25、30 DAP胚和胚乳及完整种子的观察分析, bar = 2 mm; B: 野生型籽粒(WT)在12、16、20、25、30 DAP胚和胚乳及完整种子的观察分析, bar = 2 mm。"

图6

WT和dek48籽粒在16DAP时的石蜡切片观察 A: 授粉后16 d WT籽粒的胚, bar = 200 μm; B: 授粉后16 d dek48籽粒的胚, bar = 200 μm; C: 授粉后16 d WT籽粒的糊粉层, bar = 50 μm; D: 授粉后16 d dek48籽粒的糊粉层, bar = 50 μm; AL: 糊粉层。"

表2

野生型与dek48突变籽粒分离比统计分析"

总粒数
Total kernel number		 突变体籽粒/野生型籽粒
dek48/WT		 χ2
M4 515 108/407 1.20
F2 (B73×dek48) 1019 236/783 1.84
F2 (C7-2×dek48) 1067 259/808 0.30
F2 (Mo17×dek48) 1401 326/1075 2.24

图7

多态性SNP标记的数目"

图8

SNP-index全基因组频率分布图"

图9

dek48突变体的精细定位 N: 群体大小; Recombinant: 重组单株数。"

表3

dek48定位区间的候选基因注释"

基因位点
Gene locus		 基因位置
Gene location		 基因注释
Gene annotation
Zm00001d039532 Chr. 3: 7,399,394-7,401,753 WRKY-transcription factor 56
Zm00001d039533 Chr. 3: 7,446,385-7,450,083 Plant cysteine oxidase 2
Zm00001d039534 Chr. 3: 7,451,813-7,457,371 F-box family protein
Zm00001d039535 Chr. 3: 7,479,692-7,482,979 HXXXD-type acyl-transferase family protein
Zm00001d039536 Chr. 3: 7,484,018-7,486,445 Syntaxin22
Zm00001d039537 Chr. 3: 7,487,245-7,530,464 Protein ALWAYS EARLY 3
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