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作物学报

作物学报 ›› 2023, Vol. 49 ›› Issue (11): 3122-3130.doi: 10.3724/SP.J.1006.2023.23076

• 研究简报 • 上一篇    下一篇

玉米小籽粒突变体mn-Mu的基因克隆与转录组分析

丁孟丽(), 王茹茵, 施栋晟, 李莹博, 雷洁, 陈洪宇, 申清文(), 王桂凤   

  1. 河南农业大学农学院 / 省部共建小麦玉米作物学国家重点实验室 / CIMMYT-中国(河南)小麦玉米联合研究中心, 河南郑州 450046
  • 收稿日期:2022-11-14 接受日期:2023-05-24 出版日期:2023-11-12 网络出版日期:2023-05-31
  • 通讯作者: 申清文, E-mail: shenqingwen@henau.edu.cn
  • 作者简介:E-mail: 1669279049@qq.com
  • 基金资助:
    国家自然科学基金项目(32001562);国家自然科学基金项目(U22A20466);河南农业大学科技创新基金项目(KJCX2020A04)

Map-based cloning and transcriptomic analysis of a maize miniature kernel mutant mn-Mu

DING Meng-Li(), WANG Ru-Yin, SHI Dong-Sheng, LI Ying-Bo, LEI Jie, CHEN Hong-Yu, SHEN Qing-Wen(), WANG Gui-Feng   

  1. College of Agronomy, Henan Agricultural University / State Key Laboratory of Wheat and Maize Crops Science / CIMMYT-China (Henan) Joint Research Center of Wheat and Maize, Zhengzhou 450046, Henan, China
  • Received:2022-11-14 Accepted:2023-05-24 Published:2023-11-12 Published online:2023-05-31
  • Supported by:
    National Natural Science Foundation of China(32001562);National Natural Science Foundation of China(U22A20466);Science and Technology Innovation Fund of Henan Agricultural University(KJCX2020A04)

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摘要：

玉米籽粒大小是影响产量形成的关键因素。本研究鉴定了一个转座子随机插入的小籽粒突变体mn-Mu; 该突变体相较野生型籽粒变小、种皮皱缩、淀粉含量下降, 而醇溶蛋白含量升高。石蜡切片观察发现mn-Mu突变体胚乳发育迟缓, 胚乳基底转移层细胞发育缺陷。遗传分析表明mn-Mu是由隐性单基因控制的突变, 通过图位克隆将突变基因定位在2号染色体57.83~61.91 Mb的区间, 其中已克隆的编码细胞壁转化酶的Zm00001d003776 (Miniature 1, Mn1)基因在第5个外显子上存在一个1442 bp的转座子插入。等位测试证实, mn-Mu为一个新的Mn1基因等位突变体。转录组分析表明, Mn1基因功能缺失影响碳水化合物代谢、储藏物质积累、糖基转移酶活性、细胞壁生物合成和细胞周期调控等生物学过程。本研究鉴定的新的mn1等位突变体为深入解析籽粒发育的分子调控网络提供了新的遗传材料。

关键词: 玉米, 籽粒发育, mn-Mu, 图位克隆, 细胞壁转化酶, 转录组分析

Abstract:

Kernel is the main storage organ, which accumulates nutritional compounds and determinates maize yield. In this study, we identified a miniature kernel mutant, mn-Mu, caused by a random mutator’s insertion. Compared with wild type, mn-Mu had smaller kernels, shrunken pericarp, and decreased significantly kernel weight. Compared with wild type, starch content of mn-Mu kernels decreased slightly, while zein content increased. Cytological observation showed that the development of the central endosperm and the basal endosperm transfer layer (BETL) was impaired in mn-Mu mutant compared with the wild type. Genetic analysis revealed that mn-Mu was a single gene-controlled recessive mutant. Map-based cloning indicated that the mutant gene was positioned in 57.83-61.91 Mb on chromosome 2. In this interval, Zm00001d003776 gene, which encoded CELL WALL INVERTASE 2 (INCW2, Miniature 1, Mn1) previously reported, was found that a 1442 bp transposon derived from Bronze 1 locus inserted into its 5th exon. Allelic test validated that mn-Mu was a new allelic mutant of mn1. Furthermore, transcriptomic analysis revealed that the loss of Mn1 affected gene expression in the processes of carbohydrate metabolism, storage material accumulation, glycosyltransferase activity, cell wall biosynthesis, and cell cycle regulation. In conclusion, a new allelic mutant of maize Mn1 (mn-Mu) was identified and transcriptomic analysis was further performed, which providing a novel maize germplasm and clues toward comprehensive understanding of molecular regulation mechanism of Mn1 on kernel development.

Key words: maize, kernel development, mn-Mu, gene mapping, cell wall invertase, transcriptomic analysis

表1

基因定位及克隆引物序列"

引物名称
Primer name		 正向引物
Forward sequence (5'-3')		 反向引物
Reverse sequence (5'-3')
Indel-26.11 ACTTGCCTGCTGCGACTGG TCCTGCCCGTCCCGACAG
Indel-34.89 TCGTGGCTGGCTGGAGTG CGGGAAATCGGGTTGTTGATTG
Indel-43.19 TGGTAGGCAGGGCGTTGG TGTCCGTGTAGACTGTAGGAAC
Indel-53.27 GGGAGAGCATCGACAGATACTG GACAATGACTTCGTACCTTCGG
Indel-55.89 GCCGCACACACCCTATGAAG AGCGTGCATGTATCTGGAAGTG
Indel-57.83 GCAATCCCCTTGTCCTGAAAAG CTTCCAACGCAGCAGCAATG
Indel-61.91 AGGTCAACCATCGAATCGAAGC ATCTGCATGGTTCAGAGTCTCC
Indel-83.16 TCCATCTATGGGGAGCGAGTAC GCAGTCCGTGACGACAACC
PDC1R/2F GCAACGGTGCAAGTGAACAA GCGGTACGACTACTACACCG
PDC1F/2R GTCGTAAGTTTCGCTTCGGC TCGTAGAACGTCTTGGACGC

图1

玉米mn-Mu籽粒的表型特征 A: F1自交成熟果穗; B: 野生型(上排)和mn-Mu (下排)粒长对比; C: 野生型(上排)和mn-Mu (下排)粒宽对比; D: 单粒野生型(左)和突变体(右)籽粒对比; E: 野生型(左)和mn-Mu (右)籽粒纵切; F: 野生型和mn-Mu籽粒百粒重统计。**表示Student’s t检验在0.01概率水平差异显著, 图中标尺为1 cm。"

表2

F2成熟果穗正常和突变籽粒分离比"

果穗
Ear		 正常籽粒数
Wild type		 突变籽粒数
Mutant		 χ2
1 330 89 2.9602
2 295 100 0.0076
3 297 92 0.3093
总计 Total 922 281 1.6428

图2

玉米mn-Mu突变体籽粒的组织学观察 A: 野生型籽粒, 标尺为1 mm; B: mn-Mu突变体籽粒, 标尺为1 mm; C: 野生型籽粒胚, 标尺为1 mm; D: mn-Mu突变体籽粒胚, 标尺为500 μm; E: 野生型籽粒BETL观察, 标尺为200 μm; F: mn-Mu突变体籽粒BETL观察, 标尺为200 μm。样品为授粉后18 d发育中籽粒。"

图3

玉米mn-Mu突变体籽粒的储藏物质分析 A: 野生型和mn-Mu籽粒的总淀粉含量测定; B: 野生型和mn-Mu成熟籽粒醇溶蛋白含量对比; C: 野生型和mn-Mu成熟籽粒非醇溶蛋白含量对比。M: 蛋白分子量标记; WT: 野生型; Re1/Re2/Re3代表3个生物学重复。"

图4

玉米mn-Mu的基因定位与克隆 A: 玉米mn-Mu的图位克隆。n: 定位所用的群体数目, 与InDel标记相对应的数值为交换籽粒数目; B: Mn1基因结构示意图。三角符号代表mn-Mu中转座子插入位置; C: Mn1基因DNA全长扩增。M: DNA分子量标记; WT: 野生型; Z58: 自交系郑58对照。"

图5

mn-Mu突变体与mn1突变体等位测试 A: mn-Mu/+ × mn1/+杂交果穗表型, 标尺为1 cm; B: mn1/+ × mn-Mu/+杂交果穗表型, 标尺为1 cm。"

图6

玉米mn-Mu突变体转录组测序分析 A: RNA-seq检测mn-Mu差异表达基因; B: 差异表达基因KEGG富集分析; C: 差异表达基因GO富集分析。"

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