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作物学报 ›› 2024, Vol. 50 ›› Issue (6): 1420-1434.doi: 10.3724/SP.J.1006.2024.33060

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

玉米ZmGRAS13基因的克隆及功能研究

折萌1,2(), 郑登俞2, 柯照2, 吴忠义2, 邹华文1,*(), 张中保2,*()   

  1. 1长江大学农学院, 湖北荆州 434025
    2北京市农林科学院生物技术研究所 / 农业基因资源与生物技术北京市重点实验室, 北京 100097
  • 收稿日期:2023-10-18 接受日期:2024-01-12 出版日期:2024-06-12 网络出版日期:2024-02-19
  • 通讯作者: * 邹华文, E-mail: zouhuawen@yangtzeu.edu.cn; 张中保, E-mail: zhangzhongbao@baafs.net.cn
  • 作者简介:E-mail: shemeng0403@163.com
  • 基金资助:
    北京市自然科学基金项目(6222009);北京市农林科学院创新能力建设专项(KJCX20230203);北京市农林科学院生物技术共享平台2023项目

Cloning and functional analysis of ZmGRAS13 gene in maize

SHE Meng1,2(), ZHENG Deng-Yu2, KE Zhao2, WU Zhong-Yi2, ZOU Hua-Wen1,*(), ZHANG Zhong-Bao2,*()   

  1. 1College of Agriculture, Yangtze University, Jingzhou 434025, Hubei, China
    2Institute of Biotechnology, Beijing Academy of Agriculture and Forestry Sciences / Beijing Key Laboratory of Agricultural Gene Resources and Biotechnology, Beijing 100097, China
  • Received:2023-10-18 Accepted:2024-01-12 Published:2024-06-12 Published online:2024-02-19
  • Contact: * E-mail: zouhuawen@yangtzeu.edu.cn;E-mail: zhangzhongbao@baafs.net.cn
  • Supported by:
    Beijing Natural Science Foundation(6222009);Beijing Academy of Agricultural and Forestry Sciences(KJCX20230203);Beijing Academy of Agricultural and Forestry Sciences Biotechnology Sharing Platform in 2023

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摘要：

GRAS家族是植物特有的一类转录因子, 在调控植物生长发育和响应逆境胁迫等方面发挥着重要作用。探究玉米(Zea mays L.) GRAS家族基因功能将为玉米新种质创制提供重要的基因资源。本研究克隆获得了ZmGRAS13基因(Zm00001eb401210), 利用生物信息学分析、实时荧光定量PCR (qPCR)等技术对该基因的基本特性、组织表达特性及逆境胁迫下表达模式等进行分析。生物信息学分析结果显示, 该基因编码序列全长为1638 bp, 编码545个氨基酸; ZmGRAS13蛋白不具有跨膜结构, 分子量为60.79 kD, 理论等电点为5.86, 具有GRAS家族所特有的保守结构域。对基因启动子上游2 kb序列进行分析, 发现该序列含有与逆境胁迫、激素响应及光响应等相关的顺式作用元件。qPCR分析表明, ZmGRAS13基因在玉米不同组织中均有表达, 且茎中的表达量最高; 同时该基因在不同非生物胁迫处理条件下均有不同程度的诱导表达。玉米原生质体瞬时表达实验表明, ZmGRAS13蛋白定位于细胞核。在分别含有不同浓度梯度的NaCl、甘露醇(mannitol)、脱落酸(abscisic acid, ABA)、茉莉酸(jasmonic acid, JA)和水杨酸(salicylic acid, SA)的1/2 MS固体培养基上, 转ZmGRAS13基因拟南芥株系的根长均显著长于野生型拟南芥; 在土壤中, 高盐和干旱处理下转基因拟南芥株系较野生型拟南芥生长状态更好, 且绿叶率高于野生型。转ZmGRAS13基因拟南芥与野生型相比, 抗逆生理指标MDA含量降低、叶绿素含量增加、POD和CAT活性增强, 且差异均显著。由此推测, ZmGRAS13基因可能参与玉米生长发育调控和对逆境胁迫应答及激素信号转导途径。本研究为进一步解析ZmGRAS13在玉米中的生物学功能提供了重要的参考依据。

关键词: 玉米, ZmGRAS13, 转录因子, 渗透胁迫, 盐胁迫, 异源表达

Abstract:

GRAS family is a plant-specific transcription factor, which plays an important role in regulating plant growth and development and responding to stresses. Exploring the function of GRAS family genes in maize (Zea mays L.) provides the important genetic resources for the creation of new maize germplasm. In this study, ZmGRAS13 gene (Zm00001eb401210) was cloned, and its basic characteristics, tissue expression characteristics, and the relative expression patterns under stresses were analyzed by bioinformatics and qRT-PCR. Bioinformatics showed that the full-length coding sequence of this gene was 1638 bp, encoding 545 amino acids. ZmGRAS13 protein had no transmembrane structure, and the molecular weight of 60.79 kD, the theoretical isoelectric point of 5.86, and had a conserved domain unique to the GRAS family. The analysis of 2 kb sequence upstream of the gene promoter indicated that the sequence contained cis-acting elements related to stresses, hormone response, and light response. The qRT-PCR analysis showed that ZmGRAS13 gene was expressed in different tissues of maize, and the relative expression level in stem was the highest. At the same time, the gene has different degrees of induced expression under different abiotic stress treatment conditions. The transient expression experiment of maize protoplasts demonstrated that ZmGRAS13 protein was localized in the nucleus. On 1/2 MS solid medium containing different concentrations of NaCl, mannitol, abscisic acid (ABA), jasmonic acid (JA), and salicylic acid (SA), respectively, the root length of ZmGRAS13 transgenic Arabidopsis lines was significantly longer than the control. In the soil, transgenic Arabidopsis lines grew better than the control under high salt and drought treatments, and the green leaf rate was higher than the control. Compared with the wild type, the content of stress resistance physiological index MDA decreased, the chlorophyll content increased, and the activities of POD and CAT increased in the transgenic ZmGRAS13 Arabidopsis thaliana, and the difference was significant difference. In conclusion, ZmGRAS13 gene may be involved in the regulation of maize growth and development, response to stresses and hormone signal transduction pathway. This study provides an important reference for the further analysis of the biological function of ZmGRAS13 in maize.

Key words: maize, ZmGRAS13, transcription factors, osmotic stress, salt stress, heterologous expression

表1

本实验所用的引物"

引物名称Primer name 引物序列Primer sequence (5'-3')
pZmGRAS13-FP ACCCGGGCTGCAGAATTCATGCCCATAAAATTTGCACTAG (EcoR I)
pZmGRAS13-RP CATGGTACCCTCGAGAAGCTTGCACCAGGCAGAAGATACGAC (Hind III)
pZmGRAS13RT-FP ATAGGTAGCCCTGACAGTTCCT
pZmGRAS13RT-RP TCCAACATGTAAGCTCCCAGAC
pZmGRAS13T-FP AGGCAGGTAATTGTAGCATG
pZmGRAS13T-RP TCTGTCCAACCATATCCAG
pGAPDHRT-FP
pGAPDHRT-RP		 CCCTTCATCACCACGGACTAC
AACCTTCTTGGCACCACCCT
pActin-FP
pActin-RP		 TACGAGATGCCTGATGGTCAGGTCA
TGGAGTTGTACGTGGCCTCATGGAC

图1

ZmGRAS13基因的克隆及生物信息学分析 (a): ZmGRAS13基因的克隆(M: DL2000 DNA marker; 1, 2: ZmGRAS13扩增片段); (b): 跨膜结构预测; (c): 蛋白空间结构预测; (d): 蛋白亲疏水性预测; (e): 保守结构域预测; (f): 启动子区顺式作用元件分析。"

图2

ZmGRAS13基因在玉米不同组织中的相对表达量 不同小写字母表示差异显著(P < 0.05)。"

图3

ZmGRAS13基因在玉米不同胁迫处理下的相对表达量 (a)~(d) ZmGRAS13基因分别在脱水、高盐、渗透、低温处理后的相对表达量; (e) ZmGRAS13基因分别在JA、ABA、GA、SA、2,4-D处理后的相对表达量; *: P < 0.05; **: P < 0.01。"

图4

ZmGRAS13蛋白在玉米原生质体中的亚细胞定位 GFP: 绿色荧光通道; DAPI: DAPI通道; Bright: 明场通道; Merged: GFP、DAPI和Bright通道叠加; pYBA1132:EGFP: 转入空载体的原生质体; pYBA1132:ZmGRAS13:EGFP: 转入目的基因的原生质体。"

图5

T3代ZmGRAS13转基因拟南芥鉴定 (a): T3代转基因拟南芥PCR鉴定(M: DL2000 DNA marker; P: 阳性对照; N: 阴性对照; W: 水对照); (b): T3代转基因拟南芥qPCR鉴定(WT: 野生型拟南芥; L-1~L-7: T3代转基因拟南芥株系); *: P < 0.05; **: P < 0.01。"

图6

不同盐、甘露醇浓度处理下野生型和转基因拟南芥根长比较 (A): a~d: 0、0.10、0.15和0.18 mol L-1盐处理拟南芥的生长情况; e: 不同盐浓度处理下拟南芥平均主根长度。(B): a~d: 0、0.15、0.20和0.30 mol L-1甘露醇处理拟南芥的生长情况; e: 不同甘露醇浓度处理下拟南芥平均主根长度。WT: 野生型拟南芥; L-3、L-5、L-7: 转ZmGRAS13基因拟南芥株系; *: P < 0.05; **: P < 0.01。"

图7

不同ABA、JA和SA浓度处理下野生型和转基因拟南芥根长比较 (A): a~d: 0、10、25和50 μmol L-1 ABA处理拟南芥的生长情况; e: 不同ABA浓度处理下拟南芥平均主根长度。(B): a~d: 0、50、75和100 μmol L-1 JA处理拟南芥的生长情况; e: 不同JA浓度处理下拟南芥平均主根长度。(C): a~d: 0、25、50和75 μmol L-1 SA处理拟南芥的生长情况; e: 不同SA浓度处理下拟南芥平均主根长度。缩写同图6; *: P < 0.05; **: P < 0.01。"

图8

高盐处理下转基因拟南芥表型分析 (a): 拟南芥表型; (b): 高盐处理后绿叶率; (c): MDA含量; (d): POD活性; (e): 叶绿素含量; (f): CAT活性。缩写同图6; *: P < 0.05; **: P < 0.01。"

图9

干旱处理下转基因拟南芥表型分析 (a): 拟南芥表型; (b): 干旱处理前后绿叶率; (c): MDA含量; (d): POD活性; (e): 叶绿素含量; (f): CAT活性。缩写同图6; *: P < 0.05; **: P < 0.01。"

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