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作物学报 ›› 2023, Vol. 49 ›› Issue (9): 2331-2343.doi: 10.3724/SP.J.1006.2023.23069

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

玉米ZmC2s基因家族鉴定及ZmC2-15耐热功能分析

黄钰杰(), 张啸天, 陈会丽, 王宏伟(), 丁双成()   

  1. 农业农村部长江中游作物绿色高效生产重点实验室(部省共建) / 长江大学农学院, 湖北荆州 434025
  • 收稿日期:2022-10-11 接受日期:2023-02-10 出版日期:2023-09-12 网络出版日期:2023-02-22
  • 通讯作者: *王宏伟, E-mail: wanghw@yangtzeue.edu.cn; 丁双成, E-mail: shchding@yangtzeu.edu.cn
  • 作者简介:黄钰杰, E-mail: 996802397@qq.com
  • 基金资助:
    国家自然科学基金项目(31701439)

Identification of ZmC2s gene family and functional analysis of ZmC2-15 under heat tolerance in maize

HUANG Yu-Jie(), ZHANG Xiao-Tian, CHEN Hui-Li, WANG Hong-Wei(), DING Shuang-Cheng()   

  1. Key Laboratory of Sustainable Crop Production in the Middle Reaches of the Yangtze River, Ministry of Agricultural and Rural Affairs (Co-construction by Ministry and Province), College of Agriculture, Yangtze University, Jingzhou 434025, Hubei, China
  • Received:2022-10-11 Accepted:2023-02-10 Published:2023-09-12 Published online:2023-02-22
  • Supported by:
    National Natural Science Foundation of China(31701439)

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摘要：

鉴定玉米ZmC2s基因家族成员, 分析其遗传变异与玉米耐热性的关联性, 为明确其在玉米耐热方面的功能和分子机制奠定基础。使用C2蛋白结构域PF00168, 利用hmmsearch从玉米全基因组中搜索ZmC2s基因家族成员, 分析其编码蛋白质等电点和分子量、系统进化和基因家族复制。使用候选基因关联分析的方法, 对ZmC2s基因自然变异位点与玉米苗期耐热性进行关联分析, 发现玉米ZmC2s基因家族重要的耐热候选基因。使用实时荧光定量PCR (RT-qPCR)的方法鉴定与玉米苗期耐热性最显著相关的ZmC2候选基因ZmC2-15在胁迫和激素处理下的基因相对表达水平, 在玉米原生质体瞬时表达系统中鉴定ZmC2-15-GFP的亚细胞表达部位, 以及获得过表达ZmC2-15的转基因拟南芥并分析其耐热性。从玉米参考基因组B73中共鉴定出95个玉米ZmC2s基因, 根据其物理坐标的先后顺序, 将95个玉米ZmC2s基因命名为ZmC2-1~ZmC2-95。其蛋白长度为130~2141, 等电点为4.1~10.8, 分子量为14.1~230.1。通过构建玉米、水稻和高粱基因组C2基因的进化树, 发现C2基因可以分为3个大的聚类分支, 每个聚类分支又可以细分为2个小的聚类分支。分析玉米、水稻和玉米、高粱的全基因组共线性数据, 发现了59个ZmC2s基因在水稻和高粱基因组均有一一对应的复制基因。ZmC2s基因家族关联分析表明ZmC2-15/60/91是重要的玉米耐热候选基因(P≤0.001, MLM), 其中ZmC2-15与玉米苗期耐热性最显著相关(P≤0.000,01, MLM), 且该基因的表达量在多种逆境胁迫处理下上调表达。亚细胞定位分析表明ZmC2-15表达定位于细胞质、核膜和内质网。过表达ZmC2-15的转基因拟南芥提高了植物耐热性。ZmC2-15可作为调控玉米耐热性的重要候选基因。

关键词: 玉米, 耐热, C2基因家族, 候选基因关联分析, 亚细胞定位

Abstract:

The objective of this study is to identify the members of maize ZmC2s gene family, to analyze the association between their genetic variations and heat tolerance, and to lay a foundation for clarifying its function and molecular mechanism in maize heat tolerance. Using the C2 protein domain PF00168, hmmsearch was applied to search for the members of the ZmC2s gene family from maize B73 genome. The protein isoelectric point, molecular weight, phylogenetic evolution, and gene family replication were analyzed. Using the method of candidate gene association analysis, the association between the natural variations of ZmC2s and the heat tolerance of maize seedlings was conducted, and the important heat-tolerant candidate genes of maize ZmC2s gene family were found. The relative gene expression level of the heat-tolerant candidate gene under stress was identified by Real time fluorescent quantitative PCR (RT-qPCR). The subcellular expression sites of heat-tolerant candidate gene were identified by transforming maize protoplasts. A total of 95 maize ZmC2s genes were identified from the reference genome B73 in maize. According to the order of their physical coordinates, 95 maize ZmC2s genes were named from ZmC2-1 to ZmC2-95, respectively. The length of the 95 proteins was 130-2141, the isoelectric point was 4.1-10.8, and the molecular weight was 14.1-230.1. The evolution tree of C2 gene in maize, rice, and sorghum genomes was constructed. We found that C2 genes can be divided into three major cluster branches, and each cluster branch can be subdivided into two small cluster branches. Analyzing the whole genome collinearity data of maize, rice, and sorghum, 59 ZmC2s genes were detected to have corresponding replication genes in rice and sorghum genomes. A candidate-gene based on the association analysis of ZmC2s showed that ZmC2-15/60/91 were important candidate genes for heat tolerance in maize (P ≤ 0.001, MLM), among which ZmC2-15 was the most significantly associated to heat tolerance at seedling stage (P ≤ 0.000,01, MLM), and the relative expression level of ZmC2-15 was up-regulated under various stress treatments. Subcellular localization indicated that ZmC2-15 was localized in the cytoplasm, nuclear membrane, and endoplasmic reticulum. The overexpression of ZmC2-15 improved plant heat tolerance. ZmC2-15 can be used as an important candidate gene for regulating heat tolerance in maize.

Key words: maize, heat tolerance, C2 gene family, association analysis of candidate genes, subcellular localization

图1

ZmC2s基因家族成员鉴定和蛋白理化性质分析结果图 ZmC2s基因家族成员的内含子数量(A)、外显子数量(B)、蛋白质分子量(C)、蛋白质等电点(D)、蛋白长度(E)、基因名称(F)、基因物理坐标(G)和GenBank登录号(H)。"

图2

玉米、水稻和高粱3种作物中C2结构域基因的系统发育进化树 通过比较MEGA 7中的C2结构域氨基酸序列来生成系统发生树。使用邻居连接方法, 所有的基因被分成3组, I、II和III。"

图3

玉米C2结构域基因与水稻和高粱的基因复制分析 1号、2号(A); 3号、4号(B); 5号、6号(C); 7号、8号、9号、10号(D)染色体上ZmC2s基因家族成员与水稻和高粱的基因复制分析; E: 玉米、水稻和高粱基因组C2基因复制类型和数目。"

图4

玉米C2结构域基因与玉米耐热性的关联分析 A: ZmC2s遗传多态性标记数目; ZmC2s自然变异与玉米耐热性显著关联(GLM, P ≤ 0.01); B: MLM, P ≤ 0.01; C: (MLM, P ≤ 0.001; D: MLM, P ≤ 0.000,01; E: 基因及基因内显著关联的标记数目; F: ZmC2s基因坐标; G: ZmC2s基因名称; H: ZmC2基因遗传变异与玉米苗期耐热性的关联分析QQplot; ZmC2-15 (I), ZmC2-60 (J), ZmC2-91 (K)基因遗传变异与玉米苗期耐热性的关联分析。"

图5

高温(A)、干旱(B)、低温(C)、乙烯利(D)、氯化钠(E)、过氧化氢(F)、赤霉素(G)、ABA (H)和水杨酸(I)处理下ZmC2-15基因的表达模式 柱状图代表均值±标准差。*: P ≤ 0.05; **: P ≤ 0.01; t测验。"

图6

ZmC2-15-GFP在玉米原生质体里的亚细胞定位 A: GFP蛋白在玉米原生质体亚细胞定位; B: ZmC2-15-GFP加入细胞核染料DAPI后原生质体亚细胞定位。图中ZmC2-15-GFP代指pGreenII-35SΩ-ZmC2-15-GFP, 35S:GFP代指pGreenII-35SΩ-GFP。"

图7

ZmC2-15-GFP与ERD2-mCherry (A)、ST-mCherry (B)、VHA-A1-mCherry (C)和HDEL-mCherryL (D)在玉米原生质体亚细胞共定位 HDEL-mCherry、ERD2-mCherry、ST-mCherry和VHA-A1-mCherry分别作为内质网、顺面高尔基体、高尔基体和反面高尔基体蛋白定位标记。"

图8

ZmC2-15转基因拟南芥高温胁迫表型分析 A: 高温处理前; B: 高温处理后; C: 引物及结构域示意图及半定量分析结果; D: 对照与转基因拟南芥存活率柱状图。VC代表空载pGreenII-35SΩ转化拟南芥。柱状图代表均值±标准差, 数据统计自3次生物学重复, VC、OE1和OE2各有30株。*: P ≤ 0.05; **: P ≤ 0.01; t测验。"

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