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作物学报 ›› 2022, Vol. 48 ›› Issue (6): 1312-1324.doi: 10.3724/SP.J.1006.2022.13040

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

2个玉米ZmCOP1基因的克隆及其转录丰度对不同光质处理的响应

崔连花**(), 詹为民**(), 杨陆浩, 王少瓷, 马文奇, 姜良良, 张艳培*(), 杨建平*(), 杨青华*()   

  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 / 作物分子育种国家工程中心, 河南郑州 450002
  • 收稿日期:2021-05-27 接受日期:2021-10-19 出版日期:2022-06-12 网络出版日期:2021-11-02
  • 通讯作者: 张艳培,杨建平,杨青华
  • 作者简介:崔连花, E-mail: 1970125806@qq.com;
    詹为民, E-mail: 630950832@qq.com第一联系人：

    ** 同等贡献

  • 基金资助:
    国家自然科学基金项目(31871709);河南省技术创新引导专项(182106000050);北京市自然科学基金(重点)项目(6151002)

Molecular cloning of two maize (Zea mays) ZmCOP1 genes and their transcription abundances in response to different light treatments

CUI Lian-Hua**(), ZHAN Wei-Min**(), YANG Lu-Hao, WANG Shao-Ci, MA Wen-Qi, JIANG Liang-Liang, ZHANG Yan-Pei*(), YANG Jian-Ping*(), YANG Qing-Hua*()   

  1. College of Agronomy, Henan Agricultural University / State Key Laboratory of Wheat and Maize Crops Science / Center for Crop Genome Engineering, Zhengzhou 450002, Henan, China
  • Received:2021-05-27 Accepted:2021-10-19 Published:2022-06-12 Published online:2021-11-02
  • Contact: ZHANG Yan-Pei,YANG Jian-Ping,YANG Qing-Hua
  • About author:First author contact:

    ** Contributed equally to this work

  • Supported by:
    National Natural Science Foundation of China(31871709);Henan Technology Innovation Guidance Project(182106000050);Key Project of Beijing Natural Science Foundation(6151002)

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摘要：

CONSTITUTIVE PHOTOMORPHOGENIC 1 (COP1)作为一种E3泛素连接酶, 能够介导拟南芥光信号转导中HY5、LAF1、HFR1和CO等正调控因子的泛素化降解, 从而调控幼苗光形态建成、花青素合成和开花等生物学过程。为了探索玉米COP1基因的功能, 本文以玉米自交系B73为研究材料, 利用反转录PCR (RT-PCR)方法克隆了2个ZmCOP1基因, 分别命名为ZmCOP1a和ZmCOP1b。通过生物信息学相关软件和网站对ZmCOP1进行理化性质分析、结构域预测和系统发育树分析, 采用实时荧光定量PCR (qRT-PCR)技术进一步分析2个ZmCOP1基因在不同组织中的转录丰度, 以及其在幼苗期对不同光质和光周期处理的响应。研究发现, ZmCOP1a和ZmCOP1b的开放阅读框(ORF)分别包含2082个和2061个核苷酸, 编码693个和686个氨基酸。玉米、水稻、高粱、谷子和拟南芥的COP1蛋白具有相同的结构域和较高的氨基酸序列一致性, 暗示它们可能具有相似的功能。2个ZmCOP1基因主要在玉米地上组织中表达, 均能迅速响应不同光质处理, 且ZmCOP1a的转录丰度普遍高于ZmCOP1b, 表明ZmCOP1a可能在不同光质处理下具有更重要的作用。在长日照和短日照条件下, ZmCOP1a和ZmCOP1b在黑暗阶段的转录丰度整体高于光照阶段, 有趣的是, 二者在光照阶段的表达模式相似, 但是在黑暗阶段, ZmCOP1b的转录丰度高于ZmCOP1a。这说明在不同光周期条件下, ZmCOP1b可能比ZmCOP1a更重要。综上这些研究结果表明, ZmCOP1a和ZmCOP1b具有功能冗余和分化。二者可以参与不同光信号通路, 可能在玉米光形态建成和花期调控中发挥重要作用。本研究为进一步探明ZmCOP1a和ZmCOP1b基因的功能及其在玉米育种中的应用提供了研究基础。

关键词: 玉米, ZmCOP1, 光处理, 基因克隆, 转录丰度

Abstract:

As an E3 ubiquitin ligase, CONSTITUTIVE PHOTOMORPHOGENIC 1 (COP1) mediates ubiquitination and degradation of positive regulatory factors such as HY5, LAF1, HFR1, and CO of the light signal transduction in Arabidopsis thaliana, thus mediating seedling photomorphogenesis, anthocyanin synthesis, and flowering time. To explore functional differentiation of ZmCOP1 genes, we cloned two ZmCOP1 genes, designated as ZmCOP1a and ZmCOP1b, from maize inbred line B73 by reverse transcription PCR (RT-PCR). The physicochemical properties, domain prediction and phylogenetic tree of ZmCOP1 were analyzed by bioinformatics software and website. The transcription abundances of two ZmCOP1 genes in different tissues and their responses to different light treatments at seedling stage were further analyzed by quantitative RT-PCR (qRT-PCR). We found that the open reading frames (ORFs) of ZmCOP1a and ZmCOP1b possessed 2082 and 2061 nucleotides, encoding 693 and 686 amino acids, respectively. The COP1 proteins of maize, rice, sorghum, millet, and Arabidopsis shared the same structural domains and high amino acid sequence identity, indicating that they may have similar functions. The two ZmCOP1 genes were mainly expressed in the above-ground tissues. Both of them could rapidly respond to different light treatments, while the transcription abundances of ZmCOP1a were generally higher than those of ZmCOP1b. It might suggest that ZmCOP1a may play more important roles under different light conditions. Under long-day or short-day condition, the transcription abundances of both ZmCOP1a and ZmCOP1b during the dark phase were higher than those during the light phase. Another interesting case is that both genes kept similar expression patterns during the light phase, while, transcription abundances of ZmCOP1b were higher than those of ZmCOP1a during the dark phase. It might imply that ZmCOP1b may be more important than ZmCOP1a in response to long-day or short-day treatment. In conclusion, both ZmCOP1a and ZmCOP1b had got functional redundancy and differentiation. Both genes could participate in different light signal pathways, and regulate maize photomorphogenesis and flowering period. Our results also provide a research foundation for further exploration of two ZmCOP1 genes and application in molecular breeding in maize.

Key words: Zea mays, ZmCOP1, light treatment, gene cloning, transcription abundance

表1

同源克隆所用引物"

基因名称
Gene name		 基因编号
Accession number		 正向引物序列
Forward sequence (5°-3°)		 反向引物序列
Reverse sequence (5°-3°)
ZmCOP1a Zm00001d018207 GGGGGACTCTTGACCATGGGCATGGGCGACTCCTCGGTG AAGTTCTTCTCCTTTACTAGTAGGAGCAAGTACAAGAACTTTTATGG
ZmCOP1b Zm00001d052138 GGGGGACTCTTGACCATGGGCATGGGCGACTCCTCGGTG AAGTTCTTCTCCTTTACTAGTAGGAGCAAGTACATGAACTTTTATGG

表2

qRT-PCR所用引物"

基因名称
Gene name		 基因编号
Accession number		 正向引物序列
Forward sequence (5°-3°)		 反向引物序列
Reverse sequence (5°-3°)
ZmCOP1a Zm00001d018207 GCTAACAACGTTCACTCGATAC CAATGCACATCTGATGGTTCAT
ZmCOP1b Zm00001d052138 CTGCTGCGGGCACTACCTTA CCCATTTCATTTCCCTGTTG
ZmUBQ1 Zm00001d015327 ACTTCATGCTTTCGTCCTACGCTCCA CTGGGAGGCTGGTAGTTGATTC

图1

2个ZmCOP1 cDNA片段的RT-PCR扩增及其表达载体pCAMBIA1302-ZmCOP1的双酶切鉴定 A: ZmCOP1的cDNA片段的RT-PCR扩增。M: marker III DNA ladder; 1: ZmCOP1a PCR产物; 2: ZmCOP1b PCR产物。B: pCAMBIA1302-ZmCOP1的双酶切鉴定。M: DL10000 DNA ladder; 1和2分别为pCAMBIA1302-ZmCOP1a和pCAMBIA1302-ZmCOP1b经Nco I和Spe I双酶切后获得约2082 bp和2061 bp的条带; 箭头表示ZmCOP1基因片段。"

图2

玉米与其他植物COP1蛋白在氨基酸水平的系统发育分析 在Ensembl Plants网站获得全长氨基酸序列, 使用MAFFT与MEGA软件对其进行系统发育分析, 使用DNAMAN Version 8软件进行氨基酸序列比对。ZmCOP1a: 玉米, Zm00001d018207; ZmCOP1b: 玉米, Zm00001d052138; SbCOP1: 高粱, KXG31183; SvCOP1: 狗尾草, TKW41800; SiCOP1: 谷子, KQL31700; OsCOP1: 水稻, Os02t0771100; BdCOP1: 二穗短柄草, KQK01702; TaCOP1b: 小麦, TraesCS6D02G305800; TaCOP1c: 小麦, TraesCS6B02G356400; TaCOP1a: 小麦, TraesCS6A02G326100; GmCOP1a: 大豆, KRH14801; GmCOP1b: 大豆, KRH73341; MeCOP1: 木薯, OAY35060; GrCOP1: 棉花, KJB42844; AtCOP1: 拟南芥, AT2G32950; BnCOP1a: 甘蓝型油菜, CDY52540; BnCOP1b: 甘蓝型油菜, CDY12145; BoCOP1: 甘蓝, Bo4g045420。"

图3

玉米与拟南芥、水稻、高粱和谷子的COP1蛋白的氨基酸序列比对及结构域分析 利用SMART (http://smart.embl-heidelberg.de/)和InterPro网站(https://www.ebi.ac.uk/interpro/)以及MAFFT软件进行氨基酸序列比对和结构域分析。AtCOP1: 拟南芥COP1, AT2G32950; ZmCOP1a: 玉米COP1a, Zm00001d018207; ZmCOP1b: 玉米COP1b, Zm00001d052138; OsCOP1: 水稻COP1, Os02t0771100; SbCOP1: 高粱COP1, KXG31183; SiCOP1: 谷子COP1, KQL31700。"

图4

2个ZmCOP1 qRT-PCR引物的验证及二者在不同组织中的相对表达水平 A: ZmCOP1a基因qRT-PCR的溶解曲线。B: ZmCOP1b基因qRT-PCR的溶解曲线。C: ZmCOP1a和ZmCOP1b基因qRT-PCR的相对定量标准曲线及引物扩增效率。R2: 相关系数; E: 引物扩增效率, E = 10 ^ (-1/slope) - 1。D: 以玉米自交系B73的不同组织(根、幼穗、茎、苞叶、雄花、叶枕、叶鞘、花丝和叶)为材料, 对ZmCOP1a和ZmCOP1b进行qRT-PCR分析。以根中ZmCOP1a的转录丰度为参照(设为1), 柱状图显示结果为3次生物学重复的相对平均值, 误差线代表标准差, *: P < 0.05; **: P < 0.01。"

图5

2个ZmCOP1在不同持续光质下的相对表达水平 玉米自交系B73分别在黑暗(Dk)、持续远红光(FR, 0.3 µmol m-2 s-1)、持续红光(R, 22.3 µmol m-2 s-1)、持续蓝光(B, 13.0 µmol m-2 s-1)或持续白光(WL, 17.0 µmol m-2 s-1)下生长13 d。以黑暗条件下ZmCOP1a的转录丰度(设为1)为参照, 柱状图显示的是3次生物学重复的相对平均值, 误差线代表标准差, *: P < 0.05; **: P < 0.01。"

图6

由黑暗转到不同光质后ZmCOP1a和ZmCOP1b的相对表达水平 玉米自交系B73幼苗在黑暗(Dk)条件下生长11 d后, 分别转到红光(R, 22.3 µmol m-2 s-1)、远红光(FR, 0.3 µmol m-2 s-1)、蓝光(B, 13.0 µmol m-2 s-1)和白光(WL, 17.0 µmol m-2 s-1)条件下, 并在0、0.25、0.5、1、2、4、8、12和24 h取样, 用于qRT-PCR分析。以黑暗条件下ZmCOP1a转录丰度(设为1)作为参照, 折线图显示3生物学重复的相对平均值, 误差线代表标准差。"

图7

光周期处理下ZmCOP1a和ZmCOP1b的相对表达水平 玉米自交系B73分别在长日照(LD, 16 h光照/8 h黑暗)或者短日照(SD, 8 h光照/16 h黑暗)条件下生长13 d后, 每2 h取一次样, 用于qRT-PCR分析。A和B图中数据均以长日照条件下12:00的ZmCOP1b转录丰度(设为1)为参照。折线图显示为3次生物学重复的相对平均值, 误差线代表标准差。"

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