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作物学报 ›› 2023, Vol. 49 ›› Issue (12): 3289-3301.doi: 10.3724/SP.J.1006.2023.24284

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

甘薯蔓割病(Fusarium oxysporum f. sp. batatas)相关基因IbMAPKK9的克隆与特性分析

靖小菁1,2(), 杨新笋2, 靳晓杰2, 刘意3, 雷剑2, 王连军2, 柴沙沙2, 张文英1,*(), 焦春海2,*()   

  1. 1长江大学农学院, 湖北荆州 434025
    2湖北省农业科学院粮食作物研究所, 湖北武汉 430064
    3海南大学园艺学院 / 海南省热带园艺作物品质调控重点实验室, 海南海口 570228
  • 收稿日期:2022-12-23 接受日期:2023-05-24 出版日期:2023-12-12 网络出版日期:2023-06-04
  • 通讯作者: * 张文英, E-mail: wyzhang@yangtzeu.edu.cn; 焦春海, E-mail: jiaoch@hotmail.com
  • 作者简介:E-mail: 15172397885@163.com
  • 基金资助:
    国家重点研发计划项目(2019YFD1001300);国家重点研发计划项目(2019YFD1001305);财政部和农业农村部国家现代农业产业技术体系建设专项(Sweetpotato, CARS-10)

Cloning and characterization of IbMAPKK9 gene associated with Fusarium oxysporum f. sp. batatas in sweet potato

JING Xiao-Jing1,2(), YANG Xin-Sun2, JIN Xiao-Jie2, LIU Yi3, LEI Jian2, WANG Lian-Jun2, CHAI Sha-Sha2, ZHANG Wen-Ying1,*(), JIAO Chun-Hai2,*()   

  1. 1College of Agriculture, Yangtze University, Jingzhou 434025, Hubei, China
    2Food Crops institute, Hubei Academy of Agricultural Sciences, Wuhan 430064, Hubei, China
    3Horticulture College, Hainan University / Key Laboratory for Quality Regulation of Tropical Horticultural Plants of Hainan Province, Haikou 570228, Hainan, China
  • Received:2022-12-23 Accepted:2023-05-24 Published:2023-12-12 Published online:2023-06-04
  • Contact: * E-mail: wyzhang@yangtzeu.edu.cn; E-mail: jiaoch@hotmail.com
  • Supported by:
    National Key Research and Development Program of China(2019YFD1001300);National Key Research and Development Program of China(2019YFD1001305);China Agriculture Research System of MOF and MARA(Sweetpotato, CARS-10)

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摘要：

促丝裂原活化蛋白激酶(MAPK)级联是生物体内一种重要的信号转导途径, 广泛存在于植物中。MAPKKs位于该级联反应通路中间, 对信号传递起到关键作用, 目前在甘薯中少有报道。基于前期的转录组学分析, 本研究克隆出1个与甘薯抗病相关的基因IbMAPKK9。生物信息学分析表明, IbMAPKK9包含一个987 bp开放阅读框(open reading frame, ORF), 编码328个氨基酸, 具有1个蛋白激酶家族保守结构域(PF00069), IbMAPKK9蛋白以α螺旋和无规则卷曲为主, 启动子区包含多种激素(茉莉酸甲酯、乙烯、脱落酸、赤霉素、水杨酸)相关及胁迫响应元件。进化分析表明, IbMAPKK9蛋白与三叶裂薯、日本牵牛花、番茄和马铃薯亲缘关系较近。亚细胞定位显示IbMAPKK9蛋白定位于细胞核。实时荧光定量PCR发现IbMAPKK9在甘薯根、茎、叶和叶柄中均表达, 并响应甘薯蔓割病侵染。瞬时表达分析结果表明, IbMAPKK9引起5个与水杨酸合成途径及信号转导途径相关的基因在48 h内表达上调, 推测IbMAPKK9通过介导水杨酸信号途径影响植物的抗性。本研究可为进一步解析甘薯IbMAPKK9的生物学功能提供理论依据。

关键词: 甘薯, IbMAPKK9, 蔓割病, 基因克隆, 亚细胞定位, 瞬时表达

Abstract:

Mitogen-activated protein kinase (MAPK) cascades are important signaling modules in all organisms, which widely exist in plants. Mitogen-activated protein kinase kinase (MAPKKs) are located in the middle of the cascade reaction pathway and play the key roles in signal divergence. The disease resistance gene IbMAPKK9 was screened and cloned based on transcriptome data in sweetpotato, which contained a 987 bp open reading frame, encoding 328 amino acids and 1 domain (PF00069). Its protein mainly consisted of α-helixes and random coils. Multiple hormones (methyl jasmonate, ethylene, abscisic acid, gibberellin, and salicylic acid) and stress-related response elements were detected in the promoter region. Homologous protein comparison showed that IbMAPKK9 was closely related to ItMAPKK9-like (XP_031110493.1), InMAPKK9-like (XP_019189439.1), SlMAPKK9 (NP_001234595.1), and StMAPKK9 (XP_006363984.1). Besides, IbMAPKK9 gene encoded nuclear localization protein. The relative expression levels revealed that IbMAPKK9 was expressed in roots, stems, leaves, and petioles in response to Fusarium oxysporum f. sp batatas (Fob) infection. Transient expression analysis showed that IbMAPKK9 induced up-regulation of 5 genes related to salicylic acid synthesis pathway and signal transduction pathway within 48 hours, suggesting that IbMAPKK9 affected plant resistance by mediating salicylic acid signal pathway. This study can provide a theoretical basis for further analysis of the biological function of IbMAPKK9.

Key words: sweet potato, IbMAPKK9, Fusarium wilt, gene clone, subcellular localization, transient expression

表1

本研究所用引物"

引物名称
Primer name		 序列
Primer sequence (5′-3′)
IbMAPKK9-CDS-F ATGGCACTTGTTCGTGAACGT
IbMAPKK9-CDS-R TTAATTTTGTGAATTAATTTGGGATTTG
IbMAPKK9-PRO-F CTGCCAATGTGTATCACATGTTAT
IbMAPKK9-PRO-R GATGAGAAGGTAGAAGGGGAGT
IbMAPKK9-qPCR-F GCGGTCTACGCCCTCAAAG
IbMAPKK9-qPCR-R GAATATCGGAGGTGCCGTGAC
β-Actin-F AGCAGCATGAAGATTAAGGTTGTAGCAC
β-Actin-R TGGAAAATTAGAAGCACTTCCTGTGAAC
NtPAL4-F CATTGCCACATTCAGCAACA
NtPAL4-R GGGTGGTGCTTCAACTTGTG
NtICS1-F GAGGGTGCGGAAAAAAATA
NtICS1-R CCTTGATAAGCATCGGGTT
NtNPR1-F TTCTGTATCTCTTGCTATG
NtNPR1-R ATCTACTGTTGTCCTCTGT
NtNPR3-F GGGGCTACTTACCTA
NtNPR3-R TCCCTGTCTGATTCA
NtNPR5-F AAGGATGGATGGGAT
NtNPR5-R AGTTGGGTTGTTTGC

图1

甘薯IbMAPKK9的CDS与启动子克隆 M: DL2000 marker; 1: IbMAPKK9基因CDS扩增产物; 2: IbMAPKK9启动子扩增产物。"

图2

IbMAPKK9启动子区的调控元件分布"

表2

IbMAPKK9基因启动区顺式作用元件分析"

位点名称
Site name		 序列
Sequence (5′-3′)		 功能
Function
ABRE ACGTG 脱落酸反应的顺式作用元件 Cis-acting element involved in abscisic acid response
ARE AAACCA 厌氧诱导反应的顺式作用调节元件 Cis-acting regulatory element essential for the anaerobic induction
ERE ATTTTAAA 乙烯响应元件 Ethylene responsive element
G-box CACGTG 参与光反应的顺式作用元件 Light responsive element
P-box CCTTTTG 赤霉素反应元件 Gibberellin-responsive element
MYB TAACCA 干旱诱导相关元件 MYB binding site involved in drought-inducibility
MYC CATGTG 逆境胁迫相关顺式调控元件 Stress related cis-regulatory elements
G-box CACGTG 茉莉酸甲酯应答元件 Cis-acting regulatory element involved in the MeJA-responsiveness

图3

甘薯IbMAPKK9蛋白保守结构域分析"

图4

甘薯IbMAPKK9蛋白质磷酸化位点预测结果"

表3

IbMAPKK9蛋白二级结构主要组成元件及比例"

名称
Name		 α-螺旋
Alpha helix		 β-转角
Beta turn		 无规则卷曲
Random coil		 延伸链
Extended strand
IbMAPKK9 33.54 5.18 46.95 14.33

图5

IbMAPKK9蛋白二级结构的预测 蓝色表示α-螺旋(Hh); 红色代表延伸链(Ee); 绿色表示β转角(Tt); 紫色表示无规则卷曲(Cc)。"

图6

IbMAPKK9蛋白三维结构预测"

图7

IbMAPKK9蛋白序列比对及亲缘关系 A: IbMAPKK9氨基酸序列比对图。橙线指示蛋白激酶家族保守结构域; B: IbMAPKK9蛋白进化树。IbMAPKK9用红星标注。Ib: 甘薯; It: 三叶裂薯(XP_031110493.1); In: 日本牵牛花(XP_019189439.1); Nb: 本氏烟草(BAE95414.1); Ds: 曼陀罗(MCD7449465.1); Na: 野生烟草(XP_019267338.1); St: 马铃薯(XP_006363984.1); Rs: 蔷薇(ALG02506.1); Cb: 风铃辣椒(PHT53658.1); Tw: 雷公藤(XP_038686910.1); Sl: 番茄(NP_001234595.1); La: 狭叶羽扇豆(XP_019431663.1)。"

图8

甘薯IbMAPKK9与拟南芥AtMAPKs的系统进化树"

图9

IbMAPKK9的蛋白质互作网络"

表4

STRING预测甘薯IbMAPKK9互作蛋白基本信息"

名称
Protein name		 编号
Protein ID		 注释
Protein annotation		 蛋白长度
Protein length (aa)		 置信度
Confidence
MPK3.0 A0A1J6KJD9 丝裂原活化蛋白激酶MAPK3
Mitogen-activated protein kinase 3		 471 0.999
MPK3.1 A0A1J6IKA5 丝裂原活化蛋白激酶MAPK3
Mitogen-activated protein kinase 3		 392 0.999
MPK3.2 A5H2L1 丝裂原活化蛋白激酶(Ser/Thr激酶家族)
Mitogen activated protein kinases (Ser/Thr protein kinase family)		 375 0.999
NTF4.0 A0A314KXT0 丝裂原活化蛋白激酶(Ser/Thr激酶家族)
Mitogen activated protein kinases(Ser/Thr protein kinase family)		 394 0.988
NTF4.1 A5H2L0 丝裂原活化蛋白激酶(Ser/Thr激酶家族)
Mitogen activated protein kinases (Ser/Thr protein kinase family)		 393 0.993
MEKK1.0 A0A314L1U5 丝裂原活化蛋白激酶激酶激酶MAPKKK1
Mitogen-activated protein kinase kinase kinase 1		 637 0.980
MEKK1.1 A0A1J6IHU2 丝裂原活化蛋白激酶激酶激酶MAPKKK1
Mitogen-activated protein kinase kinase kinase 1		 561 0.980
NPK1.6 A0A314L7I4 丝裂原活化蛋白激酶激酶
Mitogen-activated protein kinase kinase		 668 0.938
CTR1.1 A0A314KLJ0 丝氨酸蛋白激酶
Serine kinase		 161 0.936
CTR1.8 A0A1J6IUT7 丝氨酸蛋白激酶
Serine kinase		 827 0.936

表5

IbMAPKK9蛋白亚细胞定位预测"

网站
Software		 细胞质
Chloroplast		 细胞核
Nuclear		 线粒体
Mitochondrion		 细胞骨架
Cytoskelrtal
WoLF PSORT — 9 1 3
PSORT II 4.3% 69.6% 21.7% 4.3%

图10

IbMAPKK9的亚细胞定位 A: 空GFP合并图; B: 空GFP荧光; C: IbMAPKK9-GFP合并图; D: IbMAPKK9-GFP荧光。"

图11

IbMAPKK9基因在不同组织的表达 不同大写字母代表在P < 0.01水平差异显著。"

图12

蔓割病病原菌侵染下不同时期IbMAPKK9基因的相对表达量 不同大写字母代表在P < 0.01水平差异显著。"

图13

IbMAPKK9基因在烟草表皮细胞中的瞬时表达 A: 空GFP明场; B: 空GFP荧光; C: 空GFP DAPI荧光染料; D: 空GFP合并图; E: IbMAPKK9-GFP明场; F: IbMAPKK9-GFP荧光; G: IbMAPKK9-GFP DAPI荧光染料; H: IbMAPKK9-GFP合并图。"

图14

烟草NtPAL4、NtICS1、NtNPR1、NtNPR3和NtNPR5的表达分析"

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