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作物学报 ›› 2018, Vol. 44 ›› Issue (9): 1367-1379.doi: 10.3724/SP.J.1006.2018.01367

• 研究论文 • 上一篇    下一篇

甘蔗ScWRKY4基因的克隆与表达特性分析

王玲1(),刘峰1,戴明剑1,孙婷婷1,苏炜华1,王春风1,张旭1,毛花英1,苏亚春1,2,*(),阙友雄1,2,*()   

  1. 1 福建农林大学 / 农业部福建甘蔗生物学与遗传育种重点实验室, 福建福州 350002
    2 福建农林大学 / 教育部作物遗传育种与综合利用重点实验室, 福建福州 350002
  • 收稿日期:2018-01-28 接受日期:2018-06-12 出版日期:2018-09-10 网络出版日期:2018-07-02
  • 通讯作者: 苏亚春,阙友雄
  • 基金资助:
    本研究由国家自然科学基金项目(31671752和31101196)(31671752);本研究由国家自然科学基金项目(31671752和31101196)(31101196);福建省杰出青年基金项目(2015J06006);福建省高校杰出青年科研人才计划项目(苏亚春-2017)(SYC-2017);国家现代农业产业技术体系建设专项(CARS-17)资助

Cloning and Expression Characteristic Analysis of ScWRKY4 Gene in Sugarcane

Ling WANG1(),Feng LIU1,Ming-Jian DAI1,Ting-Ting SUN1,Wei-Hua SU1,Chun-Feng WANG1,Xu ZHANG1,Hua-Ying MAO1,Ya-Chun SU1,2,*(),You-Xiong QUE1,2,*()   

  1. 1 Key Laboratory of Sugarcane Biology and Genetic Breeding (Fujian), Ministry of Agriculture / Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
    2 Key Laboratory of Crop Genetics and Breeding and Comprehensive Utilization, Ministry of Education / Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China;
  • Received:2018-01-28 Accepted:2018-06-12 Published:2018-09-10 Published online:2018-07-02
  • Contact: Ya-Chun SU,You-Xiong QUE
  • Supported by:
    This study was supported by the National Natural Science Foundation of China(31671752);This study was supported by the National Natural Science Foundation of China(31101196);Natural Science Foundation of Fujian Province for Distinguished Young Scholars(2015J06006);Research Funds for Distinguished Young Scientists in Fujian Provincial Department of Education(SYC-2017);China Agriculture Research System (CARS-17)

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摘要：

WRKY是植物中特有的转录因子家族之一, 在植物对生物和非生物逆境胁迫的应答反应中起重要调控作用。本研究基于课题组前期构建的甘蔗(Saccharum spp.)转录组数据库, 从新台糖22 (ROC22)中成功克隆到1个WRKY基因, 命名为ScWRKY4 (GenBank登录号为MG852087)。序列分析发现, ScWRKY4基因cDNA全长1265 bp, 包含1个741 bp的完整开放读码框, 编码246个氨基酸, 该蛋白具有1个WRKYGQK保守结构域和C2H2锌指结构域, 属于IIc类WRKY转录因子。生物信息学预测分析发现, ScWRKY4蛋白为碱性的不稳定亲水性蛋白, 不存在信号肽和跨膜结构域, 蛋白二级结构元件缺少β螺旋。在本氏烟(Nicotiana benthamiana)叶片瞬时表达中, ScWRKY4蛋白定位于细胞核。酵母杂交实验结果显示, ScWRKY4不具有转录激活活性。实时荧光定量PCR分析表明, ScWRKY4基因在甘蔗的根、叶、芽和皮中的表达量无明显差异, 在蔗肉中的表达量最高, 为对照蔗根的18.38倍; 黑穗病菌侵染0~72 h, ScWRKY4在抗病品种崖城05-179中下调表达, 在感病品种ROC22中表达较稳定; 受到外源激素脱落酸、水杨酸和茉莉酸甲酯以及非生物胁迫因子氯化钠和聚乙二醇胁迫后, ScWRKY4基因均被诱导上调表达。上述研究结果表明, ScWRKY4基因可能不参与甘蔗对黑穗病的抗性反应或在该防御方面起负调控作用, 但积极响应甘蔗对盐和干旱胁迫的应答。

关键词: 甘蔗, WRKY转录因子, 生物信息学, 亚细胞定位, 转录激活活性, 实时荧光定量PCR

Abstract:

WRKY is one of the specific transcriptional regulators in plants and widely involved in the response of plants to diverse biotic and abiotic stresses. In this study, based on our previous transcriptome data of sugarcane (Saccharum spp.), we successfully cloned a WRKY gene from sugarcane variety ROC22, named as ScWRKY4 (GenBank accession number MG852087). Sequence analysis showed that the full-length cDNA of ScWRKY4 gene was 1265 bp, containing a 741 bp complete open reading frame and encoding 246 amino acids residues. ScWRKY4 protein had a conservative WRKYGQK domain and a zinc finger motif C2H2, belonging to IIc subgroup of WRKY transcription factor family. Bioinformatics analysis indicated that ScWRKY4 was an alkaline unstable hydrophilic protein with no signal peptide and transmembrane structure. The secondary structural element of ScWRKY4 protein was lack of beta helix structure. ScWRKY4 was located in the nucleus after transient expression in Nicotiana benthamiana. The result of yeast hybridization experiment demonstrated that ScWRKY4 did not possess transcriptional activity. Real-time fluorescent quantitative PCR showed that the expression of ScWRKY4 gene had no significant difference in root, leaf, bud and stem epidermis, while had the highest expression level in stem pith, which was 18.38 times higher than that in root. During inoculation with smut pathogen (Sporisorium scitamineum) at 0-72 h, the gene expression level of ScWRKY4 was down-regulated in sugarcane smut-resistant cultivar Yacheng 05-179, while was more stable in the susceptible one ROC22. The transcript of ScWRKY4 was up-regulated by exogenous hormone stresses of abscisic acid, salicylic acid, and methyl jasmonate, as well as by the abiotic treatments of sodium chloride and polyethylene glycol. These results suggest that ScWRKY4 gene may not participate in sugarcane smut resistance, or only play a negative role in this defense reaction, but positively involve in the response mechanism of salt and drought resistance in sugarcane.

Key words: sugarcane, WRKY transcription factor, bioinformatics, subcellular localization, transcriptional activity, real-time flourescent quantitative PCR

表1

引物序列及用途"

引物名称
Primer name		 引物序列
Primer sequence (5°-3°)		 用途
Usage
ScWRKY4-F TCCTCGGCATCTCCCATTCT 扩增全长
Full length amplifcation
ScWRKY4-R GTCTGGGAGCTCATGTTCGT
ScWRKY4-QF ATGAAGGTGAGGAGGAAGATG 荧光定量
qRT-PCR
ScWRKY4-QR CTTGTAGCCGTCGTCCAG
GAPDH-QF CACGGCCACTGGAAGCA 内参基因
Reference gene
GAPDH-QR TCCTCAGGGTTCCTGATGCC
ScWRKY4-Subloc-F CGGGATCCATGGAGGGGAGCA 亚细胞定位
Subcellular localization
ScWRKY4-Subloc-R TGCTCTAGAGAGCGACGTGAAAGC
ScWRKY4-BD-F GGAATTCCATATGATGGAGGGGAGCAGCCAGCT 转录激活
Transactivation
ScWRKY4-BD-R CGGGATCCGAGCGACGTGAAAGCGCAGC

图1

甘蔗ScWRKY4基因的PCR扩增 M: 15000+2000 bp DNA marker; 1: PCR产物。"

图2

PCR扩增获得的ScWRKY4基因的核酸序列及其推导的氨基酸序列(*: 终止密码子) 红色框部分为WRKYGQK基序; 黑色框部分为C2H2基序(CX4CX23HXH)。"

图3

甘蔗ScWRKY4蛋白的保守结构域分析"

图4

甘蔗ScWRKY4与其他植物WRKY蛋白的氨基酸序列比对高粱: SbWRKY57 (XP_002448267.2); 玉米: ZmWRKY36 (NP_001151912.1); II型少花古尔德草: DoWWRKY12 (OEL21317.1); 谷子: SiWRKY12 (XP_004956420.1); 二穗短柄草: BdWRKY12 (XP_014751531.1); 水稻: OsWRKY12 (XP_015635033.1)。"

图5

ScWRKY4蛋白与甘蔗及其他植物WRKY蛋白家族的系统进化树"

图6

甘蔗ScWRKY4亚细胞定位分析(A) 35S::ScWRKY4::GFP亚细胞定位重组载体的酶切验证结果。1: 15000+2000 bp DNA marker; 2: 35S::GFP/Xba I; 3: ScWRKY4 ORF PCR产物; 4: 35S::ScWRKY4::GFP/Xba I; 5: 35S::ScWRKY4::GFP/Xba I+BamH I; 6: 100 bp ladder DNA marker。(B) 农杆菌介导转化的ScWRKY4及空载体注射本氏烟叶片48 h的亚细胞定位结果。本氏烟叶片表皮细胞被用于明场、绿色荧光、蓝色荧光、明场和绿色及蓝色荧光叠加后的图像分析; 白色箭头1、2和3分别表示质膜、细胞核和细胞质, 比例尺=25 μm; 35S::GFP: 携带空载pCAMBIA 1300-GFP的农杆菌菌株; 35S::ScWRKY4::GFP: 携带重组载体pCAMBIA 1300-ScWRKY4-GFP的农杆菌菌株; DAPI: 4’,6-二脒基-2-苯基吲哚。"

图7

甘蔗ScWRKY4转录激活活性分析(A) pGBKT7-ScWRKY4重组载体的酶切验证结果。1: 15000+2000 bp DNA marker; 2: pGBKT7/Nde I; 3: ScWRKY4 ORF PCR产物; 4: pGBKT7-ScWRKY4/Nde I; 5: pGBKT7-ScWRKY4/Nde I+BamH I; 6: 2000 bp ladder DNA marker。(B) ScWRKY4转录激活活性验证。SD/-Trp: 色氨酸营养缺陷型平板培养基; SD/-Trp (+ X-α-Gal): 色氨酸营养缺陷型平板培养基(添加5-溴-4-氯-3-吲哚-α-D-半乳糖苷); SD/-Trp (+ X-α-Gal+AbA): 色氨酸营养缺陷型平板培养基(添加5-溴-4-氯-3-吲哚-α-D-半乳糖苷和金担子素A)。"

图8

ScWRKY4基因在甘蔗ROC22组织中的表达情况柱上不同的小写字母代表显著性的差异(P ≤ 0.05); n = 3; 内参基因为甘油醛-3-磷酸脱氢酶(glyceraldehyde-3-phosphate dehydrogenase, GAPDH); R: 蔗根; B: 蔗芽; L: 蔗叶; SP: 蔗肉; SE: 蔗皮。"

图9

甘蔗ScWRKY4基因在黑穗病侵染下的表达情况 Yacheng 05-179: 甘蔗抗黑穗病品种; ROC22: 甘蔗感黑穗病品种。柱上不同的小写字母代表显著性的差异(P ≤ 0.05); n = 3; 内参基因为甘油醛-3-磷酸脱氢酶(glyceraldehyde-3-phosphatedehydrogenase, GAPDH)。"

图10

甘蔗ScWRKY4基因在不同植物激素和非生物胁迫下的表达水平柱上不同的小写字母代表显著性的差异(P ≤ 0.05); n = 3; 内参基因为甘油醛-3-磷酸脱氢酶(glyceraldehyde-3-phosphate dehydrogenase, GAPDH); SA: 水杨酸(5 mmol L-1); MeJA: 茉莉酸甲酯(25 μmol L-1); NaCl: 氯化钠(模拟盐胁迫) (250 mmol L-1); PEG: 聚乙二醇(模拟干旱胁迫) (25.0%); ABA: 脱落酸(100 μmol L-1)。"

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