大白菜BrWRKY33基因及其上游调控序列的克隆与功能研究
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摘要
WRKY转录因子是一种植物特有的转录因子,WRKY33属于Ⅰ类WRKY蛋白,其WRKY结构域高度保守。大部分WRKY蛋白定位于细胞核,并可能通过蛋白之间的相互作用形成同源二聚体或异源二聚体而调控靶基因的表达。WRKY转录因子调控机制很复杂,当前研究的焦点聚集在WRKY转录因子调控防卫反应中下游抗病基因表达的机制,本实验研究大白菜BrWRKY33基因在软腐病抗病方面的作用,并对其上游调控序列功能进行了初步研究。
利用本试验室获得的大白菜抗软腐病侵染的抑制性差减杂交(Suppression SubtractiveHybridization,SSH)cDNA文库,筛选出与WRKY33具有高度同源性的7个EST,使用PCR方法从大白菜基因组中克隆了BrWRKY33基因编码区。长度为1443bp,编码480个氨基酸。
RT-PCR分析表明在Ecc侵染大白菜24h时间内,BrWRKY33基因在侵染后6h开始表达,随着侵染时间延长,BrWRKY33基因表达量在24h积累到最高。
对拟南芥突变体、野生型、BrWRKY33过表达植株进行软腐病抗病性鉴定,分别在接种后6h、12h、24h三个时间点进行病情统计,接种Ecc后6h,病情指数集中在53.6%～65.3%,12h集中在63.4%～74.8%,24h集中在90.9%～98.7%。统计结果显示,过表达株系相比突变体,组间抗病差异显著;过表达株系相比野生型,组间抗病差异不明显。
RT-PCR方法检测目的基因BrWRKY33在拟南芥中表达,SA途径的标志基因PR1、BGL-2的表达量,突变体高于野生型,说明WRKY33在SA路径中可能起负调控作用;JA途径的标志基因AOS的表达量,无明显差异;JA/ET途径的标志基因PDF1.2表达量,突变体低于野生型,野生型低于过表达株系1、3、5、6。说明BrWRKY33在拟南芥抗软腐病JA/ET介导的防御反应中,可能起正调控作用。
通过PLACE和fruitfly网站预测,结合已经克隆的BrWRKY33,确定上游调控序列分段区域,并构建植物表达载体,进行拟南芥遗传转化研究。Brw33p1、Brw33p2转基因株系GUS染色的差别,说明BrWRKY33基因启动子区的W box功能可能与AtWRKY18基因相同,起负调控作用。拟南芥转Brw33p2植株,表现出了更强的GUS活性,推断起始密码子上游-315bp区域含有BrWRKY33基因转录必需的基本元件,如CAAT box和TATA box等。拟南芥叶片接种Ecc 2h后,转Brw33p1植株接种的叶片比未接种的叶片染色更深,说明接种后诱导转基因植株gus基因表达量增加,BrWRKY33基因在植物对软腐病抗性反应中具有一定的作用。
WRKY transcription factor is a plant-specific transcription factor,and WRKY33 belongs to WRKY protein group I,which WRKY domain highly conserved.Most of WRKY protein are located in the nucleus,and can form a hetero or homologous dimers by the interactions between proteins to regulate the expression of target genes.The regulation of transcription factor WRKY are complex,and how to regulate its downstream target gene becomes the hotspot in current study,the function of transcription factor gene BrWRKY33 and its upstream regulatory sequence are analyzed and discussed.
The SSH cDNA library of Chinese cabbage(Brassica rapa subp.pekinensis cv.longbaiⅡ) inoculating Ecc(Erwinia carotovora subsp,carotovora,Ecc) is obtained,and select highly homologous,then cloned the coding region of BrWRKY33 from the Chinese cabbage genome by PCR,which have a length of 1443bp,encoding 480 amino acids.
RT-PCR analysis shows that the BrWRKY33 begins to express in 6h after Chinese cabbage inoculating Ecc,and the gene has a highest accumulation in 24hpi as the infection time grows.
Analyse the resistance identification of Arabidopsis mutant,wild-type and BrWRKY33 over-expression plants respectively,and get the disease statistics after inoculate Ecc 6hpi,12hpi, 24hpi.Disease index mainly lies in 53.6%～65.3%after inoculate with Ecc 6hpi,63.4%～74.8% after 12hpi,90.9%～98.7%after 24hpi.The statistics show that the difference obvious between transgenic plants and mutant ones in Ecc disease-resistant,and has not significant difference between transgenic plants and wild-type.
The expression of BrWRKY33 in Arabidopsis is detected through RT-PCR,PR1 and BGL-2 are defense-regulated genes which associated with SA-regulated defense responses,PR1 has an enhanced expression in mutant-1 compared to wild-type,which suggest that the WRKY33 may have a negetive effect on SA;while the expression of JA-regulated defense genes AOS,have on difference in basically expression;Interestingly PDF1.2 which associated with JA/ET-regulated defense responses,have a lower expression in mutant compared to wild-type,but have a higher expression in over-expression lines 1,3,5,6 compared to wild-type.These results show that BrWRKY33 may have a positive regulation in the Arabidopsis JA/ET-mediated defense response to Ecc.
A 1755bp up-stream regulatory sequence of transcription factor gene BrWRKY33 was amplified by PCR from Chinese cabbage(Brassica rapa subp.pekinensis cv.longbaiⅡ). Deletion mutant with a length of 315bp of the up-stream regulatory sequence was then constructed and transformed into Arabidopsis thaliana.The results indicated that the presence of some W box sequences in -1755～315bp region upstream the start codon was related to the down regulation of the gene expression,and the -315bp region contains the essential sequence for gene transcription.The difference in GUS staining in transgenic plants after and before inoculating with soft rot bacteria suggested that WRKY33 gene play a role in the resistance against soft rot disease.
利用本试验室获得的大白菜抗软腐病侵染的抑制性差减杂交(Suppression SubtractiveHybridization,SSH)cDNA文库,筛选出与WRKY33具有高度同源性的7个EST,使用PCR方法从大白菜基因组中克隆了BrWRKY33基因编码区。长度为1443bp,编码480个氨基酸。
RT-PCR分析表明在Ecc侵染大白菜24h时间内,BrWRKY33基因在侵染后6h开始表达,随着侵染时间延长,BrWRKY33基因表达量在24h积累到最高。
对拟南芥突变体、野生型、BrWRKY33过表达植株进行软腐病抗病性鉴定,分别在接种后6h、12h、24h三个时间点进行病情统计,接种Ecc后6h,病情指数集中在53.6%～65.3%,12h集中在63.4%～74.8%,24h集中在90.9%～98.7%。统计结果显示,过表达株系相比突变体,组间抗病差异显著;过表达株系相比野生型,组间抗病差异不明显。
RT-PCR方法检测目的基因BrWRKY33在拟南芥中表达,SA途径的标志基因PR1、BGL-2的表达量,突变体高于野生型,说明WRKY33在SA路径中可能起负调控作用;JA途径的标志基因AOS的表达量,无明显差异;JA/ET途径的标志基因PDF1.2表达量,突变体低于野生型,野生型低于过表达株系1、3、5、6。说明BrWRKY33在拟南芥抗软腐病JA/ET介导的防御反应中,可能起正调控作用。
通过PLACE和fruitfly网站预测,结合已经克隆的BrWRKY33,确定上游调控序列分段区域,并构建植物表达载体,进行拟南芥遗传转化研究。Brw33p1、Brw33p2转基因株系GUS染色的差别,说明BrWRKY33基因启动子区的W box功能可能与AtWRKY18基因相同,起负调控作用。拟南芥转Brw33p2植株,表现出了更强的GUS活性,推断起始密码子上游-315bp区域含有BrWRKY33基因转录必需的基本元件,如CAAT box和TATA box等。拟南芥叶片接种Ecc 2h后,转Brw33p1植株接种的叶片比未接种的叶片染色更深,说明接种后诱导转基因植株gus基因表达量增加,BrWRKY33基因在植物对软腐病抗性反应中具有一定的作用。
WRKY transcription factor is a plant-specific transcription factor,and WRKY33 belongs to WRKY protein group I,which WRKY domain highly conserved.Most of WRKY protein are located in the nucleus,and can form a hetero or homologous dimers by the interactions between proteins to regulate the expression of target genes.The regulation of transcription factor WRKY are complex,and how to regulate its downstream target gene becomes the hotspot in current study,the function of transcription factor gene BrWRKY33 and its upstream regulatory sequence are analyzed and discussed.
The SSH cDNA library of Chinese cabbage(Brassica rapa subp.pekinensis cv.longbaiⅡ) inoculating Ecc(Erwinia carotovora subsp,carotovora,Ecc) is obtained,and select highly homologous,then cloned the coding region of BrWRKY33 from the Chinese cabbage genome by PCR,which have a length of 1443bp,encoding 480 amino acids.
RT-PCR analysis shows that the BrWRKY33 begins to express in 6h after Chinese cabbage inoculating Ecc,and the gene has a highest accumulation in 24hpi as the infection time grows.
Analyse the resistance identification of Arabidopsis mutant,wild-type and BrWRKY33 over-expression plants respectively,and get the disease statistics after inoculate Ecc 6hpi,12hpi, 24hpi.Disease index mainly lies in 53.6%～65.3%after inoculate with Ecc 6hpi,63.4%～74.8% after 12hpi,90.9%～98.7%after 24hpi.The statistics show that the difference obvious between transgenic plants and mutant ones in Ecc disease-resistant,and has not significant difference between transgenic plants and wild-type.
The expression of BrWRKY33 in Arabidopsis is detected through RT-PCR,PR1 and BGL-2 are defense-regulated genes which associated with SA-regulated defense responses,PR1 has an enhanced expression in mutant-1 compared to wild-type,which suggest that the WRKY33 may have a negetive effect on SA;while the expression of JA-regulated defense genes AOS,have on difference in basically expression;Interestingly PDF1.2 which associated with JA/ET-regulated defense responses,have a lower expression in mutant compared to wild-type,but have a higher expression in over-expression lines 1,3,5,6 compared to wild-type.These results show that BrWRKY33 may have a positive regulation in the Arabidopsis JA/ET-mediated defense response to Ecc.
A 1755bp up-stream regulatory sequence of transcription factor gene BrWRKY33 was amplified by PCR from Chinese cabbage(Brassica rapa subp.pekinensis cv.longbaiⅡ). Deletion mutant with a length of 315bp of the up-stream regulatory sequence was then constructed and transformed into Arabidopsis thaliana.The results indicated that the presence of some W box sequences in -1755～315bp region upstream the start codon was related to the down regulation of the gene expression,and the -315bp region contains the essential sequence for gene transcription.The difference in GUS staining in transgenic plants after and before inoculating with soft rot bacteria suggested that WRKY33 gene play a role in the resistance against soft rot disease.
引文
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