水稻和拟南芥TFⅢA型锌指蛋白基因家族的鉴定及ZFP207基因的功能研究
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摘要
水稻是世界上最重要的粮食作物之一,低温、干旱、高盐等非生物胁迫严重制约了水稻生长和高产。水稻在受到非生物逆境胁迫时,体内发生了一系列从细胞到生理水平的反应,在这些反应过程中,主要有两类基因在起作用:一类是直接保护细胞免受逆境胁迫的功能蛋白以及渗透调节因子;另一类是参与胁迫信号传导和表达调控的蛋白激酶与转录因子。锌指蛋白是植物体内一类数量庞大的转录因子,根据其功能的不同,可以分为TFⅢA、WRKY、DOF、GATA、RING和PHD等亚家族。TFⅢA型锌指蛋白的锌指结构域具有CX2CX3FX3QALGGHX3H的序列特征。目前在植物中已经分离了近40个此类基因,这些基因主要参与调控植物的生长发育和胁迫应答反应。
利用TFⅢA型锌指蛋白家族中已知基因的氨基酸序列作为探针,搜索植物基因组数据库(tBLASTn),在水稻和拟南芥中分别鉴定获得72个和59个TFⅢA型锌指蛋白基因序列。这些基因所编码的氨基酸序列中,大部分具有单锌指和双锌指结构。染色体定位(In silicon)结果显示这些基因分别散布在水稻的12条染色体和拟南芥的5条染色体上,有些基因在染色体上串联排列。将鉴定的水稻和拟南芥TFⅢA型锌指蛋白与其他植物中已报道的TFⅢA型锌指蛋白一起进行系统发生分析,可将植物的TFⅢA型锌指蛋白划分为三个亚家族:亚家族Ⅰ中的大部分成员都具有单锌指结构,已经报道的基因大都参与调控植物的生长发育,亚家族Ⅱ中的大部分成员都具有双锌指结构,已经报道的基因大都参与植物的非生物胁迫应答反应,亚家族Ⅲ中的大部分成员具有3-5个锌指结构,已经报道的基因可能参与调控植物生长发育或其他一些代谢过程。利用cDNA芯片技术分析水稻TFⅢA型锌指蛋白基因在低温和干旱胁迫、及H_2O_2处理下的表达模式,得到其中45个基因的相关数据,45个基因中至少有27个基因在处理后表达量变化2倍以上,其中14个基因应答H_2O_2处理,8个基因受到三种处理的调控。
利用生物信息学和RT-PCR技术从水稻盐处理幼苗cDNA中分离了一个受低温和干旱胁迫诱导的,具有单个锌指结构的锌指蛋白基因ZFP207。该基因编码188个氨基酸残基,预测分子量为20.72KDa,具有一个植物特有QALGGH基序的C2H2型锌指结构域,不具有其它TFⅢA型锌指蛋白常有的结构域,如NLS,L-box,DLN-box/EAR-motif等。将ZFP207与GFP报告基因融合后,转化洋葱表皮细胞,发现ZFP207-GFP定位于洋葱表皮细胞的细胞核内;实时荧光定量PCR测定结果表明,ZFP207在水稻幼苗中的表达受到低温、高盐、干旱等多种逆境胁迫及ABA的诱导。构建了35S:ZFP207植物表达载体,利用农杆菌介导法转化到烟草,获得了ZFP207的转基因烟草植株。选取两个转基因株系(2#、4#)进行抗逆性分析,发现转ZFP207的烟草植株对低温、高盐、干旱等非生物胁迫的抗性明显增强,对ABA抑制发芽变得不敏感。进一步测定比较了转ZFP207的烟草植株与未转基因的野生型植株的脯氨酸含量,发现转基因植株中的脯氨酸含量明显高于野生型;转基因植株中烟草渗调蛋白基因Osmotin的表达量也显著增强,但转基因植株中3个LEA基因的表达没有明显变化。ZFP207是植物中第一个报道的与逆境胁迫相关的TFⅢA型单锌指基因,对ZFP207更多的研究将有助于了解植物TFⅢA型锌指蛋白结构与功能之间的关系。
Rice is one of the most important crops worldwide,but rice growth and productivity are largely affected by abiotic stresses,including low temperature,drought and high salinity. A series of changes from cellular to physiological levels occur after rice plants are stressed. These genes in responses to abiotic stresses can be classified into two groups:one encoding the function proteins and osmoprotectants directly to protect plant cells against stresses, another encoding protein kinases and transcription factors,functioning in signal transduction and regulating gene expression.The zinc finger proteins represent a large transcription factor family in plant.It could be divided into several subfamilies including TFⅢA,WRKY,DOF,GATA,RING,and PHD,based on the various functions.Recently, many researches focused on the TFⅢA-type zinc finger genes with a conserved sequence "CX2CX3FX3QALGGHX3H" in zinc finger domains.About 40 TFⅢA-type zinc finger proteins have been isolate from plant,most of them play important roles in the developmental processes or responses to abiotic stresses in plant.
In order to identify TFⅢA-type zinc finger genes from rice(Oryza sativa L.) and Arabidopsis,the amino acid sequence of several genes reported in this family,were used as query probes to search plant genome database through tBLASTn algorithm program.It was totally identified 72 and 59 of TFⅢA-type zinc finger protein genes in rice and Arabidopsis respectively.Analysis of the deduced amino acid sequences showed that most of these proteins encoded one or two zinc finger domains.These genes,in silicon,were randomly distributed on the 12 rice chromosomes and 5 Arabidopsis chromosomes respectively,with a few genes clustered.Phylogenetic analysis of these proteins and other reported proteins in plants showed that plant TFⅢA-type zinc finger proteins could he classified them as three subfamilies based on the numbers of zinc finger domains contained: SubfamilyⅠ,most of members with single zinc finger domain and the reported genes mostly involved in plant growth and development,SubfamilyⅡ,most of members with two zinc finger domains and the reported genes mostly related to responses to abiotic stresses;and SubfamilyⅢ,most of members with 3-5 zinc finger domains and the reported proteins involved in plant growth,development and other key metabolisms.Through a cDNA microarray technique the expression patterns of rice TFⅢA-type zinc finger genes were screened in rice seedlings under cold,drought and H_2O_2 treatments.We obtained the data of 45 genes.To compare with control the expressions of at least 27 genes were changed with more than 2-fold.Among them 14 were responded to H_2O_2,and 8 genes to all three treatments.
One single zinc finger protein gene,named ZFP207,was isolated from the cDNA of rice seedlings treated by salt stress by silicon cloning and RT-PCR approach.ZFP207 encodes 188 amino acids with the calculated molecular mass of 20.72 KDa.It contains a Cys2/His2-type zinc finger domain with the plant specific "QALGGH" motifs.However,in the ZFP207 there was no other domains such as NLS,L-box,DLN-box/EAR-motif discovered in most plant TFⅢA-type zinc finger proteins.ZFP207,fused with the green fluorescent protein(GFP) reporter gene,was introduced into onion epidermis cells.The ZFP207-GFP fusion protein was targeted to the nucleus.By real-time RT-PCR the inducement of ZFP207 under treatments with salt,drought,cold stress and ABA were evaluated.We constructed the 35S:ZFP207 vector and transferred it into tobacco plants by in vivo infiltration,and then we obtained the ZFP207-overexpression transgenic plants.The 2# and 4# transgenic lines were used for stresses tolerances assay.The results indicated that ZFP207-overexpression transgenic lines increased significantly cold,drought and high salt stresses tolerance compared to the WT plants,but the germination of ZFP207-overexpressed seeds is insensitive to ABA.In addition,the proline contents in transgenic plants were obviously higher than that in WT plants under normal condition. And the expression of Osmotin enhanced in transgenic lines,while the expressions of 3 LEA genes showed no change in transgenic plants.Up to date,ZFP207 is the first reported TFⅢA-type single zinc finger gene involved in abiotic stresses in plants.More analysis of ZFP207 will help to widen our understanding of the relation between the structures and functions of TFⅢA-type zinc finger proteins in plants.
利用TFⅢA型锌指蛋白家族中已知基因的氨基酸序列作为探针,搜索植物基因组数据库(tBLASTn),在水稻和拟南芥中分别鉴定获得72个和59个TFⅢA型锌指蛋白基因序列。这些基因所编码的氨基酸序列中,大部分具有单锌指和双锌指结构。染色体定位(In silicon)结果显示这些基因分别散布在水稻的12条染色体和拟南芥的5条染色体上,有些基因在染色体上串联排列。将鉴定的水稻和拟南芥TFⅢA型锌指蛋白与其他植物中已报道的TFⅢA型锌指蛋白一起进行系统发生分析,可将植物的TFⅢA型锌指蛋白划分为三个亚家族:亚家族Ⅰ中的大部分成员都具有单锌指结构,已经报道的基因大都参与调控植物的生长发育,亚家族Ⅱ中的大部分成员都具有双锌指结构,已经报道的基因大都参与植物的非生物胁迫应答反应,亚家族Ⅲ中的大部分成员具有3-5个锌指结构,已经报道的基因可能参与调控植物生长发育或其他一些代谢过程。利用cDNA芯片技术分析水稻TFⅢA型锌指蛋白基因在低温和干旱胁迫、及H_2O_2处理下的表达模式,得到其中45个基因的相关数据,45个基因中至少有27个基因在处理后表达量变化2倍以上,其中14个基因应答H_2O_2处理,8个基因受到三种处理的调控。
利用生物信息学和RT-PCR技术从水稻盐处理幼苗cDNA中分离了一个受低温和干旱胁迫诱导的,具有单个锌指结构的锌指蛋白基因ZFP207。该基因编码188个氨基酸残基,预测分子量为20.72KDa,具有一个植物特有QALGGH基序的C2H2型锌指结构域,不具有其它TFⅢA型锌指蛋白常有的结构域,如NLS,L-box,DLN-box/EAR-motif等。将ZFP207与GFP报告基因融合后,转化洋葱表皮细胞,发现ZFP207-GFP定位于洋葱表皮细胞的细胞核内;实时荧光定量PCR测定结果表明,ZFP207在水稻幼苗中的表达受到低温、高盐、干旱等多种逆境胁迫及ABA的诱导。构建了35S:ZFP207植物表达载体,利用农杆菌介导法转化到烟草,获得了ZFP207的转基因烟草植株。选取两个转基因株系(2#、4#)进行抗逆性分析,发现转ZFP207的烟草植株对低温、高盐、干旱等非生物胁迫的抗性明显增强,对ABA抑制发芽变得不敏感。进一步测定比较了转ZFP207的烟草植株与未转基因的野生型植株的脯氨酸含量,发现转基因植株中的脯氨酸含量明显高于野生型;转基因植株中烟草渗调蛋白基因Osmotin的表达量也显著增强,但转基因植株中3个LEA基因的表达没有明显变化。ZFP207是植物中第一个报道的与逆境胁迫相关的TFⅢA型单锌指基因,对ZFP207更多的研究将有助于了解植物TFⅢA型锌指蛋白结构与功能之间的关系。
Rice is one of the most important crops worldwide,but rice growth and productivity are largely affected by abiotic stresses,including low temperature,drought and high salinity. A series of changes from cellular to physiological levels occur after rice plants are stressed. These genes in responses to abiotic stresses can be classified into two groups:one encoding the function proteins and osmoprotectants directly to protect plant cells against stresses, another encoding protein kinases and transcription factors,functioning in signal transduction and regulating gene expression.The zinc finger proteins represent a large transcription factor family in plant.It could be divided into several subfamilies including TFⅢA,WRKY,DOF,GATA,RING,and PHD,based on the various functions.Recently, many researches focused on the TFⅢA-type zinc finger genes with a conserved sequence "CX2CX3FX3QALGGHX3H" in zinc finger domains.About 40 TFⅢA-type zinc finger proteins have been isolate from plant,most of them play important roles in the developmental processes or responses to abiotic stresses in plant.
In order to identify TFⅢA-type zinc finger genes from rice(Oryza sativa L.) and Arabidopsis,the amino acid sequence of several genes reported in this family,were used as query probes to search plant genome database through tBLASTn algorithm program.It was totally identified 72 and 59 of TFⅢA-type zinc finger protein genes in rice and Arabidopsis respectively.Analysis of the deduced amino acid sequences showed that most of these proteins encoded one or two zinc finger domains.These genes,in silicon,were randomly distributed on the 12 rice chromosomes and 5 Arabidopsis chromosomes respectively,with a few genes clustered.Phylogenetic analysis of these proteins and other reported proteins in plants showed that plant TFⅢA-type zinc finger proteins could he classified them as three subfamilies based on the numbers of zinc finger domains contained: SubfamilyⅠ,most of members with single zinc finger domain and the reported genes mostly involved in plant growth and development,SubfamilyⅡ,most of members with two zinc finger domains and the reported genes mostly related to responses to abiotic stresses;and SubfamilyⅢ,most of members with 3-5 zinc finger domains and the reported proteins involved in plant growth,development and other key metabolisms.Through a cDNA microarray technique the expression patterns of rice TFⅢA-type zinc finger genes were screened in rice seedlings under cold,drought and H_2O_2 treatments.We obtained the data of 45 genes.To compare with control the expressions of at least 27 genes were changed with more than 2-fold.Among them 14 were responded to H_2O_2,and 8 genes to all three treatments.
One single zinc finger protein gene,named ZFP207,was isolated from the cDNA of rice seedlings treated by salt stress by silicon cloning and RT-PCR approach.ZFP207 encodes 188 amino acids with the calculated molecular mass of 20.72 KDa.It contains a Cys2/His2-type zinc finger domain with the plant specific "QALGGH" motifs.However,in the ZFP207 there was no other domains such as NLS,L-box,DLN-box/EAR-motif discovered in most plant TFⅢA-type zinc finger proteins.ZFP207,fused with the green fluorescent protein(GFP) reporter gene,was introduced into onion epidermis cells.The ZFP207-GFP fusion protein was targeted to the nucleus.By real-time RT-PCR the inducement of ZFP207 under treatments with salt,drought,cold stress and ABA were evaluated.We constructed the 35S:ZFP207 vector and transferred it into tobacco plants by in vivo infiltration,and then we obtained the ZFP207-overexpression transgenic plants.The 2# and 4# transgenic lines were used for stresses tolerances assay.The results indicated that ZFP207-overexpression transgenic lines increased significantly cold,drought and high salt stresses tolerance compared to the WT plants,but the germination of ZFP207-overexpressed seeds is insensitive to ABA.In addition,the proline contents in transgenic plants were obviously higher than that in WT plants under normal condition. And the expression of Osmotin enhanced in transgenic lines,while the expressions of 3 LEA genes showed no change in transgenic plants.Up to date,ZFP207 is the first reported TFⅢA-type single zinc finger gene involved in abiotic stresses in plants.More analysis of ZFP207 will help to widen our understanding of the relation between the structures and functions of TFⅢA-type zinc finger proteins in plants.
引文
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