葡萄花发育相关基因的克隆与表达特性研究
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摘要
长期以来,运用传统的育种手段对多年生木本果树进行遗传改良时,由于受到童期较长等因素的影响,育种进程较为缓慢。基因工程技术的迅猛发展使其成为育种的一种重要的有巨大发展潜力的手段。花的发育是果树产量和品质的重要基础,近年来对模式植物的研究为果树花发育的研究提供了难得的机遇。
葡萄是世界上最重要的果树之一,其种植面积和总产量居世界前列。和多年生木本果树不同,葡萄具有童期较短、一年可多次开花等特点使得对葡萄花发育的研究有着重要的意义。
本文结合模式植物花发育的研究热点,对几个葡萄花发育相关的重要基因和启动子进行了研究,克隆了AGAMOUS(AG)、APATALLA3(AP3)、FLOWERING LOCUSC(FLC)、FLOWERING LOCUS T(FT)等四个同源基因的全长cDNA和LEAFY(LFY)同源基因启动子序列。对四个基因在葡萄中的表达特性进行了初步研究,以期为利用这些基因进行葡萄或其它果树的基因工程改良或表达调控提供理论依据。主要研究结果如下:
1.MADS-box转录因子在多种植物的发育过程特别是花器官和果实发育过程中发挥重要的作用.为了从葡萄中克隆出新的MADS-box基因,研究在葡萄花及果实发育中的作用,根据多种植物MADS-box基因保守区序列,设计简并引物,应用RT-PCR技术从红富士葡萄花序中分离出34条MADS-box基因cDNA片段。序列分析表明,这些片段长度在138-152 bp之间,包含基因起始密码子.推导的氨基酸序列与已登录的葡萄及其它物种的MADS-box基因同源性高达83%以上.用推导的氨基酸序列与已知的葡萄和拟南芥MADS-box家族基因进行系统发育分析,可将这些基因片段分别归入拟南芥MADS-box基因不同亚家族中。证明了葡萄中存在多种MADS-box家族基因,克隆的片段包含ABCDE模型中的各类花发育基因。
2.为探明葡萄LEAFY基因的表达调控规律,应用PCR技术从藤稔葡萄中克隆了一个长约1.8kb的DNA片段,序列全长1833bp,含有两个内含子区域,编码402个氨基酸,与葡萄VFL基因有99%的同源性。应用基因组步移法克隆了LEAFY基因的5′侧翼序列925bp,拼接后的LEAFY基因及启动子序列共2692bp(GenBank登录号EF222286)。用PLACE、PlantCARE在线启动子预测工具分析表明:该序列含有启动子的特定结构,如TATA-box,CAAT-box等,另外有一些顺式作用元件如MYB结合位点、ABA响应元件和光响应元件和一些其他的调控序列,说明葡萄LEAFY基因的表达可能受MYB、ABA和光调控.用FootPrinter在线工具对葡萄和拟南芥等其它四种植物LEAFY同源基因启动子比较,发现不同植物的启动子既有保守性,又有多样性,转录因子结合位点的分布相似,但也有区别,暗示了LEAFY基因表达调控的精确性或多样性.
3.根据获得的MADS-box基因片段设计一条特异引物,通过3'RACE的方法获得了全长cDNA,命名为VvAG(GenBank登录号:EF209334)。该片段全长978bp,包含一个681 bp的开放阅读框,编码226个氨基酸残基。通过设计特异引物RT-PCR扩增得到了编码区全长cDNA。推导的氨基酸序列与拟南芥的AGAMOUS类MADS-box基因AG、SHP1/SHP2均有较高的同源性(同源性分别为77.9%、71.2%、69.9%)。系统发育分析将VvAG基因归入MADS-box基因家族AG亚家族的euAG进化系。半定量RT-PCR分析显示,该基因在葡萄的叶片、卷须、休眠芽、花序、果实中均表达,在花器官的萼片、花瓣、雄蕊、雌蕊中也均检测到VvAG的表达,而且在这些组织中的表达丰度均较高。通过将VvAG与葡萄的一个shotgun基因组序列(AM478968)比对发现其基因组从起始密码子到终止密码子全长7534bp,由7个外显子和6个内含子组成.构建了VvAG植物双元表达载体,通过农杆菌介导法转化烟草,共获得8株hyg阳性苗.PCR检测发现8个hyg阳性株系均检测到目的条带,初步说明VvAG基因已整合到烟草基因组中;对其中5个PCR阳性株系进行RT-PCR分析,结果表明VvAG基因在5个转基因烟草株系中均发生了转录。
4.应用电子克隆技术从葡萄EST数据库中拼接到一个APETALA3(AP3)类MADS-box同源基因的全长序列,用RT-PCR的方法在红富士葡萄花序cDNA中扩增得到了该基因的编码区全长序列,命名为VvAP3(GenBank登录号EF211123).该序列编码区全长681 bp,编码226个氨基酸残基,与PMADS1、NTDEF、AP3分别有71.2%、70.4%和59.7%的同源性.该序列具有MADS-box基因典型的MADS-box和K-box结构域。系统发育分析同样将VvAP3基因归入MADS-box基因家族中AP3/PI亚家族的euAP3进化系。半定量RT-PCR分析显示,该基因在葡萄的花序及果实中表达,在叶片、卷须、休眠芽中不表达;在葡萄花器官的花瓣和雄蕊中表达,在萼片和雌蕊中不表达.通过将VvAP3与葡萄的一个shotgun基因组序列(GenBank登录号AM469149)比对发现其基因组从起始密码子到终止密码子全长7534 bp,由7个外显子和6个内含子组成,且外显子数目和每个外显子长度和拟南芥AP3基因完全相同.构建了VvAP3植物双元表达裁体,通过农杆菌介导法转化烟草,共获得7株hyg阳性苗.PCR检测发现7个hyg阳性株系均检测到目的条带,初步说明VvAP3已整合到烟草基因组中;对其中5个PCR阳性株系进行RT-PCR分析,结果表明VvAP3在这些转基因烟草株系中均发生了转录。
5.通过生物信息学在葡萄EST数据库中获得了一个FLOWERING LOCUS C(FLC)类基因全长序列,RT-PCR确认得到葡萄FLC-like MADS-box基因cDNA全长序列,命名为VvFLC(GenBank登录号EF520739)。该序列编码区全长948 bp,编码210个氨基酸残基,氨基酸序列与拟南芥的FLC、甜菜的BvFLC1-1、BVFLC1-4基因的同源性分别为43%、49%和49%。该基因具有MADS-box基因典型的MADS-box和K-box结构域。系统发育分析将VvFLC基因归入拟南芥MADS-box基因家族的FLC亚家族中。半定量RT-PCR分析结果显示,该基因在葡萄的叶片、卷须、休眠芽、花序及果实中都有表达.通过将VvFLC与葡萄基因组比对发现其由7个外显子和6个内含子组成。构建了VvFLC植物双元表达载体,通过农杆菌介导法转化烟草,共获得7株hyg阳性苗。PCR检测发现7个hyg阳性株系均检测到目的条带,初步说明VvFLC已整合到烟草基因组中;对其中4个PCR阳性株系进行RT-PCR分析,结果显示VvFLC基因在这些转基因烟草株系中均发生了转录。
6.通过生物信息学在葡萄EST数据库中获得了Flower Locus T(FT)同源基因的全长序列,RT-PCR确认得到葡萄Flower Locus T(FT)基因cDNA编码区全长序列(GenBank登录号EF203918)。该序列编码区全长525 bp,编码174个氨基酸残基,其推导的氨基酸与PnFT、FT、MdFT的同源性分别为89%、75%和74%。该序列具有FT基因典型的RKIP结构域.系统发育分析同样将VvFT聚在FT/TFL基因家族的FT组中。通过PCR从葡萄基因组中扩增得到了该基因的基因组全长(GenBank登录号EF203919),与mRNA比对结果显示该基因由4个外显子与3个内含子构成,与白杨、拟南芥的FT基因结构相同,内含子区有(TCC)_(10)、(CA)_(10)等重复序列。半定量RT-PCR分析结果表明,VvFT在葡萄花序和果实中表达,说明其不仅与开花相关,而且与果实的发育有着密切的关系。构建了VvFT植物双元表达载体,通过农杆菌介导法转化烟草,共获得8株hyg阳性苗。PCR检测发现在其中7个hyg阳性株系检测到目的条带,初步说明VvFT已整合到烟草基因组中。对其中4个PCR阳性株进行RT-PCR检测,结果表明VvFT在这些转基因烟草中均发生了转录。对转基因烟草的表型观察发现VvFT能够促进烟草开花,转基因植株出现矮化现象,但花器官形态没有发生改变.
For a long time,with traditional breeding methods,we met with many difficultics such as the long breeding period,leading to the breeding process become slowly.Genetic engineering technique has been becomeing the important potential breeding method.The floral development is the important foundation of fruit yield and qualitiy.The research of model plant flower development in recent years provided an opportunity for fruit trees.
Grape is one of the most important fruit trees in the world,its culture area and total yields are advanced in the world.Different from perennial woody fruit crops,grape has advantages of short juvenile period,multiple flowering in a year and so on.Therefore,it is of great importance to research on flower development of grape.
According to the research hotspot of the model plant flower development,we studied on several flowering related genes and the promoter from grape(Vitis vinifera×V. labrusca),cloned the full length of AGAMOUS(AG),APATALLA3(AP3),FLOWERING LOCUS C(FLC),FLOWERING LOCUS T(FT) homolog and the promoter of LEAFY gene. We studied the expression characterization of these genes,we expect that it will be used in the genetic engineering breeding or regulate gene expression.The main results are as follows:
1.As a kind of transcription factors,MADS-box genes play an important role in various plant development processes,especially in the development of floral organs and fruit.In order to isolate new MADS-box genes and find out the function of these genes in the development of grape floral organs and fruit,we designed degenerate primers according to the conservative area sequences of MADS-box family gene from various plants, obtained 34 MADS- box gene cDNA fragments amplified by RT-PCR from the inflorescence of Vitis vinifera×V.labrusca 'Red Fuji'.Sequence analysis indicated the size of these fragments is among 138-152 bp,including the initiation codon of genes,deduced amino acid sequences have a higher identify of more than 83%with MADS-box genes of grapevine and other species.By analysis of phylogeny construction,all of these gene fragments have been divided into the Arabidopsis MADS-box gene subfamily.These results showed that there are many MADS-box genes in grape,these gene fragments cover all type floral development genes of ABCDE model.
2.In order to study the rule of grape LEAFY gene expression and regulation,we amplified a 1.8kb fragment from Vitis Vinifera×V.Labrusca 'Fujiminori' by PCR.The fragment is 1833bp,deduced 402 amino acids,has two introns,which is 99%identity with VFL gene.Nine hundred and twenty five bp promoter region sequence was cloned by genome walking method.The gene and promoter are 2692bp in length(GenBank accession is EF222286).Promoter sequence analyzed by PLACE and PlantCARE showed that it has TATA-box,CAAT-box and some cis-acting element such as MYB binding site,cis-acting regulatory element involved in ABA response,some light-induced responsive elements and other transcription factor-binding sites.Compared the promoter of grape LEAFY homologue with 4 plants,by FootPrinter analysis we found that these promoter has conservatism and diversity and the distributing of transcription factor-binding sites has similarity and difference.It implied the accuracy and diversity of LEAFY gene expression and regulation.
3.As a kind of transcription factors,MADS-box genes play an important role in plant various development processes,especially in the development of floral organs.According to a MADS fragment sequence obtained by degenerate primer RT-PCR,an AG-like MADS-box gene was cloned from inflorescene by the method of 3'RACE PCR,named VvAG(GeneBank accession number EF209334),This gene is 978 bp and contained an open reading frame of 681bp coding a polypeptide of 226 amino acids.Homology analysis showed that the deduced VvAG protein was highly homologous to AG,SHP1/SHP2 of Arabidopsis thaliana,the identity is 77.9%,71.2%and 69.9%,respectively.Phylogenetic analysis also indicated that VvAG belongs to the euAG lineage of AG subfamily in MADS-box gene family.RT-PCR analysis showed that VvAG expresses at high levels in leaves,tendrils,inflorescenes,young fruits and mature fruits,and also expressed in all of 4 wheel floral organ.By blast of VvAG with a shotgun genome sequence,the result indicated that VvAG genome is consist of 7 exons and 6 introns.We constructed a overexpression vector of VvAG and transferred it into tobacco,obtained 8 hyg positive tobacco plants.PCR analysis showed that these lines are all PCR positive.Analysis of 5 PCR positive lines by RT-PCR revealed VvAG expressed in all these lines.
4.An APETALA3(AP3) homologue was isolated from grape(Vitis vinifera×V. labrusca 'Red Fuji') inflorescence by bioformatic analysis and RT-PCR.It is 681 bp, coding a polypeptide of 226 amino acids,named as VvAP3(GenBaak accession number EF211123).Homology analysis showed that the deduced VvAP3 protein was highly homologous to the MADS-box genes,PMADS1,NTDEF and AP3,the identity are 71.2%, 70.4%and 59.7%,respectively.Phylogenetic analysis also indicated that VvAP3 belongs to the euAP3 lineage of AP3 subfamily in MADS-box gene family.RT-PCR analysis showed that VvAP3 expressed in inflorescenes,young and mature fruits,and in petal and stamen, but not expressed in sepal and carpel.Blast of VvAP3 with a shotgun genome sequence indicated that VvAP3 genome is consist of 7 exons and 6 introns,the number and length of exon is the same as AP3.We constructed a overexpression vector of VvAP3 and transferred it into tobacco,obtained 7 hyg positive tobacco plants.PCR analysis showed that these lines are all PCR positive.Analysis of 5 PCR positive lines by RT-PCR revealed VvAP3 expressed in all these lines.
5.An putative FLC homologue full length cDNA was isolated from grape(Vitis vinifera×V.labrusca 'Red Fuji') inflorescence by bioformatic analysis and RT-PCR.It is 948bp in length,coding a polypeptide of 210 amino acids,named as VvFLC(GenBank accession number is EF520739).The predicted protein sequence shows homology to FLC (43%identity) and to BvFLC1-1 and BvFLC1-4(49%and 49%identity,respectively). Phylogenetic analysis also indicated that VvFLC belonged to the FLC subfamily of MADS-box gene family.VvFLC expresses in leaves,tendrils,inflorescenes,young and mature fruits.Blast of VvFLC with a genome shotgun sequence indicated that VvFLC genome is consist of 7 exons and 6 introns.We constructed a overexpression vector of VvFLC and transferred it into tobacco,obtained 7 hyg positive tobacco plants.PCR analysis showed that these lines are all PCR positive.Analysis of 4 PCR positive lines by RT-PCR revealed VvFLC expressed in all these lines.
6.An Flowering Locus T(FT) homologue was isolated from grape(Vitis vinifera×V. labrusca 'Red Fuji') inflorescence by bioformatic analysis and RT-PCR.It is 525bp in length,coding a polypeptide of 174 amino acids,named as VvFT(GenBank accession is EF203918).The deduced amino acid sequence shows a high degree of homology to PnFT1a(89%identity) and to FT and MdFT(75%and 74%identity respectively).VvFT protein has a typical RKIP domain.Phylogenetic analysis showed VvFT belongs to FT homologues of the FT/TFLl gene family.Cloning VvFT genome sequence by PCR(GenBank accession is EF203919),blast of it with VvFT mRNA showed the structure of VvFT gene is consist of 4 exons and 3 introns,the structure is the same as Arabidopsis and poplar.There has some sequence repeat in the intron of VvFT gene,such as(TCC)_(10) and(CA)_(10).VvFT is expressed in the developing inflorescene,young and mature berry showed VvFT is related with flower and fruit development.We constructed a overexpression vector of VvFT and transferred it into tobacco,obtained 8 hyg positive tobacco plants.PCR analysis showed that 7 of them are PCR positive.Analysis of 4 PCR positive lines by RT-PCR revealed VvFT expressed in all these lines.Overexpression of VvFT in tobacco hastened flowering showed that VvFT function as promoters of flowering when ectopically expressed in a heterologous plant,and the flower number of transgene plant is more than untransgene plant.A transgene plant showed a dwarfism phenotype,and no change in flower organ shape.
葡萄是世界上最重要的果树之一,其种植面积和总产量居世界前列。和多年生木本果树不同,葡萄具有童期较短、一年可多次开花等特点使得对葡萄花发育的研究有着重要的意义。
本文结合模式植物花发育的研究热点,对几个葡萄花发育相关的重要基因和启动子进行了研究,克隆了AGAMOUS(AG)、APATALLA3(AP3)、FLOWERING LOCUSC(FLC)、FLOWERING LOCUS T(FT)等四个同源基因的全长cDNA和LEAFY(LFY)同源基因启动子序列。对四个基因在葡萄中的表达特性进行了初步研究,以期为利用这些基因进行葡萄或其它果树的基因工程改良或表达调控提供理论依据。主要研究结果如下:
1.MADS-box转录因子在多种植物的发育过程特别是花器官和果实发育过程中发挥重要的作用.为了从葡萄中克隆出新的MADS-box基因,研究在葡萄花及果实发育中的作用,根据多种植物MADS-box基因保守区序列,设计简并引物,应用RT-PCR技术从红富士葡萄花序中分离出34条MADS-box基因cDNA片段。序列分析表明,这些片段长度在138-152 bp之间,包含基因起始密码子.推导的氨基酸序列与已登录的葡萄及其它物种的MADS-box基因同源性高达83%以上.用推导的氨基酸序列与已知的葡萄和拟南芥MADS-box家族基因进行系统发育分析,可将这些基因片段分别归入拟南芥MADS-box基因不同亚家族中。证明了葡萄中存在多种MADS-box家族基因,克隆的片段包含ABCDE模型中的各类花发育基因。
2.为探明葡萄LEAFY基因的表达调控规律,应用PCR技术从藤稔葡萄中克隆了一个长约1.8kb的DNA片段,序列全长1833bp,含有两个内含子区域,编码402个氨基酸,与葡萄VFL基因有99%的同源性。应用基因组步移法克隆了LEAFY基因的5′侧翼序列925bp,拼接后的LEAFY基因及启动子序列共2692bp(GenBank登录号EF222286)。用PLACE、PlantCARE在线启动子预测工具分析表明:该序列含有启动子的特定结构,如TATA-box,CAAT-box等,另外有一些顺式作用元件如MYB结合位点、ABA响应元件和光响应元件和一些其他的调控序列,说明葡萄LEAFY基因的表达可能受MYB、ABA和光调控.用FootPrinter在线工具对葡萄和拟南芥等其它四种植物LEAFY同源基因启动子比较,发现不同植物的启动子既有保守性,又有多样性,转录因子结合位点的分布相似,但也有区别,暗示了LEAFY基因表达调控的精确性或多样性.
3.根据获得的MADS-box基因片段设计一条特异引物,通过3'RACE的方法获得了全长cDNA,命名为VvAG(GenBank登录号:EF209334)。该片段全长978bp,包含一个681 bp的开放阅读框,编码226个氨基酸残基。通过设计特异引物RT-PCR扩增得到了编码区全长cDNA。推导的氨基酸序列与拟南芥的AGAMOUS类MADS-box基因AG、SHP1/SHP2均有较高的同源性(同源性分别为77.9%、71.2%、69.9%)。系统发育分析将VvAG基因归入MADS-box基因家族AG亚家族的euAG进化系。半定量RT-PCR分析显示,该基因在葡萄的叶片、卷须、休眠芽、花序、果实中均表达,在花器官的萼片、花瓣、雄蕊、雌蕊中也均检测到VvAG的表达,而且在这些组织中的表达丰度均较高。通过将VvAG与葡萄的一个shotgun基因组序列(AM478968)比对发现其基因组从起始密码子到终止密码子全长7534bp,由7个外显子和6个内含子组成.构建了VvAG植物双元表达载体,通过农杆菌介导法转化烟草,共获得8株hyg阳性苗.PCR检测发现8个hyg阳性株系均检测到目的条带,初步说明VvAG基因已整合到烟草基因组中;对其中5个PCR阳性株系进行RT-PCR分析,结果表明VvAG基因在5个转基因烟草株系中均发生了转录。
4.应用电子克隆技术从葡萄EST数据库中拼接到一个APETALA3(AP3)类MADS-box同源基因的全长序列,用RT-PCR的方法在红富士葡萄花序cDNA中扩增得到了该基因的编码区全长序列,命名为VvAP3(GenBank登录号EF211123).该序列编码区全长681 bp,编码226个氨基酸残基,与PMADS1、NTDEF、AP3分别有71.2%、70.4%和59.7%的同源性.该序列具有MADS-box基因典型的MADS-box和K-box结构域。系统发育分析同样将VvAP3基因归入MADS-box基因家族中AP3/PI亚家族的euAP3进化系。半定量RT-PCR分析显示,该基因在葡萄的花序及果实中表达,在叶片、卷须、休眠芽中不表达;在葡萄花器官的花瓣和雄蕊中表达,在萼片和雌蕊中不表达.通过将VvAP3与葡萄的一个shotgun基因组序列(GenBank登录号AM469149)比对发现其基因组从起始密码子到终止密码子全长7534 bp,由7个外显子和6个内含子组成,且外显子数目和每个外显子长度和拟南芥AP3基因完全相同.构建了VvAP3植物双元表达裁体,通过农杆菌介导法转化烟草,共获得7株hyg阳性苗.PCR检测发现7个hyg阳性株系均检测到目的条带,初步说明VvAP3已整合到烟草基因组中;对其中5个PCR阳性株系进行RT-PCR分析,结果表明VvAP3在这些转基因烟草株系中均发生了转录。
5.通过生物信息学在葡萄EST数据库中获得了一个FLOWERING LOCUS C(FLC)类基因全长序列,RT-PCR确认得到葡萄FLC-like MADS-box基因cDNA全长序列,命名为VvFLC(GenBank登录号EF520739)。该序列编码区全长948 bp,编码210个氨基酸残基,氨基酸序列与拟南芥的FLC、甜菜的BvFLC1-1、BVFLC1-4基因的同源性分别为43%、49%和49%。该基因具有MADS-box基因典型的MADS-box和K-box结构域。系统发育分析将VvFLC基因归入拟南芥MADS-box基因家族的FLC亚家族中。半定量RT-PCR分析结果显示,该基因在葡萄的叶片、卷须、休眠芽、花序及果实中都有表达.通过将VvFLC与葡萄基因组比对发现其由7个外显子和6个内含子组成。构建了VvFLC植物双元表达载体,通过农杆菌介导法转化烟草,共获得7株hyg阳性苗。PCR检测发现7个hyg阳性株系均检测到目的条带,初步说明VvFLC已整合到烟草基因组中;对其中4个PCR阳性株系进行RT-PCR分析,结果显示VvFLC基因在这些转基因烟草株系中均发生了转录。
6.通过生物信息学在葡萄EST数据库中获得了Flower Locus T(FT)同源基因的全长序列,RT-PCR确认得到葡萄Flower Locus T(FT)基因cDNA编码区全长序列(GenBank登录号EF203918)。该序列编码区全长525 bp,编码174个氨基酸残基,其推导的氨基酸与PnFT、FT、MdFT的同源性分别为89%、75%和74%。该序列具有FT基因典型的RKIP结构域.系统发育分析同样将VvFT聚在FT/TFL基因家族的FT组中。通过PCR从葡萄基因组中扩增得到了该基因的基因组全长(GenBank登录号EF203919),与mRNA比对结果显示该基因由4个外显子与3个内含子构成,与白杨、拟南芥的FT基因结构相同,内含子区有(TCC)_(10)、(CA)_(10)等重复序列。半定量RT-PCR分析结果表明,VvFT在葡萄花序和果实中表达,说明其不仅与开花相关,而且与果实的发育有着密切的关系。构建了VvFT植物双元表达载体,通过农杆菌介导法转化烟草,共获得8株hyg阳性苗。PCR检测发现在其中7个hyg阳性株系检测到目的条带,初步说明VvFT已整合到烟草基因组中。对其中4个PCR阳性株进行RT-PCR检测,结果表明VvFT在这些转基因烟草中均发生了转录。对转基因烟草的表型观察发现VvFT能够促进烟草开花,转基因植株出现矮化现象,但花器官形态没有发生改变.
For a long time,with traditional breeding methods,we met with many difficultics such as the long breeding period,leading to the breeding process become slowly.Genetic engineering technique has been becomeing the important potential breeding method.The floral development is the important foundation of fruit yield and qualitiy.The research of model plant flower development in recent years provided an opportunity for fruit trees.
Grape is one of the most important fruit trees in the world,its culture area and total yields are advanced in the world.Different from perennial woody fruit crops,grape has advantages of short juvenile period,multiple flowering in a year and so on.Therefore,it is of great importance to research on flower development of grape.
According to the research hotspot of the model plant flower development,we studied on several flowering related genes and the promoter from grape(Vitis vinifera×V. labrusca),cloned the full length of AGAMOUS(AG),APATALLA3(AP3),FLOWERING LOCUS C(FLC),FLOWERING LOCUS T(FT) homolog and the promoter of LEAFY gene. We studied the expression characterization of these genes,we expect that it will be used in the genetic engineering breeding or regulate gene expression.The main results are as follows:
1.As a kind of transcription factors,MADS-box genes play an important role in various plant development processes,especially in the development of floral organs and fruit.In order to isolate new MADS-box genes and find out the function of these genes in the development of grape floral organs and fruit,we designed degenerate primers according to the conservative area sequences of MADS-box family gene from various plants, obtained 34 MADS- box gene cDNA fragments amplified by RT-PCR from the inflorescence of Vitis vinifera×V.labrusca 'Red Fuji'.Sequence analysis indicated the size of these fragments is among 138-152 bp,including the initiation codon of genes,deduced amino acid sequences have a higher identify of more than 83%with MADS-box genes of grapevine and other species.By analysis of phylogeny construction,all of these gene fragments have been divided into the Arabidopsis MADS-box gene subfamily.These results showed that there are many MADS-box genes in grape,these gene fragments cover all type floral development genes of ABCDE model.
2.In order to study the rule of grape LEAFY gene expression and regulation,we amplified a 1.8kb fragment from Vitis Vinifera×V.Labrusca 'Fujiminori' by PCR.The fragment is 1833bp,deduced 402 amino acids,has two introns,which is 99%identity with VFL gene.Nine hundred and twenty five bp promoter region sequence was cloned by genome walking method.The gene and promoter are 2692bp in length(GenBank accession is EF222286).Promoter sequence analyzed by PLACE and PlantCARE showed that it has TATA-box,CAAT-box and some cis-acting element such as MYB binding site,cis-acting regulatory element involved in ABA response,some light-induced responsive elements and other transcription factor-binding sites.Compared the promoter of grape LEAFY homologue with 4 plants,by FootPrinter analysis we found that these promoter has conservatism and diversity and the distributing of transcription factor-binding sites has similarity and difference.It implied the accuracy and diversity of LEAFY gene expression and regulation.
3.As a kind of transcription factors,MADS-box genes play an important role in plant various development processes,especially in the development of floral organs.According to a MADS fragment sequence obtained by degenerate primer RT-PCR,an AG-like MADS-box gene was cloned from inflorescene by the method of 3'RACE PCR,named VvAG(GeneBank accession number EF209334),This gene is 978 bp and contained an open reading frame of 681bp coding a polypeptide of 226 amino acids.Homology analysis showed that the deduced VvAG protein was highly homologous to AG,SHP1/SHP2 of Arabidopsis thaliana,the identity is 77.9%,71.2%and 69.9%,respectively.Phylogenetic analysis also indicated that VvAG belongs to the euAG lineage of AG subfamily in MADS-box gene family.RT-PCR analysis showed that VvAG expresses at high levels in leaves,tendrils,inflorescenes,young fruits and mature fruits,and also expressed in all of 4 wheel floral organ.By blast of VvAG with a shotgun genome sequence,the result indicated that VvAG genome is consist of 7 exons and 6 introns.We constructed a overexpression vector of VvAG and transferred it into tobacco,obtained 8 hyg positive tobacco plants.PCR analysis showed that these lines are all PCR positive.Analysis of 5 PCR positive lines by RT-PCR revealed VvAG expressed in all these lines.
4.An APETALA3(AP3) homologue was isolated from grape(Vitis vinifera×V. labrusca 'Red Fuji') inflorescence by bioformatic analysis and RT-PCR.It is 681 bp, coding a polypeptide of 226 amino acids,named as VvAP3(GenBaak accession number EF211123).Homology analysis showed that the deduced VvAP3 protein was highly homologous to the MADS-box genes,PMADS1,NTDEF and AP3,the identity are 71.2%, 70.4%and 59.7%,respectively.Phylogenetic analysis also indicated that VvAP3 belongs to the euAP3 lineage of AP3 subfamily in MADS-box gene family.RT-PCR analysis showed that VvAP3 expressed in inflorescenes,young and mature fruits,and in petal and stamen, but not expressed in sepal and carpel.Blast of VvAP3 with a shotgun genome sequence indicated that VvAP3 genome is consist of 7 exons and 6 introns,the number and length of exon is the same as AP3.We constructed a overexpression vector of VvAP3 and transferred it into tobacco,obtained 7 hyg positive tobacco plants.PCR analysis showed that these lines are all PCR positive.Analysis of 5 PCR positive lines by RT-PCR revealed VvAP3 expressed in all these lines.
5.An putative FLC homologue full length cDNA was isolated from grape(Vitis vinifera×V.labrusca 'Red Fuji') inflorescence by bioformatic analysis and RT-PCR.It is 948bp in length,coding a polypeptide of 210 amino acids,named as VvFLC(GenBank accession number is EF520739).The predicted protein sequence shows homology to FLC (43%identity) and to BvFLC1-1 and BvFLC1-4(49%and 49%identity,respectively). Phylogenetic analysis also indicated that VvFLC belonged to the FLC subfamily of MADS-box gene family.VvFLC expresses in leaves,tendrils,inflorescenes,young and mature fruits.Blast of VvFLC with a genome shotgun sequence indicated that VvFLC genome is consist of 7 exons and 6 introns.We constructed a overexpression vector of VvFLC and transferred it into tobacco,obtained 7 hyg positive tobacco plants.PCR analysis showed that these lines are all PCR positive.Analysis of 4 PCR positive lines by RT-PCR revealed VvFLC expressed in all these lines.
6.An Flowering Locus T(FT) homologue was isolated from grape(Vitis vinifera×V. labrusca 'Red Fuji') inflorescence by bioformatic analysis and RT-PCR.It is 525bp in length,coding a polypeptide of 174 amino acids,named as VvFT(GenBank accession is EF203918).The deduced amino acid sequence shows a high degree of homology to PnFT1a(89%identity) and to FT and MdFT(75%and 74%identity respectively).VvFT protein has a typical RKIP domain.Phylogenetic analysis showed VvFT belongs to FT homologues of the FT/TFLl gene family.Cloning VvFT genome sequence by PCR(GenBank accession is EF203919),blast of it with VvFT mRNA showed the structure of VvFT gene is consist of 4 exons and 3 introns,the structure is the same as Arabidopsis and poplar.There has some sequence repeat in the intron of VvFT gene,such as(TCC)_(10) and(CA)_(10).VvFT is expressed in the developing inflorescene,young and mature berry showed VvFT is related with flower and fruit development.We constructed a overexpression vector of VvFT and transferred it into tobacco,obtained 8 hyg positive tobacco plants.PCR analysis showed that 7 of them are PCR positive.Analysis of 4 PCR positive lines by RT-PCR revealed VvFT expressed in all these lines.Overexpression of VvFT in tobacco hastened flowering showed that VvFT function as promoters of flowering when ectopically expressed in a heterologous plant,and the flower number of transgene plant is more than untransgene plant.A transgene plant showed a dwarfism phenotype,and no change in flower organ shape.
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