砂梨果实成熟软化相关基因的克隆、表达分析及其表达载体的构建
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摘要
砂梨(Pyrus pyrifolia Nakai)是我国的重要果树之一,在我国果树生产中占有重要的地位。该类型的梨果实不耐贮运,货架期短,极易造成损失。本文以砂梨果实为试材,利用RACE技术克隆了ACC合酶、ACC氧化酶、木葡聚糖转葡糖苷酶/水解酶和扩展蛋白等与果实成熟与软化相关的酶或蛋白的基因的cDNA全长,即Ppy-ACS, Ppy-ACO. Ppy-XTH和Ppy-EXP1、Ppy-EXP2、Ppy-EXP3,构建了砂梨Ppy-ACS和Ppy-ACO的正义和反义表达载体,同时对所克隆基因在果实发育过程中的表达情况和成熟果实中表达的保守的microRNAs进行了芯片分析,主要结果如下。
1.砂梨ACC合酶基因(Ppy-ACS, EF566S65)的cDNA全长为1,939bp, Ppy-ACS核苷酸序列含有5'-UTR为109bp,1,488bp完整的ORF和3'-UTR为342bp,推导编码495个氨基酸。该氨基酸序列具备ACC合酶7个保守区和组成该酶活性中心的12个氨基酸残基,即SLSKDMGFPGLJL以pYPX145载体为基础,分别将Ppy-ACS编码区序列反向和正向插入相应位点构建反义和正义表达载体,命名为pPpy-ACS(-)和pPpy-ACS(+),目的基因由双CaMV35S启动子所控制,分别将这2个表达载体导入根癌农杆菌菌株EHA105。
2.砂梨ACC氧化酶基因(Ppy-,4CO, EF451060)的cDNA全长1,225bp, Ppy-ACO核苷酸序列有一个945bp完整的ORF,5'-UTR为63bp,3'-UTR为217bp,推导编码314个氨基酸,该序列具备非血红素二价铁离子依赖型的氧化/加氧酶类的12个保守氨基酸和催化活性所需的3个氨基酸。将Ppy-ACO编码区序列反向插入pYPX145载体,构建了由双CaMV35S启动子所控制的双元表达载体,并成功将表达载体导入根癌农杆菌菌株LBA4404。
3.在砂梨果实发育期间,Ppy-ACS和Ppy-ACO的表达量随果实的生长发育而逐渐地增加,且Ppy-ACO表达量明显高于Ppy-ACS;1-MCP明显抑制果实采收后12d内的Ppy-ACS基因表达,降低果实整个采后期Ppy-ACO的表达；外源乙烯对采后果实Ppy-ACS的表达有明显的促进作用,对Ppy-ACO的表达亦有明显的促进作用,但在采后末期有所减弱。
4.砂梨木葡聚糖转葡糖苷酶／水解酶基因(Ppy-XTH, EU432411)的cDNA全长1,248bp, Ppy-XTH核苷酸序列有一个885bp完整的OEF,5’末端起始密码子ATG起始于74bp,3'-UTR为290bp,其中含有30bp的Ploy+(A)。Ppy-XTH推导编码294个氨基酸,具备XTH蛋白保守的催化活性部位DEIDFEFLG, N-末端具22个氨基酸组成的信号肽序列,定位于质外体。不同物种该家族基因的氨基酸序列聚类分析表明,Ppy-XTH属于第1类。实时荧光qRT-PCR实验结果是Ppy-XTH,在果实膨大前期积累较低,在果实膨大期大幅增加,在果实成熟期又显著下降。
5.砂梨扩展蛋白基因Ppy-EXP1(EF602031)、Ppy-EXP2(FJ619347)和Ppy-EXP1(FJ619348)的cDNA全长分别是1,395bp、1,107bp和1,136bp,分别编码252个、251个和254个氨基酸,均含有1个组氨酸(His-Phe-Asp, HFD)功能域。以氨基酸序列为基础构建了12个物种29个扩展蛋白的分子进化树,Ppy-EXP1和Ppy-EXP2聚为B类,而Ppy-EXP3聚在C类。采用实时荧光qRT-PCR分析了Ppy-EXP1、Ppy-EXP2和Ppy-EXP3在砂梨果实不同生长发育期的表达情况,Ppy-EXP1和Ppy-EXP2在盛花后0-98d的积累水平很低,在盛花后98d至果实成熟,其表达水平略有上升；Ppy-EXP3从盛花期至果实成熟的表达量一直处于较高水平,至果实成熟时达到最高水平。
6.以所有物种的6,703个microRNAs,点制的芯片为基础,对成熟果实中表达的保守microRNAs进行了检测,检测到643个阳性信号,对其中11个(8个来自植物,3个来自动物)高频表达microRNAs的靶基因进行了预测。7个来自植物microRNAs的靶基因编码转录因子,1个microRNA的靶基因参与microRNAs自身代谢,1个源自动物的miR-30也在植物上预测到靶基因,该靶基因编码异戊烯基转移酶,另外2个来自动物的miR-124和miR-181在植物上未检测到靶基因。
Sand pear (Pyrus pyrifolia Nakai) is an important fruit crop in China. However, the short shelf-life of this fruit is a serious problem and causes a high loss of yield and commercial value. Six full-length cDNAs encoding enzyme or protein genes related to fruit ripening and softening, such as ACC synthase (ACS), ACC oxidase (ACO), xyloglucan endotransglucosylase/hydrolase (XTH) and expansins (EXP) were isolated from mature fruit of sand pear using RT-PCR and RACE methods. The expression of these genes as above was detected by real-time fluorescent qRT-PCR during fruit growth and development. Three binary antisense or sense expression vectors of Ppy-ACS and Ppy-ACO were constructed for plant transformation. In addition, the effects of exogenous ethylene and1-MCP on expression of Ppy-ACS and Ppy-ACO in postharvest fruit were determined, and conserved microRNAs were detected on mature fruit by microarray. Their results were as follow.
1. The full-length cDNA (GenBank accession no. EF566865) of Ppy-ACS is1,939bp, which includes109bp of the5'-UTR, an ORF of1,488bp, and342bp of the3'-UTR. The Ppy-ACS encodes a polypeptide be composed of495amino acid residues, which includes seven conserved regions and an active site consisting of twelve essential amino acid residues, i.e. SLSKDMGFPGLR. Two binary expression vectors of Ppy-ACS, antisense named as pPpy-ACS(-) and sense named as pPpy-ACS(+), were constructed by inserting the target fragment in pYPX145. The target genes are controlled by a double CaMV35S promoter. The pPpy-ACS(-/+) was successfully introduced into Agrobacterium tumefaciens strain EHA105.
2. The Ppy-ACO full-length cDNA (EF451060) is1,225bp, which contains is945nucleotides for ORF,63bp of the5'-UTR and217bp of the3'-UTR. The deduced amino acid sequence of the Ppy-ACO is314residues and includes twelve conserved amino acid residues belonging to non-heme iron (Ⅱ) dependent family of oxygenases/oxidases and three residues essential for emzyme activation. A binary antisense expression vector with Ppy-ACO coding region was constructed by inserting the target fragment in reverse orientation in pYPX145. The expression of the antisense gene is controlled by a double CaMV35S promoter. The vector was successfully introduced into Agrobacterium tumefaciens strain LBA4404.
3. Expression of Ppy-ACS and Ppy-ACO using real-time fluorescent qRT-PCR increased with fruit growth and development, and expression of the Ppy-ACO was obviously higher than that of the Ppy-ACS during fruit growth and development.1-MCP treatment obviously inhibited the Ppy-ACS expression in12days after harvest and down-regulated the Ppy-ACO expression throughout fruit storage period. However, exogenous ethyl ene significantly up-regulated the Ppy-ACS expression at early and later stages, and evidently promoted the Ppy-ACO expression all time after harvest, only reduced at the end of fruit storage.
4. Xyloglucan endotransglucosylase/hydrolase gene (named as Ppy-XTH, EU432411) is1,248bp contained73nuleotides of the5'-UTR, an ORF consiting of885nucleotides, and290nucleotides of the3'-UTR. The deduced amino acid sequence of the Ppy-XTH is294amino acid residues, which includes a signal peptide composed of22amino acid residues in the N-timminal, and shares a conserved DEIDFEFLD motif that is likely to be the catalytic site for both xyloglucan endotransglycosylase (XET) and xyloglucan endohydrolase (XEH) activity. The result of cluster analysis of amino acid sequence for mature Ppy-XTH polypeptide with other plant such as Arabidopsis, tomato, apple and kiwifruit showed Ppy-XTH belongs to group1, which expressed in flower and fruit. The accumulation of the Ppy-XTH mRNA in fruit could be detected during the whole fruit growth and development, obviously increased from the beginning of fruit rapid growth at98days after full bloom (DAFB), steadily maintained high level from98DAFB to126DAFB, evidently decreased at the end of fruit growth phase (140DAFB).
5. Three expansin gene cDNAs, designate as Ppy-EXP1(EF602031), Ppy-EXP2(FJ619347) and Ppy-EXP3(FJ619348), were cloned from mature fruit of sand pear. The full-length cDNA of Ppy-EXP1, Ppy-EXP2and Ppy-EXP3were1,369bp,1,107bp and1,136bp, respectively. The Ppy-EXP1, Ppy-EXP1and Ppy-EXP3ordinally contain an ORF of759bp,756bp and765bp, and encode a polypeptide consisting of252,251and254amino acid residues, respectively. They all contain a histidine (His-Phe-Asp, HFD) motif as conversed amino acid domain of expansin for most plants in the central region. A phylogenetic tree was generated from the deduced amino acid alignment of three Ppy-EXPs and26other expansin homologues. Ppy-EXP1and Ppy-EXP2aligned within group B and Ppy-EXP3belonged to group C. Accumulation of Ppy-EXP1and Ppy-EXP2in the developing fruit was low level during0-98DAFB, slightly increased at the end of growth phase, and the increments of the Ppy-EXP1are more than that of the Ppy-EXP2. Accumulation of the Ppy-EXP3was higher than that of the Ppy-EXP1and the Ppy-EXP1in the developing fruit, and increased markedly with fruit growth and development, and reached to the highest level at the end of growth phase (140DAFB).
6. Conserved microRNAs were detected in mature fruit by microarray based on6,703microRNAs sequences from miRBase11.0version (http://microrna.sanger.ac.uk/). Six hundred and forty-three microRNAs were detected based on positive signal. Eleven high frequently expressed microRNAs (eight from plant, three from animal) were selected for target prediction. Seven microRNA targets belong to transcription factor families, and the left one is involved in microRNAs metabolism. Isopentenyl transferase gene was predicted as a target for miR-30, but the other two animal microRNAs, miR-124and miR-181, didn't find their targets.
1.砂梨ACC合酶基因(Ppy-ACS, EF566S65)的cDNA全长为1,939bp, Ppy-ACS核苷酸序列含有5'-UTR为109bp,1,488bp完整的ORF和3'-UTR为342bp,推导编码495个氨基酸。该氨基酸序列具备ACC合酶7个保守区和组成该酶活性中心的12个氨基酸残基,即SLSKDMGFPGLJL以pYPX145载体为基础,分别将Ppy-ACS编码区序列反向和正向插入相应位点构建反义和正义表达载体,命名为pPpy-ACS(-)和pPpy-ACS(+),目的基因由双CaMV35S启动子所控制,分别将这2个表达载体导入根癌农杆菌菌株EHA105。
2.砂梨ACC氧化酶基因(Ppy-,4CO, EF451060)的cDNA全长1,225bp, Ppy-ACO核苷酸序列有一个945bp完整的ORF,5'-UTR为63bp,3'-UTR为217bp,推导编码314个氨基酸,该序列具备非血红素二价铁离子依赖型的氧化/加氧酶类的12个保守氨基酸和催化活性所需的3个氨基酸。将Ppy-ACO编码区序列反向插入pYPX145载体,构建了由双CaMV35S启动子所控制的双元表达载体,并成功将表达载体导入根癌农杆菌菌株LBA4404。
3.在砂梨果实发育期间,Ppy-ACS和Ppy-ACO的表达量随果实的生长发育而逐渐地增加,且Ppy-ACO表达量明显高于Ppy-ACS;1-MCP明显抑制果实采收后12d内的Ppy-ACS基因表达,降低果实整个采后期Ppy-ACO的表达；外源乙烯对采后果实Ppy-ACS的表达有明显的促进作用,对Ppy-ACO的表达亦有明显的促进作用,但在采后末期有所减弱。
4.砂梨木葡聚糖转葡糖苷酶／水解酶基因(Ppy-XTH, EU432411)的cDNA全长1,248bp, Ppy-XTH核苷酸序列有一个885bp完整的OEF,5’末端起始密码子ATG起始于74bp,3'-UTR为290bp,其中含有30bp的Ploy+(A)。Ppy-XTH推导编码294个氨基酸,具备XTH蛋白保守的催化活性部位DEIDFEFLG, N-末端具22个氨基酸组成的信号肽序列,定位于质外体。不同物种该家族基因的氨基酸序列聚类分析表明,Ppy-XTH属于第1类。实时荧光qRT-PCR实验结果是Ppy-XTH,在果实膨大前期积累较低,在果实膨大期大幅增加,在果实成熟期又显著下降。
5.砂梨扩展蛋白基因Ppy-EXP1(EF602031)、Ppy-EXP2(FJ619347)和Ppy-EXP1(FJ619348)的cDNA全长分别是1,395bp、1,107bp和1,136bp,分别编码252个、251个和254个氨基酸,均含有1个组氨酸(His-Phe-Asp, HFD)功能域。以氨基酸序列为基础构建了12个物种29个扩展蛋白的分子进化树,Ppy-EXP1和Ppy-EXP2聚为B类,而Ppy-EXP3聚在C类。采用实时荧光qRT-PCR分析了Ppy-EXP1、Ppy-EXP2和Ppy-EXP3在砂梨果实不同生长发育期的表达情况,Ppy-EXP1和Ppy-EXP2在盛花后0-98d的积累水平很低,在盛花后98d至果实成熟,其表达水平略有上升；Ppy-EXP3从盛花期至果实成熟的表达量一直处于较高水平,至果实成熟时达到最高水平。
6.以所有物种的6,703个microRNAs,点制的芯片为基础,对成熟果实中表达的保守microRNAs进行了检测,检测到643个阳性信号,对其中11个(8个来自植物,3个来自动物)高频表达microRNAs的靶基因进行了预测。7个来自植物microRNAs的靶基因编码转录因子,1个microRNA的靶基因参与microRNAs自身代谢,1个源自动物的miR-30也在植物上预测到靶基因,该靶基因编码异戊烯基转移酶,另外2个来自动物的miR-124和miR-181在植物上未检测到靶基因。
Sand pear (Pyrus pyrifolia Nakai) is an important fruit crop in China. However, the short shelf-life of this fruit is a serious problem and causes a high loss of yield and commercial value. Six full-length cDNAs encoding enzyme or protein genes related to fruit ripening and softening, such as ACC synthase (ACS), ACC oxidase (ACO), xyloglucan endotransglucosylase/hydrolase (XTH) and expansins (EXP) were isolated from mature fruit of sand pear using RT-PCR and RACE methods. The expression of these genes as above was detected by real-time fluorescent qRT-PCR during fruit growth and development. Three binary antisense or sense expression vectors of Ppy-ACS and Ppy-ACO were constructed for plant transformation. In addition, the effects of exogenous ethylene and1-MCP on expression of Ppy-ACS and Ppy-ACO in postharvest fruit were determined, and conserved microRNAs were detected on mature fruit by microarray. Their results were as follow.
1. The full-length cDNA (GenBank accession no. EF566865) of Ppy-ACS is1,939bp, which includes109bp of the5'-UTR, an ORF of1,488bp, and342bp of the3'-UTR. The Ppy-ACS encodes a polypeptide be composed of495amino acid residues, which includes seven conserved regions and an active site consisting of twelve essential amino acid residues, i.e. SLSKDMGFPGLR. Two binary expression vectors of Ppy-ACS, antisense named as pPpy-ACS(-) and sense named as pPpy-ACS(+), were constructed by inserting the target fragment in pYPX145. The target genes are controlled by a double CaMV35S promoter. The pPpy-ACS(-/+) was successfully introduced into Agrobacterium tumefaciens strain EHA105.
2. The Ppy-ACO full-length cDNA (EF451060) is1,225bp, which contains is945nucleotides for ORF,63bp of the5'-UTR and217bp of the3'-UTR. The deduced amino acid sequence of the Ppy-ACO is314residues and includes twelve conserved amino acid residues belonging to non-heme iron (Ⅱ) dependent family of oxygenases/oxidases and three residues essential for emzyme activation. A binary antisense expression vector with Ppy-ACO coding region was constructed by inserting the target fragment in reverse orientation in pYPX145. The expression of the antisense gene is controlled by a double CaMV35S promoter. The vector was successfully introduced into Agrobacterium tumefaciens strain LBA4404.
3. Expression of Ppy-ACS and Ppy-ACO using real-time fluorescent qRT-PCR increased with fruit growth and development, and expression of the Ppy-ACO was obviously higher than that of the Ppy-ACS during fruit growth and development.1-MCP treatment obviously inhibited the Ppy-ACS expression in12days after harvest and down-regulated the Ppy-ACO expression throughout fruit storage period. However, exogenous ethyl ene significantly up-regulated the Ppy-ACS expression at early and later stages, and evidently promoted the Ppy-ACO expression all time after harvest, only reduced at the end of fruit storage.
4. Xyloglucan endotransglucosylase/hydrolase gene (named as Ppy-XTH, EU432411) is1,248bp contained73nuleotides of the5'-UTR, an ORF consiting of885nucleotides, and290nucleotides of the3'-UTR. The deduced amino acid sequence of the Ppy-XTH is294amino acid residues, which includes a signal peptide composed of22amino acid residues in the N-timminal, and shares a conserved DEIDFEFLD motif that is likely to be the catalytic site for both xyloglucan endotransglycosylase (XET) and xyloglucan endohydrolase (XEH) activity. The result of cluster analysis of amino acid sequence for mature Ppy-XTH polypeptide with other plant such as Arabidopsis, tomato, apple and kiwifruit showed Ppy-XTH belongs to group1, which expressed in flower and fruit. The accumulation of the Ppy-XTH mRNA in fruit could be detected during the whole fruit growth and development, obviously increased from the beginning of fruit rapid growth at98days after full bloom (DAFB), steadily maintained high level from98DAFB to126DAFB, evidently decreased at the end of fruit growth phase (140DAFB).
5. Three expansin gene cDNAs, designate as Ppy-EXP1(EF602031), Ppy-EXP2(FJ619347) and Ppy-EXP3(FJ619348), were cloned from mature fruit of sand pear. The full-length cDNA of Ppy-EXP1, Ppy-EXP2and Ppy-EXP3were1,369bp,1,107bp and1,136bp, respectively. The Ppy-EXP1, Ppy-EXP1and Ppy-EXP3ordinally contain an ORF of759bp,756bp and765bp, and encode a polypeptide consisting of252,251and254amino acid residues, respectively. They all contain a histidine (His-Phe-Asp, HFD) motif as conversed amino acid domain of expansin for most plants in the central region. A phylogenetic tree was generated from the deduced amino acid alignment of three Ppy-EXPs and26other expansin homologues. Ppy-EXP1and Ppy-EXP2aligned within group B and Ppy-EXP3belonged to group C. Accumulation of Ppy-EXP1and Ppy-EXP2in the developing fruit was low level during0-98DAFB, slightly increased at the end of growth phase, and the increments of the Ppy-EXP1are more than that of the Ppy-EXP2. Accumulation of the Ppy-EXP3was higher than that of the Ppy-EXP1and the Ppy-EXP1in the developing fruit, and increased markedly with fruit growth and development, and reached to the highest level at the end of growth phase (140DAFB).
6. Conserved microRNAs were detected in mature fruit by microarray based on6,703microRNAs sequences from miRBase11.0version (http://microrna.sanger.ac.uk/). Six hundred and forty-three microRNAs were detected based on positive signal. Eleven high frequently expressed microRNAs (eight from plant, three from animal) were selected for target prediction. Seven microRNA targets belong to transcription factor families, and the left one is involved in microRNAs metabolism. Isopentenyl transferase gene was predicted as a target for miR-30, but the other two animal microRNAs, miR-124and miR-181, didn't find their targets.
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