红掌‘阿拉巴马’低温相关基因AOX、CAT和APX的表达分析、功能验证及遗传转化体系的构建
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摘要
红掌(Anthurium andraeanum)是天南星科花烛属高档室内观赏盆花,属佛焰花序类植物。红掌由于花型奇特、颜色鲜艳多彩而深受人们喜爱,是居家装饰和馈赠亲友的最好选择,在盆花市场特别是年宵花市场上占了很大比例,因此在浙江省的种植面积正在迅速扩增。而且随着人们生活水平的不断提高,消费理念的转变,对高档盆花的需求越来越大,市场还有着巨大潜力。然而由于红掌原产热带,对温度较为敏感,生长温度要求15℃以上,低于12℃容易产生冷害,低温是限制红掌生长发育的重要逆境因子。在我省种植存在高能耗、高成本的问题,降低了市场竞争力,影响了红掌盆花产业的持续、健康的发展。要解决这个问题的最根本途径就是采用育种手段从根本上提高红掌的抗寒性,对于我省的花卉产业的做大做强则显得十分必要和紧迫。本研究采用PCR结合RACE技术,以红掌‘阿拉巴马’为材料,从其叶片中克隆得到交替氧化酶AOX,过氧化氢酶CAT和抗坏血酸过氧化物酶APX3个基因全长,并对其进行生物信息学分析；运用实时定量PCR技术分析各基因在红掌不同组织器官和不同低温胁迫下的表达情况：构建了红掌抗坏血酸过氧化物酶基因的植物表达载体,并转化烟草进行功能验证；最后,将脂肪酸去饱和酶基因转化红掌并获得了抗性植株。取得的研究结果如下：
1、红掌交替氧化酶基因AnAOX全长基因的克隆与序列分析
从红掌叶片中克隆与低温胁迫相关的基因AOX,其全长序列为1170bp,命名为AnAOX (GenBank登录号：JX163297),该基因含有1038bp的开放阅读框,编码345个氨基酸,AnAOX编码蛋白预测的等电点(pI)为8.51,分子量大小约为38.0kD。利用DNAMAN5.2.2软件,分析AnAOX基因编码的氨基酸序列,认为该基因具有完整的3’和5’末端,是花烛属红掌交替氧化酶基因的全长序列。红掌AOX推测的氨基酸序列与马铃薯(Solanum tuberosum)、水芭蕉(Lysichiton camtschatcensis)、羽叶蔓绿绒(Philodendron bipinnatifidum)和肾叶臭菘(Symplocarpus renifolius)的AOX的同源性分别为82%、80%、78%和77%。所有的序列中都存在完全保守的铁离子结合区域和组氨酸残基,它们可能与AOX基因在植物中的功能密切相关。
2、红掌过氧化氢酶基因AnCAT基因的克隆与序列分析
以红掌‘阿拉巴马’品种叶片提取的总RNA为模板,通过RT-PCR与RACE扩增,获得一个1704bp的过氧化氢酶(Catalase, CAT)基因的cDNA序列,其基因编码区共1476bp,编码492个氨基酸,命名为AnCAT(GenBank登录号：JX163298)。AnCAT编码蛋白预测的等电点(pI)为6.89,相对分子量约为57.1kD。红掌CAT推测的氨基酸序列与与马蹄莲(Zantedeschia aethiopica)、油棕(Elaeis guineensis)、小果野蕉(Musa acuminata)和籼稻(Oryza sativa Indica)的CAT的同源性分别为91%、87%、87%和81%。
3、红掌抗坏血酸过氧化物酶基因AnAPX基因的克隆与序列分析
以天南星利花烛属红掌‘阿拉巴马’品种叶片提取的总RNA为模板,通过RT-PCR与RACE扩增,获得一个888bp的抗坏血酸过氧化物酶(Ascorbate peroxidase, APX)基因的cDNA序列,其基因编码区共750bp,编码250个氨基酸,命名为AnAPX(GenBank登录号：JQ8385071)。根据Protparam在线软件预测红掌抗坏血酸过氧化物酶蛋白的理化性质,结果表明推测APX蛋白的分子式为C1232H1904N330O363S7,相对分子量约为27.4kD,等电点(pI)为5.40；理论推导半衰期约30h,不稳定参数为32.55,属于稳定蛋白。红掌APX推测的氨基酸序列与马蹄莲(Zantedeschia aethiopica)、华东葡萄(Vitis pseucdoreticulata)、棉花(Gossypium hirsutum)、油棕(Elaeis guineensis)和玉米(Zea mays)的APX的同源性分别为93%、87%、87%和86%。
4、红掌inAOX、AnCAT和AnAPX基因的表达分析
通过定量PCR研究上述三个基因在转录水平上表达与红掌叶片低温胁迫之间的关系发现,6℃处理后,,红掌叶片AnAOX, AnCAT和AnAPX基因均上调表达。其中AnAOX基因在低温处理12h后上调表达不明显,处理24h后开始明显上调表达,且处理24h表达量显著高于12h；低温处理超出24h后,表达量略有下降,36～48h之间表达量没有明显差异。AnCAI基因在低温处理12h后表达明显上调,超过12h表达量开始下降,当低温处理时间达到48h时,表达量与对照接近甚至略低于对照；AnAPX基因在低温处理12h明显上调表达,且随着低温处理时间的延长,上调表达逐渐上升,当处理48h后上调表达达到顶峰。由此推测,红掌AnAPX基因表达量的变化与低温及低温持续时间关系最密切,其次是AnAOX基因、AnCAT基因。
5、红掌AnAPX植物表达载体的构建
根据AnAPX基因全长序列设计特异引物,以红掌‘阿拉巴马’叶片为试材,得到AnAPX基因完整的开放阅读框。将扩增到的片段与pMD18-T载体连接并进行序列测定,表明插入片段正确、克降片段经双酶切消化插入到植物表达载体pCAMBIA2300中,构建了AnAPX基因的表达载体。在成功构建AnAPX表达载体后,采用电击法将重组质粒转化到根癌农杆菌菌株中,为后续AnAPX功能验证莫定了基础。
6、汀掌基因转化
采用根癌农杆菌介导法,以红掌愈伤组织为侵染材料,将绿色荧光蛋白基因GFP和脂肪酸去饱和酶基因FAD3转化红掌,通过G418和卡那霉素筛选,获得了转基因红掌植株。用PCR反应对具卡那霉素抗性的转基因红掌植株进一步鉴定,结果表明目的基因FAD3已经整合到部分转基因植株的基因组中,成功地获得了转FAD3基因的红掌植株。
Anthurium andraeanum is the Araceae Anthurium high-grade indoor ornamental potted plants, belonging to the spadix plants. A. andraeanum is deeply loved by many people due to its beautiful shape and bright colour. It is the best choice for home decoration and gift for relatives and friends. It takes up a great percentage of flower market especially in the Lunar New Year Flower Market. Therefore, its planting area has been rapidly increasing in our province. Moreover, with the improvement of people's living standard and the change of comsumption concept, the demand for high-grade potted flower is becoming greater and greater and it has huge potential in the market. However, as A. andraeanum is native to the tropical area and sensitive to temperature, it requires the growth temperature above15℃and cold damage could easily take place below12℃. Low temperature is one of the most important factors which constrain the growth of A. andraeanum. The production of Anthurium in our province is facing the problems of high energy cost, which reduces the market competitiveness and affects the sustainable and healthy development of the industry of A. andraeanum potted flower. The most effective way to solve this problem is to use the breeding method to improve the cold resistance of A. andraeanum plants. In this study, using A. andraeanum 'alabama' as material, three full-length genes of alternative oxidase, catalase and ascorbate peroxidase were cloned by PCR combined with RACE technology and analyzed by bioinformatic tools. The real-time PCR technique was used to analyze the gene expression in different organs of A. andraeanum and under different low temperature stresses. The plant expression vector of ascorbate peroxidase gene from A. andraeanum was constructed and APX gene was transformed into tobacco plants in order to indentify its function. Finally, the fatty acid desaturase gene was transferred to A. andraeanum and the resistant plants were obtained. The results are as follows:1. The cloning and sequence analysis of a full length alternative oxidase gene (AnAOX) from A. andraeanum
An alternative oxidase gene was cloned from A. andraeanum leaves. The full-length AOX cDNA sequence was1170bp, which contained an open reading frame of1038bp. This gene was named as AnAOX and the genbank accession number is JX163297. The sequence encoded a protein of345amino acid residues with a calculated molecular weight of38.0kD consisting of an isoelectric point of8.51. The amino acid sequence of AnAOX gene code was analyzed using DNAMAN5.2.2software. The results showed that this gene has the whole3and5 terminal and it is the full length sequence of alternative oxidase gene of A. andraeanum.2. The cloning and sequence analysis of a full length hydrogen peroxide oxidase gene (AnCAT) of A. andraeanum
A hydrogen peroxide gene was cloned from A. andraeanum 'alabama' leaves. The full-length CAT cDNA sequence was1704bp, which contained an open reading frame of1476bp. This gene was named as AnCAT and the genbank accession number is JX163298. The sequence encoded a protein of492amino acid residues with a calculated molecular weight of57.1kD consisting of an isoelectric point of6.89. The homology between the CAT of tentative amino acid sequence of A. andraeanum and that of Zantedeschia aethiopica, Elaeis guineensis, Musa acuminata and Oryza sativa Indica are91%,87%,87%and81%, respectively.3. The cloning and sequence analysis of a full length ascorbate peroxidase gene(AnAPX) of A. andraeanum
An ascorbate peroxidase gene was cloned from A. andraeanum'alabama'leaves. The full-length APX cDNA sequence was888bp, which contained an open reading frame of750bp. This gene was named as AnAPX and the genbank accession number is JQ8385071. The sequence encoded a protein of250amino acid residues with a calculated molecular weight of27.4kD consisting of an isoelectric point of5.4. According to the prediction of physicochemical property of AnAPX protein by ProtParam online software, it suggested that the tentative APX protein molecular formula was C1232H1904N330O363S7. The molecular weight was about27.4kD with an isoelectric point of5.4, half-life period of about30h and the unstable parameter of32.55, belonging to stable protein. The homology between the AnAPX of tentative amino acid sequence and those of Zantedeschia aethiopica, Vitis pseudoreticulata, Gossypium hirsutum, Elaeis guineensis and Zea mays were93%,87%,87%and86%, respectively.
4. The gene expression analysis of AnAOX, AnCAT and AnAPX under cold stress
Using the quantitative real time PCR, the gene expression levels of the three genes in the A. andraeanum leaves were determined after different time of low temperature stress (6℃), we observed that the expression of AnAOX, AnCAT and AnAPX genes were all up-regulated. The AnAOX gene upregulation of the expression was not obvious after12h of low temperature treatment, and became obvious after24h low temperature treatment. The expression level of24h is obviously higher than that of12h. After more than24h low temperature treatment, the expression level slightly went down and there was no obvious difference during the period of36～48h. The AnCAT gene expression upregulation was obvious after12h low temperature treatment and its expression level began to drop after more than12h. The expression level closed to or even slightly below to the control when low temperature treatment reached48h. The AnAPX gene expression upregulation was obvious after12h low temperature treatment. Along with the time of the low temperature treatment, its expression rose gradually and reached to the peak when treated48h with low temperature. It is speculated that the change of the expression level of AnAPX gene was most closely related to the low temperature and its time course, followed by AnAOX, and finally AnCAT gene.
5. The building of plant expression vector of AnAPX.
Design the peculiar primer on the basis of AnAPX gone full length sequence, taking the A. andraeanum'alabama'as test material, we got the whole open reading frame of AnAPX gene. The obtained section was connected with the pMD18-T vector and measured, and the results suggested that the inserted section was correct. After double enzyme digestion cut, the clone section has been inserted into the plant expression vector pCAMBIA2300and the expression vector of AnAPX gene has been built. Having built the expression vector of AnAPX successfully, we used electroporation to transform recombinant plasmid into the bacterial strain of agrobacterium tumefaciens, which established the basis of the follow-up function identification of AnAPX.
6. The gene transformation of A. andraeanum
Using meloidogynosis sonication-assisted Agrobacterium-mediated transformation, taking callus of A. andraeanum as infection material, we converted the green fluorescent protein (GFP) gene and fatty acid desaturase (FAD3) gene into A. andraeanum. By screening with G418and kanamycin, we have gotten the transgenic plants of A. andreanum. Further identification of the transgenic plants showed that the target gene FAD3had been integrated into the genome of partial transgenic plants. Therefore, we have successfully obtained the transgenic plants of A. andreanum with FAD3gene.
1、红掌交替氧化酶基因AnAOX全长基因的克隆与序列分析
从红掌叶片中克隆与低温胁迫相关的基因AOX,其全长序列为1170bp,命名为AnAOX (GenBank登录号：JX163297),该基因含有1038bp的开放阅读框,编码345个氨基酸,AnAOX编码蛋白预测的等电点(pI)为8.51,分子量大小约为38.0kD。利用DNAMAN5.2.2软件,分析AnAOX基因编码的氨基酸序列,认为该基因具有完整的3’和5’末端,是花烛属红掌交替氧化酶基因的全长序列。红掌AOX推测的氨基酸序列与马铃薯(Solanum tuberosum)、水芭蕉(Lysichiton camtschatcensis)、羽叶蔓绿绒(Philodendron bipinnatifidum)和肾叶臭菘(Symplocarpus renifolius)的AOX的同源性分别为82%、80%、78%和77%。所有的序列中都存在完全保守的铁离子结合区域和组氨酸残基,它们可能与AOX基因在植物中的功能密切相关。
2、红掌过氧化氢酶基因AnCAT基因的克隆与序列分析
以红掌‘阿拉巴马’品种叶片提取的总RNA为模板,通过RT-PCR与RACE扩增,获得一个1704bp的过氧化氢酶(Catalase, CAT)基因的cDNA序列,其基因编码区共1476bp,编码492个氨基酸,命名为AnCAT(GenBank登录号：JX163298)。AnCAT编码蛋白预测的等电点(pI)为6.89,相对分子量约为57.1kD。红掌CAT推测的氨基酸序列与与马蹄莲(Zantedeschia aethiopica)、油棕(Elaeis guineensis)、小果野蕉(Musa acuminata)和籼稻(Oryza sativa Indica)的CAT的同源性分别为91%、87%、87%和81%。
3、红掌抗坏血酸过氧化物酶基因AnAPX基因的克隆与序列分析
以天南星利花烛属红掌‘阿拉巴马’品种叶片提取的总RNA为模板,通过RT-PCR与RACE扩增,获得一个888bp的抗坏血酸过氧化物酶(Ascorbate peroxidase, APX)基因的cDNA序列,其基因编码区共750bp,编码250个氨基酸,命名为AnAPX(GenBank登录号：JQ8385071)。根据Protparam在线软件预测红掌抗坏血酸过氧化物酶蛋白的理化性质,结果表明推测APX蛋白的分子式为C1232H1904N330O363S7,相对分子量约为27.4kD,等电点(pI)为5.40；理论推导半衰期约30h,不稳定参数为32.55,属于稳定蛋白。红掌APX推测的氨基酸序列与马蹄莲(Zantedeschia aethiopica)、华东葡萄(Vitis pseucdoreticulata)、棉花(Gossypium hirsutum)、油棕(Elaeis guineensis)和玉米(Zea mays)的APX的同源性分别为93%、87%、87%和86%。
4、红掌inAOX、AnCAT和AnAPX基因的表达分析
通过定量PCR研究上述三个基因在转录水平上表达与红掌叶片低温胁迫之间的关系发现,6℃处理后,,红掌叶片AnAOX, AnCAT和AnAPX基因均上调表达。其中AnAOX基因在低温处理12h后上调表达不明显,处理24h后开始明显上调表达,且处理24h表达量显著高于12h；低温处理超出24h后,表达量略有下降,36～48h之间表达量没有明显差异。AnCAI基因在低温处理12h后表达明显上调,超过12h表达量开始下降,当低温处理时间达到48h时,表达量与对照接近甚至略低于对照；AnAPX基因在低温处理12h明显上调表达,且随着低温处理时间的延长,上调表达逐渐上升,当处理48h后上调表达达到顶峰。由此推测,红掌AnAPX基因表达量的变化与低温及低温持续时间关系最密切,其次是AnAOX基因、AnCAT基因。
5、红掌AnAPX植物表达载体的构建
根据AnAPX基因全长序列设计特异引物,以红掌‘阿拉巴马’叶片为试材,得到AnAPX基因完整的开放阅读框。将扩增到的片段与pMD18-T载体连接并进行序列测定,表明插入片段正确、克降片段经双酶切消化插入到植物表达载体pCAMBIA2300中,构建了AnAPX基因的表达载体。在成功构建AnAPX表达载体后,采用电击法将重组质粒转化到根癌农杆菌菌株中,为后续AnAPX功能验证莫定了基础。
6、汀掌基因转化
采用根癌农杆菌介导法,以红掌愈伤组织为侵染材料,将绿色荧光蛋白基因GFP和脂肪酸去饱和酶基因FAD3转化红掌,通过G418和卡那霉素筛选,获得了转基因红掌植株。用PCR反应对具卡那霉素抗性的转基因红掌植株进一步鉴定,结果表明目的基因FAD3已经整合到部分转基因植株的基因组中,成功地获得了转FAD3基因的红掌植株。
Anthurium andraeanum is the Araceae Anthurium high-grade indoor ornamental potted plants, belonging to the spadix plants. A. andraeanum is deeply loved by many people due to its beautiful shape and bright colour. It is the best choice for home decoration and gift for relatives and friends. It takes up a great percentage of flower market especially in the Lunar New Year Flower Market. Therefore, its planting area has been rapidly increasing in our province. Moreover, with the improvement of people's living standard and the change of comsumption concept, the demand for high-grade potted flower is becoming greater and greater and it has huge potential in the market. However, as A. andraeanum is native to the tropical area and sensitive to temperature, it requires the growth temperature above15℃and cold damage could easily take place below12℃. Low temperature is one of the most important factors which constrain the growth of A. andraeanum. The production of Anthurium in our province is facing the problems of high energy cost, which reduces the market competitiveness and affects the sustainable and healthy development of the industry of A. andraeanum potted flower. The most effective way to solve this problem is to use the breeding method to improve the cold resistance of A. andraeanum plants. In this study, using A. andraeanum 'alabama' as material, three full-length genes of alternative oxidase, catalase and ascorbate peroxidase were cloned by PCR combined with RACE technology and analyzed by bioinformatic tools. The real-time PCR technique was used to analyze the gene expression in different organs of A. andraeanum and under different low temperature stresses. The plant expression vector of ascorbate peroxidase gene from A. andraeanum was constructed and APX gene was transformed into tobacco plants in order to indentify its function. Finally, the fatty acid desaturase gene was transferred to A. andraeanum and the resistant plants were obtained. The results are as follows:1. The cloning and sequence analysis of a full length alternative oxidase gene (AnAOX) from A. andraeanum
An alternative oxidase gene was cloned from A. andraeanum leaves. The full-length AOX cDNA sequence was1170bp, which contained an open reading frame of1038bp. This gene was named as AnAOX and the genbank accession number is JX163297. The sequence encoded a protein of345amino acid residues with a calculated molecular weight of38.0kD consisting of an isoelectric point of8.51. The amino acid sequence of AnAOX gene code was analyzed using DNAMAN5.2.2software. The results showed that this gene has the whole3and5 terminal and it is the full length sequence of alternative oxidase gene of A. andraeanum.2. The cloning and sequence analysis of a full length hydrogen peroxide oxidase gene (AnCAT) of A. andraeanum
A hydrogen peroxide gene was cloned from A. andraeanum 'alabama' leaves. The full-length CAT cDNA sequence was1704bp, which contained an open reading frame of1476bp. This gene was named as AnCAT and the genbank accession number is JX163298. The sequence encoded a protein of492amino acid residues with a calculated molecular weight of57.1kD consisting of an isoelectric point of6.89. The homology between the CAT of tentative amino acid sequence of A. andraeanum and that of Zantedeschia aethiopica, Elaeis guineensis, Musa acuminata and Oryza sativa Indica are91%,87%,87%and81%, respectively.3. The cloning and sequence analysis of a full length ascorbate peroxidase gene(AnAPX) of A. andraeanum
An ascorbate peroxidase gene was cloned from A. andraeanum'alabama'leaves. The full-length APX cDNA sequence was888bp, which contained an open reading frame of750bp. This gene was named as AnAPX and the genbank accession number is JQ8385071. The sequence encoded a protein of250amino acid residues with a calculated molecular weight of27.4kD consisting of an isoelectric point of5.4. According to the prediction of physicochemical property of AnAPX protein by ProtParam online software, it suggested that the tentative APX protein molecular formula was C1232H1904N330O363S7. The molecular weight was about27.4kD with an isoelectric point of5.4, half-life period of about30h and the unstable parameter of32.55, belonging to stable protein. The homology between the AnAPX of tentative amino acid sequence and those of Zantedeschia aethiopica, Vitis pseudoreticulata, Gossypium hirsutum, Elaeis guineensis and Zea mays were93%,87%,87%and86%, respectively.
4. The gene expression analysis of AnAOX, AnCAT and AnAPX under cold stress
Using the quantitative real time PCR, the gene expression levels of the three genes in the A. andraeanum leaves were determined after different time of low temperature stress (6℃), we observed that the expression of AnAOX, AnCAT and AnAPX genes were all up-regulated. The AnAOX gene upregulation of the expression was not obvious after12h of low temperature treatment, and became obvious after24h low temperature treatment. The expression level of24h is obviously higher than that of12h. After more than24h low temperature treatment, the expression level slightly went down and there was no obvious difference during the period of36～48h. The AnCAT gene expression upregulation was obvious after12h low temperature treatment and its expression level began to drop after more than12h. The expression level closed to or even slightly below to the control when low temperature treatment reached48h. The AnAPX gene expression upregulation was obvious after12h low temperature treatment. Along with the time of the low temperature treatment, its expression rose gradually and reached to the peak when treated48h with low temperature. It is speculated that the change of the expression level of AnAPX gene was most closely related to the low temperature and its time course, followed by AnAOX, and finally AnCAT gene.
5. The building of plant expression vector of AnAPX.
Design the peculiar primer on the basis of AnAPX gone full length sequence, taking the A. andraeanum'alabama'as test material, we got the whole open reading frame of AnAPX gene. The obtained section was connected with the pMD18-T vector and measured, and the results suggested that the inserted section was correct. After double enzyme digestion cut, the clone section has been inserted into the plant expression vector pCAMBIA2300and the expression vector of AnAPX gene has been built. Having built the expression vector of AnAPX successfully, we used electroporation to transform recombinant plasmid into the bacterial strain of agrobacterium tumefaciens, which established the basis of the follow-up function identification of AnAPX.
6. The gene transformation of A. andraeanum
Using meloidogynosis sonication-assisted Agrobacterium-mediated transformation, taking callus of A. andraeanum as infection material, we converted the green fluorescent protein (GFP) gene and fatty acid desaturase (FAD3) gene into A. andraeanum. By screening with G418and kanamycin, we have gotten the transgenic plants of A. andreanum. Further identification of the transgenic plants showed that the target gene FAD3had been integrated into the genome of partial transgenic plants. Therefore, we have successfully obtained the transgenic plants of A. andreanum with FAD3gene.
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