麻疯树pepc1基因的克隆及其调控烟草蛋白质和油脂合成的作用分析
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摘要
麻疯树主要分布于热带和亚热带地区,在我国栽培或半野生于热带地区和干热河谷地区。麻疯树种子种仁中含有丰富的油脂成分,因此它很有可能成为未来替代化石能源的具有巨大开发潜力的木本油料树种之一。但适宜麻疯树种植的土地面积十分有限,麻疯树种仁含油量的提高可提高麻疯树单位面积产油量,可以缓解在有限适宜土地上发展麻疯树生物柴油的矛盾。通过基因工程技术导入调控油脂合成的基因是提高麻疯树种子含油率,培育高油新品种的有效方法之一。近年来研究发现磷酸烯醇式丙酮酸羧化酶(PEPCase)参与了植物种子油脂合成的调控,因此通过反义抑制技术控制麻疯树pepc基因的表达,可能是提高麻疯树种子含油量的有效途径。
本研究以麻疯树未成熟种子为材料,采用RT-PCR和RACE法从中克隆了麻疯树pepc基因的一个家族成员pepc1,采用生物信息学分析了其氨基酸序列,构建pepc1基因的正义和反义表达载体,将它们导入烟草,通过转基因烟草的实时荧光定量PCR、PEPC酶活性测定、蛋白质、脂肪酸含量变化,研究pepc1基因在转录水平、翻译水平对烟草脂肪酸合成的影响,通过pepc1基因在麻疯树不同组织、种子发育不同阶段的表达模式研究pepc1基因在麻疯树中的转录水平表达情况,鉴定麻疯树pepc1基因对油脂合成的调控作用,为定向遗传改良提高麻疯树种子油含量建立技术体系和提供科学依据。主要研究结果如下：
(1)通过RT-PCR法克隆了麻疯树vepc基因的cDNA片段,在此基础上又通过5'RACE法克隆了pepc基因的5’端,该cDNA全长3124bp,其中开放阅读框(ORF)长为2898bp,编码了965个氨基酸。
(2)通过对麻疯树PEPCase蛋白的生物信息学分析表明,该蛋白的分子量为110.6kD,该蛋白酸性氨基酸占总氨基酸的14.6%,碱性氨基酸占总氨基酸的13.5%,理论等电点6.18,稳定系数为46.5,单条肽链属于不稳定蛋白。二级结构主要以无规卷曲为主,间或α螺旋和β直链。麻疯树PEPCase蛋白是亲水性的,无跨膜区及信号肽。本研究克隆的pepc基因编码的蛋白属于基因家族中的PEPC1,基因为pepc1。与蓖麻的氨基酸同源性最高,在系统进化树中也与蓖麻先聚合,它与烟草的同源性也很高。通过NCBI的保守功能域(CDD)搜索,发现在11～965aa之间是一个典型的大Ppc结构域,搜索蛋白质的活性位点,发现46个功能位点,分为7种模式。麻疯树PEPC1的结构特征与大部分PEPC的结构类似,功能位点略有差异。
(3)利用克隆获得的麻疯树管家基因actin作为参照。对麻疯树pepc1基因进行了时空表达模式研究,发现在开花到果实成熟过程中,在前4个阶段pepc1基因的表达量是增加的,在后5个阶段表达量明显下降,在叶中表达量不太高。
(4)根据植物表达载体pCAMBIA2300-35S、pBI121的多克隆位点特征和麻疯树pepc1基因序列特征,分别构建了pepc1基因的正义表达载体pCAMBIA-KSpepc和反义表达载体pBI-BXpepc。采用农杆菌介导法将正义、反义pepc1基因转化烟草,经卡那霉素筛选、PCR和PCR-Southern鉴定,结果获得了3株转正义pepc1基因的烟草和10株转反义pepc1基因的烟草。在对转麻疯树pepc1基因烟草的实时荧光定量PCR结果中发现,转正义pepc1基因的烟草,烟草内源pepc基因表达量变化不大,外源pepc1增加。10株转反义pepc1基因的烟草,有7株烟草内源pepc基因和外源pepc1基因都降低。
(5)通过对转基因烟草叶片PEPCase酶活性、蛋白质含量、总脂及脂肪酸含量等指标的测定,结果表明转正义pepc1基因的3株PEPCase酶活性、蛋白质含量均比野生型对照高,这3株植株中有1株总脂含量较对照高,其余两株较对照低。转反义pepc1基因的10株植株有9株酶活性比对照低,这9株中有5株其蛋白质含量比对照低,其中4株的总脂含量比对照高。转基因烟草的脂肪酸组份发生了一定的变化,但各成分含量主次变化不大。通过对蛋白质/总脂比率与野生型对照相比较,最终获得2株转正义pepc1基因和4株转反义pepc1基因的烟草与预期结果一致,其外源的麻疯树pepc1基因参与了油脂合成的调控。
Jatropha curcas L. is widely distributed in tropic and subtropic. In China, it is existed or cultivated or semi-wide in tropic and dry-hot valley region. Jatropha curcas L. has an huge development potential as an oil tree substituting fossil fuels in future, because of its high oil content in seeds. But sutibale lands for planting is so limited, and improving oil content in seeds is one of breeding goals. PEPCase is reported involved in oil synthesis payway. Therefore, down-regulated expression of pepc may enhance oil content in seeds antisense approach.
In this study, pepcl was cloned an a member of pepc gene family using RT-PCR and RACE techniques from development seeds of Jatropha curcas L.. Its amino acid sequence was analyzed. Expression vectors with sense and antisense of gene were contructed. These vectors were introduced into Agrobacteria and used to transform to tobacco. Transgenic taboccos were confirmed by Southern blot, real-time PCR, and further characterized by determining the activity of PEPC, content of protein and fatty acid. The expression patterns were analyzed in different tissues and different stages in developing seeds. Those results reflected expression of pepcl in level of transtription and translation. The main results are as followed:
1. The 3'-fragment of pepc from Jatropha curcas L. development seeds was isolated via RT-PCR, and 5'-fragment was amplified by RACE. The cDNA length of pepc is 3124 bp. The open read frame (ORF) contains 2898 bp,965 amino acids.
2. The molecular weight of PEPC is 110.6 kD with the the acidic amino acid account for 14.6%, alkali amino acid 13.5% of the total, and pⅠat 6.18. The single peptide chain was not stabily because of its low stability coefficient. The amino acid secondary structure is coil mainly, a-helical andβ-strand in occasion. The hydrophilic protein has not transmembrane region and signal peptide. The protein belongs to PEPC1 in pepc gene family, and the gene named pepcl. The homology of PEPC 1 of Jatropha curcas L. and Ricinus communis is the highest, and higher with tobacco.A typical large Ppc functional domain span 11-965aa was also found using the CDD program an NCBI. In addition,46 activity sites were found belonged to 7 types by searching in website at http://cn.expasy.org/prosite/.
3. The housekeeping gene of Jatropha curcas L., actin was cloned. The temporal and spatial expression patterns of pepcl Jatropha curcas L. were studied using real-time PCR. During different stages from flower to mature fruit, the pepcl expression shows increasing in first four stages, and decreasing in late five stages. The expression level of pepcl is low in leaves.
4. The sense expression vector pCAMBIA-KSpepc and antisense expression pBI-BXpepc were constructed based on the character of pepcl sequence. The sense pepcl and antisense pepcl gene were transformed into tobacco by Agrobacterium-mediated method. Three plants with sense pepcl and ten plants with antisense pepcl were obtained after kanamycin selection, PCR and PCR-Southern analysis. The transgenic tobacco plants were analyzed by real-time PCR. The expression level of pepc of tobacco endogenous changed little, but up-regulated of Jatropha curcas L. in sense pepcl plant. Of the ten antisense pepcl plants, seven plants have the expression level of both tobacco pepc and Jatropha curcas L. pepcl down-regulated.
5. The activity of PEPCase, contents of total protein and fat were determined from leaves of transgene tobacco. Three sense pepcl tobacco plants were found higher than wild type. Two of the three transgenic plants were lower than wild type in fat content. In ten antisense pepcl gene plants, nine plants were lower than wild type in activity of PEPCase, but five plants decreased in total protein content and four plants increased in fat content. The fatty acid composition of transgene plants were found changed little. Therefore, two sense pepcl gene tobacco and four antisense pepcl tobacco lines were obtained with expected result.
本研究以麻疯树未成熟种子为材料,采用RT-PCR和RACE法从中克隆了麻疯树pepc基因的一个家族成员pepc1,采用生物信息学分析了其氨基酸序列,构建pepc1基因的正义和反义表达载体,将它们导入烟草,通过转基因烟草的实时荧光定量PCR、PEPC酶活性测定、蛋白质、脂肪酸含量变化,研究pepc1基因在转录水平、翻译水平对烟草脂肪酸合成的影响,通过pepc1基因在麻疯树不同组织、种子发育不同阶段的表达模式研究pepc1基因在麻疯树中的转录水平表达情况,鉴定麻疯树pepc1基因对油脂合成的调控作用,为定向遗传改良提高麻疯树种子油含量建立技术体系和提供科学依据。主要研究结果如下：
(1)通过RT-PCR法克隆了麻疯树vepc基因的cDNA片段,在此基础上又通过5'RACE法克隆了pepc基因的5’端,该cDNA全长3124bp,其中开放阅读框(ORF)长为2898bp,编码了965个氨基酸。
(2)通过对麻疯树PEPCase蛋白的生物信息学分析表明,该蛋白的分子量为110.6kD,该蛋白酸性氨基酸占总氨基酸的14.6%,碱性氨基酸占总氨基酸的13.5%,理论等电点6.18,稳定系数为46.5,单条肽链属于不稳定蛋白。二级结构主要以无规卷曲为主,间或α螺旋和β直链。麻疯树PEPCase蛋白是亲水性的,无跨膜区及信号肽。本研究克隆的pepc基因编码的蛋白属于基因家族中的PEPC1,基因为pepc1。与蓖麻的氨基酸同源性最高,在系统进化树中也与蓖麻先聚合,它与烟草的同源性也很高。通过NCBI的保守功能域(CDD)搜索,发现在11～965aa之间是一个典型的大Ppc结构域,搜索蛋白质的活性位点,发现46个功能位点,分为7种模式。麻疯树PEPC1的结构特征与大部分PEPC的结构类似,功能位点略有差异。
(3)利用克隆获得的麻疯树管家基因actin作为参照。对麻疯树pepc1基因进行了时空表达模式研究,发现在开花到果实成熟过程中,在前4个阶段pepc1基因的表达量是增加的,在后5个阶段表达量明显下降,在叶中表达量不太高。
(4)根据植物表达载体pCAMBIA2300-35S、pBI121的多克隆位点特征和麻疯树pepc1基因序列特征,分别构建了pepc1基因的正义表达载体pCAMBIA-KSpepc和反义表达载体pBI-BXpepc。采用农杆菌介导法将正义、反义pepc1基因转化烟草,经卡那霉素筛选、PCR和PCR-Southern鉴定,结果获得了3株转正义pepc1基因的烟草和10株转反义pepc1基因的烟草。在对转麻疯树pepc1基因烟草的实时荧光定量PCR结果中发现,转正义pepc1基因的烟草,烟草内源pepc基因表达量变化不大,外源pepc1增加。10株转反义pepc1基因的烟草,有7株烟草内源pepc基因和外源pepc1基因都降低。
(5)通过对转基因烟草叶片PEPCase酶活性、蛋白质含量、总脂及脂肪酸含量等指标的测定,结果表明转正义pepc1基因的3株PEPCase酶活性、蛋白质含量均比野生型对照高,这3株植株中有1株总脂含量较对照高,其余两株较对照低。转反义pepc1基因的10株植株有9株酶活性比对照低,这9株中有5株其蛋白质含量比对照低,其中4株的总脂含量比对照高。转基因烟草的脂肪酸组份发生了一定的变化,但各成分含量主次变化不大。通过对蛋白质/总脂比率与野生型对照相比较,最终获得2株转正义pepc1基因和4株转反义pepc1基因的烟草与预期结果一致,其外源的麻疯树pepc1基因参与了油脂合成的调控。
Jatropha curcas L. is widely distributed in tropic and subtropic. In China, it is existed or cultivated or semi-wide in tropic and dry-hot valley region. Jatropha curcas L. has an huge development potential as an oil tree substituting fossil fuels in future, because of its high oil content in seeds. But sutibale lands for planting is so limited, and improving oil content in seeds is one of breeding goals. PEPCase is reported involved in oil synthesis payway. Therefore, down-regulated expression of pepc may enhance oil content in seeds antisense approach.
In this study, pepcl was cloned an a member of pepc gene family using RT-PCR and RACE techniques from development seeds of Jatropha curcas L.. Its amino acid sequence was analyzed. Expression vectors with sense and antisense of gene were contructed. These vectors were introduced into Agrobacteria and used to transform to tobacco. Transgenic taboccos were confirmed by Southern blot, real-time PCR, and further characterized by determining the activity of PEPC, content of protein and fatty acid. The expression patterns were analyzed in different tissues and different stages in developing seeds. Those results reflected expression of pepcl in level of transtription and translation. The main results are as followed:
1. The 3'-fragment of pepc from Jatropha curcas L. development seeds was isolated via RT-PCR, and 5'-fragment was amplified by RACE. The cDNA length of pepc is 3124 bp. The open read frame (ORF) contains 2898 bp,965 amino acids.
2. The molecular weight of PEPC is 110.6 kD with the the acidic amino acid account for 14.6%, alkali amino acid 13.5% of the total, and pⅠat 6.18. The single peptide chain was not stabily because of its low stability coefficient. The amino acid secondary structure is coil mainly, a-helical andβ-strand in occasion. The hydrophilic protein has not transmembrane region and signal peptide. The protein belongs to PEPC1 in pepc gene family, and the gene named pepcl. The homology of PEPC 1 of Jatropha curcas L. and Ricinus communis is the highest, and higher with tobacco.A typical large Ppc functional domain span 11-965aa was also found using the CDD program an NCBI. In addition,46 activity sites were found belonged to 7 types by searching in website at http://cn.expasy.org/prosite/.
3. The housekeeping gene of Jatropha curcas L., actin was cloned. The temporal and spatial expression patterns of pepcl Jatropha curcas L. were studied using real-time PCR. During different stages from flower to mature fruit, the pepcl expression shows increasing in first four stages, and decreasing in late five stages. The expression level of pepcl is low in leaves.
4. The sense expression vector pCAMBIA-KSpepc and antisense expression pBI-BXpepc were constructed based on the character of pepcl sequence. The sense pepcl and antisense pepcl gene were transformed into tobacco by Agrobacterium-mediated method. Three plants with sense pepcl and ten plants with antisense pepcl were obtained after kanamycin selection, PCR and PCR-Southern analysis. The transgenic tobacco plants were analyzed by real-time PCR. The expression level of pepc of tobacco endogenous changed little, but up-regulated of Jatropha curcas L. in sense pepcl plant. Of the ten antisense pepcl plants, seven plants have the expression level of both tobacco pepc and Jatropha curcas L. pepcl down-regulated.
5. The activity of PEPCase, contents of total protein and fat were determined from leaves of transgene tobacco. Three sense pepcl tobacco plants were found higher than wild type. Two of the three transgenic plants were lower than wild type in fat content. In ten antisense pepcl gene plants, nine plants were lower than wild type in activity of PEPCase, but five plants decreased in total protein content and four plants increased in fat content. The fatty acid composition of transgene plants were found changed little. Therefore, two sense pepcl gene tobacco and four antisense pepcl tobacco lines were obtained with expected result.
引文
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