油菜籽油脂合成途径上游ACCase和PEPCase基因的克隆及功能研究
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摘要
植物油是日常生活中的必需品,是能量聚集的自然平台,与消费者的生活息息相关。种子是植物油的主要贮藏器官,改良植物种子脂肪酸组成和提高含油量是油料作物研究的热点之一。在脂肪酸生物合成途径中,乙酰辅酶A羧化酶(Acetyl CoA Carboxylase, ACCase)是其中的关键酶之一,其主要作用是催化乙酰辅酶A羧化形成丙二酰辅酶A,为脂肪酸和许多次生代谢产物的合成提供底物。另外,脂肪酸和蛋白质的合成有关,两个代谢过程均需要利用糖酵解产物磷酸烯醇式丙酮酸(Phosphoenolpyruvate, PEP)这一相同底物。PEP在丙酮酸激酶作用下转化为丙酮酸,丙酮酸生成乙酰辅酶A后在ACCase作用下进入脂肪酸合成途径,而磷酸烯醇式丙酮酸羧化酶(Phosphoenolpyruvate carboxylase, PEPC)催化PEP合成草酰乙酸最终进入蛋白质代谢。PEPC是植物中具有多种生理功能的细胞质酶,大量研究表明PEPC参与脂肪酸调控、碳／氮吸收并与盐和干旱胁迫反应有关,但对其功能认识仍很有限,尤其是细菌型PEPC。本研究分别利用正向遗传学和反向遗传学方法研究了脂肪酸代谢过程中相关酶ACCase和PEPC基因的调控机制。
由于accD基因编码的β-CT亚基是ACCase基因表达的限制因子,本研究通过在油菜籽粒中超量表达大肠杆菌异质型ACCase基因,加速乙酰CoA羧化形成丙二酸单酰CoA这一关键进程,提高油菜籽脂肪酸的起始合成能力。本研究克隆了甘蓝型油菜及近缘种中异质型ACCase的β-CT亚基accD编码基因,采用生物信息学的方法分析了氨基酸序列,构建了受种子特异表达启动子驱动的异质型大肠杆菌ACCase β-CT亚基accD基因的植物表达载体以及大肠杆菌ACCase四个亚基基因分别受种子特异性表达启动子驱动的植物串联表达载体,将这两个载体分别用于转化甘蓝型油菜“超油2号”。通过转基因油菜的半定量PCR、蛋白质、脂肪酸含量变化,研究了异质型大肠杆菌基因accD与其它三个细胞核基因的互作对油菜脂肪酸合成的影响,为定向遗传改良油菜种子含油量建立了有效的技术体系。本研究还利用人工小RNA (artifical mRNA, amiRNA)技术敲除拟南芥细菌型Atppc4基因进行功能分析,研究该基因在脂肪酸代谢以及耐盐胁迫等方面的作用,以探讨拟南芥中细菌型Atppc4基因敲除对植物型PEPC的影响,为今后改良油菜品质提供理论依据。主要结果如下：
1.以甘蓝型油菜(Brassica napus)、甘蓝(Brassica oleracea)、芥菜型油菜(Brassica juncea)和白菜型油菜(Brassica rapa)幼嫩叶片的cDNA为模板,对异质型乙酰辅酶A羧化酶p-CT亚基(羧基转移酶)的accD编码基因进行扩增,得到长度分别为1470、1470、1464和1464bp的编码序列,分别可编码489、489、487和487个氨基酸的蛋白质。白菜型油菜、芥菜型油菜和甘蓝的accD序列登录号分别为EU410075、EU410076和EU410077。结果表明来自4个油菜近缘种的accD基因高度同源,编码的蛋白质都具有相似的锌指结构和C-末端5个相同的基元,其中基元I (GSMGSVVG)和基元Ⅱ(PLIIVCASGGARMQE)在所有植物及大肠杆菌(Escherichia coli)的β-CT亚基中普遍存在。Southern杂交显示该基因在甘蓝型油菜及其近缘种的基因组中呈单拷贝。
2.本研究E. coli ACCase的β-CT亚基accD (eaccD)基因与拟南芥Rubisco小亚基(rbcS)转运肽基因融合,构建以油菜Napin启动子驱动的种子特异表达载体。利用根癌农杆菌(Agrobacterium tumefaciens)介导法,将该表达载体转入“超油2号”,获得了7个阳性转基因株系。RT-PCR检测其中3株的结果证明eaccD基因可在转基因株系的种子中转录,但在叶片中不转录,说明该表达载体转入油菜中能引起植株种子eaccD基因的特异表达。通过转基因与对照植株种子的油脂和蛋白质含量的比较分析,发现对照种子的平均含油量为48.01%、蛋白质平均含量为25.10%；而7个转基因株系油菜种子的平均含油量为52.08%,比对照含油量相对增加8.45%；但转基因株系的蛋白质含量则有不同程度的降低,平均比对照植株下降约6.69%。转基因油菜千粒重比对照植株有所提高。因此大肠杆菌accD基因在油菜中的表达能够增加种子重量和含油量。
3.采用同尾酶反复酶切的方法,构建了拟南芥Rubisco小亚基转运肽与ACCase各亚基基因的融合串联表达载体,用A. tumefaciens介导的下胚轴转化对照品种“超油2号”。经过筛选获得了11株转化再生植株。经PCR. Southern和GUS染色检测证明4个目的基因已整合到T0再生植株油菜基因组中。T2转基因株系平均含油量为49.59%,转基因株系的含油量比对照(48.01%)提高了3.30%。
4.构建了人工小RNA表达载体Atppc4-amiRNA,利用农杆菌介导花絮浸染法(in floral dip)转化拟南芥,对转基因株系进行了分子检测。在拟南芥转基因株系内检测到成熟小RNA的表达,拟南芥转基因株系根中Atppc4的表达水平明显下降,然而其它3个植物型PEPC基因Atppc1、Atppc2和Atppc3在根中的表达明显得到上调。转基因株系根中的PEPC酶活性提高了5.1倍,表明植物中细菌型和植物型PEPC基因具有互作,细菌型PEPC基因Atppc4可以调控植物型PEPC基因的表达。分析拟南芥株系和对照植株的脂肪酸含量和组分的结果表明,转基因株系和对照的脂肪酸含量和脂肪酸组分未发生明显变化。
5.atppc4的下调表达提高拟南芥的耐盐能力。在非盐处理培养基上,Atppc4-amiRNA转基因株系比野生型的根系生长更慢。发现7日龄转基因株系的长度比野生型株系短29.0%,将这些植株移到含有150mM NaCl的1/2MS培养基上培养7天,转基因株系长度比野生型株系要短16.3%。说明转基因株系的根系由于Atppc4基因表达被抑制；盐处理后转基因株系根的生长能得到部分缓解,从而缩短了与野生型根长的距离。该结果说明特异性敲除细菌型Atppc4基因提高了拟南芥的耐盐性。
Vegetable oil is an essential product in our daily life and constitutes a major source of energy its consumers. The seed is the main storage organ of the oil plants. The improvement in the quality and quantity of seed fatty acid have currently become major research topic. Fatty acid biosynthesis in plants takes place within plastids. Acetyl-coenzyme A carboxylase (ACCase) catalyzes the first committed step of fatty acid synthesis, the carboxylastion of acetyl-CoA to malonyl-CoA which supplies the substrates for fatty acids and many other secondary metabolites synthesis. On the other hand, fatty acids synthesis is closely related to protein synthesis. Phosphoenolpyruvate (PEP) from the glycolysis is a common substrate for fatty acid and protein synthesis. PEP can be converted into pyruvate acid by pyruvate kinase. The pyruvate acid generates acetyl-CoA through the catalysis of pyruvate dehydrogenase. The resulting acetyl-CoA is the key precursor of fatty acid synthesis. Phosphoenolpyruvate carboxylase (PEPC) catalyzes the irreversible β-carboxylation of phosphoenolpyruvate (PEP) to form oxaloacetate into protein metabolism. PEPC is a cytoplasmic enzyme with versatile functions in plants. Although PEPC are reported to be involved in fatty acid accumulation and nitrogen assimilation, as well as salt and drought stresses, knowledge on the function of PEPC is still limited, particularly on the bacterial-type PEPC. In this paper, the regulation mechanism on fatty acid metabolism by ACCase and PEPC genes was investigated with forward and reverse genetic methods.
Firstly, the E.coli heteromeric ACCase was overexpressed in Brassica seeds to accelerate the formation of malonic acid by acetyl-CoA carboxylation, the key step in fatty acid synthesis. The heteromeric ACCase subunit encoding gene accD is a restriction factor. The accD genes from four close-related species of Brassica were cloned and their sequences were analyzed. Then two plant expressed vectors containing the heterogeneous-type E. coli ACCase accD gene (eaccD) and the genes encoding the four subunits of E.coli ACCase were tandemly constructed. The two vectors were further transformed into the "Chaoyou2" rapeseed to investigate the heterogeneity of eaccD and the other three nuclear genes assembled in fatty acid content. The data indicated that an efficient technique for directed genetic improvement of rapeseed oil content was established. In this research, the gene Atppc4which encodes the bacterial-type PEPC in Arabidopsis was knocked out with the artificial mRNA (amiRNA) technology to investigate the in vivo function. The effect of Atppc4on fatty acid metabolism and resistance to salt stress were elucidated. The study expanded our knowledge on the function of the bacterial-type PEPC in Arabidopsis. The main results are as follows:
1. The cDNA fragments of accD gene were amplified from young leaves of Brassica napus, Brassica oleracea, Brassica juncea and Brassica rapa by RT-PCR. The lengths of the amplified accD genes were1470,1470,1464and1464bp, which encode489,489,487and487amino acids, respectively. Sequence analysis showed that the cDNA sequences of the four accD genes were highly homologous. The amino acid sequences deduced from the four accD genes contained a similar zinc finger structure and five identical motifs in C-terminal regions. Among them, motif I (GSMGSVVG) and motif II (PLIIVCASGGARMQE) were present in β-carboxyltransferases of all plant species and E coli. Southern blotting analysis demonstrated that there was only a single copy of accD gene in the genomes of the four close-related species of Brassica.
2. To investigate the influence of eaccD gene on Brassica fatty acid content, an expression construct was constructed by fusing the Arabidopsis rubisco small subunit (rbcS) transit peptide and the Brassica seed specific promoter napin to eaccD gene. The expression construct was transformed into B. napus via Agrobacterium-mediated transformation and seven transgenic plants were obtained. The eaccD transcript was confirmed by RT-PCR analysis. The data showed that eaccD was specifically transcripted in seeds. This demonstrated that the expression construct can result in the specific expression of eaccD in immature seeds of Brassica. Seed oil and protein contents of the seeds were detected both in the transgenic lines and the control plants. The total oil content of the control seeds was48.01%and the average protein content was25.10%. Whereas the average fatty acid content in the transgenic lines seeds was52.08%, which was approximately8.45%higher than that of the wild plants. The seed oil content increase was followed by the decrease in protein content. The average protein content was6.69%lower than the wild plants. The weight of1000grains also increased in the transgenic plants. These results suggest that the expression of eaccD in Brassica could significantly increase the seeds oil content and seed weight.
3. To increase the content of fatty acid of Brassica by transgenic method, a plant expression vector was constructed by adding an Arabidopsis Rubisco SSU chloroplast signal peptide to the5'end of the target gene ACCase and hooked with the seed special expression promoter Napin in series. Transgenic Brasscia(Brasscia napus L.) plants were obtained via Agrobacterium-mediated hypocotyle.11positively plants were obtained. Four target genes were transformed into To Brassica genome. The seed oil contents of the control and the transgenic plants were48.01%and49.59%, respectively, with the transgenic plants being about3.30%higher than the control plants.
4. Plant expressed vector Atppc4-amiRNA was constructed and transformed into Arabidopsis with the in floral dip method via Agrobacterium-mediated transformation. The molecular detection was conducted in Atppc4-amiRNA transgenic lines. The maturedamiRNA was dramatically overexpressed in the transgenic plants. The transgenic plants with constitutively expressed Atppc4-amiRNA exhibited substantially decreased accumulation of Atppc4transcripts, whereas other three plant-type PEPC genes, Atppcl, Atppc2and Atppc3were significantly up-regulated in roots. PEPC activity improved about5.1times in roots of Atppc4-amiRNA transgenic lines. This result indicates that transcription of bacterial-type and plant-type PEPC genes in plants interact with each other in plants. The bacterial-type PEPC genes, Atppc4, may play an important role in modulating the transcription of plant-type PEPC genes. There was no significant difference in fatty acid content between the wild-type plants and transgenic lines. Moreover, fatty acid compositions also had no significant changes in the seeds of wild-type plants and transgenic lines. This research suggested that the transcription of Atppc4might be independent of the lipid content in Arabidopsis.
5. Down-regulation of Atppc4improved salt tolerance in Arabidopsis. Studies on7-d-old seedling roots showed that transgenic plants exhibited approximately29.0%less growth in the root system than that of wild-type plants. However when these7-d-old seedlings were moved to a1/2MS medium containing150mM NaCl and incubated under the same conditions for another week, the transgenic plants showed approximately16.3%less root growth than of the wild-type plants. These results showed that salt treatment could partially relieve the inhibition of root growth caused by Atppc4suppression.
由于accD基因编码的β-CT亚基是ACCase基因表达的限制因子,本研究通过在油菜籽粒中超量表达大肠杆菌异质型ACCase基因,加速乙酰CoA羧化形成丙二酸单酰CoA这一关键进程,提高油菜籽脂肪酸的起始合成能力。本研究克隆了甘蓝型油菜及近缘种中异质型ACCase的β-CT亚基accD编码基因,采用生物信息学的方法分析了氨基酸序列,构建了受种子特异表达启动子驱动的异质型大肠杆菌ACCase β-CT亚基accD基因的植物表达载体以及大肠杆菌ACCase四个亚基基因分别受种子特异性表达启动子驱动的植物串联表达载体,将这两个载体分别用于转化甘蓝型油菜“超油2号”。通过转基因油菜的半定量PCR、蛋白质、脂肪酸含量变化,研究了异质型大肠杆菌基因accD与其它三个细胞核基因的互作对油菜脂肪酸合成的影响,为定向遗传改良油菜种子含油量建立了有效的技术体系。本研究还利用人工小RNA (artifical mRNA, amiRNA)技术敲除拟南芥细菌型Atppc4基因进行功能分析,研究该基因在脂肪酸代谢以及耐盐胁迫等方面的作用,以探讨拟南芥中细菌型Atppc4基因敲除对植物型PEPC的影响,为今后改良油菜品质提供理论依据。主要结果如下：
1.以甘蓝型油菜(Brassica napus)、甘蓝(Brassica oleracea)、芥菜型油菜(Brassica juncea)和白菜型油菜(Brassica rapa)幼嫩叶片的cDNA为模板,对异质型乙酰辅酶A羧化酶p-CT亚基(羧基转移酶)的accD编码基因进行扩增,得到长度分别为1470、1470、1464和1464bp的编码序列,分别可编码489、489、487和487个氨基酸的蛋白质。白菜型油菜、芥菜型油菜和甘蓝的accD序列登录号分别为EU410075、EU410076和EU410077。结果表明来自4个油菜近缘种的accD基因高度同源,编码的蛋白质都具有相似的锌指结构和C-末端5个相同的基元,其中基元I (GSMGSVVG)和基元Ⅱ(PLIIVCASGGARMQE)在所有植物及大肠杆菌(Escherichia coli)的β-CT亚基中普遍存在。Southern杂交显示该基因在甘蓝型油菜及其近缘种的基因组中呈单拷贝。
2.本研究E. coli ACCase的β-CT亚基accD (eaccD)基因与拟南芥Rubisco小亚基(rbcS)转运肽基因融合,构建以油菜Napin启动子驱动的种子特异表达载体。利用根癌农杆菌(Agrobacterium tumefaciens)介导法,将该表达载体转入“超油2号”,获得了7个阳性转基因株系。RT-PCR检测其中3株的结果证明eaccD基因可在转基因株系的种子中转录,但在叶片中不转录,说明该表达载体转入油菜中能引起植株种子eaccD基因的特异表达。通过转基因与对照植株种子的油脂和蛋白质含量的比较分析,发现对照种子的平均含油量为48.01%、蛋白质平均含量为25.10%；而7个转基因株系油菜种子的平均含油量为52.08%,比对照含油量相对增加8.45%；但转基因株系的蛋白质含量则有不同程度的降低,平均比对照植株下降约6.69%。转基因油菜千粒重比对照植株有所提高。因此大肠杆菌accD基因在油菜中的表达能够增加种子重量和含油量。
3.采用同尾酶反复酶切的方法,构建了拟南芥Rubisco小亚基转运肽与ACCase各亚基基因的融合串联表达载体,用A. tumefaciens介导的下胚轴转化对照品种“超油2号”。经过筛选获得了11株转化再生植株。经PCR. Southern和GUS染色检测证明4个目的基因已整合到T0再生植株油菜基因组中。T2转基因株系平均含油量为49.59%,转基因株系的含油量比对照(48.01%)提高了3.30%。
4.构建了人工小RNA表达载体Atppc4-amiRNA,利用农杆菌介导花絮浸染法(in floral dip)转化拟南芥,对转基因株系进行了分子检测。在拟南芥转基因株系内检测到成熟小RNA的表达,拟南芥转基因株系根中Atppc4的表达水平明显下降,然而其它3个植物型PEPC基因Atppc1、Atppc2和Atppc3在根中的表达明显得到上调。转基因株系根中的PEPC酶活性提高了5.1倍,表明植物中细菌型和植物型PEPC基因具有互作,细菌型PEPC基因Atppc4可以调控植物型PEPC基因的表达。分析拟南芥株系和对照植株的脂肪酸含量和组分的结果表明,转基因株系和对照的脂肪酸含量和脂肪酸组分未发生明显变化。
5.atppc4的下调表达提高拟南芥的耐盐能力。在非盐处理培养基上,Atppc4-amiRNA转基因株系比野生型的根系生长更慢。发现7日龄转基因株系的长度比野生型株系短29.0%,将这些植株移到含有150mM NaCl的1/2MS培养基上培养7天,转基因株系长度比野生型株系要短16.3%。说明转基因株系的根系由于Atppc4基因表达被抑制；盐处理后转基因株系根的生长能得到部分缓解,从而缩短了与野生型根长的距离。该结果说明特异性敲除细菌型Atppc4基因提高了拟南芥的耐盐性。
Vegetable oil is an essential product in our daily life and constitutes a major source of energy its consumers. The seed is the main storage organ of the oil plants. The improvement in the quality and quantity of seed fatty acid have currently become major research topic. Fatty acid biosynthesis in plants takes place within plastids. Acetyl-coenzyme A carboxylase (ACCase) catalyzes the first committed step of fatty acid synthesis, the carboxylastion of acetyl-CoA to malonyl-CoA which supplies the substrates for fatty acids and many other secondary metabolites synthesis. On the other hand, fatty acids synthesis is closely related to protein synthesis. Phosphoenolpyruvate (PEP) from the glycolysis is a common substrate for fatty acid and protein synthesis. PEP can be converted into pyruvate acid by pyruvate kinase. The pyruvate acid generates acetyl-CoA through the catalysis of pyruvate dehydrogenase. The resulting acetyl-CoA is the key precursor of fatty acid synthesis. Phosphoenolpyruvate carboxylase (PEPC) catalyzes the irreversible β-carboxylation of phosphoenolpyruvate (PEP) to form oxaloacetate into protein metabolism. PEPC is a cytoplasmic enzyme with versatile functions in plants. Although PEPC are reported to be involved in fatty acid accumulation and nitrogen assimilation, as well as salt and drought stresses, knowledge on the function of PEPC is still limited, particularly on the bacterial-type PEPC. In this paper, the regulation mechanism on fatty acid metabolism by ACCase and PEPC genes was investigated with forward and reverse genetic methods.
Firstly, the E.coli heteromeric ACCase was overexpressed in Brassica seeds to accelerate the formation of malonic acid by acetyl-CoA carboxylation, the key step in fatty acid synthesis. The heteromeric ACCase subunit encoding gene accD is a restriction factor. The accD genes from four close-related species of Brassica were cloned and their sequences were analyzed. Then two plant expressed vectors containing the heterogeneous-type E. coli ACCase accD gene (eaccD) and the genes encoding the four subunits of E.coli ACCase were tandemly constructed. The two vectors were further transformed into the "Chaoyou2" rapeseed to investigate the heterogeneity of eaccD and the other three nuclear genes assembled in fatty acid content. The data indicated that an efficient technique for directed genetic improvement of rapeseed oil content was established. In this research, the gene Atppc4which encodes the bacterial-type PEPC in Arabidopsis was knocked out with the artificial mRNA (amiRNA) technology to investigate the in vivo function. The effect of Atppc4on fatty acid metabolism and resistance to salt stress were elucidated. The study expanded our knowledge on the function of the bacterial-type PEPC in Arabidopsis. The main results are as follows:
1. The cDNA fragments of accD gene were amplified from young leaves of Brassica napus, Brassica oleracea, Brassica juncea and Brassica rapa by RT-PCR. The lengths of the amplified accD genes were1470,1470,1464and1464bp, which encode489,489,487and487amino acids, respectively. Sequence analysis showed that the cDNA sequences of the four accD genes were highly homologous. The amino acid sequences deduced from the four accD genes contained a similar zinc finger structure and five identical motifs in C-terminal regions. Among them, motif I (GSMGSVVG) and motif II (PLIIVCASGGARMQE) were present in β-carboxyltransferases of all plant species and E coli. Southern blotting analysis demonstrated that there was only a single copy of accD gene in the genomes of the four close-related species of Brassica.
2. To investigate the influence of eaccD gene on Brassica fatty acid content, an expression construct was constructed by fusing the Arabidopsis rubisco small subunit (rbcS) transit peptide and the Brassica seed specific promoter napin to eaccD gene. The expression construct was transformed into B. napus via Agrobacterium-mediated transformation and seven transgenic plants were obtained. The eaccD transcript was confirmed by RT-PCR analysis. The data showed that eaccD was specifically transcripted in seeds. This demonstrated that the expression construct can result in the specific expression of eaccD in immature seeds of Brassica. Seed oil and protein contents of the seeds were detected both in the transgenic lines and the control plants. The total oil content of the control seeds was48.01%and the average protein content was25.10%. Whereas the average fatty acid content in the transgenic lines seeds was52.08%, which was approximately8.45%higher than that of the wild plants. The seed oil content increase was followed by the decrease in protein content. The average protein content was6.69%lower than the wild plants. The weight of1000grains also increased in the transgenic plants. These results suggest that the expression of eaccD in Brassica could significantly increase the seeds oil content and seed weight.
3. To increase the content of fatty acid of Brassica by transgenic method, a plant expression vector was constructed by adding an Arabidopsis Rubisco SSU chloroplast signal peptide to the5'end of the target gene ACCase and hooked with the seed special expression promoter Napin in series. Transgenic Brasscia(Brasscia napus L.) plants were obtained via Agrobacterium-mediated hypocotyle.11positively plants were obtained. Four target genes were transformed into To Brassica genome. The seed oil contents of the control and the transgenic plants were48.01%and49.59%, respectively, with the transgenic plants being about3.30%higher than the control plants.
4. Plant expressed vector Atppc4-amiRNA was constructed and transformed into Arabidopsis with the in floral dip method via Agrobacterium-mediated transformation. The molecular detection was conducted in Atppc4-amiRNA transgenic lines. The maturedamiRNA was dramatically overexpressed in the transgenic plants. The transgenic plants with constitutively expressed Atppc4-amiRNA exhibited substantially decreased accumulation of Atppc4transcripts, whereas other three plant-type PEPC genes, Atppcl, Atppc2and Atppc3were significantly up-regulated in roots. PEPC activity improved about5.1times in roots of Atppc4-amiRNA transgenic lines. This result indicates that transcription of bacterial-type and plant-type PEPC genes in plants interact with each other in plants. The bacterial-type PEPC genes, Atppc4, may play an important role in modulating the transcription of plant-type PEPC genes. There was no significant difference in fatty acid content between the wild-type plants and transgenic lines. Moreover, fatty acid compositions also had no significant changes in the seeds of wild-type plants and transgenic lines. This research suggested that the transcription of Atppc4might be independent of the lipid content in Arabidopsis.
5. Down-regulation of Atppc4improved salt tolerance in Arabidopsis. Studies on7-d-old seedling roots showed that transgenic plants exhibited approximately29.0%less growth in the root system than that of wild-type plants. However when these7-d-old seedlings were moved to a1/2MS medium containing150mM NaCl and incubated under the same conditions for another week, the transgenic plants showed approximately16.3%less root growth than of the wild-type plants. These results showed that salt treatment could partially relieve the inhibition of root growth caused by Atppc4suppression.
引文
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