小麦品种陕253α-醇溶蛋白基因的克隆、原核表达及功能鉴定
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摘要
小麦贮藏蛋白的主要类型包括麦谷蛋白(glutenin)与麦醇溶蛋白(gliadin),约占小麦籽粒蛋白质的80%,并共同构成了面筋的主要成分,其数量与比例关系决定着面筋的质量。醇溶蛋白是组成面筋的主要成分之一,约占蛋白质总量的40%～50%,其所具有的品质效应一直是争论的焦点与研究的难点,其原因不仅归结于该类蛋白亚基在分离上的难度,也与该基因家族的多样性及蛋白质空间重构有关。虽然不同学者得出了正负相关的矛盾结论,但大多数的研究结果认为醇溶蛋白降低面团的稳定性。
本研究根据已知的α-醇溶蛋白基因的两端保守序列来设计特异引物,以基因组DNA为模板通过PCR扩增目的基因,克隆测序后比较分析克隆基因的分子结构及其同源关系;然后利用设计的不扩增信号肽的表达引物,扩增的目的片段,回收的目的片段后与表达载体连接,利用E. coli体外表达α-醇溶蛋白,经柱层析及低温冷冻干燥获得目的蛋白亚基,通过氧化-还原反应将蛋白亚基整合到基础弱筋面粉中,并利用4g粉质仪研究α-醇溶蛋白亚基对面团流变学特性的影响,结果如下:
1.从强筋小麦品种陕253中克隆得到7条序列,分别命名为α-Agli-1~7,GenBank登录号为GQ891681~ GQ891687,其中α-Agli-3和α-Agli-5为真基因,其余5个基因由于编码区存在提前终止密码子,因此推测为假基因。序列分析表明,α-Agli-3和α-Agli-5的编码区的相似性为97.90%,推导的氨基酸序列的相似性为97.33%,其余5个序列的相似性达到85%以上。从提前终止密码子的分布区域来看,主要分布在重复区。提前终止密码子出现的位点和类型主要有CAA/CAG→TAA和CAA/CAG/GAG→TAG,其使用频率依次为: TAA (66%)> TAG (34%)。
2.进一步的分析表明,克隆得到的7条序列都具有α-醇溶蛋白基因的典型结构,包括保守的信号肽、N-端重复区、多聚谷氨酰胺I区、N-端非重复区(特征区I)、多聚谷氨酰胺II区和C-端非重复区(特征区II)。重复区具有基本的重复单元PQPQPFP和PQQPY,多聚谷氨酰胺区的主要差异在于长度不同和非谷氨酰胺残基的出现。α-Agli-4,α-Agli-6和α-Agli-7均有6个半胱氨酸残基。α-Agli-1和α-Agli-2分别含有5个和9个半胱氨酸残基,可能是由于移码突变导致的;α-Agli-3和α-Agli-5含有7个半胱氨酸残基,是由于在Uniq domain II区编码第6位点氨基酸的碱基发生C→G转换,导致丝氨酸(Ser)→半胱氨酸(Cys),这一额外的半胱氨酸残基将有可能与相邻储藏蛋白亚基形成一个分子间二硫键,从而参与面筋的聚合。
3.分析α-Agli-3和α-Agli-5的序列,将α-Agli-5进行了原核表达,根据其编码区序列设计特异的表达引物,扩增目的基因,构建了原核表达载体pET32a-α-Agli-5,将含有重组质粒pET32a-α-Agli-5的阳性克隆经过夜培养转化到表达宿主菌BL21 (DE3)感受态细胞中,经过1mmoL的IPTG诱导,28℃培养6 h ,提取细菌总蛋白上清液,经12 % SDS-PAGE电泳和western blot检测后,发现均产生了一个55 kD大小的蛋白条带,即为表达的目的蛋白,结果与预测的分子量相符。
4.大量制备表达的目的蛋白,经?KTA purifier 100蛋白层析系统(Amersham Biosciences, NJ)纯化获得目的蛋白,通过氧化-还原反应将亚基整合到基础面粉中,利用4 g粉质仪研究α-醇溶蛋白亚基对面团流变学特性的影响,结果表明α-醇溶蛋白α-Agli-5的添加增强了面筋的弹性但是降低了面筋的强度。
Wheat storage protein mainly included glutenin and gliadin , about 80% of wheat seed protein, and constituted the main ingredient of gluten. Quality of gluten was determined by quantity and scale of glutenin and gliadin. Gliadin was one of the main ingredient of gluten, about 40%~50% of total amount of protein, the quality of effects had been issue and difficulty of research.The reasons was not only to the difficulty of separation, but also with rich diversity of the genes family and reconsitution of the protein space. Though different scholars had severally conflicting conclusion, most of the research results showed that gliadin reduced stability of the dough.
Specific primers were designed according to conserved ends of the coding region of knownα-gliadin genes, the genomic DNA was a template for PCR, the molecular structure of gene and its kinship were analyzed;Target fragment was PCR with primers of expression of which didn’t PCR signal peptide, then recovered target fragment and was connected with express vector. The subunit ofα-gliadin was expressed by E. coli effectively and was got by purifying, then integrated into the flour by the oxidation reduction reaction, rheological properties were tested by micro 4g farinograph in order to research the effect ofα-gliadin to dough, the results were as follows:
1. Seven DNA sequences were cloned from a strong gluten wheat variety Shaan 253, namedα-Agli-1~7(GenBank NO. was GQ891681~ GQ891687)respectively, except forα-Agli-3 andα-Agli-5, the remaining five genes were pseudogene of which genetic coding region had stop codons.The sequence analysis showed that the identity ofα-Agli-3 andα-Agli-5 was 97.90%, the remaining five sequences was above 85%. The analysis of the distribution region of stop codons showed that they were mainly in repeated sequence region. The sites and type were CAA/CAG→TAA and CAA/CAG/GAG→TAG,the frequency of each stop codons was TAA (66%)> TAG (34%)of which CAA mutate to TAA was the highest.
2. Further analysis showed that the seven cloned sequences had typical structure ofα-gliadin genes of which included: a conservative signal peptide, N-terminal repeats, poly-glutamine I region, N-terminal non-repeat region (unique domain I), poly-glutamine II region and C-terminal non-repeat region(unique domain II). Repeated region had basic repeat unit PQPQPFP and PQQPY. The differences of poly-glutamine region were length and appearance of non-glutamine.α-Agli-4,α-Agli-6 andα-Agli-7 had six cysteine residues,α-Agli-1 andα-Agli-2 had five and nine cysteine residues respectively, may be caused by frameshift mutations;α-Agli-3 andα-Agli-5 had seven cysteine residues of which in the unique domain II cysteine was instead of serine due to C→G in the codon of the sixth amino acid, which maybe form intermolecular disulfide bond.
3. Analyzedα-Agli-3 andα-Agli-5 sequence,α-Agli-5 was expressed in E.coli. Specific expressed primers were designed according to its coding sequence to amplify the target fragment.Prokaryotic expression vector pET32a-α-Agli-5 was constructed, the positive clones of recombinant plasmid pET32a-α-Agli-5 was transformed into expression host strain BL21(DE3)competent cells, after inducting by IPTG of 1mmoL, 28℃6 h culture , supernatant of the total bacterial proteins was extracted then 12% SDS-PAGE electrophoresis and western blot were used to test the product, they both appeared a 55 kD protein band of which was the purpose expression protein, and the results was in accord with the predicted molecular weight.
4. A large number of expressed protein was prepared and purified by ?KTA purifier 100 protein chromatography system.Purified protein was integrated into the flour by the oxidation reduction reaction, rheological properties were tested by micro 4 g farinograph in order to research the effect ofα-gliadin to dough. The results show that flexibility of gluten increased but strength of gluten decreaded whenα-gliadinα-Agli-5 was added.
本研究根据已知的α-醇溶蛋白基因的两端保守序列来设计特异引物,以基因组DNA为模板通过PCR扩增目的基因,克隆测序后比较分析克隆基因的分子结构及其同源关系;然后利用设计的不扩增信号肽的表达引物,扩增的目的片段,回收的目的片段后与表达载体连接,利用E. coli体外表达α-醇溶蛋白,经柱层析及低温冷冻干燥获得目的蛋白亚基,通过氧化-还原反应将蛋白亚基整合到基础弱筋面粉中,并利用4g粉质仪研究α-醇溶蛋白亚基对面团流变学特性的影响,结果如下:
1.从强筋小麦品种陕253中克隆得到7条序列,分别命名为α-Agli-1~7,GenBank登录号为GQ891681~ GQ891687,其中α-Agli-3和α-Agli-5为真基因,其余5个基因由于编码区存在提前终止密码子,因此推测为假基因。序列分析表明,α-Agli-3和α-Agli-5的编码区的相似性为97.90%,推导的氨基酸序列的相似性为97.33%,其余5个序列的相似性达到85%以上。从提前终止密码子的分布区域来看,主要分布在重复区。提前终止密码子出现的位点和类型主要有CAA/CAG→TAA和CAA/CAG/GAG→TAG,其使用频率依次为: TAA (66%)> TAG (34%)。
2.进一步的分析表明,克隆得到的7条序列都具有α-醇溶蛋白基因的典型结构,包括保守的信号肽、N-端重复区、多聚谷氨酰胺I区、N-端非重复区(特征区I)、多聚谷氨酰胺II区和C-端非重复区(特征区II)。重复区具有基本的重复单元PQPQPFP和PQQPY,多聚谷氨酰胺区的主要差异在于长度不同和非谷氨酰胺残基的出现。α-Agli-4,α-Agli-6和α-Agli-7均有6个半胱氨酸残基。α-Agli-1和α-Agli-2分别含有5个和9个半胱氨酸残基,可能是由于移码突变导致的;α-Agli-3和α-Agli-5含有7个半胱氨酸残基,是由于在Uniq domain II区编码第6位点氨基酸的碱基发生C→G转换,导致丝氨酸(Ser)→半胱氨酸(Cys),这一额外的半胱氨酸残基将有可能与相邻储藏蛋白亚基形成一个分子间二硫键,从而参与面筋的聚合。
3.分析α-Agli-3和α-Agli-5的序列,将α-Agli-5进行了原核表达,根据其编码区序列设计特异的表达引物,扩增目的基因,构建了原核表达载体pET32a-α-Agli-5,将含有重组质粒pET32a-α-Agli-5的阳性克隆经过夜培养转化到表达宿主菌BL21 (DE3)感受态细胞中,经过1mmoL的IPTG诱导,28℃培养6 h ,提取细菌总蛋白上清液,经12 % SDS-PAGE电泳和western blot检测后,发现均产生了一个55 kD大小的蛋白条带,即为表达的目的蛋白,结果与预测的分子量相符。
4.大量制备表达的目的蛋白,经?KTA purifier 100蛋白层析系统(Amersham Biosciences, NJ)纯化获得目的蛋白,通过氧化-还原反应将亚基整合到基础面粉中,利用4 g粉质仪研究α-醇溶蛋白亚基对面团流变学特性的影响,结果表明α-醇溶蛋白α-Agli-5的添加增强了面筋的弹性但是降低了面筋的强度。
Wheat storage protein mainly included glutenin and gliadin , about 80% of wheat seed protein, and constituted the main ingredient of gluten. Quality of gluten was determined by quantity and scale of glutenin and gliadin. Gliadin was one of the main ingredient of gluten, about 40%~50% of total amount of protein, the quality of effects had been issue and difficulty of research.The reasons was not only to the difficulty of separation, but also with rich diversity of the genes family and reconsitution of the protein space. Though different scholars had severally conflicting conclusion, most of the research results showed that gliadin reduced stability of the dough.
Specific primers were designed according to conserved ends of the coding region of knownα-gliadin genes, the genomic DNA was a template for PCR, the molecular structure of gene and its kinship were analyzed;Target fragment was PCR with primers of expression of which didn’t PCR signal peptide, then recovered target fragment and was connected with express vector. The subunit ofα-gliadin was expressed by E. coli effectively and was got by purifying, then integrated into the flour by the oxidation reduction reaction, rheological properties were tested by micro 4g farinograph in order to research the effect ofα-gliadin to dough, the results were as follows:
1. Seven DNA sequences were cloned from a strong gluten wheat variety Shaan 253, namedα-Agli-1~7(GenBank NO. was GQ891681~ GQ891687)respectively, except forα-Agli-3 andα-Agli-5, the remaining five genes were pseudogene of which genetic coding region had stop codons.The sequence analysis showed that the identity ofα-Agli-3 andα-Agli-5 was 97.90%, the remaining five sequences was above 85%. The analysis of the distribution region of stop codons showed that they were mainly in repeated sequence region. The sites and type were CAA/CAG→TAA and CAA/CAG/GAG→TAG,the frequency of each stop codons was TAA (66%)> TAG (34%)of which CAA mutate to TAA was the highest.
2. Further analysis showed that the seven cloned sequences had typical structure ofα-gliadin genes of which included: a conservative signal peptide, N-terminal repeats, poly-glutamine I region, N-terminal non-repeat region (unique domain I), poly-glutamine II region and C-terminal non-repeat region(unique domain II). Repeated region had basic repeat unit PQPQPFP and PQQPY. The differences of poly-glutamine region were length and appearance of non-glutamine.α-Agli-4,α-Agli-6 andα-Agli-7 had six cysteine residues,α-Agli-1 andα-Agli-2 had five and nine cysteine residues respectively, may be caused by frameshift mutations;α-Agli-3 andα-Agli-5 had seven cysteine residues of which in the unique domain II cysteine was instead of serine due to C→G in the codon of the sixth amino acid, which maybe form intermolecular disulfide bond.
3. Analyzedα-Agli-3 andα-Agli-5 sequence,α-Agli-5 was expressed in E.coli. Specific expressed primers were designed according to its coding sequence to amplify the target fragment.Prokaryotic expression vector pET32a-α-Agli-5 was constructed, the positive clones of recombinant plasmid pET32a-α-Agli-5 was transformed into expression host strain BL21(DE3)competent cells, after inducting by IPTG of 1mmoL, 28℃6 h culture , supernatant of the total bacterial proteins was extracted then 12% SDS-PAGE electrophoresis and western blot were used to test the product, they both appeared a 55 kD protein band of which was the purpose expression protein, and the results was in accord with the predicted molecular weight.
4. A large number of expressed protein was prepared and purified by ?KTA purifier 100 protein chromatography system.Purified protein was integrated into the flour by the oxidation reduction reaction, rheological properties were tested by micro 4 g farinograph in order to research the effect ofα-gliadin to dough. The results show that flexibility of gluten increased but strength of gluten decreaded whenα-gliadinα-Agli-5 was added.
引文
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