小麦低分子量麦谷蛋白亚基分离及密码子使用特征分析
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摘要
根据不同分子量蛋白质在十二烷基硫酸钠-聚丙烯酰胺凝胶电泳(SDS-PAGE)迁移率不同,小麦(Triticum aestivum L.)麦谷蛋白亚基分为高分子量麦谷蛋白亚基(HMW-GS)和低分子量麦谷蛋白亚基(LMW-GS)。小麦HMW-GS和LMW-GS互作形成空间网格状结构决定小麦面包烘烤品质。然而小麦LMW-GS研究相对滞后。因此,获得我国独立的自主知识产权的LMW-GS功能基因,并用于改良我国小麦面包烘烤品质已迫在眉睫。
该论文首先以国内优质强筋小麦中优9507为实验材料,通过优化SDS-PAGE技术蛋白质分离条件,分离小麦LMW-GS区段的特异条带,质谱鉴定获得肽段;进一步分析小麦麦谷蛋白亚基基因密码子使用特性,采用RT-PCR和RACE技术克隆目的基因核酸片段,建立小麦LMW-GS基因克隆技术体系。在此基础之上,以新疆本地主栽强筋小麦新春8为实验材料,锁定小麦面包烘烤品质主效LMW-GS,质谱鉴定获得肽段,为后续该小麦面包烘烤品质主效LMW-GS基因克隆、小麦HMW-GS和LMW-GS互作研究提供科学的数据参考。该论文主要研究结果如下:
含0.3mol/LNaI和1.4%4-VP的LMW-GS提取液能有效提取小麦LMW-GS成分。在此基础上,当分离胶浓度为T=14%/C=3%、分离时间为40h、电极缓冲液为Tris-硼酸或Tris-甘氨酸时,能更加有效、准确、且重复性高地分离数目庞大的小麦LMW-GS。该技术体系为小麦LMW-GS深入研究的第一个限速步骤提供了良好的解决方案。
从国内优质强筋小麦中优9507中锁定1个小麦面包烘烤品质可能相关的LMW-GS,命名为ZY9507-L1,其表观分子量为34kD。质谱鉴定获得5个肽段,分别命名为ZL1-1、ZL1-2、ZL1-3、ZL1-4、ZL1-5。其中,ZL1-1(肽段序列:GIIQPQQPAQLEVIR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ACJ76961.1)的覆盖率(Query coverage)为66%,ZL1-2(肽段序列:SVVHSIIMQQEQR)与小麦已知LMW-GS氨基酸序列(GenBank登录号:ABB04903.1)的Query coverage为92%。其余3个质谱鉴定获得的小肽段均与NCBI数据库中小麦已知LMW-GS氨基酸序列不存在相似性。
为了解小麦麦谷蛋白亚基基因密码子使用特性,该文运用CodonW1.4.2和SPSS16.0软件对13个来源于小麦麦谷蛋白亚基基因的密码子用法特性进行了分析。结果显示,除1Dy10基因有效密码子数(ENC值)小于40外,其余亚基基因ENC值均介于40-50之间;密码子第三位GC含量(GC3)均在0.38-0.45之间;1Bx14基因偏向于使用A或T结尾的密码子,其余12个亚基基因偏向于使用G或C结尾的密码子;1Dy10与其余12个亚基基因间距离系数较大,然而各组染色体麦谷蛋白x-型或y-型亚基基因间距离系数较小。结果表明,小麦麦谷蛋白亚基基因具有较强的密码子使用偏爱性,各基因密码子用法上具有相似性,每一种氨基酸均存在1-2个偏爱密码子。在密码子用法差异上表现为,除1By9和1Dy10之外,无论是A染色体组、B染色体组,还是D染色体组,各组染色体上麦谷蛋白x-型或y-型亚基基因间密码子使用频率差异最小。这一分析结果对小麦HMW-GS功能基因资源的高效利用和LMW-GS新基因的克隆具有一定的指导意义。
根据小麦面包烘烤品质相关LMW-GS ZY9507-L1质谱鉴定结果和小麦麦谷蛋白亚基基因密码子使用特性,该文以国内优质强筋小麦中优9507为实验材料,利用ZL1-1肽段设计3’端上游引物,采用RT-PCR和RACE技术获得目的基因ZL1-13’端序列。结果显示,分离获得2个不同的LMW-GS基因序列,分别命名为ZL1-1P和ZL1-1I,其中,ZL1-1P基因长度为620bp,可编码包含186个氨基酸残基的成熟蛋白质。该基因与NCBI数据库中许多小麦已知LMW-GS基因均具有同源性,其与小麦已知LMW-GS基因(登录号:FJ441109.1)的覆盖率为19%(见图5-11);ZL1-1I基因长度为693bp,可编码包含209个氨基酸残基的成熟蛋白质。该基因与NCBI数据库中许多小麦已知LMW-GS基因均具有同源性,该基因序列与已知LMW-GS基因(登录号:FJ441109.1)的覆盖率为17%。初步证实上述2个基因可能均属于小麦LMW-GS基因的一部分,而且也证实了该小麦LMW-GS基因克隆技术体系是完全可行的。
在上述已建立的基因克隆技术体系基础之上,根据面包烘烤品质的决定机制特征,从新疆本地主栽强筋小麦新春8中锁定1个小麦面包烘烤品质主效LMW-GS,命名为XC8-L1,其表观分子量为46kD。质谱鉴定获得215个肽段,分别命名为XL1-1、XL1-2、XL1-3、XL1-4、XL1-5、……、XL1-215。其中,XL1-1(肽段序列:AIIYSIVLQEQQQVR)、XL1-2(肽段序列:VNVPLYR)、XL1-3(肽段序列:TTTSVPFGVGTGVGAY)和XL1-4(肽段序列:LEVMTSIALR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ACZ59817.1 )的Query coverage为100% , XL1-5 (肽段序列:TTTNVPFGVGTGVGSY)和XL1-6(肽段序列:VNVPLYR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ACA63873.1)的Query coverage为100%,XL1-7(肽段序列:QIAQLEVMTSIALR)和XL1-8(肽段序列:QIPEQSR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ADX68840.1)的Query coverage为100%,XL1-9(肽段序列:GTFLQPHQIAR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ADX68849.1)的Query coverage为100%。其余206个质谱鉴定后获得的小肽段均与NCBI数据库中小麦已知LMW-GS氨基酸序列不存在相似性。
该论文不仅为小麦面包烘烤品质相关功能基因克隆提供技术体系,而且为获得我国独立的自主知识产权的小麦面包烘烤品质主效LMW-GS基因克隆提供有效的氨基酸序列支持,还为深入探讨小麦HMW-GS和LMW-GS互作研究、加快我国面包小麦改良步伐并实现优良面包粉的自主生产奠定一定的理论基础。
Glutenin polymers are formed by high (HMW-GS) and low molecular weight glutenins subunits(LMW-GS) on the basis of their mobilities in the sodium dodecyl sulfate polyacrylamide gel electrophoresis(SDS-PAGE) in wheat(Triticum aestivum L.). It has been demonstrated that the variation in HMW-GS and LMW-GS composition has a strong impact on wheat bread-making quality. However, the deeply study on LMW-GS is far lower than HMW-GS. So, it’s very importace to develop our own intellectual property about LMW-GS.
In this paper, the SDS-PAGE separation condition was optimazed, then interrelated subunit may be for wheat bread-making quality was isolated in Zhongyou 9507, and identified them by LC-MS. In addition, we analyzed the codon usage of glutenin subunits genes in wheat, to explore HMW-GS genetic resources efficiently and clone new LMW-GS genes. Soon after, it isolated nucleotide sequence of interrelated subunit by RT-PCR and RACE. On this basis, we choose Xinchun 8 as the experimental material, then isolated the major subunit on wheat bread-making quality by optimized SDS-PAGE technonlgy, and identified them by LC-MS. The main results are as follows:
The LMW-GS extraction buffer containing 0.3mol/L NaI and 1.4% 4-VP can obtain LMW-GS availably, it's the best condition under T=14%C=3%、tris-boric acid buffer or tris-glycine buffer to separate glutenin subunits effectively. It provide a theoretical basic step for studying LMW-GS in breeding quality wheat cultivar.
It locked 1 interrelated subunit in Zhongyou9507, named ZY9507-L1, its apparent molecular weight is 34kDa. It obtained 5 peptides by LC-MS, named ZL1-1、ZL1-2、ZL1-3、ZL1-4、ZL1-5 orderly. At the same time, Query coverage is 66% between ZL1-1(Peptide is GIIQPQQPAQLEVIR)and amino acid sequence of known wheat LMW-GS(ACJ76961.1) in NCBI. Query coverage is 92% between ZL1-2 (Peptide is SVVHSIIMQQEQR)and amino acid sequence of known wheat LMW-GS(ABB04903.1) in NCBI. There is little similarity between other two peptides and known LMW-GS in NCBI.
The value of Effective Numberof Codons (ENC) of the other 12 glutenin subunits genes was between 40-50, except the 1Dy10 gene was less than 40.The GC3 contents were all between 0.38-0.45. The 1Bx14 Gene preferred to use A or T at the end of codon and the remaining 12 subunit genes preferred to use G or C . The distance coefficient of 1Dy10 was larger with rest of 12 genes, but it was small between x-type or y-type subunit genes of each chromosomes. The results showed that the glutenin subunit genes of wheat possessed a strong codon usage bias clearly. The codon usage of glutenin subunits genes was similar, and there were optimal(one or two) for each amino acid codon. Except for 1By9 and 1Dy10, the other 12 genes had the smallest differences in frequency of utilizing codon. The results would help to explore HMW-GS genetic resources efficiently and clone new LMW-GS genes.
According to the LC-MS and characterization on codon usage of glutenin subunits genes in Wheat, we designed the specific primers on the basis of ZL1-1 peptide, it extracted the total RNA in Zhongyou 9507 by Trizol, then it gained first strand cDNA by RT-PCR.And It gained 3’nucleotide sequence of ZL1-1 finally. It showed that, it gained two new genes, named ZL1-1P and ZL1-1I respectively. The coding regions of ZL1-1P is 620bp, encoding the proteins with 186 amino acid residues. The coding regions of ZL1-1I is 693bp, encoding the proteins with 209 amino acid residues. There have homology among ZL1-1P、ZL1-1I and the known wheat LMW-GS in NCBI. It indicated that the genes is the LMW-GS preliminary.
It locked 1 major subunit which is about wheat bread-making quality in Xinchun 8, named XC8-L1, molecular weight is 46kD. It obtained 215 peptides by LC-MS, named XL1-1、XL1-2、XL1-3、XL1-4、XL1-5、……、XL1-215 orderly. At the same time, Query coverage is 100% among XL1-1(Peptide is AIIYSIVLQEQQQVR)、XL1-2(Peptide is VNVPLYR)、XL1-3(Peptide is TTTSVPFGVGTGVGAY)、(Peptide is LEVMTSIALR)and amino acid sequence of wheat LMW-GS(ACZ59817.1) in NCBI, Query coverage is 100% among XL1-5(Peptide is TTTNVPFGVGTGVGSY) and XL1-6(Peptide is VNVPLYR)and amino acid sequence of wheat LMW-GS(ACA63873.1) in NCBI, Query coverage is 100% among XL1-7(Peptide is QIAQLEVMTSIALR) and XL1-8(Peptide is QIPEQSR)and amino acid sequence of wheat LMW-GS(ADX68840.1) in NCBI, Query coverage is 100% between XL1-9(Peptide is GTFLQPHQIAR)and amino acid sequence of wheat LMW-GS(ADX68849.1) in NCBI.
It will laid a theorwtical foundation for establishing relevant functional gene cloning system、obtaining our own intellectual property LMW-GS genes.
该论文首先以国内优质强筋小麦中优9507为实验材料,通过优化SDS-PAGE技术蛋白质分离条件,分离小麦LMW-GS区段的特异条带,质谱鉴定获得肽段;进一步分析小麦麦谷蛋白亚基基因密码子使用特性,采用RT-PCR和RACE技术克隆目的基因核酸片段,建立小麦LMW-GS基因克隆技术体系。在此基础之上,以新疆本地主栽强筋小麦新春8为实验材料,锁定小麦面包烘烤品质主效LMW-GS,质谱鉴定获得肽段,为后续该小麦面包烘烤品质主效LMW-GS基因克隆、小麦HMW-GS和LMW-GS互作研究提供科学的数据参考。该论文主要研究结果如下:
含0.3mol/LNaI和1.4%4-VP的LMW-GS提取液能有效提取小麦LMW-GS成分。在此基础上,当分离胶浓度为T=14%/C=3%、分离时间为40h、电极缓冲液为Tris-硼酸或Tris-甘氨酸时,能更加有效、准确、且重复性高地分离数目庞大的小麦LMW-GS。该技术体系为小麦LMW-GS深入研究的第一个限速步骤提供了良好的解决方案。
从国内优质强筋小麦中优9507中锁定1个小麦面包烘烤品质可能相关的LMW-GS,命名为ZY9507-L1,其表观分子量为34kD。质谱鉴定获得5个肽段,分别命名为ZL1-1、ZL1-2、ZL1-3、ZL1-4、ZL1-5。其中,ZL1-1(肽段序列:GIIQPQQPAQLEVIR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ACJ76961.1)的覆盖率(Query coverage)为66%,ZL1-2(肽段序列:SVVHSIIMQQEQR)与小麦已知LMW-GS氨基酸序列(GenBank登录号:ABB04903.1)的Query coverage为92%。其余3个质谱鉴定获得的小肽段均与NCBI数据库中小麦已知LMW-GS氨基酸序列不存在相似性。
为了解小麦麦谷蛋白亚基基因密码子使用特性,该文运用CodonW1.4.2和SPSS16.0软件对13个来源于小麦麦谷蛋白亚基基因的密码子用法特性进行了分析。结果显示,除1Dy10基因有效密码子数(ENC值)小于40外,其余亚基基因ENC值均介于40-50之间;密码子第三位GC含量(GC3)均在0.38-0.45之间;1Bx14基因偏向于使用A或T结尾的密码子,其余12个亚基基因偏向于使用G或C结尾的密码子;1Dy10与其余12个亚基基因间距离系数较大,然而各组染色体麦谷蛋白x-型或y-型亚基基因间距离系数较小。结果表明,小麦麦谷蛋白亚基基因具有较强的密码子使用偏爱性,各基因密码子用法上具有相似性,每一种氨基酸均存在1-2个偏爱密码子。在密码子用法差异上表现为,除1By9和1Dy10之外,无论是A染色体组、B染色体组,还是D染色体组,各组染色体上麦谷蛋白x-型或y-型亚基基因间密码子使用频率差异最小。这一分析结果对小麦HMW-GS功能基因资源的高效利用和LMW-GS新基因的克隆具有一定的指导意义。
根据小麦面包烘烤品质相关LMW-GS ZY9507-L1质谱鉴定结果和小麦麦谷蛋白亚基基因密码子使用特性,该文以国内优质强筋小麦中优9507为实验材料,利用ZL1-1肽段设计3’端上游引物,采用RT-PCR和RACE技术获得目的基因ZL1-13’端序列。结果显示,分离获得2个不同的LMW-GS基因序列,分别命名为ZL1-1P和ZL1-1I,其中,ZL1-1P基因长度为620bp,可编码包含186个氨基酸残基的成熟蛋白质。该基因与NCBI数据库中许多小麦已知LMW-GS基因均具有同源性,其与小麦已知LMW-GS基因(登录号:FJ441109.1)的覆盖率为19%(见图5-11);ZL1-1I基因长度为693bp,可编码包含209个氨基酸残基的成熟蛋白质。该基因与NCBI数据库中许多小麦已知LMW-GS基因均具有同源性,该基因序列与已知LMW-GS基因(登录号:FJ441109.1)的覆盖率为17%。初步证实上述2个基因可能均属于小麦LMW-GS基因的一部分,而且也证实了该小麦LMW-GS基因克隆技术体系是完全可行的。
在上述已建立的基因克隆技术体系基础之上,根据面包烘烤品质的决定机制特征,从新疆本地主栽强筋小麦新春8中锁定1个小麦面包烘烤品质主效LMW-GS,命名为XC8-L1,其表观分子量为46kD。质谱鉴定获得215个肽段,分别命名为XL1-1、XL1-2、XL1-3、XL1-4、XL1-5、……、XL1-215。其中,XL1-1(肽段序列:AIIYSIVLQEQQQVR)、XL1-2(肽段序列:VNVPLYR)、XL1-3(肽段序列:TTTSVPFGVGTGVGAY)和XL1-4(肽段序列:LEVMTSIALR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ACZ59817.1 )的Query coverage为100% , XL1-5 (肽段序列:TTTNVPFGVGTGVGSY)和XL1-6(肽段序列:VNVPLYR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ACA63873.1)的Query coverage为100%,XL1-7(肽段序列:QIAQLEVMTSIALR)和XL1-8(肽段序列:QIPEQSR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ADX68840.1)的Query coverage为100%,XL1-9(肽段序列:GTFLQPHQIAR)与NCBI数据库中小麦已知LMW-GS氨基酸序列(GenBank登录号:ADX68849.1)的Query coverage为100%。其余206个质谱鉴定后获得的小肽段均与NCBI数据库中小麦已知LMW-GS氨基酸序列不存在相似性。
该论文不仅为小麦面包烘烤品质相关功能基因克隆提供技术体系,而且为获得我国独立的自主知识产权的小麦面包烘烤品质主效LMW-GS基因克隆提供有效的氨基酸序列支持,还为深入探讨小麦HMW-GS和LMW-GS互作研究、加快我国面包小麦改良步伐并实现优良面包粉的自主生产奠定一定的理论基础。
Glutenin polymers are formed by high (HMW-GS) and low molecular weight glutenins subunits(LMW-GS) on the basis of their mobilities in the sodium dodecyl sulfate polyacrylamide gel electrophoresis(SDS-PAGE) in wheat(Triticum aestivum L.). It has been demonstrated that the variation in HMW-GS and LMW-GS composition has a strong impact on wheat bread-making quality. However, the deeply study on LMW-GS is far lower than HMW-GS. So, it’s very importace to develop our own intellectual property about LMW-GS.
In this paper, the SDS-PAGE separation condition was optimazed, then interrelated subunit may be for wheat bread-making quality was isolated in Zhongyou 9507, and identified them by LC-MS. In addition, we analyzed the codon usage of glutenin subunits genes in wheat, to explore HMW-GS genetic resources efficiently and clone new LMW-GS genes. Soon after, it isolated nucleotide sequence of interrelated subunit by RT-PCR and RACE. On this basis, we choose Xinchun 8 as the experimental material, then isolated the major subunit on wheat bread-making quality by optimized SDS-PAGE technonlgy, and identified them by LC-MS. The main results are as follows:
The LMW-GS extraction buffer containing 0.3mol/L NaI and 1.4% 4-VP can obtain LMW-GS availably, it's the best condition under T=14%C=3%、tris-boric acid buffer or tris-glycine buffer to separate glutenin subunits effectively. It provide a theoretical basic step for studying LMW-GS in breeding quality wheat cultivar.
It locked 1 interrelated subunit in Zhongyou9507, named ZY9507-L1, its apparent molecular weight is 34kDa. It obtained 5 peptides by LC-MS, named ZL1-1、ZL1-2、ZL1-3、ZL1-4、ZL1-5 orderly. At the same time, Query coverage is 66% between ZL1-1(Peptide is GIIQPQQPAQLEVIR)and amino acid sequence of known wheat LMW-GS(ACJ76961.1) in NCBI. Query coverage is 92% between ZL1-2 (Peptide is SVVHSIIMQQEQR)and amino acid sequence of known wheat LMW-GS(ABB04903.1) in NCBI. There is little similarity between other two peptides and known LMW-GS in NCBI.
The value of Effective Numberof Codons (ENC) of the other 12 glutenin subunits genes was between 40-50, except the 1Dy10 gene was less than 40.The GC3 contents were all between 0.38-0.45. The 1Bx14 Gene preferred to use A or T at the end of codon and the remaining 12 subunit genes preferred to use G or C . The distance coefficient of 1Dy10 was larger with rest of 12 genes, but it was small between x-type or y-type subunit genes of each chromosomes. The results showed that the glutenin subunit genes of wheat possessed a strong codon usage bias clearly. The codon usage of glutenin subunits genes was similar, and there were optimal(one or two) for each amino acid codon. Except for 1By9 and 1Dy10, the other 12 genes had the smallest differences in frequency of utilizing codon. The results would help to explore HMW-GS genetic resources efficiently and clone new LMW-GS genes.
According to the LC-MS and characterization on codon usage of glutenin subunits genes in Wheat, we designed the specific primers on the basis of ZL1-1 peptide, it extracted the total RNA in Zhongyou 9507 by Trizol, then it gained first strand cDNA by RT-PCR.And It gained 3’nucleotide sequence of ZL1-1 finally. It showed that, it gained two new genes, named ZL1-1P and ZL1-1I respectively. The coding regions of ZL1-1P is 620bp, encoding the proteins with 186 amino acid residues. The coding regions of ZL1-1I is 693bp, encoding the proteins with 209 amino acid residues. There have homology among ZL1-1P、ZL1-1I and the known wheat LMW-GS in NCBI. It indicated that the genes is the LMW-GS preliminary.
It locked 1 major subunit which is about wheat bread-making quality in Xinchun 8, named XC8-L1, molecular weight is 46kD. It obtained 215 peptides by LC-MS, named XL1-1、XL1-2、XL1-3、XL1-4、XL1-5、……、XL1-215 orderly. At the same time, Query coverage is 100% among XL1-1(Peptide is AIIYSIVLQEQQQVR)、XL1-2(Peptide is VNVPLYR)、XL1-3(Peptide is TTTSVPFGVGTGVGAY)、(Peptide is LEVMTSIALR)and amino acid sequence of wheat LMW-GS(ACZ59817.1) in NCBI, Query coverage is 100% among XL1-5(Peptide is TTTNVPFGVGTGVGSY) and XL1-6(Peptide is VNVPLYR)and amino acid sequence of wheat LMW-GS(ACA63873.1) in NCBI, Query coverage is 100% among XL1-7(Peptide is QIAQLEVMTSIALR) and XL1-8(Peptide is QIPEQSR)and amino acid sequence of wheat LMW-GS(ADX68840.1) in NCBI, Query coverage is 100% between XL1-9(Peptide is GTFLQPHQIAR)and amino acid sequence of wheat LMW-GS(ADX68849.1) in NCBI.
It will laid a theorwtical foundation for establishing relevant functional gene cloning system、obtaining our own intellectual property LMW-GS genes.
引文
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