小麦HMW-GS缺失种质资源的筛选鉴定及其品质效应研究
摘要
高分子量谷蛋白亚基(high molecular weight glutenin subunits, HMW-GS)是面团粘弹性的主要决定因素,HMW-GS的组成和变异直接影响到小麦的加工品质。本研究在对小麦种质资源HMW-GS组成分析的基础上,选择HMW-GS不同缺失类型的材料与长江下游地区主栽弱筋小麦品种扬麦13、扬麦18杂交组配,衍生回交群体与自交高代姊妹系群体,对HMW-GS不同缺失类型的遗传与品质效应进行了研究,探讨HMW-GS不同缺失类型在弱筋小麦品质改良上的应用价值。主要实验结果如下：
1、小麦HMW-GS缺失种质资源的筛选和鉴定：采用SDS-PAGE电泳检测方法,对261份小麦微核心种质和54份应用核心种质、97份CIMMYT人工合成小麦、20份普通六倍体小麦和2份转基因小麦进行了HMW-GS组成分析。供试的小麦微核心种质和应用核心种质在Glu-Al、 Glu-B1禾(?)Glu-D1三个位点的组成比较单一,主要由Null、7+8和2+12亚基组成。部分微核心种质表现出HMW-GS的缺失现象。其中,5份材料Glu-B1位点缺失1By亚基,表达1Bx7或1Bx22亚基；3份材料Glu-D1位点缺失1Dx或1Dy亚基,表达1Dyl2或lDx2亚基。在人工合成小麦中,HMW-GS存在着丰富的等位变异,2份材料在Glu-A1位点表达稀有亚基1Axl’,在Glu-B1位点和Glu-D1位点分别发现一个未知的亚基(暂命名为Y亚基和X亚基),并发现了一些新的亚基组合；此外,1份材料Glu-B1位点缺失lBy型亚基,表达1Bx7亚基：1份材料表现为Glu-A1禾WGlu-B1位点共同缺失；2份材料在Glu-D1位点缺失1Dx亚基,表达1Dyl0亚基。在供试的普通六倍体小麦中,3份材料表现为Glu-D1位点完全缺失,3份材料表现为Glu-A1和Glu-D1位点共同缺失,其中拉花9650在Glu-B1位点同时表达4种不同亚基类型。2份转基因小麦分别表现为Glu-D1位点1Dx5亚基缺失和Glu-1位点HMW-GS完全缺失。
2、 HMW-GS不同缺失类型的遗传：普通六倍体小麦2GS0414-10Glu-A1和Glu-D1位点共同缺失、拉花9650Glu-D1位点缺失,在Fl代均表现为隐性；在F2代表现为3：1或9：3：3：1的分离比例；BC1F1代不表达HMW-GS缺失。人工合成小麦SYN351Glu-A1和Glu-B1位点共同缺失在F1代表现为隐性；在F2代不符合3：1或9：3：3：1的理论比例,出现1By8缺失现象；BC1F1代未表达HMW-GS缺失。2份转基因小麦1Dx5缺失和HMW-GS完全缺失在F1代均表现为显性遗传；由于外源小RNA基因对1Dx5亚基的表达有特异性抑制作用,导致1Dx5缺失在F2代表现出13(不表达1Dx5)：3(表达1Dx5)的分离比例,在BC1F1代表现出3(不表达1Dx5)：1(表达1Dx5)的分离比例；HMW-GS完全缺失在F2代表现出3(不表达HMW-GS)1(表达HMW-GS)的分离比例,在BC1F1代表现出1(不表达HMW-GS)1(表达HMW-GS)的分离比例。
3、供体亲本的品质指标：在普通六倍体小麦2GS0414-10、拉花9650、转基因小麦Glu-1Dx5-RNAi和Glu-1-RNAi中,均表现较低的麦谷蛋白含量和较高的醇溶蛋白含量；较低的SDS沉淀值和水SRC,较短的面团形成时间和稳定时间,较低的粉质质量指数；与转化受体品种Bobwhite相比较,转基因小麦蛋白质含量显著增加,饼干直径增大。人工合成小麦SYN351具有较低的湿面筋含量和较高的面筋指数。
4、 HMW-GS不同缺失类型杂交后代的品质指标：对以Glu-A1和Glu-D1位点共同缺失的材料2GS0414-10、HMW-GS完全缺失的材料Glu-1-RNAi为供体亲本,以扬麦13和扬麦18为受体亲本衍生的回交后代BC2F3、BC1F3和两个自交高代姊妹系F5的品质测试结果表明：与HMW-GS正常表达的籽粒品质比较,当Glu-A1位点缺失时SDS沉淀值及水SRC均无显著变化；当Glu-D1位点缺失时SDS沉淀值显著降低,水SRC无显著变化；当Glu-A1和Glu-D1位点共同缺失或Glu-1位点HMW-GS完全缺失时,SDS沉淀值和水溶剂保持力(SRC)均显著降低；另外在普通六倍体小麦2GS0414-10的杂交后代中,当Glu-A1缺失、Glu-D1缺失或Glu-A1和Glu-D1位点共同缺失,籽粒蛋白质含量差异不显著。
5、综合本研究结果,HMW-GS缺失降低了麦谷蛋白含量,增加了醇溶蛋白含量；降低了SDS沉淀值和水溶剂保持力,缩短了面团形成时间和稳定时间,这有利于降低面筋含量和面团筋力强度,增大面团的延展性,增加饼干的直径。表明,HMW-GS缺失对改善弱筋小麦的加工品质可能有一定的贡献,可以考虑将不同HMW-GS缺失材料应用于弱筋小麦的品质改良工作。
HMW-GS (high molecular weight glutenin subunits) is important in determining the viscoelastic properties of wheat. Distribution and variation of HMW-GS are also related with the processing quality of wheat. In this study, HMW-GS distribution in wheat germplasm resources was firstly analyzed. Then, wheat lines with different types of HMW-GS deletion were selected and crossed with two soft wheat varieties Yangmai13and Yangmai18to generate advanced backcross and inbred population. To investigate the potential of HMW-GS deletion in soft wheat breeding, genetic behavior and quality effect of different types of HMW-GS deletion were analyzed. The results were described in detail as followings.
1. Screening of wheat germplasm resources with HMW-GS deletion. The HMW-GS distribution of432wheat materials, including a mini core set of wheat lines with261members and54application-core collection germplasms,97synthetic wheat lines from CIMMYT,20common wheat lines and two transgenetic wheat lines, were analyzed by the SDS-PAGE technique. The Glu-Al, Glu-B1and Glu-D1loci in a mini core set of wheat lines mainly consist of Null,7+8and2+12subunits. However, some wheat core lines show a certain HMW-GS deletion. Five lines possess1Bx7type or1Bx22subunits but lack1By type subunits at the Glu-B1loci. Other three lines have1Dx2or1Dy12type subunits but lack1Dy or1Dx type subunits at the Glu-D1loci. High variation of HMW-GS constitution was observed in synthetic wheat lines. A rare subunit temporarily named1Ax1' was identified. Two unknown subunits temporarily named Y and X type subunits were also found in synthetic wheat lines. Additionally, some new subunit combinations were detected in synthetic wheat lines. One synthetic wheat line contains1Bx7type subunit but lacks1By type subunit at the Glu-B1locus. Two synthetic wheat lines possess1Dy10type subunits but lack1Dx type subunits at the Glu-D1loci. Both Glu-Al and Glu-B1loci were absent in one synthetic wheat line. The Glu-Dl loci were lack in three common wheat lines. Both Glu-A1and Glu-D1loci were lost from three common wheat lines. The Glu-B1loci contains four type subunits in Lahua9650. The1Dx5subunit at the Glu-D1locus and whole Glu-D1locus were lack in two transgenetic wheat lines, respectively.
2. Genetic behavior of different types of HMW-GS deletion. The double-deletion of Glu-A1and Glu-D1loci of HMW-GS in common wheat line2GS0414-10and Glu-D1loci absence in Lahua9650all inherit recessively in F1progeny. HMW-GS composition in F2progenies is coincident with the3(HMW-GS expression):1(HMW-GS silence) ratio of Mendelian inheritance. HMW-GS absence was not observed in BC1F1lines. The double-deletion of Glu-A1and Glu-B1loci of HMW-GS in synthetic wheat line SYN351inherits recessively in F1progeny. HMW-GS distribution in F2progenies is inconsistent with the theoretical ratio3:1or9:3:3:1. HMW-GS absence was not observed in BC1F1plants. The silent effect of HMW-GS is dominant in two transgenetic wheat. Due to the suppression of1Dx5subunit by small RNA, HMW-GS distribution in F2and BC1F1progenies shows13(1Dx5subunit silence):3HMW-GS (1Dx5subunit expression) and3(1Dx5subunit silence):1(1Dx5subunit expression) ratios, respectively. The whole HMW-GS silent effect in F2and BC1F1progenies shows3(HMW-GS silence):1(HMW-GS expression) and1(HMW-GS silence):1(HMW-GS expression) ratios, respectively.
3. Quality evaluation of donor parents. Donor parents2GS0414-10, Lahua9605and Glu-1Dx5-RNA,Glu-1-RNAi all have comparatively low gluten content and high gliadin content, have relatively low SDS sedimentation value, water solvent retention capacity, development time, stability time and the index of flour quality. Compared with that of receptor line Bobwhite, Protein content increases significantly in transgenetic lines. Additionally, diameter of biscuits increases significantly due to all HMW-GS silence. Lower wet gluten content and higher gluten index due to the double-deletion of Glu-A1and Glu-B1loci in SYN351.
4. Quality analysis of progenies derived from different types of HMW-GS deletion. The wheat line2GS0414-10with the double-deletion of Glu-A1and Glu-Dl loci, and HMW-GS silence in transgenetic wheat lines were adopted as donor parent. Two soft wheat varieties Yangmai13and Yangmai18were used as receptor parents and crossed to generate BC2F3、BC1F3and F5progenies. Quality of parents and their progenies were tested. Results indicated that Glu-A1absence has little effect on protein content, SDS sedimentation value and water solvent retention capacity. The Glu-D1absence does not influence protein content and water solvent retention capacity but significantly influences the SDS sedimentation value. Intriguingly, double-deletion of Glu-A1and Glu-D1loci has little effect on protein content but significantly influence the SDS sedimentation value and water solvent retention capacity. Both the SDS sedimentation value and water solvent retention capacity decrease significantly owing to whole HMW-GS silence.
5. Results presented in this study indicated that wheat lines with HMW-GS deletion have comparatively low gluten content and high gliadin content, which is benefical to decrease the dough strength and increase the dough extensibility. Wheat lines with HMW-GS deletion also have relatively low SDS sedimentation value, water solvent retention capacity, development time and stability time. The HMW-GS deletion effect may be useful in quality improvement in soft wheat.
1、小麦HMW-GS缺失种质资源的筛选和鉴定：采用SDS-PAGE电泳检测方法,对261份小麦微核心种质和54份应用核心种质、97份CIMMYT人工合成小麦、20份普通六倍体小麦和2份转基因小麦进行了HMW-GS组成分析。供试的小麦微核心种质和应用核心种质在Glu-Al、 Glu-B1禾(?)Glu-D1三个位点的组成比较单一,主要由Null、7+8和2+12亚基组成。部分微核心种质表现出HMW-GS的缺失现象。其中,5份材料Glu-B1位点缺失1By亚基,表达1Bx7或1Bx22亚基；3份材料Glu-D1位点缺失1Dx或1Dy亚基,表达1Dyl2或lDx2亚基。在人工合成小麦中,HMW-GS存在着丰富的等位变异,2份材料在Glu-A1位点表达稀有亚基1Axl’,在Glu-B1位点和Glu-D1位点分别发现一个未知的亚基(暂命名为Y亚基和X亚基),并发现了一些新的亚基组合；此外,1份材料Glu-B1位点缺失lBy型亚基,表达1Bx7亚基：1份材料表现为Glu-A1禾WGlu-B1位点共同缺失；2份材料在Glu-D1位点缺失1Dx亚基,表达1Dyl0亚基。在供试的普通六倍体小麦中,3份材料表现为Glu-D1位点完全缺失,3份材料表现为Glu-A1和Glu-D1位点共同缺失,其中拉花9650在Glu-B1位点同时表达4种不同亚基类型。2份转基因小麦分别表现为Glu-D1位点1Dx5亚基缺失和Glu-1位点HMW-GS完全缺失。
2、 HMW-GS不同缺失类型的遗传：普通六倍体小麦2GS0414-10Glu-A1和Glu-D1位点共同缺失、拉花9650Glu-D1位点缺失,在Fl代均表现为隐性；在F2代表现为3：1或9：3：3：1的分离比例；BC1F1代不表达HMW-GS缺失。人工合成小麦SYN351Glu-A1和Glu-B1位点共同缺失在F1代表现为隐性；在F2代不符合3：1或9：3：3：1的理论比例,出现1By8缺失现象；BC1F1代未表达HMW-GS缺失。2份转基因小麦1Dx5缺失和HMW-GS完全缺失在F1代均表现为显性遗传；由于外源小RNA基因对1Dx5亚基的表达有特异性抑制作用,导致1Dx5缺失在F2代表现出13(不表达1Dx5)：3(表达1Dx5)的分离比例,在BC1F1代表现出3(不表达1Dx5)：1(表达1Dx5)的分离比例；HMW-GS完全缺失在F2代表现出3(不表达HMW-GS)1(表达HMW-GS)的分离比例,在BC1F1代表现出1(不表达HMW-GS)1(表达HMW-GS)的分离比例。
3、供体亲本的品质指标：在普通六倍体小麦2GS0414-10、拉花9650、转基因小麦Glu-1Dx5-RNAi和Glu-1-RNAi中,均表现较低的麦谷蛋白含量和较高的醇溶蛋白含量；较低的SDS沉淀值和水SRC,较短的面团形成时间和稳定时间,较低的粉质质量指数；与转化受体品种Bobwhite相比较,转基因小麦蛋白质含量显著增加,饼干直径增大。人工合成小麦SYN351具有较低的湿面筋含量和较高的面筋指数。
4、 HMW-GS不同缺失类型杂交后代的品质指标：对以Glu-A1和Glu-D1位点共同缺失的材料2GS0414-10、HMW-GS完全缺失的材料Glu-1-RNAi为供体亲本,以扬麦13和扬麦18为受体亲本衍生的回交后代BC2F3、BC1F3和两个自交高代姊妹系F5的品质测试结果表明：与HMW-GS正常表达的籽粒品质比较,当Glu-A1位点缺失时SDS沉淀值及水SRC均无显著变化；当Glu-D1位点缺失时SDS沉淀值显著降低,水SRC无显著变化；当Glu-A1和Glu-D1位点共同缺失或Glu-1位点HMW-GS完全缺失时,SDS沉淀值和水溶剂保持力(SRC)均显著降低；另外在普通六倍体小麦2GS0414-10的杂交后代中,当Glu-A1缺失、Glu-D1缺失或Glu-A1和Glu-D1位点共同缺失,籽粒蛋白质含量差异不显著。
5、综合本研究结果,HMW-GS缺失降低了麦谷蛋白含量,增加了醇溶蛋白含量；降低了SDS沉淀值和水溶剂保持力,缩短了面团形成时间和稳定时间,这有利于降低面筋含量和面团筋力强度,增大面团的延展性,增加饼干的直径。表明,HMW-GS缺失对改善弱筋小麦的加工品质可能有一定的贡献,可以考虑将不同HMW-GS缺失材料应用于弱筋小麦的品质改良工作。
HMW-GS (high molecular weight glutenin subunits) is important in determining the viscoelastic properties of wheat. Distribution and variation of HMW-GS are also related with the processing quality of wheat. In this study, HMW-GS distribution in wheat germplasm resources was firstly analyzed. Then, wheat lines with different types of HMW-GS deletion were selected and crossed with two soft wheat varieties Yangmai13and Yangmai18to generate advanced backcross and inbred population. To investigate the potential of HMW-GS deletion in soft wheat breeding, genetic behavior and quality effect of different types of HMW-GS deletion were analyzed. The results were described in detail as followings.
1. Screening of wheat germplasm resources with HMW-GS deletion. The HMW-GS distribution of432wheat materials, including a mini core set of wheat lines with261members and54application-core collection germplasms,97synthetic wheat lines from CIMMYT,20common wheat lines and two transgenetic wheat lines, were analyzed by the SDS-PAGE technique. The Glu-Al, Glu-B1and Glu-D1loci in a mini core set of wheat lines mainly consist of Null,7+8and2+12subunits. However, some wheat core lines show a certain HMW-GS deletion. Five lines possess1Bx7type or1Bx22subunits but lack1By type subunits at the Glu-B1loci. Other three lines have1Dx2or1Dy12type subunits but lack1Dy or1Dx type subunits at the Glu-D1loci. High variation of HMW-GS constitution was observed in synthetic wheat lines. A rare subunit temporarily named1Ax1' was identified. Two unknown subunits temporarily named Y and X type subunits were also found in synthetic wheat lines. Additionally, some new subunit combinations were detected in synthetic wheat lines. One synthetic wheat line contains1Bx7type subunit but lacks1By type subunit at the Glu-B1locus. Two synthetic wheat lines possess1Dy10type subunits but lack1Dx type subunits at the Glu-D1loci. Both Glu-Al and Glu-B1loci were absent in one synthetic wheat line. The Glu-Dl loci were lack in three common wheat lines. Both Glu-A1and Glu-D1loci were lost from three common wheat lines. The Glu-B1loci contains four type subunits in Lahua9650. The1Dx5subunit at the Glu-D1locus and whole Glu-D1locus were lack in two transgenetic wheat lines, respectively.
2. Genetic behavior of different types of HMW-GS deletion. The double-deletion of Glu-A1and Glu-D1loci of HMW-GS in common wheat line2GS0414-10and Glu-D1loci absence in Lahua9650all inherit recessively in F1progeny. HMW-GS composition in F2progenies is coincident with the3(HMW-GS expression):1(HMW-GS silence) ratio of Mendelian inheritance. HMW-GS absence was not observed in BC1F1lines. The double-deletion of Glu-A1and Glu-B1loci of HMW-GS in synthetic wheat line SYN351inherits recessively in F1progeny. HMW-GS distribution in F2progenies is inconsistent with the theoretical ratio3:1or9:3:3:1. HMW-GS absence was not observed in BC1F1plants. The silent effect of HMW-GS is dominant in two transgenetic wheat. Due to the suppression of1Dx5subunit by small RNA, HMW-GS distribution in F2and BC1F1progenies shows13(1Dx5subunit silence):3HMW-GS (1Dx5subunit expression) and3(1Dx5subunit silence):1(1Dx5subunit expression) ratios, respectively. The whole HMW-GS silent effect in F2and BC1F1progenies shows3(HMW-GS silence):1(HMW-GS expression) and1(HMW-GS silence):1(HMW-GS expression) ratios, respectively.
3. Quality evaluation of donor parents. Donor parents2GS0414-10, Lahua9605and Glu-1Dx5-RNA,Glu-1-RNAi all have comparatively low gluten content and high gliadin content, have relatively low SDS sedimentation value, water solvent retention capacity, development time, stability time and the index of flour quality. Compared with that of receptor line Bobwhite, Protein content increases significantly in transgenetic lines. Additionally, diameter of biscuits increases significantly due to all HMW-GS silence. Lower wet gluten content and higher gluten index due to the double-deletion of Glu-A1and Glu-B1loci in SYN351.
4. Quality analysis of progenies derived from different types of HMW-GS deletion. The wheat line2GS0414-10with the double-deletion of Glu-A1and Glu-Dl loci, and HMW-GS silence in transgenetic wheat lines were adopted as donor parent. Two soft wheat varieties Yangmai13and Yangmai18were used as receptor parents and crossed to generate BC2F3、BC1F3and F5progenies. Quality of parents and their progenies were tested. Results indicated that Glu-A1absence has little effect on protein content, SDS sedimentation value and water solvent retention capacity. The Glu-D1absence does not influence protein content and water solvent retention capacity but significantly influences the SDS sedimentation value. Intriguingly, double-deletion of Glu-A1and Glu-D1loci has little effect on protein content but significantly influence the SDS sedimentation value and water solvent retention capacity. Both the SDS sedimentation value and water solvent retention capacity decrease significantly owing to whole HMW-GS silence.
5. Results presented in this study indicated that wheat lines with HMW-GS deletion have comparatively low gluten content and high gliadin content, which is benefical to decrease the dough strength and increase the dough extensibility. Wheat lines with HMW-GS deletion also have relatively low SDS sedimentation value, water solvent retention capacity, development time and stability time. The HMW-GS deletion effect may be useful in quality improvement in soft wheat.
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