野生二粒麦（T.dicoccoides）谷蛋白亚基双向电泳分析及其LMW-GS 编码基因的分子克隆
|
摘要
小麦贮藏蛋白中的高、低分子量谷蛋白亚基是决定小麦面粉品质优劣的重要因素。野生二粒小麦(Triticum dicoccoides，AABB，2n=4x=28)是普通小麦基因组AABB的供体，具有较高的蛋白含量和丰富的遗传变异，是小麦品质改良的重要基因资源，因此对其谷蛋白亚基进行鉴定及其编码基因的分子克隆对于定向改良小麦品质以及了解贮藏蛋白基因家族结构、分子进化关系具有重要的意义。本研究利用十二烷基硫酸钠聚丙烯酰胺凝胶电泳(SDS-PAGE)，酸性聚丙烯酰胺凝胶电泳(A-PAGE)和双向聚丙烯酰胺凝胶电泳(2-D PAGE：A-PAGE×SDS-PAGE)三种电泳方法对28份野生二粒小麦的高分子量谷蛋白亚基(HMW-GS)和低分子量谷蛋白亚基(LMW-GS)进行分离鉴定，然后采用等位基因特异PCR(Allele SpecificPCR，AS-PCR)方法对低分子量谷蛋白亚基(LMW-GS)基因进行分子克隆、序列测定和比较分析。主要研究结果如下：
●高、低分子量谷蛋白亚基的凝胶电泳分离与鉴定
利用SDS-PAGE方法，对28份野生二粒小麦高分子量谷蛋白亚基进行初步分离鉴定，结果发现：Glu-A1位点编码Null、1、1~*、2~*四种类型的亚基；Glu-B1位点编码12种亚基组合类型，分别是6+8，7+8，13+19，6+16，6+19，6~*+8，6.1+22.1，13+22.1，13+22~*，14+15，13+Null，Null+16，其中除了前三种外均为新鉴定的亚基类型。在此基础上采用SDS-PAGE，A-PAGE和A-PAGE×SDS-PAGE三种不同的方法对9个具有新型HMW-GS的材料进行进一步分离鉴定，结果发现：部分HMW-GS在SDS-PAGE和A-PAGE中的迁移率存在差异。除了Y22的亚基1~*分成两个点以外，其余在SDS-PAGE中鉴定的HMW-GS在A-PAGE×SDS-PAGE中均为清晰单一的蛋白质点，由各自特定的基因编码。
利用SDS-PAGE和A-PAGE×SDS-PAGE两种不同方法，对野生二粒小麦LMW-GS进行了鉴定。根据1996年Jackson等提出的LMW-GS等位基因命名方法，对4个野生二粒小麦LMW-GS等位基因进行了鉴定，结果如下：Y6(g,f)，Y7(d,f)，Y22(g,f)，Y25(d,f)。此外，以LMW-2型标准品种Simeto(Null，7+8)作对照，对28份野生二粒小麦进行筛选，结果发现11份具有LMW-2型电泳图谱，表明它们可能象硬粒小麦一样具有较好的加工品质，因此野生二粒小麦可能具有丰富的优质谷蛋白亚基基因资源。
●LMW-GS基因的分子克隆、序列比较与分子进化分析
利用一对特异的LMW-GS基因引物X13306-F595和X13306-R2103进行AS-PCR扩增，从三个野生二粒小麦中均获得约1600bp的单一特异强扩增条带，分别回收克隆后进行DNA序列测定，得到基因全长分别为1649bp、1472bp、1754bp
首都师范大学硕士学位论文
的包括上下游和编码区的LMW-GS完整基因序列。根据N一端氨基酸序列，发现这
三个基因编码的成熟蛋白的第一个氨基酸是异亮氨酸，因此它们编码的蛋白是
LMW-i型亚基。与11个已知LM私GS基因进行序列比较，克隆的三个基因具有
LMW-GS典型的结构特点，编码区依次由信号肤、区域n、区域111、区域W和
区域V组成，缺失了区域I。在这三个基因中，LMW-GS特有的8个半肤氨酸残
基全部位于C一端区域，与LMW-m和LMW-s型亚基存在较大差异。分子进化树
分析表明:LMW-i型与LMW-m型和LMW-s型基因各自形成两个独立的分支，
LMW-m型和LMW-s型基因在一个分支内也形成两个亚组，但具有较高的同源性。
可以推测不同LMW-GS类型的基因经遗传分化后沿着各自不同的方向进化，随着
DNA重复、缺失、插入、突变等事件发生，LMW-i型基因比LMW-m型和LMW-s
型基因积累了更多的变异，从而形成一组具有独特特征的基因类型。对上述基因
的转化表达以及对面包品质的影响正在进一步研究之中。
Both high- and low-molecular-weight glutenin subunits are the important compnents of wheat seed storage proteins and play the major role in determining bread-making quality. The AABB genome donor of bread wheat - wild emmer wheat (Triticum dicoccoides, AABB,2n=4x=28 ) was found to possess high protein content and extensive glutenin variation, therefore it is expected to serve as an important gene resource for bread wheat quality improvement. In this study, HMW- and LMW-glutenin subunits from 28 wild emmer accessions were separated and identified by SDS-PAGE, A-PAGE and A-PAGE X SDS-PAGE. A pair of allele specific PCR primers was used to amplify and clone some LMW-GS genes. The main results were as the followings:
Separation and identification of HMW-GS and LMW-GS in Triticum dicoccoides by PAGE
HMW-glutenin subunits of 28 wild emmer accessions were analyzed using SDS-PAGE and extensive variations at Glu-3 loci were detected. Four allelic variants (Null, 1, 1*, 2*) at Glu-A1 locus and twelve allelic variants (6+8, 7+8, 13+19, 6+ 16, 6+19, 6*+8, 6. 1+22. 1, 13+22. 1,13+22*, 14+15, 13, 16) at Glu-Bl locus were found. Except for 6+8, 7+8 and 13+19, HMW-glutenin patterns controlled by Glu-Bl locus were new allelic variants. Nine wild emmer accessions with novel HMW glutenin subunits were further characterized and confirmed by A-PAGE and two-dimensional gel electrophoresis (A-PAGE x SDS-PAGE). Some HMW-GS separated by A-PAGE showed different mobilities comparing to SDS-PAGE.
LMW-glutenin subunits in Triticum dicoccoides were analyzed by SDS-PAGE, A-PAGE X SDS-PAGE. The allelic variants of four wild emmer accessions were identified based on the nomenclature of alleles proposed by Jackson et al.(1996). The results showed that eleven wild emmer accessions possessed the LMW-2 detected in cultivated durum wheat, which was found to have positive effect on bread-making quality.
Cloning, characterization and molecular evolutional analysis of LMW-GS genes
A pair of LMW-GS gene AS-PCR primers X13306-F595 and X13306-R2103 was used to amplify the LMW-GS genes from Triticum dicoccoides accessions. Single strongly amplified band with about 1600bp from three accessions were obtained, and then the amplified products were cloned and sequenced. The complete nucleotide sequences of three LMW-GS genes have 1649bp, 1472bp and 1754bp, respectively, containing an open reading frame, 5'-upstream sequence and 3'-downstream sequence.
First amino acid of the deduced mature proteins of three LMW-GS genes was isoleucine amino acid residues, so they were all classed to LMW-i type glutenins. Three LMW-GS gene sequences showed a clear structural organization of the polypeptide comparing with other LMW-glutenin DNA sequences previously published. Except the domain I, three deduced proteins are all composed of signal peptide, domain II, domain III, domain IV and domain V. The expected number of eight cysteine residues encoded by three genes was all located in the C-terminal region, which are different from LMW-m and LMW-s type subunits. The phenetic trees produced by nucleotide and protein sequences showed that the LMW-i type subunit genes and LMW-m, LMW-s type subunit genes were clustered together respectively. LMW-m and LMW-s type genes were also separated into two subclasses. This suggested that different LMW genes have its own evolutionary way after genetic differentiation. But the LMW-i type subunit genes have accumulated more variations than LMW-m and LMW-s type genes during the evolutional process.
●高、低分子量谷蛋白亚基的凝胶电泳分离与鉴定
利用SDS-PAGE方法，对28份野生二粒小麦高分子量谷蛋白亚基进行初步分离鉴定，结果发现：Glu-A1位点编码Null、1、1~*、2~*四种类型的亚基；Glu-B1位点编码12种亚基组合类型，分别是6+8，7+8，13+19，6+16，6+19，6~*+8，6.1+22.1，13+22.1，13+22~*，14+15，13+Null，Null+16，其中除了前三种外均为新鉴定的亚基类型。在此基础上采用SDS-PAGE，A-PAGE和A-PAGE×SDS-PAGE三种不同的方法对9个具有新型HMW-GS的材料进行进一步分离鉴定，结果发现：部分HMW-GS在SDS-PAGE和A-PAGE中的迁移率存在差异。除了Y22的亚基1~*分成两个点以外，其余在SDS-PAGE中鉴定的HMW-GS在A-PAGE×SDS-PAGE中均为清晰单一的蛋白质点，由各自特定的基因编码。
利用SDS-PAGE和A-PAGE×SDS-PAGE两种不同方法，对野生二粒小麦LMW-GS进行了鉴定。根据1996年Jackson等提出的LMW-GS等位基因命名方法，对4个野生二粒小麦LMW-GS等位基因进行了鉴定，结果如下：Y6(g,f)，Y7(d,f)，Y22(g,f)，Y25(d,f)。此外，以LMW-2型标准品种Simeto(Null，7+8)作对照，对28份野生二粒小麦进行筛选，结果发现11份具有LMW-2型电泳图谱，表明它们可能象硬粒小麦一样具有较好的加工品质，因此野生二粒小麦可能具有丰富的优质谷蛋白亚基基因资源。
●LMW-GS基因的分子克隆、序列比较与分子进化分析
利用一对特异的LMW-GS基因引物X13306-F595和X13306-R2103进行AS-PCR扩增，从三个野生二粒小麦中均获得约1600bp的单一特异强扩增条带，分别回收克隆后进行DNA序列测定，得到基因全长分别为1649bp、1472bp、1754bp
首都师范大学硕士学位论文
的包括上下游和编码区的LMW-GS完整基因序列。根据N一端氨基酸序列，发现这
三个基因编码的成熟蛋白的第一个氨基酸是异亮氨酸，因此它们编码的蛋白是
LMW-i型亚基。与11个已知LM私GS基因进行序列比较，克隆的三个基因具有
LMW-GS典型的结构特点，编码区依次由信号肤、区域n、区域111、区域W和
区域V组成，缺失了区域I。在这三个基因中，LMW-GS特有的8个半肤氨酸残
基全部位于C一端区域，与LMW-m和LMW-s型亚基存在较大差异。分子进化树
分析表明:LMW-i型与LMW-m型和LMW-s型基因各自形成两个独立的分支，
LMW-m型和LMW-s型基因在一个分支内也形成两个亚组，但具有较高的同源性。
可以推测不同LMW-GS类型的基因经遗传分化后沿着各自不同的方向进化，随着
DNA重复、缺失、插入、突变等事件发生，LMW-i型基因比LMW-m型和LMW-s
型基因积累了更多的变异，从而形成一组具有独特特征的基因类型。对上述基因
的转化表达以及对面包品质的影响正在进一步研究之中。
Both high- and low-molecular-weight glutenin subunits are the important compnents of wheat seed storage proteins and play the major role in determining bread-making quality. The AABB genome donor of bread wheat - wild emmer wheat (Triticum dicoccoides, AABB,2n=4x=28 ) was found to possess high protein content and extensive glutenin variation, therefore it is expected to serve as an important gene resource for bread wheat quality improvement. In this study, HMW- and LMW-glutenin subunits from 28 wild emmer accessions were separated and identified by SDS-PAGE, A-PAGE and A-PAGE X SDS-PAGE. A pair of allele specific PCR primers was used to amplify and clone some LMW-GS genes. The main results were as the followings:
Separation and identification of HMW-GS and LMW-GS in Triticum dicoccoides by PAGE
HMW-glutenin subunits of 28 wild emmer accessions were analyzed using SDS-PAGE and extensive variations at Glu-3 loci were detected. Four allelic variants (Null, 1, 1*, 2*) at Glu-A1 locus and twelve allelic variants (6+8, 7+8, 13+19, 6+ 16, 6+19, 6*+8, 6. 1+22. 1, 13+22. 1,13+22*, 14+15, 13, 16) at Glu-Bl locus were found. Except for 6+8, 7+8 and 13+19, HMW-glutenin patterns controlled by Glu-Bl locus were new allelic variants. Nine wild emmer accessions with novel HMW glutenin subunits were further characterized and confirmed by A-PAGE and two-dimensional gel electrophoresis (A-PAGE x SDS-PAGE). Some HMW-GS separated by A-PAGE showed different mobilities comparing to SDS-PAGE.
LMW-glutenin subunits in Triticum dicoccoides were analyzed by SDS-PAGE, A-PAGE X SDS-PAGE. The allelic variants of four wild emmer accessions were identified based on the nomenclature of alleles proposed by Jackson et al.(1996). The results showed that eleven wild emmer accessions possessed the LMW-2 detected in cultivated durum wheat, which was found to have positive effect on bread-making quality.
Cloning, characterization and molecular evolutional analysis of LMW-GS genes
A pair of LMW-GS gene AS-PCR primers X13306-F595 and X13306-R2103 was used to amplify the LMW-GS genes from Triticum dicoccoides accessions. Single strongly amplified band with about 1600bp from three accessions were obtained, and then the amplified products were cloned and sequenced. The complete nucleotide sequences of three LMW-GS genes have 1649bp, 1472bp and 1754bp, respectively, containing an open reading frame, 5'-upstream sequence and 3'-downstream sequence.
First amino acid of the deduced mature proteins of three LMW-GS genes was isoleucine amino acid residues, so they were all classed to LMW-i type glutenins. Three LMW-GS gene sequences showed a clear structural organization of the polypeptide comparing with other LMW-glutenin DNA sequences previously published. Except the domain I, three deduced proteins are all composed of signal peptide, domain II, domain III, domain IV and domain V. The expected number of eight cysteine residues encoded by three genes was all located in the C-terminal region, which are different from LMW-m and LMW-s type subunits. The phenetic trees produced by nucleotide and protein sequences showed that the LMW-i type subunit genes and LMW-m, LMW-s type subunit genes were clustered together respectively. LMW-m and LMW-s type genes were also separated into two subclasses. This suggested that different LMW genes have its own evolutionary way after genetic differentiation. But the LMW-i type subunit genes have accumulated more variations than LMW-m and LMW-s type genes during the evolutional process.
引文
[美]D．R．马歇克，J．T．门永，R．R．布格斯，M．W．克努特，W．A．布伦南，S-H林著，朱厚础等译(1999)蛋白质纯化与鉴定实验指南，科学出版社，北京：249-262
郭尧君主编(1999) 蛋白质电泳实验技术，科学出版社，北京：54-68
J．萨母布鲁克，E．F．弗里奇，T．曼尼阿蒂斯(1998)分子克隆实验指南(第二版)．科学出版社，北京：19-22，55-56，880—886
姜怡(2003)面包小麦及其近源种HMW谷蛋白亚基鉴定及其编码基因的分子克隆，首都师范大学硕士论文
李保云，解超杰，尤明山，毛善锋，刘广田(2002) 野生二粒小麦高分子量谷蛋白亚基的多态性分析．麦类作物学报22(3)：21—24
汪家政，范明主编(2000) 蛋白质技术手册，科学出版社，北京：
王罡，季静，胡含(1995)小麦种子贮藏蛋白的遗传学研究进展．遗传17(1)：45—48
夏其昌主编(1997) 蛋白质化学研究技术与进展．科学出版社，北京：99—107
晏月明，刘广田，Prodanovic S，Mominovic G S，Zorie D，Perovie D(1998)小麦谷蛋白亚基的凝胶电泳分离及其品种鉴定．中国粮油学报13(6)：1—5
余建中，晏月明，蔡民华，胡英考 (2002) 毛细管电泳技术及其在作物贮藏蛋白研究中的应用．麦类作物学报22(3)：78—81
张发云，晏月明，胡英考，胡赞民，周奕华(2002) 小麦HMW谷蛋白亚基基因克隆研究进展．中国生物工程杂志22(6)：33-40
Anderson OD, Green FC, Vip RE, Halford NG, Shewry PR, Malpica-Romero JM (1989) Nucleotide sequences of the two high-molecular-weight glutenin genes from the D-genome of a hexaploid bread wheat, Tritivum aestivum L. cv Cheyenne. Nucleic Acids Res 17:461-462
Bartels D, Thompson RD (1983) The characterization of cDNA clones coding for wheat storage proteins. Nucleic Acids Res 11: 2961-2977
Bean SR, Lookhart GL (2001a) Recent developments in high-performance capillary electrophoresis of cereal proteins. Electrophoresis 22:1503-1509
Bean SR, Lookhart GL (2001b) High-performance capillary electrophoresis of meat, dairy, and cereal proteins. Electrophoresis 22:4207-4215
Benmoussa M, Vezina LP, Page M, Yelle S,Laberge S (2000) Genetic polymorphism in low-molecular-weight glutenin genes from Triticum aestivum, variety Chinese Spring. Theor Appl Genet 100:789-793
Bietz JA, Shepherd KW, Wall JS (1975) Single kernel analysis of glutenin:use in wheat genetics and breeding. Cereal Chem 52:513-532
Bietz JA, Schmalzried E (1992) Capillary electrophoresis of wheat proteins: optimization and use for varietal identification. Cereal Foods World 37:555-560
Brown JWS,Flavell RB (1981a) Fractionation of wheat gliadin and glutenin subunits by two-dimensional electrophoresis and the role of group 6 and group 2 chromosomes in gliadin synthesis. Theor Appl Genet 59:349-359
Brown JWS, Law CN, Worland AJ, Flavell RB (1981b) Genetic variation in wheat endosperm proteins:an analysis by two-dimensional electrophoresis using intervarietal chromosomal substitution lines. Theor Appl Genet 59:361-371
Bunce NAC, White RP, Shewry PR (1985) Variation in estimates of molecular weight of cereal prolamins by SDS-PAGE. Journal of Cereal Science 3:131-142
Caballero L,Martin LM,Alvarez JB (2004) Genetic Variability of the low-molecular-weight glutenin subunits in spelt wheat (Triticum aestivum ssp.spelta L.em Thell.). Theor Appl Genet 108:914-919
Cassidy BG, Dvorak J (1991) Molecular characterization of a low-molecular-weight glutenin cDNA clone from Triticum durum. Theor Appl Genet 81:653-660
Cassidy BG, Dvorak J, Anderson OD (1998) The wheat low-molecular-weight glutenin genes: characterization of six new genes and progress in understanding gene family structure. Theor Appl.Genet 96:743-750
Ciaffi M, Dominici L, Lafiandra D, Porceddu E (1992) Seed storage proteins of wild wheat progenitors and their relationships with technological properties Hereditas 116:315-322
Ciaffi M,Lafiandra D, Porceddu E, Benedettelli S (1993) Storage-protein variation in wild emmer wheat (Triticum turgidum ssp. dicoccoides) from Jordan and Turkey. Ⅰ. Electrophoretie characterization of genotypes. Theor Appl Genet 86:474-480
Ciaffi M, Lee YK, Tamas L, Gupta R, Skerritt J, Appels R (1999) The low-molecular-weight glutenin subunit proteins of primitive wheats. Ⅲ. The genes from D-genome species. Theor Appl Genet 98:135-148
Cloutier S, Rampitsch C, Penner GA, Lukow OM (2001) Cloning and expression of a LMW-i glutenin gene. Journal of Cereal Science 33: 143-154
Colot V, Bartels D, Thompson R, Flavell R (1989) Molecular characterization of an active wheat LMW glutenin gene and its relation to other wheat and barley prolamin genes. Mol Gen Genet 216:81-90
Dachkevitch T, Redaelli R, Biancardi AM, Metakovsky EV, Pogna NE (1993) Genetics of gliadins coded by the group-1 chromosomes in the high-quality bread wheat cultivar Neepawa. Theor Appl Genet 86:389-399
D'Ovidio R,Tanzarella OA, Porceddu E (1992) Nucleotide sequence of a low-molecular-weight glutenin from Triticum durum. Plant Mol Biol 18:781-784
D'Ovidio R, Simeone M, Masci S, Porceddu E (1997) Molecular characterization of a LMW-GS gene located on chromosome 1B and the development of primers specific for the Glu-B3 complex locus in durum wheat. Theor Appl Genet
95:1119-1126
D'Ovidio R, Marchitelli C, Ercoli CL, Porceddu E (1999) Sequence similarity between allelic Glu-B3 genes related to quality properties of durum wheat. Theor Appl Genet 98:455-461
Draper SR (1987) ISTA Variety Committee Report of the working group for Biochemical Tests for Cultivar Identification 1983-1986. Seed Sci. & Technol. 15: 431-434
Galili G, Feldman M (1983) Genetic control of endosperm proteins in wheat. 2. Variation of HMW glutenin and gliadin subunits of Triticum aestivum. Theor Appl Genet 66:77-86
Galili G, Felsenburg T, Levy AA, Altschuler Y, Feldman M (1988) Inactivity of high-molecular-weight glutenin genes in wild diploid and tetraploid wheats. In: Proceedings of the 7th international wheat genetics symposium: 81-87. Eds. T. E. Miller and R. M. D. Koebner. Cambridge, UK: Institute of Plant Science Research.
Gianibelli MC, Larroque OR, Maeritchie F, Wrigley CW (2001) Biochemical, genetic,and molecular characterization of wheat glutenin and its component subunits. Cereal Chem 78(6): 635-646
Gianibelli MC, Masci S, Larroque R, Lafiandra D (2002a) Biochemical characterization of a novel polymeric protein subunit from bread wheat (Triticum aestivum L.). Journal of Cereal Science 35:265-276
Gianibelli MC, Wrigley CW, Macritchie F (2002b) Polymorphism of low M_r glutenin subunits in Triticum tauschii. Journal of Cereal Science 35:277-286
Gianibelli MC, Solomon RG (2003) A Novel y-type High Mr Glutenin Subunit (12.4~t) Present in Triticum tauschii. Journal of Cereal Science 37:253-256
Gupta RB, Singh NK, Shepherd KW (1989) The cumulative effect of allelic variation in LMW and HMW glutenin subunits on dough properties in the progeny of two bread wheats. Theor Appl Genet 77:57-64
Gupta RB, Shepherd KW (1990) Two-step one-dimensional SDS-PAGE analysis of LMW subunits of glutenin. Theor Appl Genet 80:65-74
Gupta RB, Macritchie F (1991) Rapid communication a rapid one-step one-dimensional SDS-PAGE procedure for analysis of subunit composition of glutenin in wheat. Journal of Cereal Science 14:105-109
Gupta RB, Macritchie F (1994) Allelic variation at glutenin subunit and gliadin loci, Glu-1, Glu-3 and Gli-1 of common wheats. Ⅱ. Biochemical basis of the allelic effects on dough properties. Journal of Cereal Science 19:19-29
Gupta RB, Popineau Y, Lefebvre J, Cornec M, Lawrence GJ, MacRitchie F (1995) Biochemical basis of flour properties in bread wheats.Ⅱ.Changes in polymeric
protein formation and dough/gluten properties associated with the loss of low M_r or high M_r glutenin subunits. Journal of Cereal Science 21: 103-116
Halford NG, Forde J, Shewry PR, Kreis M (1989) Functional analysis of the upstream regions of a silent and an expressed member of a family of wheat seed protein genes in transgenic tobacco. Plant Sci 62:207-216
Herbert Wieser (2000) Comparative investigations of gluten proteins from different wheat species. 1. Qualitative and quantitative composition of gluten protein types. Eur Food Res Technol 211: 262-268
Ikeda TM, Nagamine T, Fukupka H, Yano H (2002) Identification of new low-molecular-weight glutenin subunit genes in wheat. Theor Appl Genet 104: 680-687
Jackson EA, Holt LM,Payne PI (1983) Characterization of high-molecular-weight gliadin and low-molecular-weight subunits of wheat endosperm by two-dimensional eleetrophoresis and the chromosomal location of their controlling genes. Theor Appl Genet 66:29-37
Jackson EA, Morel MH, Strohm TS, Branlard G; Metakovsky EV, Redaelli R (1996) Proposal for combining the classification systems of alleles of Gli-1 and Glu-3 loci in bread wheat (Triticum aestivum L.). J Genet & Breed 50:321-336
Kasarda DD (1989) Glutenin structure in relation to wheat quality In: Pomeranz Y(ed) wheat is unique. AACC, St Paul, Minnesota: 299-302
Kolster P, Krechting CF, Gelder WMJ van (1988) Variation in high molecular weight glutenin subunits of Triticura aestivum and T.turgidum ssp.dicoccoides.Euphytica S: 141-145
Klose J (1975) Protein mapping by combined isoelectric focusing and electrophoresis of mouse tissues. A novel approach to testing for induced point mutations in mammals. Human Genetics 26:231-243
Lee YK, Bekes F, Gupta R, Appels R, Morell MK (1999a) The low-molecular-weight glutenin subunit proteins of primitive wheats.Ⅰ.Variation in A-genome species. Theor Appl Genet 98:119-125
Lee YK, Ciaffi M, Appels R, Morell MK (1999b) The low-molecular-weight glutenin subunit proteins of primitive wheats. Ⅱ. The genes from A-genome species. Theor Appl Genet 98:126-134
Lee YK, Bekes F, Gras P, Ciaffi M, Morell MK, Appels R (1999c) The low-molecular-weight glutenin subunit proteins of primitive wheats.IV.Functional properties of products from individual genes. Theor Appl Genet 98:149-155
Lew EJL, Kuzmicky DD, Kasarda DD (1992) Characterization of low molecular weight glutenin subunits by reversed-phase high-performance liquid chromatography, sodium dodecyl sulfate-polyacrylamide gel electrophoresis, and
N-terminal amino acid sequencing. Cereal Chem 69(5): 508-515
Liu CY (1995) Research note identification of a new low Mr glutenin subunit locus on chromosome 1B of durum wheat. Journal of Cereal Science 21: 209-213
Luo C, Griffin WB, Branlard G, Mcneil DL (2001) Comparison of low-and high molecular-weight wheat glutenin allele effects on flour quality. Theor Appl Genet 102:1088-1098
Mackie AM, Sharp P J, Lagudah ES (1996) The Nucleotide and Derived Amino Acid Sequence of a HMW Glutenin Gene from Triticum tauschii and Comparison with those from the D Genome of Bread Wheat. Journal of Cereal Science 24:73-78
Masci S, Lafiandra D, Porceddu E, Lew EJL, Tao HP, Kasarda DD (1993) D-glutenin subunits: N-terminal sequences and evidence for the presence of cystein. Cereal Chem 70(5): 581-585
Masci S, Lew EJL,Lafiandra D, Porceddu E, Kasarda DD (1995) Characterization of low molecular weight glutenin subunits in durum wheat by reversed-phase high-performance liquid chromatography and N-terminal sequencing. Cereal Chem 72(1): 100-104
Masci S, D'Ovidio R, Lafiandra D, Kasarda DD (1998) Characterization of a low-molecular-weight glutenin subunit gene from bread wheat and the corresponding protein that represents a major subunit of the glutenin polymer. Plant Physiol 118:1147-1158
Masci S, D'Ovidio R, Lafiandra D, Kasarda DD (2000) A 1B-coded lowmolecular-weight glutenin subunits associated with quality in durum wheat shows strong similarity to a subunit present in some bread wheat cultivars. Theor Appl Genet 100:396-400
Masci S, Rovelli L,Kasarda DD, Vensel WH (2002) Characterisation and chromosomal localization of C-type low-molecular-weight glutenin subunits in the bread wheat cultivar Chinese Spring. Theor Appl Genet 104:422-428
Mcdonald MB, Elliot LJ, Sweeney PM (1994) Research note DNA extraction from dry seeds for RAPD analyses in varietal identification studies. Seed Sci & Technol 22:171-176
Mecham DK, Kasarda DD, Qualset CO (1978) Genetic aspects of wheat gliadin proteins. Biochem Genet 16:831-853
Metakovsky EV, Branlard G, Chernakov VM, Upelniek VP, Redaelli R, Pogna NE (1997) Recombination mapping of some chromosome 1A-,1B-,1D-and 6B-controlled gliadins and low-molecular-weight glutenin subunits in common wheat. Theor Appl Genet 94:788-795
Morel MH (1994) Acid-polyacrylamide gel electrophoresis of wheat glutenins: a new tool for the separation of high and low molecular weight subunits. Cereal Chem
71(3): 238-242
Nagamine T, Kai Y, Takayama T, Yanagisawa T, Taya S (2000) Allelic variations of glutenin subunit loci Glu-1 and Glu-3 in southern Japanese wheat and their effects on dough and gluten properties. J Cereal Sci 32:129-135
Nieto-Taladriz MT, Ruiz M, Martinez MC, Vazquez JF, Carrillo JM (1997) Variation and classification of B-low-molecular-weight glutenin subunit alleles in durum wheat. Theor Appl Genet 95:1155-1160
O'Farrel PH (1975) High resolution two-dimensional electrophoresis of proteins. J Biol Chem 250:4007-4021
Okita TW (1984) Identification and DNA sequence analysis of a γ-gliadin cDNA plasmid from winter wheat. Plant Mol Biol 3:325-332
Okita TW, Cheesbrough V, Reeves CD (1985) Evolution and heterogeneity of the α/β-type and γ-type gliadin DNA sequences. J Biol Chem 260:8203-8213
Payne PI, Corfield KG (1979) Subunit composition of wheat glutenin proteins, isolated by gel filtration in a dissociating medium. Planta 145:83-88
Payne PI, Law CN, Mudd EE (1980) Control by homoeologous group 1 chromosomes of the high-molecular-weight subunits of glutenin,a major protein of wheat endosperm. Theor Appl Genet 58:113-120
Payne PI, Holt LM, Lawrence GJ, Law CN (1982) The genetics of gliadin and glutenin, the major storage proteins of the wheat endosperm. Qual Plant Foods Hum Nutr 31: 229-241
Payne PI, Lawrence GJ (1983) Catalogue of alleles for the complex gene loci, Glu-A1,Glu-B1, and Glu-D1 which code for high-molecular-weight subunits of glutenin in hexaploid wheat. Cereal Research Communications 11(1): 29-35
Payne PI, Jackson EA, Holt LM, Law CN (1984a) Genetic linkage between endosperm storage protein genes on each of the short arms of chromosomes 1A and 1B in wheat. Theor Appl Genet 67:235-243
Payne PI, Jackson EA,Holt LM (1984b) The association between γ-gliadin 45 and gluten strength in durum wheat varieties:a direct causal effect or the result of genetic linkage? Journal of Cereal Science 2:73-81
Payne PI, Roberts MS, Holt LM (1986) Location of genes controlling the D group of LMW glutenin subunits on chromosome 1D of bread wheat. Genet Res Camb 47:175-179
Payne PI (1987) Genetics of wheat storage proteins and the effect of allelic variation on bread-making quality. Plant Physiol 38:141-153
Pfluger LA, Martin LM, Alvarez JB (2001) Variation in the HMW and LMW glutenin subunits from Spanish accessions of emmer wheat (Triticum turgidum ssp. dicoccum Schrank. Theor Appl Genet 102:767-772
Pitts EG., Rafalski JA, Hedgcoth C (1988) Nucleotide sequence and encoded amino acid sequence of a genomic gene region for a low molecular weight glutenin. Nucleic Acids Res 16:11376
Pogna N, Lafiandra D, Feillet P, Autran JC (1988) Evidence for a direct causal effect of low molecular weight glutenin subunits on gluten viscoelasticity in durum wheats. Journal of Cereal Science 7:211-214
Redaelli R, Morel MH, Autran JC, Pogna NE (1995) Genetic analysis of low Mr glutenin subunits fractionated by two-dimensional electrophoresis (A-PAGE× SDS-PAGE). Journal of Cereal Science 21: 5-13
Rodriguez-Quijano M, Nieto-Taladriz MT, Carrillo JM (1997) Variation in B-LMW glutenin subunits in Einkorn wheats. Genetic Resources and Crop Evolution 44: 539-543
Shewry PR, Halford NG, Tatham AS (1989) The high molecular weight subunits of wheat, barley and rye: genetics, molecular biology, chemistry and role in wheat gluten structure and functionality. In: Oxford Surveys of Plant and Molecular Cell Biology, Vol.6, Oxford University Press, London:163-219
Singh NK, Shepherd KW (1988) Linkage mapping of the genes controlling endosperm proteins in wheat. 1.Genes on the short arms of group 1 chromosomes. Theor Appl Genet 75:628-641
Singh NK, Shepherd KW (1991) Rapid communication a simplified SDS-PAGE procedure for separating LMW subunits of glutenin. J Cereal Sci 14:203-208
Sissons MJ, Bekes F, Skerritt JH (1998) Isolation and Functionality testing of low molecular weight glutenin subunits. Cereal Chem 75(1): 30-36
Sreeramulu G, Singh NK (1997) Genetic and biochemical characterization of novel low molecular weight glutenin subunits in wheat (Triticum aestivum L.). Genome 40:41-48
Tao HP, Kasarda DD (1989) Two-dimensional gel mapping and N-terminal sequencing of LMW-glutenin subunits. Journal of Experimental Botany 40(218): 1015-1020
Tosi P, D'Ovidio R, Napier JA, Bekes F, Shewry PR (2004) Expression of epitope-tagged LMW glutenin subunits in the starchy endosperm of transgenic wheat and their incorporation into glutenin polymers. Theor Appl Genet 108: 468-476
Van Campenhout S, Stappen JV, Sagi L, Volckaert G (1995) Locus specific primers for LMW glutenin genes on each of the group 1 chromosomes of hexaploid wheat. Theor Appl Genet 91: 313-319
Vensel WH, Tarr GE, Kasarda DD (1995) C-terminal and internal sequences of a low molecular weight (LMW-s) type of glutenin subunit. Cereal Chem 72(4): 356-359
Wrigley CW (1970) Protein mapping by combined gel electrofocusing and electrophoresis: application to the study of genotypic variations in wheat gliadins. Biochem Genet 4:509-516
Wrigley CW (1996) Giant proteins with flour power. Nature 381:738-739
Xu SS, Khan K, Klindworth DL, Faris JD (2004) Chromosomal location of genes for novel glutenin subunits and gliadins in wild emmer wheat (Triticum turgidum L.var.dicoccoides). Theor Appl Genet 108:1221-1228
Yan Y, Prodanovic S, Mladenov N, Milovanovic M (1999a) Genetic control of low molecular weight glutenin subunits in wheat by A-PAGE analysis. Cereal Research Communications 27:251-257
Yan Y, Surlan-Momirovic G,. Prodanovic S, Zoric D, and Liu G. (1999b) Capillary zone electrophoresis analysis of gliadin proteins from Chinese and Yugoslav winter wheat cultivars. Euphytica, 105:197-204
Yan Y, Hsam SLK, Yu JZ, Jiang Y, Zeller FJ (2003a) Genetic polymorphisms at Gli-D~t gliadin loci in Aegilops tauschii as revealed by acid polyacrylamide gel and capillary electrophoresis. Plant Breed 122:120-124
Yan Y, Hsam SLK, Yu JZ, Jiang Y, Zeller FJ (2003b) Allelic variation of the HMW glutenin subunits in Aegilops tauschii accessions detected by Sodium Dodecyl Sulphate (SDS-PAGE), Acid Polyacrylamide Gel (A-PAGE) and Capillary Electrophoresis. Euphytica 130:377-385
Yan Y, Yu JZ, Jiang Y, Hu Y, Cai M, Hsam SLK, Zeller FJ (2003c) Capillary electrophoresis separation of high molecular weight glutenin subunits in bread wheat (Triticum aestivum L.) and related species with phosphate-based buffers. Electrophoresis 24:1429-1436
Yan Y, Hsam SLK, Yu JZ, Jiang Y, Ohtsuka I, Zeller FJ (2003d) HMW and LMW glutenin alleles among putative tetraploid and hexaploid European spelt wheat (Triticum spelta L.) progenitors. Theor Appl Genet 107:1321-1330
Yan Y, Zheng J, Xiao Y, Yu J, Hu Y, Cai M, Li, Y, Hsam SLK and Zeller FJ (2004) Identification and molecular characterization of a novel y-type Glu-1D~t1 glutenin gene of Aegilops tauschii. Theor Appl Genet 108:1349-1358
Yates Ⅲ JR (1998) Special feature: tutorial —Mass spectrometry and the age of the proteome. Journal of Mass Spectrometry 33: 1-19
郭尧君主编(1999) 蛋白质电泳实验技术，科学出版社，北京：54-68
J．萨母布鲁克，E．F．弗里奇，T．曼尼阿蒂斯(1998)分子克隆实验指南(第二版)．科学出版社，北京：19-22，55-56，880—886
姜怡(2003)面包小麦及其近源种HMW谷蛋白亚基鉴定及其编码基因的分子克隆，首都师范大学硕士论文
李保云，解超杰，尤明山，毛善锋，刘广田(2002) 野生二粒小麦高分子量谷蛋白亚基的多态性分析．麦类作物学报22(3)：21—24
汪家政，范明主编(2000) 蛋白质技术手册，科学出版社，北京：
王罡，季静，胡含(1995)小麦种子贮藏蛋白的遗传学研究进展．遗传17(1)：45—48
夏其昌主编(1997) 蛋白质化学研究技术与进展．科学出版社，北京：99—107
晏月明，刘广田，Prodanovic S，Mominovic G S，Zorie D，Perovie D(1998)小麦谷蛋白亚基的凝胶电泳分离及其品种鉴定．中国粮油学报13(6)：1—5
余建中，晏月明，蔡民华，胡英考 (2002) 毛细管电泳技术及其在作物贮藏蛋白研究中的应用．麦类作物学报22(3)：78—81
张发云，晏月明，胡英考，胡赞民，周奕华(2002) 小麦HMW谷蛋白亚基基因克隆研究进展．中国生物工程杂志22(6)：33-40
Anderson OD, Green FC, Vip RE, Halford NG, Shewry PR, Malpica-Romero JM (1989) Nucleotide sequences of the two high-molecular-weight glutenin genes from the D-genome of a hexaploid bread wheat, Tritivum aestivum L. cv Cheyenne. Nucleic Acids Res 17:461-462
Bartels D, Thompson RD (1983) The characterization of cDNA clones coding for wheat storage proteins. Nucleic Acids Res 11: 2961-2977
Bean SR, Lookhart GL (2001a) Recent developments in high-performance capillary electrophoresis of cereal proteins. Electrophoresis 22:1503-1509
Bean SR, Lookhart GL (2001b) High-performance capillary electrophoresis of meat, dairy, and cereal proteins. Electrophoresis 22:4207-4215
Benmoussa M, Vezina LP, Page M, Yelle S,Laberge S (2000) Genetic polymorphism in low-molecular-weight glutenin genes from Triticum aestivum, variety Chinese Spring. Theor Appl Genet 100:789-793
Bietz JA, Shepherd KW, Wall JS (1975) Single kernel analysis of glutenin:use in wheat genetics and breeding. Cereal Chem 52:513-532
Bietz JA, Schmalzried E (1992) Capillary electrophoresis of wheat proteins: optimization and use for varietal identification. Cereal Foods World 37:555-560
Brown JWS,Flavell RB (1981a) Fractionation of wheat gliadin and glutenin subunits by two-dimensional electrophoresis and the role of group 6 and group 2 chromosomes in gliadin synthesis. Theor Appl Genet 59:349-359
Brown JWS, Law CN, Worland AJ, Flavell RB (1981b) Genetic variation in wheat endosperm proteins:an analysis by two-dimensional electrophoresis using intervarietal chromosomal substitution lines. Theor Appl Genet 59:361-371
Bunce NAC, White RP, Shewry PR (1985) Variation in estimates of molecular weight of cereal prolamins by SDS-PAGE. Journal of Cereal Science 3:131-142
Caballero L,Martin LM,Alvarez JB (2004) Genetic Variability of the low-molecular-weight glutenin subunits in spelt wheat (Triticum aestivum ssp.spelta L.em Thell.). Theor Appl Genet 108:914-919
Cassidy BG, Dvorak J (1991) Molecular characterization of a low-molecular-weight glutenin cDNA clone from Triticum durum. Theor Appl Genet 81:653-660
Cassidy BG, Dvorak J, Anderson OD (1998) The wheat low-molecular-weight glutenin genes: characterization of six new genes and progress in understanding gene family structure. Theor Appl.Genet 96:743-750
Ciaffi M, Dominici L, Lafiandra D, Porceddu E (1992) Seed storage proteins of wild wheat progenitors and their relationships with technological properties Hereditas 116:315-322
Ciaffi M,Lafiandra D, Porceddu E, Benedettelli S (1993) Storage-protein variation in wild emmer wheat (Triticum turgidum ssp. dicoccoides) from Jordan and Turkey. Ⅰ. Electrophoretie characterization of genotypes. Theor Appl Genet 86:474-480
Ciaffi M, Lee YK, Tamas L, Gupta R, Skerritt J, Appels R (1999) The low-molecular-weight glutenin subunit proteins of primitive wheats. Ⅲ. The genes from D-genome species. Theor Appl Genet 98:135-148
Cloutier S, Rampitsch C, Penner GA, Lukow OM (2001) Cloning and expression of a LMW-i glutenin gene. Journal of Cereal Science 33: 143-154
Colot V, Bartels D, Thompson R, Flavell R (1989) Molecular characterization of an active wheat LMW glutenin gene and its relation to other wheat and barley prolamin genes. Mol Gen Genet 216:81-90
Dachkevitch T, Redaelli R, Biancardi AM, Metakovsky EV, Pogna NE (1993) Genetics of gliadins coded by the group-1 chromosomes in the high-quality bread wheat cultivar Neepawa. Theor Appl Genet 86:389-399
D'Ovidio R,Tanzarella OA, Porceddu E (1992) Nucleotide sequence of a low-molecular-weight glutenin from Triticum durum. Plant Mol Biol 18:781-784
D'Ovidio R, Simeone M, Masci S, Porceddu E (1997) Molecular characterization of a LMW-GS gene located on chromosome 1B and the development of primers specific for the Glu-B3 complex locus in durum wheat. Theor Appl Genet
95:1119-1126
D'Ovidio R, Marchitelli C, Ercoli CL, Porceddu E (1999) Sequence similarity between allelic Glu-B3 genes related to quality properties of durum wheat. Theor Appl Genet 98:455-461
Draper SR (1987) ISTA Variety Committee Report of the working group for Biochemical Tests for Cultivar Identification 1983-1986. Seed Sci. & Technol. 15: 431-434
Galili G, Feldman M (1983) Genetic control of endosperm proteins in wheat. 2. Variation of HMW glutenin and gliadin subunits of Triticum aestivum. Theor Appl Genet 66:77-86
Galili G, Felsenburg T, Levy AA, Altschuler Y, Feldman M (1988) Inactivity of high-molecular-weight glutenin genes in wild diploid and tetraploid wheats. In: Proceedings of the 7th international wheat genetics symposium: 81-87. Eds. T. E. Miller and R. M. D. Koebner. Cambridge, UK: Institute of Plant Science Research.
Gianibelli MC, Larroque OR, Maeritchie F, Wrigley CW (2001) Biochemical, genetic,and molecular characterization of wheat glutenin and its component subunits. Cereal Chem 78(6): 635-646
Gianibelli MC, Masci S, Larroque R, Lafiandra D (2002a) Biochemical characterization of a novel polymeric protein subunit from bread wheat (Triticum aestivum L.). Journal of Cereal Science 35:265-276
Gianibelli MC, Wrigley CW, Macritchie F (2002b) Polymorphism of low M_r glutenin subunits in Triticum tauschii. Journal of Cereal Science 35:277-286
Gianibelli MC, Solomon RG (2003) A Novel y-type High Mr Glutenin Subunit (12.4~t) Present in Triticum tauschii. Journal of Cereal Science 37:253-256
Gupta RB, Singh NK, Shepherd KW (1989) The cumulative effect of allelic variation in LMW and HMW glutenin subunits on dough properties in the progeny of two bread wheats. Theor Appl Genet 77:57-64
Gupta RB, Shepherd KW (1990) Two-step one-dimensional SDS-PAGE analysis of LMW subunits of glutenin. Theor Appl Genet 80:65-74
Gupta RB, Macritchie F (1991) Rapid communication a rapid one-step one-dimensional SDS-PAGE procedure for analysis of subunit composition of glutenin in wheat. Journal of Cereal Science 14:105-109
Gupta RB, Macritchie F (1994) Allelic variation at glutenin subunit and gliadin loci, Glu-1, Glu-3 and Gli-1 of common wheats. Ⅱ. Biochemical basis of the allelic effects on dough properties. Journal of Cereal Science 19:19-29
Gupta RB, Popineau Y, Lefebvre J, Cornec M, Lawrence GJ, MacRitchie F (1995) Biochemical basis of flour properties in bread wheats.Ⅱ.Changes in polymeric
protein formation and dough/gluten properties associated with the loss of low M_r or high M_r glutenin subunits. Journal of Cereal Science 21: 103-116
Halford NG, Forde J, Shewry PR, Kreis M (1989) Functional analysis of the upstream regions of a silent and an expressed member of a family of wheat seed protein genes in transgenic tobacco. Plant Sci 62:207-216
Herbert Wieser (2000) Comparative investigations of gluten proteins from different wheat species. 1. Qualitative and quantitative composition of gluten protein types. Eur Food Res Technol 211: 262-268
Ikeda TM, Nagamine T, Fukupka H, Yano H (2002) Identification of new low-molecular-weight glutenin subunit genes in wheat. Theor Appl Genet 104: 680-687
Jackson EA, Holt LM,Payne PI (1983) Characterization of high-molecular-weight gliadin and low-molecular-weight subunits of wheat endosperm by two-dimensional eleetrophoresis and the chromosomal location of their controlling genes. Theor Appl Genet 66:29-37
Jackson EA, Morel MH, Strohm TS, Branlard G; Metakovsky EV, Redaelli R (1996) Proposal for combining the classification systems of alleles of Gli-1 and Glu-3 loci in bread wheat (Triticum aestivum L.). J Genet & Breed 50:321-336
Kasarda DD (1989) Glutenin structure in relation to wheat quality In: Pomeranz Y(ed) wheat is unique. AACC, St Paul, Minnesota: 299-302
Kolster P, Krechting CF, Gelder WMJ van (1988) Variation in high molecular weight glutenin subunits of Triticura aestivum and T.turgidum ssp.dicoccoides.Euphytica S: 141-145
Klose J (1975) Protein mapping by combined isoelectric focusing and electrophoresis of mouse tissues. A novel approach to testing for induced point mutations in mammals. Human Genetics 26:231-243
Lee YK, Bekes F, Gupta R, Appels R, Morell MK (1999a) The low-molecular-weight glutenin subunit proteins of primitive wheats.Ⅰ.Variation in A-genome species. Theor Appl Genet 98:119-125
Lee YK, Ciaffi M, Appels R, Morell MK (1999b) The low-molecular-weight glutenin subunit proteins of primitive wheats. Ⅱ. The genes from A-genome species. Theor Appl Genet 98:126-134
Lee YK, Bekes F, Gras P, Ciaffi M, Morell MK, Appels R (1999c) The low-molecular-weight glutenin subunit proteins of primitive wheats.IV.Functional properties of products from individual genes. Theor Appl Genet 98:149-155
Lew EJL, Kuzmicky DD, Kasarda DD (1992) Characterization of low molecular weight glutenin subunits by reversed-phase high-performance liquid chromatography, sodium dodecyl sulfate-polyacrylamide gel electrophoresis, and
N-terminal amino acid sequencing. Cereal Chem 69(5): 508-515
Liu CY (1995) Research note identification of a new low Mr glutenin subunit locus on chromosome 1B of durum wheat. Journal of Cereal Science 21: 209-213
Luo C, Griffin WB, Branlard G, Mcneil DL (2001) Comparison of low-and high molecular-weight wheat glutenin allele effects on flour quality. Theor Appl Genet 102:1088-1098
Mackie AM, Sharp P J, Lagudah ES (1996) The Nucleotide and Derived Amino Acid Sequence of a HMW Glutenin Gene from Triticum tauschii and Comparison with those from the D Genome of Bread Wheat. Journal of Cereal Science 24:73-78
Masci S, Lafiandra D, Porceddu E, Lew EJL, Tao HP, Kasarda DD (1993) D-glutenin subunits: N-terminal sequences and evidence for the presence of cystein. Cereal Chem 70(5): 581-585
Masci S, Lew EJL,Lafiandra D, Porceddu E, Kasarda DD (1995) Characterization of low molecular weight glutenin subunits in durum wheat by reversed-phase high-performance liquid chromatography and N-terminal sequencing. Cereal Chem 72(1): 100-104
Masci S, D'Ovidio R, Lafiandra D, Kasarda DD (1998) Characterization of a low-molecular-weight glutenin subunit gene from bread wheat and the corresponding protein that represents a major subunit of the glutenin polymer. Plant Physiol 118:1147-1158
Masci S, D'Ovidio R, Lafiandra D, Kasarda DD (2000) A 1B-coded lowmolecular-weight glutenin subunits associated with quality in durum wheat shows strong similarity to a subunit present in some bread wheat cultivars. Theor Appl Genet 100:396-400
Masci S, Rovelli L,Kasarda DD, Vensel WH (2002) Characterisation and chromosomal localization of C-type low-molecular-weight glutenin subunits in the bread wheat cultivar Chinese Spring. Theor Appl Genet 104:422-428
Mcdonald MB, Elliot LJ, Sweeney PM (1994) Research note DNA extraction from dry seeds for RAPD analyses in varietal identification studies. Seed Sci & Technol 22:171-176
Mecham DK, Kasarda DD, Qualset CO (1978) Genetic aspects of wheat gliadin proteins. Biochem Genet 16:831-853
Metakovsky EV, Branlard G, Chernakov VM, Upelniek VP, Redaelli R, Pogna NE (1997) Recombination mapping of some chromosome 1A-,1B-,1D-and 6B-controlled gliadins and low-molecular-weight glutenin subunits in common wheat. Theor Appl Genet 94:788-795
Morel MH (1994) Acid-polyacrylamide gel electrophoresis of wheat glutenins: a new tool for the separation of high and low molecular weight subunits. Cereal Chem
71(3): 238-242
Nagamine T, Kai Y, Takayama T, Yanagisawa T, Taya S (2000) Allelic variations of glutenin subunit loci Glu-1 and Glu-3 in southern Japanese wheat and their effects on dough and gluten properties. J Cereal Sci 32:129-135
Nieto-Taladriz MT, Ruiz M, Martinez MC, Vazquez JF, Carrillo JM (1997) Variation and classification of B-low-molecular-weight glutenin subunit alleles in durum wheat. Theor Appl Genet 95:1155-1160
O'Farrel PH (1975) High resolution two-dimensional electrophoresis of proteins. J Biol Chem 250:4007-4021
Okita TW (1984) Identification and DNA sequence analysis of a γ-gliadin cDNA plasmid from winter wheat. Plant Mol Biol 3:325-332
Okita TW, Cheesbrough V, Reeves CD (1985) Evolution and heterogeneity of the α/β-type and γ-type gliadin DNA sequences. J Biol Chem 260:8203-8213
Payne PI, Corfield KG (1979) Subunit composition of wheat glutenin proteins, isolated by gel filtration in a dissociating medium. Planta 145:83-88
Payne PI, Law CN, Mudd EE (1980) Control by homoeologous group 1 chromosomes of the high-molecular-weight subunits of glutenin,a major protein of wheat endosperm. Theor Appl Genet 58:113-120
Payne PI, Holt LM, Lawrence GJ, Law CN (1982) The genetics of gliadin and glutenin, the major storage proteins of the wheat endosperm. Qual Plant Foods Hum Nutr 31: 229-241
Payne PI, Lawrence GJ (1983) Catalogue of alleles for the complex gene loci, Glu-A1,Glu-B1, and Glu-D1 which code for high-molecular-weight subunits of glutenin in hexaploid wheat. Cereal Research Communications 11(1): 29-35
Payne PI, Jackson EA, Holt LM, Law CN (1984a) Genetic linkage between endosperm storage protein genes on each of the short arms of chromosomes 1A and 1B in wheat. Theor Appl Genet 67:235-243
Payne PI, Jackson EA,Holt LM (1984b) The association between γ-gliadin 45 and gluten strength in durum wheat varieties:a direct causal effect or the result of genetic linkage? Journal of Cereal Science 2:73-81
Payne PI, Roberts MS, Holt LM (1986) Location of genes controlling the D group of LMW glutenin subunits on chromosome 1D of bread wheat. Genet Res Camb 47:175-179
Payne PI (1987) Genetics of wheat storage proteins and the effect of allelic variation on bread-making quality. Plant Physiol 38:141-153
Pfluger LA, Martin LM, Alvarez JB (2001) Variation in the HMW and LMW glutenin subunits from Spanish accessions of emmer wheat (Triticum turgidum ssp. dicoccum Schrank. Theor Appl Genet 102:767-772
Pitts EG., Rafalski JA, Hedgcoth C (1988) Nucleotide sequence and encoded amino acid sequence of a genomic gene region for a low molecular weight glutenin. Nucleic Acids Res 16:11376
Pogna N, Lafiandra D, Feillet P, Autran JC (1988) Evidence for a direct causal effect of low molecular weight glutenin subunits on gluten viscoelasticity in durum wheats. Journal of Cereal Science 7:211-214
Redaelli R, Morel MH, Autran JC, Pogna NE (1995) Genetic analysis of low Mr glutenin subunits fractionated by two-dimensional electrophoresis (A-PAGE× SDS-PAGE). Journal of Cereal Science 21: 5-13
Rodriguez-Quijano M, Nieto-Taladriz MT, Carrillo JM (1997) Variation in B-LMW glutenin subunits in Einkorn wheats. Genetic Resources and Crop Evolution 44: 539-543
Shewry PR, Halford NG, Tatham AS (1989) The high molecular weight subunits of wheat, barley and rye: genetics, molecular biology, chemistry and role in wheat gluten structure and functionality. In: Oxford Surveys of Plant and Molecular Cell Biology, Vol.6, Oxford University Press, London:163-219
Singh NK, Shepherd KW (1988) Linkage mapping of the genes controlling endosperm proteins in wheat. 1.Genes on the short arms of group 1 chromosomes. Theor Appl Genet 75:628-641
Singh NK, Shepherd KW (1991) Rapid communication a simplified SDS-PAGE procedure for separating LMW subunits of glutenin. J Cereal Sci 14:203-208
Sissons MJ, Bekes F, Skerritt JH (1998) Isolation and Functionality testing of low molecular weight glutenin subunits. Cereal Chem 75(1): 30-36
Sreeramulu G, Singh NK (1997) Genetic and biochemical characterization of novel low molecular weight glutenin subunits in wheat (Triticum aestivum L.). Genome 40:41-48
Tao HP, Kasarda DD (1989) Two-dimensional gel mapping and N-terminal sequencing of LMW-glutenin subunits. Journal of Experimental Botany 40(218): 1015-1020
Tosi P, D'Ovidio R, Napier JA, Bekes F, Shewry PR (2004) Expression of epitope-tagged LMW glutenin subunits in the starchy endosperm of transgenic wheat and their incorporation into glutenin polymers. Theor Appl Genet 108: 468-476
Van Campenhout S, Stappen JV, Sagi L, Volckaert G (1995) Locus specific primers for LMW glutenin genes on each of the group 1 chromosomes of hexaploid wheat. Theor Appl Genet 91: 313-319
Vensel WH, Tarr GE, Kasarda DD (1995) C-terminal and internal sequences of a low molecular weight (LMW-s) type of glutenin subunit. Cereal Chem 72(4): 356-359
Wrigley CW (1970) Protein mapping by combined gel electrofocusing and electrophoresis: application to the study of genotypic variations in wheat gliadins. Biochem Genet 4:509-516
Wrigley CW (1996) Giant proteins with flour power. Nature 381:738-739
Xu SS, Khan K, Klindworth DL, Faris JD (2004) Chromosomal location of genes for novel glutenin subunits and gliadins in wild emmer wheat (Triticum turgidum L.var.dicoccoides). Theor Appl Genet 108:1221-1228
Yan Y, Prodanovic S, Mladenov N, Milovanovic M (1999a) Genetic control of low molecular weight glutenin subunits in wheat by A-PAGE analysis. Cereal Research Communications 27:251-257
Yan Y, Surlan-Momirovic G,. Prodanovic S, Zoric D, and Liu G. (1999b) Capillary zone electrophoresis analysis of gliadin proteins from Chinese and Yugoslav winter wheat cultivars. Euphytica, 105:197-204
Yan Y, Hsam SLK, Yu JZ, Jiang Y, Zeller FJ (2003a) Genetic polymorphisms at Gli-D~t gliadin loci in Aegilops tauschii as revealed by acid polyacrylamide gel and capillary electrophoresis. Plant Breed 122:120-124
Yan Y, Hsam SLK, Yu JZ, Jiang Y, Zeller FJ (2003b) Allelic variation of the HMW glutenin subunits in Aegilops tauschii accessions detected by Sodium Dodecyl Sulphate (SDS-PAGE), Acid Polyacrylamide Gel (A-PAGE) and Capillary Electrophoresis. Euphytica 130:377-385
Yan Y, Yu JZ, Jiang Y, Hu Y, Cai M, Hsam SLK, Zeller FJ (2003c) Capillary electrophoresis separation of high molecular weight glutenin subunits in bread wheat (Triticum aestivum L.) and related species with phosphate-based buffers. Electrophoresis 24:1429-1436
Yan Y, Hsam SLK, Yu JZ, Jiang Y, Ohtsuka I, Zeller FJ (2003d) HMW and LMW glutenin alleles among putative tetraploid and hexaploid European spelt wheat (Triticum spelta L.) progenitors. Theor Appl Genet 107:1321-1330
Yan Y, Zheng J, Xiao Y, Yu J, Hu Y, Cai M, Li, Y, Hsam SLK and Zeller FJ (2004) Identification and molecular characterization of a novel y-type Glu-1D~t1 glutenin gene of Aegilops tauschii. Theor Appl Genet 108:1349-1358
Yates Ⅲ JR (1998) Special feature: tutorial —Mass spectrometry and the age of the proteome. Journal of Mass Spectrometry 33: 1-19
© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号 地址：北京市海淀区学院路29号 邮编：100083 电话：办公室：(+86 10)66554848；文献借阅、咨询服务、科技查新：66554700 |