离子束诱导普通小麦小偃81 HMW-GS 1Ax1缺失突变家系突变机制研究
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摘要
河南省离子束生物工程重点实验室以低能N+注入普通小麦小偃81干种子,诱变处理后经检测共发现1Axl,1Bxl4+1Byl5,1Dyl2和1Dx2+1Dyl2四种类型的高分子量麦谷蛋白亚基缺失突变体,经过四代选育获得遗传稳定的高分子量麦谷蛋白1Axl亚基缺失突变群体。本研究对1Axl亚基缺失M3代群体部分农艺性状进行了观察并对M4代亚基缺失性状的遗传稳定性以及该缺失家系的突变机理进行了分析。
1、高分子量麦谷蛋白1Axl亚基缺失M3代群体部分农艺性状调查
高分子量麦谷蛋白1Axl亚基缺失M3代群体共有25个株系,大田观察发现各株系内部各个性状稳定一致。M3代盛花期及成熟期均比野生型早,在株高、穗长、分蘖、旗叶叶面积、株型、千粒重、穗粒重及籽粒性状等方面与野生型存在一定的差异,而在叶毛、叶鞘、叶耳、叶舌、茎叶蜡质、叶型、穗型、穗色、芒性等方面基本都与野生型一致。
2、高分子量麦谷蛋白lAxl亚基缺失M4代群体SDS-PAGE电泳分析
每个单株随机取5粒种子做SDS-PAGE电泳,对各单株种子高分子量麦谷蛋白亚基组成进行分析,结果表明高分子量麦谷蛋白1 Axl亚基缺失突变家系遗传稳定,不存在性状分离。
3.高分子量谷蛋白亚基1Axl沉默基因的分离与测序
为了研究离子束诱导普通小麦小偃81 HMW-GS 1Ax1缺失家系突变机制,我们根据已知的Glu-1Ax1基因序列信息设计了一对引物对该突变家系Glu-1Ax1基因的编码区进行扩增,经克隆测序后与野生型序列相比较。4.普通小麦沉默的高分子量谷蛋白亚基1Axl的序列特征
分析了3个突变家系的1Axl亚基基因序列后发现,变异类型包括基因片段缺失、碱基的转换、颠换、缺失及插入,其中主要的变异类型为碱基转换,占总变异的70%以上,颠换约占总变异的20%,由我们的实验结果可以看出低能离子束有较广诱变谱。
5.普通小麦高分子量谷蛋白亚基1Axl基因沉默的分子机制
在3个突变家系的1Axl亚基基因序列中,由于基因突变,主要包括碱基置换和移码突变等,使得突变型1Axl基因序列中出现了提前终止密码子,因此突变型1Axl基因序列编码的蛋白提前终止,从而造成了1Axl亚基的缺失。
The Henan Province Key Laboratory of Ion Beam Bioengineering irradiated the triticum aestivum XiaoYan 81 dry seeds with low energy N+, and then we found four types of high molecular weight glutenin subunit deletion mutations, such as lAxl, 1Bx14+1By15, 1Dy12 and 1Dx2+lDyl2 subunit deletion, undergoes four generation of select and breeding, we obtained the high molecular weight glutenin subunit lAxl deletion mutation groups with genetic stability. In this study, the field character of the M3 of high molecular weight glutenin subunit lAxl deletion mutation groups has been carried on the observation, we also analyzed the genetic stability of M4 of the deletion mutation family their mutation mechanisms.
1. The agronomic traits of M3
There are 25 lines altogether in our High molecular weight glutenin subunits lAxl deletion mutation groups, and the agronomic traits are same as each other within a specific line. Both the full flowering time and the mature time of the M3 are earlier than the wild-type. There are some difference between the M3 and the wild-type in plant height, panicle length, tiller, flag leaf area, plant type, grain weight, grain weight per spike and the grain characteristics. And there are some traits of M3, such as leaf ears, leaf tongue, leaf wax, leaf type, spike color, which are basically consistent with the wild type.
2. SDS-PAGE electrophoresis analyze of M4
Five seeds per plant were taken randomly to do SDS-PAGE electrophoresis, the HMW-GSs composition analysis of every single plant indicated that the HMW-GS lAxl deletion mutation family are genetic stability, and there is no characters separation.
3. Isolation and sequencing of the silence HMW-GS 1Ax1 gene
In order to study the mutation mechanism of the XiaoYan 81 HMW-GS 1Ax1 deletion mutation family which induced by Ion Beam, we have designed a pair of primers according to the Glu-lAxl gene sequencing information to amplify the Glu-lAxl mutation gene coding region, then post-clone and sequencing and compared with the Glu-lAxl gene sequence of the wild-type.
4. sequence characteristics of the silence HMW-GS 1Ax1 gene
The central repeat region of the 1Ax1 subunit mutant gene has a high degree of variation, while the nucleotide variation of the 5'terminal and the 3'terminal non-repeat region is relatively small. The mutation types include gene fragment deletion, base conversion, transversions and deletion, and the main mutation type is base coversion which accounting foe more than 70% of the totle variance, then is base trabsversion which accounting for about 20% of the totle variance. Through the results of our experiment, it can be seen that low energy ion beam inducing mutagenesis have a wider spectrum.
5. The molecular mechanism of the HMW-GS 1Ax1 gene silencing
In these three mutation families, because of the mutations, there appeared several premature termination codons which allowing the protein encoded by the mutant gene early termination in the silencing HMW-GS 1Ax1 gene, resulting in the lack of 1Ax1 subunits.
1、高分子量麦谷蛋白1Axl亚基缺失M3代群体部分农艺性状调查
高分子量麦谷蛋白1Axl亚基缺失M3代群体共有25个株系,大田观察发现各株系内部各个性状稳定一致。M3代盛花期及成熟期均比野生型早,在株高、穗长、分蘖、旗叶叶面积、株型、千粒重、穗粒重及籽粒性状等方面与野生型存在一定的差异,而在叶毛、叶鞘、叶耳、叶舌、茎叶蜡质、叶型、穗型、穗色、芒性等方面基本都与野生型一致。
2、高分子量麦谷蛋白lAxl亚基缺失M4代群体SDS-PAGE电泳分析
每个单株随机取5粒种子做SDS-PAGE电泳,对各单株种子高分子量麦谷蛋白亚基组成进行分析,结果表明高分子量麦谷蛋白1 Axl亚基缺失突变家系遗传稳定,不存在性状分离。
3.高分子量谷蛋白亚基1Axl沉默基因的分离与测序
为了研究离子束诱导普通小麦小偃81 HMW-GS 1Ax1缺失家系突变机制,我们根据已知的Glu-1Ax1基因序列信息设计了一对引物对该突变家系Glu-1Ax1基因的编码区进行扩增,经克隆测序后与野生型序列相比较。4.普通小麦沉默的高分子量谷蛋白亚基1Axl的序列特征
分析了3个突变家系的1Axl亚基基因序列后发现,变异类型包括基因片段缺失、碱基的转换、颠换、缺失及插入,其中主要的变异类型为碱基转换,占总变异的70%以上,颠换约占总变异的20%,由我们的实验结果可以看出低能离子束有较广诱变谱。
5.普通小麦高分子量谷蛋白亚基1Axl基因沉默的分子机制
在3个突变家系的1Axl亚基基因序列中,由于基因突变,主要包括碱基置换和移码突变等,使得突变型1Axl基因序列中出现了提前终止密码子,因此突变型1Axl基因序列编码的蛋白提前终止,从而造成了1Axl亚基的缺失。
The Henan Province Key Laboratory of Ion Beam Bioengineering irradiated the triticum aestivum XiaoYan 81 dry seeds with low energy N+, and then we found four types of high molecular weight glutenin subunit deletion mutations, such as lAxl, 1Bx14+1By15, 1Dy12 and 1Dx2+lDyl2 subunit deletion, undergoes four generation of select and breeding, we obtained the high molecular weight glutenin subunit lAxl deletion mutation groups with genetic stability. In this study, the field character of the M3 of high molecular weight glutenin subunit lAxl deletion mutation groups has been carried on the observation, we also analyzed the genetic stability of M4 of the deletion mutation family their mutation mechanisms.
1. The agronomic traits of M3
There are 25 lines altogether in our High molecular weight glutenin subunits lAxl deletion mutation groups, and the agronomic traits are same as each other within a specific line. Both the full flowering time and the mature time of the M3 are earlier than the wild-type. There are some difference between the M3 and the wild-type in plant height, panicle length, tiller, flag leaf area, plant type, grain weight, grain weight per spike and the grain characteristics. And there are some traits of M3, such as leaf ears, leaf tongue, leaf wax, leaf type, spike color, which are basically consistent with the wild type.
2. SDS-PAGE electrophoresis analyze of M4
Five seeds per plant were taken randomly to do SDS-PAGE electrophoresis, the HMW-GSs composition analysis of every single plant indicated that the HMW-GS lAxl deletion mutation family are genetic stability, and there is no characters separation.
3. Isolation and sequencing of the silence HMW-GS 1Ax1 gene
In order to study the mutation mechanism of the XiaoYan 81 HMW-GS 1Ax1 deletion mutation family which induced by Ion Beam, we have designed a pair of primers according to the Glu-lAxl gene sequencing information to amplify the Glu-lAxl mutation gene coding region, then post-clone and sequencing and compared with the Glu-lAxl gene sequence of the wild-type.
4. sequence characteristics of the silence HMW-GS 1Ax1 gene
The central repeat region of the 1Ax1 subunit mutant gene has a high degree of variation, while the nucleotide variation of the 5'terminal and the 3'terminal non-repeat region is relatively small. The mutation types include gene fragment deletion, base conversion, transversions and deletion, and the main mutation type is base coversion which accounting foe more than 70% of the totle variance, then is base trabsversion which accounting for about 20% of the totle variance. Through the results of our experiment, it can be seen that low energy ion beam inducing mutagenesis have a wider spectrum.
5. The molecular mechanism of the HMW-GS 1Ax1 gene silencing
In these three mutation families, because of the mutations, there appeared several premature termination codons which allowing the protein encoded by the mutant gene early termination in the silencing HMW-GS 1Ax1 gene, resulting in the lack of 1Ax1 subunits.
引文
[1]刘广田,李保云.小麦品质遗传改良的目标和方法[M].北京：中国农大出版社,2003.
[2]Osborne TB. The proteins of the wheat kernel. Washington DC:Carnegie Institution of Washington,1907.
[3]Bietz JA, Wall JS. The effect of various extractants on the subunit composition and associations of wheat glutenin[J]. Cereal Chemistry.1975,52:145-155.
[4]Payne PI, Law CN, Mudd EE. Control by homoeologous group 1 chromosomes of the high-molecular-weight subunits of glutenin, a major protein of wheat endosperm[J]. Theoretical and Applied Genetics,1980,58:113-120.
[5]Lawrence G J, Shepherd KW. Inheritance of glutenin protein subunits of wheat[J]. Theoretical and Applied Genetics,1981,60:333-337.
[6]Payne PI, Lawrence GJ. Catalogue of alleles for the complex gene loci,Glu-A1,Glu-B1,and Glu-D1 which code for the high-molecular-weight subunits of glutenin in hexaploid wheat[J]. Cereal research communications,1983,11:29-35.
[7]赵和,卢少源,李宗智.小麦高分子量麦谷蛋白亚基遗传变异及其与品质和其它农艺性状关系的研究[J].作物学报,1994,20(1)：67-75.
[8]栗站稳,王学路,卢少源,等.小麦主要贮藏蛋白亚基组分在我国种质资源中的分布[J].北京农业科学,1995,13(6)：21-24.
[9]张学勇,庞斌双,游光霞,等.中国小麦品种资源Glu1位点组成概况及遗传多样性分析[J].中国农业科学,2002,35(11)：1302-1310.
[10]Xu SS, Khan K, Klindworth DL, et al. Chromosomal location of genes for novel glutenin subunits and gliadins in wild emmer wheat(Triticum turgidum L var Dicoccoides)[J]. Theoretical and Applied Genetics,2004,108:1221-1228.
[11]Forde J, Malpica JM, Halford NG,e t al. The Nucleotide Sequence of a HMW Glutenin Subunit Gene Located on Chromosome 1A of Wheat(Triticum aestivum L)[J]. Nucleic Acids Research,1985,13:6817-6832.
[12]Anderson OD, Green C, Yip PE, et al. Nucleotide Sequences of the High Molecular Weight Glutenin Genes from the D-Genome of Hexaploid Wheat,Triticum aestivum L.cv Cheyenne[J]. Nucleic Acids Research,1989,17:461-462.
[13]Halford NG, Forde J, Anderson OD. The Nucleotide and Deduced Amino Acid Sequences of an HMW Glutenin Subunit Gene from Chromosome 1B of Bread Wheat(Triticum aesfvum L.) and Comparison with Those of Genes from Chromosomes 1A and 1D[J]. Theoretical and Applied Genetics,1987,75:117-126.
[14]Shewry PR, Halford NG. Cereal seed proteins:structures,properties and role in grain utilization [J]. Journal of Experimental Botany,2002,53:947-958.
[15]Shewry PR, Tatham AS. Disulphide bonds in wheat gluten Proteins[J]. Journal of Cereal Science,1997,25:207-227.
[16]Kasarda DD, Nimmo CC, Bernardin JE. Structural aspects and genetic relationships of gliadins [M]. In Coelic Disease:Proc.2nd Int,1974,25-36.
[17]Ewart JAD. Glutenin structure[J]. Journal of the Science of Food and Agriculture,1978, 30(5):482-492.
[18]Gilbert SM, Wellner N, Belton PS, et al. Expression and characterization of a highly repetitive peptide derived from a Wheat seed storage protein[J]. Biochimica et Biophysica Acta,2000,1479:135-146.
[19]Belton PS. On the elasticity of wheat gluten[J]. Journal of Cereal Science,1999,29: 103-109.
[20]Tilley KA, Benjamin RE, Bagorogoza KE, et al. Tyrosine cross-links molecular basis of gluten structure and function[J]. Journal of Agricultural and Food Chemistry,2001,49: 2627-2632.
[21]Payne PI. Genetics of wheat storage Proteins and the effect of allelic variationon breadmaking quality[J]. Annual Review of Plant Physiology,1987,38:141-153.
[22]毛沛.高分子量麦谷蛋白亚基组成及其与面包烘烤品质关系的研究[D].保定：河北农业大学.1992.
[23]Ammar K, Warren E, Kronstad, et al. Morris Breadmaking quality of selected durum wheat geno-types and its relationship with high molecular weight glutenin subunits allelic variation and gluten proteinpolymeric composition[J]. Cereal Chemistry,2000,77(2):230-236.
[24]卫增泉.离子束生物效应及应用中值得研究的几个问题[J].激光生物学报,2006,15(4)：383-387.
[25]余增亮.离子束生物技术引论[M].安徽：安徽科学技术出版社,1998.
[26]王卫东,王燕,王雪青,等.离子注入对小麦当代形态生理影响的初步研究[J].激光生物学报,2003,12(5)：360-363.
[27]王卫东,王燕,王雪青,等.氮离子注入后拟南芥种子表皮的扫描电镜观察[J].电子显微学报,2004,23(2)：148-151.
[28]袁世斌,薛建明,王宇刚,等.碳离子束辐照拟南芥介导外源基因转移的研究[J].激光生物学报,2003,12(5)：326-331.
[29]余增亮,霍裕平.离子注入生物学研究述评[J].安徽农业大学学报,1994,21(3)：221-225.
[30]邵春林,许安,余增亮.离子注入生物分子的电荷交换效应[J].核技术,1997,20(2)：70-73.
[31]张冬梅,孙素琴,马志龙,等.低能离子束辐照氨基酸结构变化的傅里叶变换红外光谱法研究[J].核技术,2000,23(4)：213-215.
[32]梁剑平.重离子辐照对喹烯酮的影响和重离子放射增敏药物的开发[D].兰州：中国科学院近代物理研究所,2000.
[33]袁世斌.低能重离子束对生物分子的辐照效应研究[D].兰州：中国科学院近代物理研究所,2002.
[34]石怀斌,邵春林,王相勤,等.低能氮离子对腺嘌呤水溶液损伤作用的光谱分析[J].光谱学与光谱分析,2002,22(2)：278-280.
[35]谷运红,王卫东,秦广雍,等.低能N+离子注入胸腺嘧啶的XPS研究[J].光谱学与光谱分析,2007,2：400-403.
[36]徐家萍,刘明辉,汪泰初.离子束诱变桑品种与亲本的同工酶和RAPD比较分析[J].安徽农业大学学报,2002,29(3)：286-288.
[37]焦浈,秦广雍,霍裕平.低能铁离子注入烟草叶片的过氧化物酶同工酶和全蛋白分析[J].河南农业科学,2004,6：23-25.
[38]赵俊杰,黄中文,常胜合.N+离子注入对不同小麦品种同工酶的影响[J].河南农业科学,2006,8：25-27.
[39]张从宇,王敏,张子学,等.60Co-γ和Ar+处理对小麦的诱变效应及POD同工酶分析[J].激光生物学报,2008,17(6)：780-786.
[40]李文建,卫增泉,颉红梅,等.中能氮离子束辐照小麦种子产生自由基的研究[J].核技术,1995,18(7)：415-418.
[41]孔福全.重离子致DNA损伤及修复的研究[D].天津：河北工业大学,2005.
[42]颉红梅,郝冀方,卫增泉,等.重离子束辐照牧草的细胞学研究[J].激光生物学报,2003,12(5)：346-349.
[43]杨美英,武志海,崔秋华,等.粳稻氩离子(Ar+)束诱变育种的细胞学效应研究[J].吉林农业大学学报,2002,24(5)：35-38.
[44]王崇英,王如娟,杨汉民.低能N+重离子注入处理后小麦种胚内的DNA期外合成[J].辐射研究与辐射工艺学报,1994,22(2)：205-208.
[45]杨剑波,吴李君,李莉,等.离子束辐照M13mp18-DNA(RF1)诱导lacZ突变的序列分析[J].中国科学(B辑),1995,25(12)：1273-1278.
[46]常凤启,刘选明,李银心,等.低能N+辐照拟南芥诱导基因组DNA碱基变异分析[J].中国科学(C辑),2003,33(2)：117-124.
[47]吴丽芳,李红.离子束生物工程应用研究进展[J].物理,1999,28(12)：708-712.
[48]杨赞林,甘斌杰,余增亮.离子注入小麦诱变育种的回顾与展望[J].安徽农业科学,2002,30(5)：639-641,648.
[49]李兴林,王晓娟,李文建,等.重离子辐照春麦稳定后代品质和贮藏蛋白的变化[J].核农学报,2000,14(5)：311-314.
[50]颉红梅,王浩瀚,王菊芳,等.重离子束定点诱变育种初探[J].原子核物理评论,2001,18(3)：174-176.
[51]陈秀兰,柳学余.利用离子注入诱变创造小麦新种质[J].江苏农学院学报,1998,19(2)： 62-64.
[52]李兰真,秦广雍,霍裕平,等.离子注入在小麦诱变育种上的应用研究初报[J].河南农业大学学报,2001,35(1)：9-12.
[53]张金莲,胡钺,卫增泉,等.低能重离子注入春小麦甘种子生物效应的初步研究[J].甘肃农业科技,1995,2：2-4.
[54]张利华.N2激光和氮离子注入对小麦一些性状的影响[J].种子,1999,3：15-17.
[55]宣云,王会峰,黄群策,等.离子束生物技术在小麦改良中的应用[J].农业生物技术科学,2004,20(1)：32-34,76.
[56]Yu ZL, Yang JB, Wu YJ, et al. Transferring Gus gene into intact rice cells by low energy ion beam[J]. Nuclear Instruments and Methods in Physics Research,1993, B80:1328-1331.
[57]吴丽芳,李红,宋道军,等.低能氩离子束介导将绿色荧光蛋白基因导入小麦的研究[J].南京农业大学学报,2000,23(3)：17-20.
[58]聂利红,崔党群,闻捷,等.离子束介导大豆789转入普通小麦后代的麦谷蛋白和醇溶蛋白分析[J].麦类作物学报,2005,25(3)：125-127.
[59]焦浈.低能离子束介导大豆全DNA导入小麦后其蛋白质遗传特性研究[D].郑州：郑州大学,2003.
[60]吴丽芳,余增亮.离子注入法获得大豆-小麦分子远缘杂种及后代的变异分析[J].核农学报,2000,14(4)：206-211.
[61]姬生栋,李吉学,徐存拴,等.离子束介导大豆DNA小麦后代的蛋白质变异性分析[J].麦类作物学报,2001,21(3)：18-21.
[62]杨随庄,尚勋武,叶春雷.小麦高分子量谷蛋白亚基基因分子育种研究进展[J].麦类作物学报,2007,27(2)：358-363.
[63]Halford NG, Field JM, Blair H, et al. Analysis of HMW glutenin subunits encoded by chromosome 1A of bread wheat(Triticum aestivum L.) indicates quantitative effects on grain quality[J]. Theoretical and Applied Genetics,1992,83:373-378.
[64]张金锐,刘勇,林刚,等.小麦高分子量谷蛋白亚基序列及其结构与加工品质的关系[J].中国生物工程杂志,2006,26(8)：103-110.
[65]Shewry PR. The synthesis,processing and deposition of gluten proteins in the Developing wheat grain[J]. Cereal Foods World,1999,44:587-589
[66]Kohler P, Keck B, Muller S, et al. Disulphide bonds in wheat gluten[C]. In:Wheat Kernel Proteins,Molecular and functional aspects (Universita degli studi della Tuscia consiglio nazionale delle ricerche, ed). Viterbo, Italy,1995,45-53.
[67]王关林,方宏筠.植物基因工程原理与技术[M].北京：科学出版社,1998,33.
[68]Altpeter F, Vimla V, Vibha S, et al. Integration and expression of the high-molecular-weight glutenin subunit 1Ax1 gene into wheat[J]. Nature Biotechnology,1996,14:155-159.
[69]Vasil IK, Bean S, Zhao JM,et al. Evalution of baking properties and gluten composition of filed grown transgenic wheat lines expressing high molecular weight glutenin gene 1Ax1[J]. Journal of Plant Physiology,2001,158(4):521-528.
[70]Popineau Y, Deshayes G, Lefebvre J, et al. Prolamin aggregation,gluten viscoelasticity,and mixing properties of transgenic wheat lines expressing 1Ax and 1Dx high molecular weight glutenin subunit transgenes[J]. Journal of Agricultural and Food Chemistry,2001,49(1): 395-401
[71]Barro F, Rooks L, Bekes F, et al. Transformation of wheat with high molecular weight subunit genes results in improved functional properties[J]. Nature Biotechnology,1997,15: 1295-1299.
[72]Barro F, Barcelo P, Lazzeri PA, et al. Functional properties and agronomic performance of Transgenic Tritordeum expressing high molecular weight glutenin subunit genes 1Ax1 and 1Dx5[J]. Journal of Cereal Science,2003,37:65-70.
[73]Alvarez ML, Gomezm M, Carrillo JM,et al. Analysis of dough functionallity of flours from transgenic wheat[J]. Molecular Breeding,2001,8(1):103-108.
[74]M Rakszegi, G Pastori, HD Jones,et al. Technological quality of field grown transgenic lines of commercial wheat cultivars expressing the 1Ax1 glutenin subunit gene[J]. Journal of Cereal Science,2008,47:310-321.
[75]Field JM, Bhandari D, Bonet A, et al. Introgression of transgenes into a commercial cultivar confrms differential effects of HMW subunits 1Ax1 and 1Dx5 on gluten properties[J]. Journal of Cereal science,2008,48:457-463.
[76]Eckardt N A. RNA goes mobile [J]. Plant cell,2002,101:543-553.
[77]Margiotta B, Urbano M, Colaprico G, et al. Detection of y-type subunit at the Glu-A1 locus in some Swedish bread wheat lines [J]. Journal of Cereal Science,1996,23:203-211.
[78]He ZH, Pena RJ, Rajaram S. High molecular weight glutenin subunit composition of Chinese bread wheat [J]. Euphytica,1992,64:11-20.
[79]Morgunov AL, Rogers WJ, Sayers EG, et al. The high molecular weight glutenin composition of Soviet wheat varieties [J]. Euphytica,1990,51:41-52.
[80]Lawrence GJ. The high-molecular-weight glutenin subunit composition of Australian wheat cultivars [J]. Australian Journal of Agricultural Research,1986,37:123-125.
[81]Payne PI, Nightingale MA, Krattinger AF, et al. The relationship between HMW Glutenin Subunit composition and the bread-making quality of British grown wheat varieties [J]. Journal of the Science of Food and Agriculture,1987,40:51-65.
[82]Lukow OM, Payne PI, Tkachuk R.The HMW glutenin subunit composition of Canadian wheat cultivars and their association with bread-making quality [J]. Journal of the Science of Food and Agriculture,1989,46(4):451-460.
[83]Vallega V, Waines J G. High molecular weight glutenin subunit variation in Triticum turgidum var. dicoccum [J]. Theoretical and Applied Genetics,1987,74:706-710.
[84]董永梅,杨欣明,柴守诚,等.中国小麦代表性地方品种高分子量麦谷蛋白亚基组成分 析[J].麦类作物学报,2007,27(5)：820-824.
[85]刘悦,杨足君,李光蓉,等.四川南部和重庆地区小麦地方品种的高分子量谷蛋白亚基组成分析[J].麦类作物学报,2006,26(4)：47-50.
[86]杨恩年,张洁,杨武云,等.六倍体普通小麦高分子量谷蛋白亚基Glu-A1和Glu-B1共同缺失材料研究初报[J].西南农业学报,2007,20(2)：293-295.
[87]Lawrence GJ. Dough and baking quality of wheat line deficient in glutenin subunits controlled by the Glu-A1, Glu-B1 and Glu-D1 loci [J]. Journal of Cereal Science,1988,7: 109-112.
[88]Payne P I. Genetics of wheat storage proteins and the effect of allelic variation on breadmaking quality [J]. Annual Review of Plant Physiology and Plant Molecular Biology, 1987,38:141-153.
[89]Upelniek V, Novoselskaya A, Sutka J,et al. Genetic variation at storage protein-coding loci of common wheat (cv'Chinese Spring') induced by nitrosoethylurea and by the cultivation of immature embryos in vitro [J]. Theoretical and Applied Genetics,1995,90:372-379.
[90]Zhu YF, Li YW, Chen Y, et al. Generation and characterization of a high molecular weight glutenin 1Bx14 deficient mutant in common wheat[J]. Plant Breeding,2005,124:421-427.
[91]张怀渝,宋云,畅志坚,等.低能N+注入诱导小麦种子高分子量麦谷蛋白亚基和醇溶蛋白的变异[J].核农学报,2005,19(4)：245-250.
[92]Altpeter F, Vasil V, Srivastava V,et al. Integration and expression of the high molecular weight glutenin subunit lAxl gene into wheat [J]. Nature Biotechnology,1996,14: 1155-1159.
[93]Alvarez ML, Guelman S, Halford NG, et al. Silencing of HMW glutenins in transgenic wheat expressing extra HMW subunits[J]. Theoretical and Applied Genetics,2000,100:319-327.
[94]Uthayakumaran S, Lukow OM. Functional and multiple end-use characterisation of Canadian wheat using a reconstituted dough system [J]. Journal of the Science of Food and Agriculture, 2003,83:889-898.
[95]霍裕平,秦广雍,苏明杰,等.离子束诱变小麦遗传新规律的研究[C].第三次全国离子束生物工程学大会暨第一次国际研讨会论文集,2002.
[96]赵勇.低能重离子引起DNA链断裂和诱发基因突变谱的研究[D].武汉：武汉大学,2003.
[97]潘多.低能N+离子束诱变的机理研究：突变谱分析[D].哈尔滨：东北林业大学,2005.
[98]辛庆国.7Li离子束核反应生物诱变效应与作用机制研究[D].泰安：山东农业大学,2007.
[99]钱时权.合成PHA相关基因离子束诱变及其高产突变体分子机制研究[D].合肥：安徽农业大学,2007.
[100]李珂,张东正,赵瑾,等.重离子注入与生物体的相互作用及遗传诱变的分子机制[J].原子核物理评论,2007,24(2)：117-123.
[101]Gu YQ,Salse J, Coleman D, et al. Types and rates of sequence evolution at the high molecular weight glutenin locus in hexaploid wheat and its ancestral genomes[J]. Genetics, 2006,174:1493-1504.
[102]Harberd NP, Flavell RB, Thompson RD. Identification of a transposon like insertion in a Glu-1 allele of wheat[J]. Molecular Genetics and Genomics,1987,209:326-332.
[103]Bustos AD, Rubio P, Jouve N. Molecular characterisation of the inactive allele of the gene Glu-A1 and the development of a set of AS-PCR markers for HMW glutenins of wheat[J]. Theoretical and Applied Genetics,2000,100:1085-1094.
[104]Yang ZJ, Li GR, Liu C, et al. Molecular Characterization of a HMW Glutenin subunit allele providing evidence for silencing of x-type gene on Glu-B1 [J]. Acta Genetica Sinica,2006, 33 (10):929-936.
[105]Forde J, Malpica JM, Halford NG, et al. The nucleotide sequence of a HMW subunit gene located on chromosome 1A of wheat (Triticum eastivum L.)[J]. Nucleic Acids Research, 1985,13:6817-6832.
[106]Liu SW, Gao X, Xia GM. Characterizing HMW-GS alleles of decaploid Agropyron elongatum in relation to evolution and wheat breeding[J]. Theoretical and Applied Genetics, 2008,116:325-334.
[107]Wang JR, Yan ZH, Wei YM, et al. Characterization of high-molecular-weight glutenin subunit genes from Elytrigia elongata(Host) Nevski[J]. Plant Breeding,2006,125:89-95.
[108]Feng DS, Chen FG, Zhao SY, et al. High-molecular-weight glutenin subunit genes in decaploid Agropyron elongatum [J]. Acta Botanica Sinica,2004,46:489-496.
[109]Wan YF, Wang DW, Shewry PR, et al. Isolation and characterization of five high-molecular-weight subunit of glutenin genes from Triticum timopheevi and Aegilops cylindrica[J]. Theoretical and Applied Genetics,2002,1041:828-839.
[110]Chen FG, Zhu XY, Xia GM. Sequence of a new HMW glutenin subunit gene in Haynaldia villosa[J]. Acta Botanica Boreali-Occidentalia Sinica,2005,25:1410-1414.
[111]Guo ZF,Yan ZH,Wang JR, et al. Characterization of HMW prolamines and their coding sequences from Crithopsis delileana[J]. Hereditas,2005,142:56-64.
[112]Sun M, Yan Y, Jiang Y, et al. Molecular cloning and comparative analysis of a y-type inactive HMW glutenin subunit gene from cultivated emmer wheat (Triticum dicoccum L)[J]. Hereditas,2004,141(1):46-54.
[113]Yuan ZW,Chen QJ,Zhang LQ, et al. Molecular characterization of two silenced y-type genes for Glu-B1 in Triticum aestivum ssp. yunnanese and ssp. Tibetanum[J]. Journal of Integrative Plant Biology,2009,51(1):93-99.
[114]Halford NG, Forde J, Shewry PR, et al. Functional analysis of the upstream regions of a silent and an expressed member of a family of wheat seed protein genes in transgenic tobacco[J]. Plant Science,1989,62:207-216.
[115]Bai JR., Jia X, Liu KF,et al. Cloning and characterization of the coding sequences of the 1 Ay high molecular weight glutenin subunit genes from Triticum urartu[J]. Acta Botanica Sinica, 2004,46(4):463-471.
[116]Jiang QT, Wei YM, Wang F, et al. Characterization and comparative analysis of HMW glutenin lAy alleles with differential expressions [J]. BMC Plant Biology,2009,9(1):16.
[117]郑雯.野生二粒小麦1Ay高分子量谷蛋白亚基的分子克隆及其“表达-沉默”机理探讨[D].雅安：四川农业大学,2008,24-26.
[2]Osborne TB. The proteins of the wheat kernel. Washington DC:Carnegie Institution of Washington,1907.
[3]Bietz JA, Wall JS. The effect of various extractants on the subunit composition and associations of wheat glutenin[J]. Cereal Chemistry.1975,52:145-155.
[4]Payne PI, Law CN, Mudd EE. Control by homoeologous group 1 chromosomes of the high-molecular-weight subunits of glutenin, a major protein of wheat endosperm[J]. Theoretical and Applied Genetics,1980,58:113-120.
[5]Lawrence G J, Shepherd KW. Inheritance of glutenin protein subunits of wheat[J]. Theoretical and Applied Genetics,1981,60:333-337.
[6]Payne PI, Lawrence GJ. Catalogue of alleles for the complex gene loci,Glu-A1,Glu-B1,and Glu-D1 which code for the high-molecular-weight subunits of glutenin in hexaploid wheat[J]. Cereal research communications,1983,11:29-35.
[7]赵和,卢少源,李宗智.小麦高分子量麦谷蛋白亚基遗传变异及其与品质和其它农艺性状关系的研究[J].作物学报,1994,20(1)：67-75.
[8]栗站稳,王学路,卢少源,等.小麦主要贮藏蛋白亚基组分在我国种质资源中的分布[J].北京农业科学,1995,13(6)：21-24.
[9]张学勇,庞斌双,游光霞,等.中国小麦品种资源Glu1位点组成概况及遗传多样性分析[J].中国农业科学,2002,35(11)：1302-1310.
[10]Xu SS, Khan K, Klindworth DL, et al. Chromosomal location of genes for novel glutenin subunits and gliadins in wild emmer wheat(Triticum turgidum L var Dicoccoides)[J]. Theoretical and Applied Genetics,2004,108:1221-1228.
[11]Forde J, Malpica JM, Halford NG,e t al. The Nucleotide Sequence of a HMW Glutenin Subunit Gene Located on Chromosome 1A of Wheat(Triticum aestivum L)[J]. Nucleic Acids Research,1985,13:6817-6832.
[12]Anderson OD, Green C, Yip PE, et al. Nucleotide Sequences of the High Molecular Weight Glutenin Genes from the D-Genome of Hexaploid Wheat,Triticum aestivum L.cv Cheyenne[J]. Nucleic Acids Research,1989,17:461-462.
[13]Halford NG, Forde J, Anderson OD. The Nucleotide and Deduced Amino Acid Sequences of an HMW Glutenin Subunit Gene from Chromosome 1B of Bread Wheat(Triticum aesfvum L.) and Comparison with Those of Genes from Chromosomes 1A and 1D[J]. Theoretical and Applied Genetics,1987,75:117-126.
[14]Shewry PR, Halford NG. Cereal seed proteins:structures,properties and role in grain utilization [J]. Journal of Experimental Botany,2002,53:947-958.
[15]Shewry PR, Tatham AS. Disulphide bonds in wheat gluten Proteins[J]. Journal of Cereal Science,1997,25:207-227.
[16]Kasarda DD, Nimmo CC, Bernardin JE. Structural aspects and genetic relationships of gliadins [M]. In Coelic Disease:Proc.2nd Int,1974,25-36.
[17]Ewart JAD. Glutenin structure[J]. Journal of the Science of Food and Agriculture,1978, 30(5):482-492.
[18]Gilbert SM, Wellner N, Belton PS, et al. Expression and characterization of a highly repetitive peptide derived from a Wheat seed storage protein[J]. Biochimica et Biophysica Acta,2000,1479:135-146.
[19]Belton PS. On the elasticity of wheat gluten[J]. Journal of Cereal Science,1999,29: 103-109.
[20]Tilley KA, Benjamin RE, Bagorogoza KE, et al. Tyrosine cross-links molecular basis of gluten structure and function[J]. Journal of Agricultural and Food Chemistry,2001,49: 2627-2632.
[21]Payne PI. Genetics of wheat storage Proteins and the effect of allelic variationon breadmaking quality[J]. Annual Review of Plant Physiology,1987,38:141-153.
[22]毛沛.高分子量麦谷蛋白亚基组成及其与面包烘烤品质关系的研究[D].保定：河北农业大学.1992.
[23]Ammar K, Warren E, Kronstad, et al. Morris Breadmaking quality of selected durum wheat geno-types and its relationship with high molecular weight glutenin subunits allelic variation and gluten proteinpolymeric composition[J]. Cereal Chemistry,2000,77(2):230-236.
[24]卫增泉.离子束生物效应及应用中值得研究的几个问题[J].激光生物学报,2006,15(4)：383-387.
[25]余增亮.离子束生物技术引论[M].安徽：安徽科学技术出版社,1998.
[26]王卫东,王燕,王雪青,等.离子注入对小麦当代形态生理影响的初步研究[J].激光生物学报,2003,12(5)：360-363.
[27]王卫东,王燕,王雪青,等.氮离子注入后拟南芥种子表皮的扫描电镜观察[J].电子显微学报,2004,23(2)：148-151.
[28]袁世斌,薛建明,王宇刚,等.碳离子束辐照拟南芥介导外源基因转移的研究[J].激光生物学报,2003,12(5)：326-331.
[29]余增亮,霍裕平.离子注入生物学研究述评[J].安徽农业大学学报,1994,21(3)：221-225.
[30]邵春林,许安,余增亮.离子注入生物分子的电荷交换效应[J].核技术,1997,20(2)：70-73.
[31]张冬梅,孙素琴,马志龙,等.低能离子束辐照氨基酸结构变化的傅里叶变换红外光谱法研究[J].核技术,2000,23(4)：213-215.
[32]梁剑平.重离子辐照对喹烯酮的影响和重离子放射增敏药物的开发[D].兰州：中国科学院近代物理研究所,2000.
[33]袁世斌.低能重离子束对生物分子的辐照效应研究[D].兰州：中国科学院近代物理研究所,2002.
[34]石怀斌,邵春林,王相勤,等.低能氮离子对腺嘌呤水溶液损伤作用的光谱分析[J].光谱学与光谱分析,2002,22(2)：278-280.
[35]谷运红,王卫东,秦广雍,等.低能N+离子注入胸腺嘧啶的XPS研究[J].光谱学与光谱分析,2007,2：400-403.
[36]徐家萍,刘明辉,汪泰初.离子束诱变桑品种与亲本的同工酶和RAPD比较分析[J].安徽农业大学学报,2002,29(3)：286-288.
[37]焦浈,秦广雍,霍裕平.低能铁离子注入烟草叶片的过氧化物酶同工酶和全蛋白分析[J].河南农业科学,2004,6：23-25.
[38]赵俊杰,黄中文,常胜合.N+离子注入对不同小麦品种同工酶的影响[J].河南农业科学,2006,8：25-27.
[39]张从宇,王敏,张子学,等.60Co-γ和Ar+处理对小麦的诱变效应及POD同工酶分析[J].激光生物学报,2008,17(6)：780-786.
[40]李文建,卫增泉,颉红梅,等.中能氮离子束辐照小麦种子产生自由基的研究[J].核技术,1995,18(7)：415-418.
[41]孔福全.重离子致DNA损伤及修复的研究[D].天津：河北工业大学,2005.
[42]颉红梅,郝冀方,卫增泉,等.重离子束辐照牧草的细胞学研究[J].激光生物学报,2003,12(5)：346-349.
[43]杨美英,武志海,崔秋华,等.粳稻氩离子(Ar+)束诱变育种的细胞学效应研究[J].吉林农业大学学报,2002,24(5)：35-38.
[44]王崇英,王如娟,杨汉民.低能N+重离子注入处理后小麦种胚内的DNA期外合成[J].辐射研究与辐射工艺学报,1994,22(2)：205-208.
[45]杨剑波,吴李君,李莉,等.离子束辐照M13mp18-DNA(RF1)诱导lacZ突变的序列分析[J].中国科学(B辑),1995,25(12)：1273-1278.
[46]常凤启,刘选明,李银心,等.低能N+辐照拟南芥诱导基因组DNA碱基变异分析[J].中国科学(C辑),2003,33(2)：117-124.
[47]吴丽芳,李红.离子束生物工程应用研究进展[J].物理,1999,28(12)：708-712.
[48]杨赞林,甘斌杰,余增亮.离子注入小麦诱变育种的回顾与展望[J].安徽农业科学,2002,30(5)：639-641,648.
[49]李兴林,王晓娟,李文建,等.重离子辐照春麦稳定后代品质和贮藏蛋白的变化[J].核农学报,2000,14(5)：311-314.
[50]颉红梅,王浩瀚,王菊芳,等.重离子束定点诱变育种初探[J].原子核物理评论,2001,18(3)：174-176.
[51]陈秀兰,柳学余.利用离子注入诱变创造小麦新种质[J].江苏农学院学报,1998,19(2)： 62-64.
[52]李兰真,秦广雍,霍裕平,等.离子注入在小麦诱变育种上的应用研究初报[J].河南农业大学学报,2001,35(1)：9-12.
[53]张金莲,胡钺,卫增泉,等.低能重离子注入春小麦甘种子生物效应的初步研究[J].甘肃农业科技,1995,2：2-4.
[54]张利华.N2激光和氮离子注入对小麦一些性状的影响[J].种子,1999,3：15-17.
[55]宣云,王会峰,黄群策,等.离子束生物技术在小麦改良中的应用[J].农业生物技术科学,2004,20(1)：32-34,76.
[56]Yu ZL, Yang JB, Wu YJ, et al. Transferring Gus gene into intact rice cells by low energy ion beam[J]. Nuclear Instruments and Methods in Physics Research,1993, B80:1328-1331.
[57]吴丽芳,李红,宋道军,等.低能氩离子束介导将绿色荧光蛋白基因导入小麦的研究[J].南京农业大学学报,2000,23(3)：17-20.
[58]聂利红,崔党群,闻捷,等.离子束介导大豆789转入普通小麦后代的麦谷蛋白和醇溶蛋白分析[J].麦类作物学报,2005,25(3)：125-127.
[59]焦浈.低能离子束介导大豆全DNA导入小麦后其蛋白质遗传特性研究[D].郑州：郑州大学,2003.
[60]吴丽芳,余增亮.离子注入法获得大豆-小麦分子远缘杂种及后代的变异分析[J].核农学报,2000,14(4)：206-211.
[61]姬生栋,李吉学,徐存拴,等.离子束介导大豆DNA小麦后代的蛋白质变异性分析[J].麦类作物学报,2001,21(3)：18-21.
[62]杨随庄,尚勋武,叶春雷.小麦高分子量谷蛋白亚基基因分子育种研究进展[J].麦类作物学报,2007,27(2)：358-363.
[63]Halford NG, Field JM, Blair H, et al. Analysis of HMW glutenin subunits encoded by chromosome 1A of bread wheat(Triticum aestivum L.) indicates quantitative effects on grain quality[J]. Theoretical and Applied Genetics,1992,83:373-378.
[64]张金锐,刘勇,林刚,等.小麦高分子量谷蛋白亚基序列及其结构与加工品质的关系[J].中国生物工程杂志,2006,26(8)：103-110.
[65]Shewry PR. The synthesis,processing and deposition of gluten proteins in the Developing wheat grain[J]. Cereal Foods World,1999,44:587-589
[66]Kohler P, Keck B, Muller S, et al. Disulphide bonds in wheat gluten[C]. In:Wheat Kernel Proteins,Molecular and functional aspects (Universita degli studi della Tuscia consiglio nazionale delle ricerche, ed). Viterbo, Italy,1995,45-53.
[67]王关林,方宏筠.植物基因工程原理与技术[M].北京：科学出版社,1998,33.
[68]Altpeter F, Vimla V, Vibha S, et al. Integration and expression of the high-molecular-weight glutenin subunit 1Ax1 gene into wheat[J]. Nature Biotechnology,1996,14:155-159.
[69]Vasil IK, Bean S, Zhao JM,et al. Evalution of baking properties and gluten composition of filed grown transgenic wheat lines expressing high molecular weight glutenin gene 1Ax1[J]. Journal of Plant Physiology,2001,158(4):521-528.
[70]Popineau Y, Deshayes G, Lefebvre J, et al. Prolamin aggregation,gluten viscoelasticity,and mixing properties of transgenic wheat lines expressing 1Ax and 1Dx high molecular weight glutenin subunit transgenes[J]. Journal of Agricultural and Food Chemistry,2001,49(1): 395-401
[71]Barro F, Rooks L, Bekes F, et al. Transformation of wheat with high molecular weight subunit genes results in improved functional properties[J]. Nature Biotechnology,1997,15: 1295-1299.
[72]Barro F, Barcelo P, Lazzeri PA, et al. Functional properties and agronomic performance of Transgenic Tritordeum expressing high molecular weight glutenin subunit genes 1Ax1 and 1Dx5[J]. Journal of Cereal Science,2003,37:65-70.
[73]Alvarez ML, Gomezm M, Carrillo JM,et al. Analysis of dough functionallity of flours from transgenic wheat[J]. Molecular Breeding,2001,8(1):103-108.
[74]M Rakszegi, G Pastori, HD Jones,et al. Technological quality of field grown transgenic lines of commercial wheat cultivars expressing the 1Ax1 glutenin subunit gene[J]. Journal of Cereal Science,2008,47:310-321.
[75]Field JM, Bhandari D, Bonet A, et al. Introgression of transgenes into a commercial cultivar confrms differential effects of HMW subunits 1Ax1 and 1Dx5 on gluten properties[J]. Journal of Cereal science,2008,48:457-463.
[76]Eckardt N A. RNA goes mobile [J]. Plant cell,2002,101:543-553.
[77]Margiotta B, Urbano M, Colaprico G, et al. Detection of y-type subunit at the Glu-A1 locus in some Swedish bread wheat lines [J]. Journal of Cereal Science,1996,23:203-211.
[78]He ZH, Pena RJ, Rajaram S. High molecular weight glutenin subunit composition of Chinese bread wheat [J]. Euphytica,1992,64:11-20.
[79]Morgunov AL, Rogers WJ, Sayers EG, et al. The high molecular weight glutenin composition of Soviet wheat varieties [J]. Euphytica,1990,51:41-52.
[80]Lawrence GJ. The high-molecular-weight glutenin subunit composition of Australian wheat cultivars [J]. Australian Journal of Agricultural Research,1986,37:123-125.
[81]Payne PI, Nightingale MA, Krattinger AF, et al. The relationship between HMW Glutenin Subunit composition and the bread-making quality of British grown wheat varieties [J]. Journal of the Science of Food and Agriculture,1987,40:51-65.
[82]Lukow OM, Payne PI, Tkachuk R.The HMW glutenin subunit composition of Canadian wheat cultivars and their association with bread-making quality [J]. Journal of the Science of Food and Agriculture,1989,46(4):451-460.
[83]Vallega V, Waines J G. High molecular weight glutenin subunit variation in Triticum turgidum var. dicoccum [J]. Theoretical and Applied Genetics,1987,74:706-710.
[84]董永梅,杨欣明,柴守诚,等.中国小麦代表性地方品种高分子量麦谷蛋白亚基组成分 析[J].麦类作物学报,2007,27(5)：820-824.
[85]刘悦,杨足君,李光蓉,等.四川南部和重庆地区小麦地方品种的高分子量谷蛋白亚基组成分析[J].麦类作物学报,2006,26(4)：47-50.
[86]杨恩年,张洁,杨武云,等.六倍体普通小麦高分子量谷蛋白亚基Glu-A1和Glu-B1共同缺失材料研究初报[J].西南农业学报,2007,20(2)：293-295.
[87]Lawrence GJ. Dough and baking quality of wheat line deficient in glutenin subunits controlled by the Glu-A1, Glu-B1 and Glu-D1 loci [J]. Journal of Cereal Science,1988,7: 109-112.
[88]Payne P I. Genetics of wheat storage proteins and the effect of allelic variation on breadmaking quality [J]. Annual Review of Plant Physiology and Plant Molecular Biology, 1987,38:141-153.
[89]Upelniek V, Novoselskaya A, Sutka J,et al. Genetic variation at storage protein-coding loci of common wheat (cv'Chinese Spring') induced by nitrosoethylurea and by the cultivation of immature embryos in vitro [J]. Theoretical and Applied Genetics,1995,90:372-379.
[90]Zhu YF, Li YW, Chen Y, et al. Generation and characterization of a high molecular weight glutenin 1Bx14 deficient mutant in common wheat[J]. Plant Breeding,2005,124:421-427.
[91]张怀渝,宋云,畅志坚,等.低能N+注入诱导小麦种子高分子量麦谷蛋白亚基和醇溶蛋白的变异[J].核农学报,2005,19(4)：245-250.
[92]Altpeter F, Vasil V, Srivastava V,et al. Integration and expression of the high molecular weight glutenin subunit lAxl gene into wheat [J]. Nature Biotechnology,1996,14: 1155-1159.
[93]Alvarez ML, Guelman S, Halford NG, et al. Silencing of HMW glutenins in transgenic wheat expressing extra HMW subunits[J]. Theoretical and Applied Genetics,2000,100:319-327.
[94]Uthayakumaran S, Lukow OM. Functional and multiple end-use characterisation of Canadian wheat using a reconstituted dough system [J]. Journal of the Science of Food and Agriculture, 2003,83:889-898.
[95]霍裕平,秦广雍,苏明杰,等.离子束诱变小麦遗传新规律的研究[C].第三次全国离子束生物工程学大会暨第一次国际研讨会论文集,2002.
[96]赵勇.低能重离子引起DNA链断裂和诱发基因突变谱的研究[D].武汉：武汉大学,2003.
[97]潘多.低能N+离子束诱变的机理研究：突变谱分析[D].哈尔滨：东北林业大学,2005.
[98]辛庆国.7Li离子束核反应生物诱变效应与作用机制研究[D].泰安：山东农业大学,2007.
[99]钱时权.合成PHA相关基因离子束诱变及其高产突变体分子机制研究[D].合肥：安徽农业大学,2007.
[100]李珂,张东正,赵瑾,等.重离子注入与生物体的相互作用及遗传诱变的分子机制[J].原子核物理评论,2007,24(2)：117-123.
[101]Gu YQ,Salse J, Coleman D, et al. Types and rates of sequence evolution at the high molecular weight glutenin locus in hexaploid wheat and its ancestral genomes[J]. Genetics, 2006,174:1493-1504.
[102]Harberd NP, Flavell RB, Thompson RD. Identification of a transposon like insertion in a Glu-1 allele of wheat[J]. Molecular Genetics and Genomics,1987,209:326-332.
[103]Bustos AD, Rubio P, Jouve N. Molecular characterisation of the inactive allele of the gene Glu-A1 and the development of a set of AS-PCR markers for HMW glutenins of wheat[J]. Theoretical and Applied Genetics,2000,100:1085-1094.
[104]Yang ZJ, Li GR, Liu C, et al. Molecular Characterization of a HMW Glutenin subunit allele providing evidence for silencing of x-type gene on Glu-B1 [J]. Acta Genetica Sinica,2006, 33 (10):929-936.
[105]Forde J, Malpica JM, Halford NG, et al. The nucleotide sequence of a HMW subunit gene located on chromosome 1A of wheat (Triticum eastivum L.)[J]. Nucleic Acids Research, 1985,13:6817-6832.
[106]Liu SW, Gao X, Xia GM. Characterizing HMW-GS alleles of decaploid Agropyron elongatum in relation to evolution and wheat breeding[J]. Theoretical and Applied Genetics, 2008,116:325-334.
[107]Wang JR, Yan ZH, Wei YM, et al. Characterization of high-molecular-weight glutenin subunit genes from Elytrigia elongata(Host) Nevski[J]. Plant Breeding,2006,125:89-95.
[108]Feng DS, Chen FG, Zhao SY, et al. High-molecular-weight glutenin subunit genes in decaploid Agropyron elongatum [J]. Acta Botanica Sinica,2004,46:489-496.
[109]Wan YF, Wang DW, Shewry PR, et al. Isolation and characterization of five high-molecular-weight subunit of glutenin genes from Triticum timopheevi and Aegilops cylindrica[J]. Theoretical and Applied Genetics,2002,1041:828-839.
[110]Chen FG, Zhu XY, Xia GM. Sequence of a new HMW glutenin subunit gene in Haynaldia villosa[J]. Acta Botanica Boreali-Occidentalia Sinica,2005,25:1410-1414.
[111]Guo ZF,Yan ZH,Wang JR, et al. Characterization of HMW prolamines and their coding sequences from Crithopsis delileana[J]. Hereditas,2005,142:56-64.
[112]Sun M, Yan Y, Jiang Y, et al. Molecular cloning and comparative analysis of a y-type inactive HMW glutenin subunit gene from cultivated emmer wheat (Triticum dicoccum L)[J]. Hereditas,2004,141(1):46-54.
[113]Yuan ZW,Chen QJ,Zhang LQ, et al. Molecular characterization of two silenced y-type genes for Glu-B1 in Triticum aestivum ssp. yunnanese and ssp. Tibetanum[J]. Journal of Integrative Plant Biology,2009,51(1):93-99.
[114]Halford NG, Forde J, Shewry PR, et al. Functional analysis of the upstream regions of a silent and an expressed member of a family of wheat seed protein genes in transgenic tobacco[J]. Plant Science,1989,62:207-216.
[115]Bai JR., Jia X, Liu KF,et al. Cloning and characterization of the coding sequences of the 1 Ay high molecular weight glutenin subunit genes from Triticum urartu[J]. Acta Botanica Sinica, 2004,46(4):463-471.
[116]Jiang QT, Wei YM, Wang F, et al. Characterization and comparative analysis of HMW glutenin lAy alleles with differential expressions [J]. BMC Plant Biology,2009,9(1):16.
[117]郑雯.野生二粒小麦1Ay高分子量谷蛋白亚基的分子克隆及其“表达-沉默”机理探讨[D].雅安：四川农业大学,2008,24-26.