~(60)Co-γ射线诱变小麦M_3代品质性状的遗传变异分析
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摘要
为探讨~(60)Co-γ射线辐射处理对郑麦9023的诱变效应,用200 Gy的~(60)Co-γ射线辐射处理郑麦9023干种子,对M_3代群体166个株系的籽粒蛋白质含量、湿面筋含量、硬度、沉降值、高分子量谷蛋白(GMP)含量、脂肪氧化酶(LOX)活性和多酚氧化酶(PPO)活性共7个品质性状进行了遗传变异分析及高分量麦谷蛋白亚基(HMW-GS)鉴定。结果表明,M_3代群体籽粒的蛋白质含量、湿面筋含量、LOX活性均值高于亲本,硬度、沉降值、GMP含量和PPO活性均值低于亲本。以超过群体均值±1.96×标准差[P(|u|≥1.96σ)≤0.05]为标准确定变异株系,蛋白质含量和湿面筋含量仅发生正向变异,发生变异的株系分别为5个和3个;硬度没有产生正向变异,发生负向变异的株系有7个。M_3代群体中发生高分子量谷蛋白亚基(HMW-GS)变异株系6个,亚基组成从郑麦9023的Null/7+8/2+12突变为1/7+8/2+12,品质得分提高了2分。在M_3代群体中筛选出13个强筋小麦株系,无弱筋小麦突变体。说明~(60)Co-γ射线可以诱导蛋白质亚基发生变异,拓宽小麦的遗传基础。
To clarify the mutagenic effects of radiation on quality characters of Zhengmai 9023, dry seeds of wheat were radiated by ~(60)Co-γ ray with the dose of 200 Gy. The genetic variation of 166 M_3 generation on protein content, wet gluten content, hardness index, sedimentation, GMP content, LOX activity and PPO activity were analyzed, as well as their HMW-GS were identified by SDS-PAGE. The results showed that the mean values of M_3 population in protein content, wet gluten content and LOX activity were higher than those of parents. Hardness, sedimentation, GMP content and PPO activity of progenies were lower than those of parents. Depending on the criterion of over mean±1.96×SD, protein content and wet gluten content only generated positive variation, no negative variation, accounting for five lines and three lines in positive variation, respectively. There were seven lines had negative variation in hardness. Six mutant lines of HMW-GS were identified in M_3 by SDS-PAGE. The subunit composition changed from Null/7+8/2+12 to 1/7+8/2+12,with 2 points quality scores increased. There were 13 mutant lines with high gluten and no lower gluten mutant line in M_3 generation. The analysis of mutant lines variation indicated that subunit composition variations could be obtained, and the genetic basis in original wheat varieties could be broadened.
To clarify the mutagenic effects of radiation on quality characters of Zhengmai 9023, dry seeds of wheat were radiated by ~(60)Co-γ ray with the dose of 200 Gy. The genetic variation of 166 M_3 generation on protein content, wet gluten content, hardness index, sedimentation, GMP content, LOX activity and PPO activity were analyzed, as well as their HMW-GS were identified by SDS-PAGE. The results showed that the mean values of M_3 population in protein content, wet gluten content and LOX activity were higher than those of parents. Hardness, sedimentation, GMP content and PPO activity of progenies were lower than those of parents. Depending on the criterion of over mean±1.96×SD, protein content and wet gluten content only generated positive variation, no negative variation, accounting for five lines and three lines in positive variation, respectively. There were seven lines had negative variation in hardness. Six mutant lines of HMW-GS were identified in M_3 by SDS-PAGE. The subunit composition changed from Null/7+8/2+12 to 1/7+8/2+12,with 2 points quality scores increased. There were 13 mutant lines with high gluten and no lower gluten mutant line in M_3 generation. The analysis of mutant lines variation indicated that subunit composition variations could be obtained, and the genetic basis in original wheat varieties could be broadened.
引文
[1]杨震,彭选明,彭伟正.作物诱变育种研究进展[J].激光生物学报,2016,25(4):306.YANG Z,PENG X M,PENG W Z.Progress of study on crop mutation breeding [J].Acta Laser Biology Sinica,2016,25(4):306.
[2]范家霖,张建伟,杨保安,等.河南省小麦诱变育种成就及发展对策[J].麦类作物学报,1999(6):17.FAN J L,ZHANG J W,YANG B A,et al.Achievements and development strategies of mutation breeding of wheat in Henan Province [J].Journal of Triticeae Crops,1999(6):17.
[3]王彩萍,左联忠,许琦,等.优质小麦“农大179”辐射诱变M2 代RVA参数的变异分析[J].麦类作物学报,2007,27(2):237.WANG C P,ZUO L Z,XU Q,et al.RVA parameter variation among M2 population of radiated wheat variety Nongda 179 [J].Journal of Triticeae Crops,2007,27(2):237.
[4]赵林姝,刘录祥.农作物辐射诱变育种研究进展[J].激光生物学报,2017,26(6):484.ZHAO L S,LIU L X.Research progresses in irradiation-induced mutation breeding in crops [J].Acta Laser Biology Sinica,2017,26(6):484.
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[6]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.YANG S Z,SHANG X W,YE C L.Advance on improvement of wheat high-molecular-weight glutenin subunits via molecular breeding [J].Journal of Triticeae Crops,2007,27(2):358.
[7]LIU S W.Genetic variation of high molecular weight glutenin subunits associated with processing quality improvements in wheat [J].Plant Physiology Journal,2011,47(6):531.
[8]胡琳,盖钧镒,许为钢,等.小麦不同高分子量谷蛋白亚基对面筋数量和质量的影响[J].麦类作物学报,2006,26(6):50.HU L,GAI J Y,XU W G,et al.Effects of high molecular weight glutenin subunits on quantity and quality of wheat gluten [J].Journal of Triticeae Crops,2006,26(6):50.
[9]FUERST E P,ANDERSON J V,MORRIS C F.Delineating the role of polyphenol oxidase in the darkening of alkaline wheat noodles [J].Journal of Agricultural & Food Chemistry,2006,54(6):2378.
[10]张钰玉,王晓龙,张晓科,等.陕西小麦品种(系)脂肪氧化酶活性基因的遗传多态性分析[J].麦类作物学报,2012,32(4):620.ZHANG Y Y,WANG X L,ZHANG X K,et al.Genetic polymorphism of lipoxygenase genes in Shaanxi wheat cultivars and lines [J].Journal of Triticeae Crops,2012,32(4):620.
[11]LEENHARDT F,LYAN B,ROCK E,et al.Genetic variability of carotenoid concentration,and lipoxygenase and peroxidase activities among cultivated wheat species and bread wheat varieties [J].European Journal of Agronomy,2006,25(2):170.
[12]BORRELLI G M,LEONARDIS A M D,PLATANI C,et al.Distribution along durum wheat kernel of the components involved in semolina colour [J].Journal of Cereal Science,2008,48(2):494.
[13]冯毅,闵东红,孙道杰,等.黄淮麦区部分推广小麦品种高分子量麦谷蛋白亚基组成分析[J].麦类作物学报,2006,26(2):69.FENG Y,MIN D H,SUN D J,et al.Composition of HMW-GS of wheat cultivars grown in Huanghuai wheat production area [J].Journal of Triticeae Crops,2006,26(2):69.
[14]WEEGELS P L,PIJPEKAMP A M V D,GRAVELAND A,et al.Depolymerisation and re-polymerisation of wheat glutenin during dough processing.I.Relationships between glutenin macropolymer content and quality parameters [J].Journal of Cereal Science,1996,23(2):104.
[15]孙辉,姚大年,李宝云,等.普通小麦谷蛋白大聚合体的含量与烘焙品质相关关系[J].中国粮油学报,1998(6):14.SUN H,YAO D N,LI B Y,et al.Correlation between content of glutenin macropolymer(GMP) in wheat and baking quality [J].Journal of the Chinese Cereals and Oils Association,1998(6):14.
[16]CATO L,HALMOS A L,SMALL D M.Measurement of lipoxygenase in Australian white wheat flour:The effect of lipoxygenase on the quality properties of white salted noodles [J].Journal of the Science of Food & Agriculture,2006,86(11):1670.
[17]ANDERSON J V,MORRIS C F.An improved whole-seed assay for screening wheat germplasm for polyphenol oxidase activity [J].Crop Science,2001,41(6):1704.
[18]LI L,HE Z,YAN J,et al.Allelic variation at the Glu-1 and Glu-3 loci,presence of the 1B·1R translocation,and their effects on mixographic properties in Chinese bread wheats [J].Euphytica,2005,142(3):197.
[19]SINGH N K,SHEPHERD K W,CORNISH G B.A simplified SDS-PAGE procedure for separating LMW subunits of glutenin [J].Journal of Cereal Science,1991,14(3):203.
[20]PAYNE P I.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 [J].Cereal Research Communications,1983,11(1):29.
[21] MAGO R,SPIELMEYER W,LAWRENCE G J,et al.Resistance genes for rye stem rust(SrR) and barley powdery midew(Mla) are located in syntenic regions on short arm of chromosome [J].Genome,2004,47:112.
[22]张跃强,樊哲儒,李剑锋,等.60Co-γ 射线诱变选育早熟、优质、高产小麦新品种新春30号[J].核农学报,2010,24(5):978.ZHANG Y Q,FAN Z R,LI J F,et al.Breeding of Xinchun 30 with early maturity,good quality and high yield by hybridization and 60Co-γ-rays irradiation [J].Journal of Nuclear Agricultural Sciences,2010,24(5):978.
[23]张从宇,王敏,李文阳,等.60Co-γ诱变皖麦50 HMW-GS变异的筛选及其M3代品质分析[J].激光生物学报,2014,23(2):130.ZHANG C Y,WANG M,LI W Y,et al.Mutation screening of HMW-GS composition of Wanmai 50 irradiated by 60Co-γ and analysis on its quality of M3 generation [J].Acta Laser Biology Sinica,2014,23(2):130.
[24]朱守晶,王敏,张从宇.60Co-γ辐射诱变小麦M3代品质性状的遗传变异分析[J].激光生物学报,2010,19(6):764.ZHU S J,WANG M,ZHANG C Y.Genetic variation analysis of quality characters of M3 radiated by 60Co-γ in wheat [J].Acta Laser Biology Sinica,2010,19(6):764.
[25]王峰,王敏,张从宇.重庆面包麦辐射诱变后代性状变异分析[J].麦类作物学报,2013,33(6):1134.WANG F,WANG M,ZHANG C Y.Mutational analysis on Chongqing bread wheat induced by irradiation [J].Journal of Triticeae Crops,2013,33(6):1134.
[26]李秀贞,吕善勇.辐射对麦类作物蛋白质含量的影响[J].核农学通报,1994(2):53.LI X Z,Lü S Y.The protein content effect on wheat crops by irradiation [J].Journal of Nuclear Agricultural Sciences,1994(2):53.
[27]程西永,吴少辉,李海霞,等.小麦高、低分子量麦谷蛋白亚基对品质性状的影响[J].麦类作物学报,2014,34(4):487.CHENG X Y,WU S H,LI H X,et al.Effects of HMW and LMW glutenin subunits on wheat quality traits [J].Journal of Triticeae Crops,2014,34(4):487.
[28]杨芳萍,王生荣.小麦高分子量谷蛋白亚基与品质性状的关系[J].麦类作物学报,2003,23(4):35.YANG F P,WANG S R.Relationship between the subunits of high molecular weight gluten and quality characters of wheat [J].Journal of Triticeae Crops,2003,23(4):35.
[2]范家霖,张建伟,杨保安,等.河南省小麦诱变育种成就及发展对策[J].麦类作物学报,1999(6):17.FAN J L,ZHANG J W,YANG B A,et al.Achievements and development strategies of mutation breeding of wheat in Henan Province [J].Journal of Triticeae Crops,1999(6):17.
[3]王彩萍,左联忠,许琦,等.优质小麦“农大179”辐射诱变M2 代RVA参数的变异分析[J].麦类作物学报,2007,27(2):237.WANG C P,ZUO L Z,XU Q,et al.RVA parameter variation among M2 population of radiated wheat variety Nongda 179 [J].Journal of Triticeae Crops,2007,27(2):237.
[4]赵林姝,刘录祥.农作物辐射诱变育种研究进展[J].激光生物学报,2017,26(6):484.ZHAO L S,LIU L X.Research progresses in irradiation-induced mutation breeding in crops [J].Acta Laser Biology Sinica,2017,26(6):484.
[5]中华人民共和国国家质量监督检验检疫总局.GB/T1789221999,优质强筋弱筋小麦[S].北京:中国标准出版社,1999.General administration of quality supervision,inspection and quarantine of the People’s Republic of China:GB/T178922 1999,High quality wheat with strong gluten and weak gluten [S].Beijing:China Zhijian Publishing House,1999.
[6]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.YANG S Z,SHANG X W,YE C L.Advance on improvement of wheat high-molecular-weight glutenin subunits via molecular breeding [J].Journal of Triticeae Crops,2007,27(2):358.
[7]LIU S W.Genetic variation of high molecular weight glutenin subunits associated with processing quality improvements in wheat [J].Plant Physiology Journal,2011,47(6):531.
[8]胡琳,盖钧镒,许为钢,等.小麦不同高分子量谷蛋白亚基对面筋数量和质量的影响[J].麦类作物学报,2006,26(6):50.HU L,GAI J Y,XU W G,et al.Effects of high molecular weight glutenin subunits on quantity and quality of wheat gluten [J].Journal of Triticeae Crops,2006,26(6):50.
[9]FUERST E P,ANDERSON J V,MORRIS C F.Delineating the role of polyphenol oxidase in the darkening of alkaline wheat noodles [J].Journal of Agricultural & Food Chemistry,2006,54(6):2378.
[10]张钰玉,王晓龙,张晓科,等.陕西小麦品种(系)脂肪氧化酶活性基因的遗传多态性分析[J].麦类作物学报,2012,32(4):620.ZHANG Y Y,WANG X L,ZHANG X K,et al.Genetic polymorphism of lipoxygenase genes in Shaanxi wheat cultivars and lines [J].Journal of Triticeae Crops,2012,32(4):620.
[11]LEENHARDT F,LYAN B,ROCK E,et al.Genetic variability of carotenoid concentration,and lipoxygenase and peroxidase activities among cultivated wheat species and bread wheat varieties [J].European Journal of Agronomy,2006,25(2):170.
[12]BORRELLI G M,LEONARDIS A M D,PLATANI C,et al.Distribution along durum wheat kernel of the components involved in semolina colour [J].Journal of Cereal Science,2008,48(2):494.
[13]冯毅,闵东红,孙道杰,等.黄淮麦区部分推广小麦品种高分子量麦谷蛋白亚基组成分析[J].麦类作物学报,2006,26(2):69.FENG Y,MIN D H,SUN D J,et al.Composition of HMW-GS of wheat cultivars grown in Huanghuai wheat production area [J].Journal of Triticeae Crops,2006,26(2):69.
[14]WEEGELS P L,PIJPEKAMP A M V D,GRAVELAND A,et al.Depolymerisation and re-polymerisation of wheat glutenin during dough processing.I.Relationships between glutenin macropolymer content and quality parameters [J].Journal of Cereal Science,1996,23(2):104.
[15]孙辉,姚大年,李宝云,等.普通小麦谷蛋白大聚合体的含量与烘焙品质相关关系[J].中国粮油学报,1998(6):14.SUN H,YAO D N,LI B Y,et al.Correlation between content of glutenin macropolymer(GMP) in wheat and baking quality [J].Journal of the Chinese Cereals and Oils Association,1998(6):14.
[16]CATO L,HALMOS A L,SMALL D M.Measurement of lipoxygenase in Australian white wheat flour:The effect of lipoxygenase on the quality properties of white salted noodles [J].Journal of the Science of Food & Agriculture,2006,86(11):1670.
[17]ANDERSON J V,MORRIS C F.An improved whole-seed assay for screening wheat germplasm for polyphenol oxidase activity [J].Crop Science,2001,41(6):1704.
[18]LI L,HE Z,YAN J,et al.Allelic variation at the Glu-1 and Glu-3 loci,presence of the 1B·1R translocation,and their effects on mixographic properties in Chinese bread wheats [J].Euphytica,2005,142(3):197.
[19]SINGH N K,SHEPHERD K W,CORNISH G B.A simplified SDS-PAGE procedure for separating LMW subunits of glutenin [J].Journal of Cereal Science,1991,14(3):203.
[20]PAYNE P I.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 [J].Cereal Research Communications,1983,11(1):29.
[21] MAGO R,SPIELMEYER W,LAWRENCE G J,et al.Resistance genes for rye stem rust(SrR) and barley powdery midew(Mla) are located in syntenic regions on short arm of chromosome [J].Genome,2004,47:112.
[22]张跃强,樊哲儒,李剑锋,等.60Co-γ 射线诱变选育早熟、优质、高产小麦新品种新春30号[J].核农学报,2010,24(5):978.ZHANG Y Q,FAN Z R,LI J F,et al.Breeding of Xinchun 30 with early maturity,good quality and high yield by hybridization and 60Co-γ-rays irradiation [J].Journal of Nuclear Agricultural Sciences,2010,24(5):978.
[23]张从宇,王敏,李文阳,等.60Co-γ诱变皖麦50 HMW-GS变异的筛选及其M3代品质分析[J].激光生物学报,2014,23(2):130.ZHANG C Y,WANG M,LI W Y,et al.Mutation screening of HMW-GS composition of Wanmai 50 irradiated by 60Co-γ and analysis on its quality of M3 generation [J].Acta Laser Biology Sinica,2014,23(2):130.
[24]朱守晶,王敏,张从宇.60Co-γ辐射诱变小麦M3代品质性状的遗传变异分析[J].激光生物学报,2010,19(6):764.ZHU S J,WANG M,ZHANG C Y.Genetic variation analysis of quality characters of M3 radiated by 60Co-γ in wheat [J].Acta Laser Biology Sinica,2010,19(6):764.
[25]王峰,王敏,张从宇.重庆面包麦辐射诱变后代性状变异分析[J].麦类作物学报,2013,33(6):1134.WANG F,WANG M,ZHANG C Y.Mutational analysis on Chongqing bread wheat induced by irradiation [J].Journal of Triticeae Crops,2013,33(6):1134.
[26]李秀贞,吕善勇.辐射对麦类作物蛋白质含量的影响[J].核农学通报,1994(2):53.LI X Z,Lü S Y.The protein content effect on wheat crops by irradiation [J].Journal of Nuclear Agricultural Sciences,1994(2):53.
[27]程西永,吴少辉,李海霞,等.小麦高、低分子量麦谷蛋白亚基对品质性状的影响[J].麦类作物学报,2014,34(4):487.CHENG X Y,WU S H,LI H X,et al.Effects of HMW and LMW glutenin subunits on wheat quality traits [J].Journal of Triticeae Crops,2014,34(4):487.
[28]杨芳萍,王生荣.小麦高分子量谷蛋白亚基与品质性状的关系[J].麦类作物学报,2003,23(4):35.YANG F P,WANG S R.Relationship between the subunits of high molecular weight gluten and quality characters of wheat [J].Journal of Triticeae Crops,2003,23(4):35.