大麦品质性状的基因型和环境变异及其机理研究
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摘要
大麦的主要用途是酿造啤酒和加工饲料,其品质优劣直接影响其最终产品的商业用途及价值,因此倍受重视。大麦品质性状大多为多基因控制的数量性状,不仅受品种本身遗传特性的制约,同时受外界环境条件的影响。本文主要研究了大麦籽粒蛋白总量、蛋白组分含量、β-淀粉酶活性,极限糊精酶活性及β-葡聚糖含量等主要啤用品质性状的基因型与环境变异及不同氮肥运筹对部分品质性状的影响,并利用大麦突变体TL43及其亲本Triumph研究了β-淀粉酶活性的调控机理,利用DH群体鉴定与定位大麦品质相关性状的QTLs。主要结果如下:
1.栽培大麦品质性状的基因型差异及灌浆后期的温度效应
研究了148个栽培大麦品种籽粒β-淀粉酶活性、蛋白质总量及蛋白组分含量的基因型变异,并利用离体穗培养技术研究了两个大麦品种在不同灌浆后期温度下的籽粒品质变化。β-淀粉酶活性、蛋白质总量及蛋白组分含量的基因型变异显著,β-淀粉酶活性与蛋白质总量及四种蛋白组分含量呈显著正相关,以与醇溶蛋白的相关性最高。与灌浆后期适温(22℃白天/16℃黑夜)相比,高温(32℃白天/26℃黑夜)显著提高β-淀粉酶活性及蛋白质组分含量,清蛋白含量变化较小,而谷蛋白含量影响最大。高温影响A型淀粉粒形状和B型淀粉粒数量,启示出淀粉结构变化可能是高温恶化麦芽品质的原因之一。
2.西藏野生大麦β-淀粉酶活性、极限糊精酶活性、β-葡聚糖及蛋白组分含量的基因型与环境变异
为研究西藏野生大麦籽粒品质性状的基因型与环境变异,测定了76个西藏野生大麦基因型在杭州、武汉两地的β-淀粉酶活性、β-葡聚糖和蛋白质组分含量。籽粒β-淀粉酶活性、β-葡聚糖和蛋白组分含量均存在显著的基因型差异,杭州2006-2007年度种植条件下的β-淀粉酶活性均值和变异幅度最高,且β-淀粉酶活性和β-葡聚糖含量的变异系数和变化趋势恰好相反。四种蛋白质组分中,醇溶蛋白含量与β-淀粉酶活性呈显著正相关。研究鉴定和筛选到β-淀粉酶活性和β-葡聚糖含量表现独特的种质材料,显示出可用于栽培大麦麦芽品质性状的改良。
3.氮肥运筹对栽培大麦粒重、蛋白组分含量和β-淀粉酶活性影响
研究了在总用氮量相同条件下四种氮肥运筹处理对栽培大麦品种花30粒重、蛋白质组分含量和β-淀粉酶活性的影响。不同氮肥运筹处理之间,大麦粒重无显著差异;增加后期用氮比例显著增加籽粒的醇溶蛋白、谷蛋白含量和β-淀粉酶活性,但对清蛋白和球蛋白含量无显著影响,说明提高大麦生育后期的用氮比例对啤用品质的影响是双重的,即在提高蛋白质含量的同时增加了β-淀粉酶活性。
4.孕穗期氮肥对大麦籽粒蛋白质组分和激素含量动态的影响
以欧洲大麦栽培品种Triumph及其ABA突变体TL43为材料,研究孕穗肥对籽粒灌浆期蛋白质组分和激素含量动态变化的影响。不同氮肥处理下,均表现TL43粒重较低但蛋白质含量较高;孕穗期高氮处理显著提高籽粒蛋白质含量,但对蛋白质组分含量的影响,两供试材料表现不同,TL43四种蛋白组分含量都显著提高,而Triumph仅为谷蛋白含量显著增加;籽粒灌浆期,两供试材料之间以及不同氮肥处理之间,GA_3含量变化差异较小;ABA和ZR含量的变化两供试材料之间差异明显,孕穗期施用氮肥显著降低灌浆后期的ABA含量。
5.大麦灌浆期H_2O_2与ABA互作对β-淀粉酶累积的调控机理
利用离体穗培养试验研究外源ABA及PEG处理对大麦品种Triumph及其突变体TL43灌浆期籽粒H_2O_2、ABA含量动态变化及β-淀粉酶的积累。两个基因型的籽粒ABA含量动态变化一致,而H_2O_2与β-淀粉酶的积累差异较大。β-淀粉酶的积累与H_2O_2含量呈正相关,与ABA含量呈负相关且与高ABA:H_2O_2持续时间有关。PEG提高β-淀粉酶活性主要与H_2O_2有关,而与ABA无关。外源添加H_2O_2和AsA显著提高Triumph的β-淀粉酶活性,但降低TL43的β-淀粉酶活性。籽粒内源H_2O_2含量动态始终与β-淀粉酶累积一致。结果显示出,胁迫条件下H_2O_2和ABA参与β-淀粉酶合成的调节。
6.大麦β-淀粉酶、极限糊精酶活性、β-葡聚糖和蛋白组分含量的QTL分析
利用六棱大麦品种CM72和二棱品种Gairdner为亲本,杂交F1代经花药培养获得的93个加倍单倍体(DH群体)株系,通过SSR标记构建连锁图谱,进行β-淀粉酶、极限糊精酶活性、β-葡聚糖和蛋白组分含量的QTL分析。8个品质性状共定位到9个QTLs,分别位于3H,4H,5H及7H染色体上,其中3H染色体上定位到5个QTLs,说明3H染色体对于麦芽品质有重要作用。各个QTL的表型变异范围在8.08%-30.25%之间。β-葡聚糖含量与极限糊精酶活性分别定位在4H、5H染色体上。醇溶蛋白QTL与已报道的硅流蛋白(Silica Eluate Protein)位点重合。β-淀粉酶活性的QTL定位在3H染色体上,而谷蛋白含量的QTL则定位在5H与7H染色体上。
Barley is mainly used as the materials for beer and feed production,and its quality is of important for the commercial values of the final product,thus the improvement of barley quality has been placed into a major objective of barley breeding and agronomy studies.The traits of barley quality belong to the quantitative characters, indicating that they are not only controlled by complex genetic factors,but also greatly affected by environmental factors.In the current study,the genotypic and environmental variation ofβ-amylase and limit dextrinase activities,β-glucan and protein fraction contents in barley grains and the influence of timing of nitrogen application on these malt quality characters were investigated;In addition,the possible mechanisms of P-amylase activity regulation was illustrated by using a barley mutant TL43 and its wild parent Triumph as experimental materials,and QTL analysis of these quality traits was conducted by using a double haploid population.
The main results were as follows:
1.The variation ofβ-amylase activity and protein fractions in barley grains as affected by genotypes and post-anthesis temperatures
The variation ofβ-amylase activity and protein fractions in barley grains was evaluated among 148 barley genotypes grown in the same field condition and using in vitro culture of two barley cultivars under different temperatures during grain development.The results showed that there were considerable genotypic variations inβ-amylase activity and protein fractions.Correlation analysis indicated thatβ-amylase activity was positively correlated with total protein and all protein fraction contents, with the highest correlation coefficient for hordeins.Furthermore,higher post-anthesis temperature(32/26℃,day/night) significantly enhancedβ-amylase activity and protein fractions,with albumins and glutelins being least and most affected respectively,in comparison with the plants under lower temperature(22/16℃).The high temperature also altered the morphology of A granules and number of B granules, suggesting starch structure change may also be a reason for deteriorated malting quality under high temperature.
2.Genotypic and environmental variation ofβ-amylase,limit dextrinase activity,β-glucan and protein fraction contents in Tibetan wild barley
The effects of genotypes and environments(locations and years) onβ-amylase,limit dextrinase activity,β-glucan and protein fraction contents in Tibetan wild barley grown in Hangzhou for two years and Wuhan for one year were investigated in this study.Significant genotypic differences and great variations were detected in grainβ-amylase,limit dextrinase activity,β-glucan and protein fraction contents.The largest variation and highest average value ofβ-amylase activity were found in Hangzhou 06-07,while the trend of coefficient of variation was opposite to the mean content ofβ-glucan over different environments.Hordein content had the highest correlation coefficient withβ-amylase activity among four protein fractions.The great variation of these traits in Tibetan wild barley will be helpful for improving modern malting barley or utilization of barley as health food in breeding.
3.Effects of the timing of nitrogen application on grain weight,protein fraction content andβ-amylase activity in barley
The effects of four timing treatments of nitrogen fertilizer application with the same total nitrogen rate on grain weight,protein fraction contents andβ-amylase activity in barley cultivar Hua-30 were investigated.The results showed that there was no significant difference in grain weight among the four treatments.However the increasing proportion of nitrogen application at later growth stage caused the dramatic increase of hordein and glutelin contents andβ-amylase activity,while did not alter the albumin and globulin contents.It may be suggested that the influence of increased nitrogen application at later growth stage on malt barley quality is dual,i.e.improving quality through enhancingβ-amylase activity and deteriorating quality through increasing protein content.
4.The influence of nitrogen application at booting stage on protein fraction and hormonal changes during grain filling
The influence of nitrogen application at booting stage on protein fraction and hormone content change during grain filling was investigated using a European barley cultivar Triumph and its ABA mutant TL43 as the experimental materials.The results showed that TL43 had lower grain weight and higher protein content in comparison with Triumph in all nitrogen application treatments.The higher rate of nitrogen application at booting stage resulted in significant increase of protein content in grains. However the influence of the nitrogen application treatment on protein fraction contents varied with genotype,all four fraction contents being significantly increased for TL43,and only glutelin content being significantly affected for Triumph.There was little difference in GA3 content between the genotypes and among the nitrogen treatments.However significant difference was detected in ABA and ZA contents between the two genotypes.Nitrogen application at booting stage caused a marked reduction of ABA content at later grain filling stage.
5.The interaction between H_2O_2 and ABA onβ-amylase accumulation under osmotic stress during seed development in barley
The effects of exogenous ABA and PEG treatments on grain H_2O_2,ABA andβ-amylase accumulation were studied during seed development in Triumph/TL43 using a spike cultural experiment.Triumph and TL43 showed the similar pattern of ABA changes,but different patterns of H_2O_2 andβamylase accumulation during grain filling.Theβamylase accumulation was positively correlated with H_2O_2, negatively correlated with ABA,and mainly determined by the time of high ABA: H_2O_2 stage.The PEG treatment enhancedβamylase synthesis,which was associated with H_2O_2 but not closely related to ABA.Exogenous application of H_2O_2 and AsA enhanced theβamylase activity in Triumph,but reduced that in TL43.However,the endogenous H_2O_2 content in grains was consistent withβamylase accumulation.A novel model may be speculated according to the results that H_2O_2 and ABA are participated in the synthesis ofβ-amylase when barley plants are exposed to temperature stress.
6.Genetic Mapping of Quantitative Trait Loci Associated withβ-amylase,limit dextrinase activities,β-glucan and protein fraction contents in Barley
Interval mapping was utilized to detect quantitative trait loci(QTLs) controlling some malt quality traits using a doubled-haploid population developed from a cross of a six-rowed(cv CM 72) and two-rowed(cv Gairdner) cultivars.A total of nine QTLs for eight traits were mapped to chromosomes 3H,4H,5H and 7H,respectively,with five QTLs on chromosome 3H,indicating an important role of 3H on malt quality. The phenotypic variation accounted by individual QTL ranged from 8.08%to 30.25%. The loci of QTLs forβ-glucan and limit dextrinase were identified on chromosome 4H and 5H,respectively.QTL for hordeins was coincident with the region of silica eluate(SE) protein on 3HS.One locus on 3H was found to be related toβ-amylase and two loci on 5H and 7H were found to be associated with glutelins.
1.栽培大麦品质性状的基因型差异及灌浆后期的温度效应
研究了148个栽培大麦品种籽粒β-淀粉酶活性、蛋白质总量及蛋白组分含量的基因型变异,并利用离体穗培养技术研究了两个大麦品种在不同灌浆后期温度下的籽粒品质变化。β-淀粉酶活性、蛋白质总量及蛋白组分含量的基因型变异显著,β-淀粉酶活性与蛋白质总量及四种蛋白组分含量呈显著正相关,以与醇溶蛋白的相关性最高。与灌浆后期适温(22℃白天/16℃黑夜)相比,高温(32℃白天/26℃黑夜)显著提高β-淀粉酶活性及蛋白质组分含量,清蛋白含量变化较小,而谷蛋白含量影响最大。高温影响A型淀粉粒形状和B型淀粉粒数量,启示出淀粉结构变化可能是高温恶化麦芽品质的原因之一。
2.西藏野生大麦β-淀粉酶活性、极限糊精酶活性、β-葡聚糖及蛋白组分含量的基因型与环境变异
为研究西藏野生大麦籽粒品质性状的基因型与环境变异,测定了76个西藏野生大麦基因型在杭州、武汉两地的β-淀粉酶活性、β-葡聚糖和蛋白质组分含量。籽粒β-淀粉酶活性、β-葡聚糖和蛋白组分含量均存在显著的基因型差异,杭州2006-2007年度种植条件下的β-淀粉酶活性均值和变异幅度最高,且β-淀粉酶活性和β-葡聚糖含量的变异系数和变化趋势恰好相反。四种蛋白质组分中,醇溶蛋白含量与β-淀粉酶活性呈显著正相关。研究鉴定和筛选到β-淀粉酶活性和β-葡聚糖含量表现独特的种质材料,显示出可用于栽培大麦麦芽品质性状的改良。
3.氮肥运筹对栽培大麦粒重、蛋白组分含量和β-淀粉酶活性影响
研究了在总用氮量相同条件下四种氮肥运筹处理对栽培大麦品种花30粒重、蛋白质组分含量和β-淀粉酶活性的影响。不同氮肥运筹处理之间,大麦粒重无显著差异;增加后期用氮比例显著增加籽粒的醇溶蛋白、谷蛋白含量和β-淀粉酶活性,但对清蛋白和球蛋白含量无显著影响,说明提高大麦生育后期的用氮比例对啤用品质的影响是双重的,即在提高蛋白质含量的同时增加了β-淀粉酶活性。
4.孕穗期氮肥对大麦籽粒蛋白质组分和激素含量动态的影响
以欧洲大麦栽培品种Triumph及其ABA突变体TL43为材料,研究孕穗肥对籽粒灌浆期蛋白质组分和激素含量动态变化的影响。不同氮肥处理下,均表现TL43粒重较低但蛋白质含量较高;孕穗期高氮处理显著提高籽粒蛋白质含量,但对蛋白质组分含量的影响,两供试材料表现不同,TL43四种蛋白组分含量都显著提高,而Triumph仅为谷蛋白含量显著增加;籽粒灌浆期,两供试材料之间以及不同氮肥处理之间,GA_3含量变化差异较小;ABA和ZR含量的变化两供试材料之间差异明显,孕穗期施用氮肥显著降低灌浆后期的ABA含量。
5.大麦灌浆期H_2O_2与ABA互作对β-淀粉酶累积的调控机理
利用离体穗培养试验研究外源ABA及PEG处理对大麦品种Triumph及其突变体TL43灌浆期籽粒H_2O_2、ABA含量动态变化及β-淀粉酶的积累。两个基因型的籽粒ABA含量动态变化一致,而H_2O_2与β-淀粉酶的积累差异较大。β-淀粉酶的积累与H_2O_2含量呈正相关,与ABA含量呈负相关且与高ABA:H_2O_2持续时间有关。PEG提高β-淀粉酶活性主要与H_2O_2有关,而与ABA无关。外源添加H_2O_2和AsA显著提高Triumph的β-淀粉酶活性,但降低TL43的β-淀粉酶活性。籽粒内源H_2O_2含量动态始终与β-淀粉酶累积一致。结果显示出,胁迫条件下H_2O_2和ABA参与β-淀粉酶合成的调节。
6.大麦β-淀粉酶、极限糊精酶活性、β-葡聚糖和蛋白组分含量的QTL分析
利用六棱大麦品种CM72和二棱品种Gairdner为亲本,杂交F1代经花药培养获得的93个加倍单倍体(DH群体)株系,通过SSR标记构建连锁图谱,进行β-淀粉酶、极限糊精酶活性、β-葡聚糖和蛋白组分含量的QTL分析。8个品质性状共定位到9个QTLs,分别位于3H,4H,5H及7H染色体上,其中3H染色体上定位到5个QTLs,说明3H染色体对于麦芽品质有重要作用。各个QTL的表型变异范围在8.08%-30.25%之间。β-葡聚糖含量与极限糊精酶活性分别定位在4H、5H染色体上。醇溶蛋白QTL与已报道的硅流蛋白(Silica Eluate Protein)位点重合。β-淀粉酶活性的QTL定位在3H染色体上,而谷蛋白含量的QTL则定位在5H与7H染色体上。
Barley is mainly used as the materials for beer and feed production,and its quality is of important for the commercial values of the final product,thus the improvement of barley quality has been placed into a major objective of barley breeding and agronomy studies.The traits of barley quality belong to the quantitative characters, indicating that they are not only controlled by complex genetic factors,but also greatly affected by environmental factors.In the current study,the genotypic and environmental variation ofβ-amylase and limit dextrinase activities,β-glucan and protein fraction contents in barley grains and the influence of timing of nitrogen application on these malt quality characters were investigated;In addition,the possible mechanisms of P-amylase activity regulation was illustrated by using a barley mutant TL43 and its wild parent Triumph as experimental materials,and QTL analysis of these quality traits was conducted by using a double haploid population.
The main results were as follows:
1.The variation ofβ-amylase activity and protein fractions in barley grains as affected by genotypes and post-anthesis temperatures
The variation ofβ-amylase activity and protein fractions in barley grains was evaluated among 148 barley genotypes grown in the same field condition and using in vitro culture of two barley cultivars under different temperatures during grain development.The results showed that there were considerable genotypic variations inβ-amylase activity and protein fractions.Correlation analysis indicated thatβ-amylase activity was positively correlated with total protein and all protein fraction contents, with the highest correlation coefficient for hordeins.Furthermore,higher post-anthesis temperature(32/26℃,day/night) significantly enhancedβ-amylase activity and protein fractions,with albumins and glutelins being least and most affected respectively,in comparison with the plants under lower temperature(22/16℃).The high temperature also altered the morphology of A granules and number of B granules, suggesting starch structure change may also be a reason for deteriorated malting quality under high temperature.
2.Genotypic and environmental variation ofβ-amylase,limit dextrinase activity,β-glucan and protein fraction contents in Tibetan wild barley
The effects of genotypes and environments(locations and years) onβ-amylase,limit dextrinase activity,β-glucan and protein fraction contents in Tibetan wild barley grown in Hangzhou for two years and Wuhan for one year were investigated in this study.Significant genotypic differences and great variations were detected in grainβ-amylase,limit dextrinase activity,β-glucan and protein fraction contents.The largest variation and highest average value ofβ-amylase activity were found in Hangzhou 06-07,while the trend of coefficient of variation was opposite to the mean content ofβ-glucan over different environments.Hordein content had the highest correlation coefficient withβ-amylase activity among four protein fractions.The great variation of these traits in Tibetan wild barley will be helpful for improving modern malting barley or utilization of barley as health food in breeding.
3.Effects of the timing of nitrogen application on grain weight,protein fraction content andβ-amylase activity in barley
The effects of four timing treatments of nitrogen fertilizer application with the same total nitrogen rate on grain weight,protein fraction contents andβ-amylase activity in barley cultivar Hua-30 were investigated.The results showed that there was no significant difference in grain weight among the four treatments.However the increasing proportion of nitrogen application at later growth stage caused the dramatic increase of hordein and glutelin contents andβ-amylase activity,while did not alter the albumin and globulin contents.It may be suggested that the influence of increased nitrogen application at later growth stage on malt barley quality is dual,i.e.improving quality through enhancingβ-amylase activity and deteriorating quality through increasing protein content.
4.The influence of nitrogen application at booting stage on protein fraction and hormonal changes during grain filling
The influence of nitrogen application at booting stage on protein fraction and hormone content change during grain filling was investigated using a European barley cultivar Triumph and its ABA mutant TL43 as the experimental materials.The results showed that TL43 had lower grain weight and higher protein content in comparison with Triumph in all nitrogen application treatments.The higher rate of nitrogen application at booting stage resulted in significant increase of protein content in grains. However the influence of the nitrogen application treatment on protein fraction contents varied with genotype,all four fraction contents being significantly increased for TL43,and only glutelin content being significantly affected for Triumph.There was little difference in GA3 content between the genotypes and among the nitrogen treatments.However significant difference was detected in ABA and ZA contents between the two genotypes.Nitrogen application at booting stage caused a marked reduction of ABA content at later grain filling stage.
5.The interaction between H_2O_2 and ABA onβ-amylase accumulation under osmotic stress during seed development in barley
The effects of exogenous ABA and PEG treatments on grain H_2O_2,ABA andβ-amylase accumulation were studied during seed development in Triumph/TL43 using a spike cultural experiment.Triumph and TL43 showed the similar pattern of ABA changes,but different patterns of H_2O_2 andβamylase accumulation during grain filling.Theβamylase accumulation was positively correlated with H_2O_2, negatively correlated with ABA,and mainly determined by the time of high ABA: H_2O_2 stage.The PEG treatment enhancedβamylase synthesis,which was associated with H_2O_2 but not closely related to ABA.Exogenous application of H_2O_2 and AsA enhanced theβamylase activity in Triumph,but reduced that in TL43.However,the endogenous H_2O_2 content in grains was consistent withβamylase accumulation.A novel model may be speculated according to the results that H_2O_2 and ABA are participated in the synthesis ofβ-amylase when barley plants are exposed to temperature stress.
6.Genetic Mapping of Quantitative Trait Loci Associated withβ-amylase,limit dextrinase activities,β-glucan and protein fraction contents in Barley
Interval mapping was utilized to detect quantitative trait loci(QTLs) controlling some malt quality traits using a doubled-haploid population developed from a cross of a six-rowed(cv CM 72) and two-rowed(cv Gairdner) cultivars.A total of nine QTLs for eight traits were mapped to chromosomes 3H,4H,5H and 7H,respectively,with five QTLs on chromosome 3H,indicating an important role of 3H on malt quality. The phenotypic variation accounted by individual QTL ranged from 8.08%to 30.25%. The loci of QTLs forβ-glucan and limit dextrinase were identified on chromosome 4H and 5H,respectively.QTL for hordeins was coincident with the region of silica eluate(SE) protein on 3HS.One locus on 3H was found to be related toβ-amylase and two loci on 5H and 7H were found to be associated with glutelins.
引文
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