普通小麦重要品质性状的QTL定位
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摘要
品质改良是我国当前小麦育种的主要目标之一。利用分子数量遗传学方法研究品质相关性状、进行基因定位,可为分子标记辅助选择和品质改良提供理论基础。1B.1R易位系在我国小麦育种中曾发挥过重要作用,但鉴于其对小麦加工品质的负面影响,明确其对小麦品质遗传效应,对1B.1R易位系在育种中的应用具有重要的指导意义。此外,籽粒硬度、高低分子量谷蛋白亚基和面粉颜色对小麦磨粉品质和加工品质也有重要影响。本研究以PH82-2(硬质、非1B.1R)为母本与内乡188(软质、1B.1R)杂交,通过一粒传获得240个F_(5:6)重组自交系,于2004-05及2005-06年度种植于安阳、焦作和泰安,对重要的品质性状,即面粉和中国白盐面条颜色参数及PPO活性、黄色素含量(YPC)、容重(TW)、千粒重(TKW)、面粉蛋白含量(FPC)、mixograph参数及面粉糊化参数(RVA参数)和面条品质性状进行QTL定位,并分析1B.1R易位、高低分子量谷蛋白亚基及籽粒硬度对这些品质性状的遗传效应。主要结果如下:
     1.对面粉和面条的11个颜色参数及其相关性状的QTL结果表明,这些性状主要受5D染色体上籽粒硬度位点(Ha)、1B染色体短臂(short arm ofchromosome 1B,SA1B)或1RS(Sec1)及7A染色体上Psy-A1基因位点影响。
     籽粒硬度位点(Ha)是影响面粉颜色参数、FPC和面粉PPO活性的主要因素之一,尤其对面粉亮度(FL~*)、黄色度(Fb~*)、面粉颜色指标(FCI)和面粉PPO活性,影响最大,贡献率分别为80.3%、57.9%、73.2%和55.1%,但对面粉a~*(Fa~*)影响很小,贡献率仅为2.0%。Ha位点对籽粒硬度的贡献率为75%。
     1RS和Psy-A1基因位点是造成面粉黄色素含量高(YPC)及导致面粉、面条偏黄(Fb~*和Nb~*)的主要因素;1RS对它们的贡献率为31.9%、7.8%和13.7%,而Psy-A1基因位点为33.9%、12.6%和22.0%。此外,1RS和Psy-A1位点对面粉a~*(Fa~*)和Kent-Jones颜色等级(KJ)影响也非常大,对Fa~*的贡献率分别为26.1%和35.9%,对KJ的贡献率17.2%和19.3%;但对面条亮度(NL~*)影响较小,贡献率仅为4.5%和4.3%;1RS对FCI和面条a~*(Na~*)没有影响,Psy-A1基因对它们的影响也非常小。
     2A染色体的Ppo-A1基因位点对PPO活性的贡献率为20.2%;除此之外,籽粒硬度位点对PPO活性表现出更大的贡献率,达55.1%。;在3A和3B染色体上分别检测到一个控制面粉蛋白含量的QTL,贡献率为5.9%和4.9%;在1A、3B和4A上发现影响YPC、Fa~*和KJ的QTL,其中4A上影响YPC的QTL在各环境中均可稳定检测到;在7B染色体上与Psy-A1基因的同源位置上也发现一个影响FCI的QTL。
     2.对15个品质性状参数进行QTL分析,结果表明麦谷蛋白亚基位点(Glu-B1、Glu-D1和Glu-A3)、SA1B或1RS和Ha位点是影响这些品质参数的主效位点。
     控制容重(TW)和千粒重(TKW)的QTL主要位于小麦染色体的第4、5、6同源群以及7A染色体,但效应都比较低。共检测到4个QTL影响面粉蛋白含量,分别位于3A、3B和5D(2个),其中Ha位点影响最大,贡献率达27.1%。在控制Zeleny沉降值(ZS)的6个位点中,1RS和Glu-D1位点影响最大,贡献率为19.3%和17.6%。
     Mixograph参数主要受麦谷蛋白亚基位点及SA1B的影响。Glu-D1位点对峰值时间(MPT)和弱化值(Weakening slope,WS)贡献率最大,分别为43.1%和39.7%,同时对峰值宽度(MEW)和8min带宽(MTxW)影响也较大,贡献率分别为16.2%和24.2%。SA1B对8min带宽影响最大,贡献率为42.2%,而对峰值宽度和WS影响则较小,贡献率为9.8%和11.1%。Glu-B1位点对峰值时间影响较大,贡献率为18.0%,而对峰值宽度、8min带宽和WS贡献率为10.3%、11.0%和14.2%。Glu-A3位点对这些mixograph参数影响都较小,贡献率为5.9%~10.0%。此外,Ha对峰值宽度也有较大,贡献率为21.8%。
     Ha对RVA参数,峰值黏度(RPV)、最终黏度(RFV)、反弹值(RSb)都有重要影响,尤其对糊化温度(RPT)影响最大,贡献率达71.5%。此外,在1A、5B和7D上都检测到控制峰值黏度的QTL;在1B、3B、5B、6D和7A上都发现控制最终黏度的QTL,其中,1B上的QTL影响最大,贡献率为21.6%。在1B、3B、4B和7A上检测到影响反弹值的QTL,但效应均较小。对稀澥值(RBd)只检测到两个QTL,分别位于SA1B和6D上,贡献率分别为5.0%和6.3%。
     控制面条黏着性的QTL位于1A、1B和4D染色体,其中1B染色体上的QTL与Glu-B1有关,贡献率为8.8%。影响面条弹性的2个QTL分别位于1B和5D(Ha)位点上,贡献率分别为9.4%和8.1%。
Improvement of wheat quality is very important for vavious wheat-based end-products,and has become a main objective of wheat breeding in China.QTL mapping for quality-related traits would provide a theoretical basis for molecular marker-assisted selection in wheat breeding program.Although the 1B.1R transloction has positive effects on many agronomic traits,it largely reduces wheat quality.Therefore,it is important to study the genetic bases of quality-related traits and estimate the genetic effects of the 1B.1R translocation on yield traits for developing wheat cultivars with high yield and good quality.In addition,grain hardness,high- and low-molecular-weight glutenin subunits and flour color showed large influences on wheat processing quality.The objectives of this study were to map quantitative trait loci(QTL) for color parameters of flour and Chinese white salted noodle(CWSN),polyphenol oxidase(PPO) acitivity,yellow pigment content(YPC), milling traits,gluten quality,flour pasting properties,and Chinese white salted noodle (CWSN) qualities,and investigate the genetic effects of 1B.1R translocation and grain hardness,using a recombinant inbred line(RIL) population derived from a cross between the Chinese wheat cultivars 'PH82-2'(hard and non-1B.1R translocation) and 'Neixiang 188'(soft and 1B.1R translocation) planted in Anyang,Jiaozuo and Taian in 2004-05 and 2005-06 cropping seasons.The main results are summarized below.
     1.The color parameters of flour and noodles,PPO and YPC were mainly controlled by three genomic regions or loci,grain hardness loci(Ha) on chromosome 5DS,short arm of chromosome 1B(SA1B) or 1RS(Sec1) and Psy-A1 genes on chromosome 7AL.
     Ha locus had large influences on flour brightness(FL~*),flour yellowness(Fb~*), flour color index(FCI) and PPO activity,explaining 80.3%,57.9%,73.2%and 55.1% of the phenotypic variance,respectively and additionally on grain hardness,accounted for 75.2%of the phenotypic variance.
     SA1B(1RS) and Psy-A1 genes showed large effects on YPC,Fb~* and Nb~*, accounting for 31.9%and 33.9%,7.8%and 12.6%,and 13.7%and 22.0%of the phenotypic variance,respectively.They also had large effects on flour color parameter a~*(Fa~*) and Kent-Jones color grade(KJ),explaining 26.1%and 35.9%,and 17.2% and 19.3%of the phenotypic variance,respectively,and on noodle brightness(NL~*), accounting for 4.5%and 4.3%of the phenotypic variance,respectively.However, 1RS had no effect on FCI and noodle color paremeter a~*(Na~*),and Psy-A1 genes indicated a small effects on them.
     The major QTL for PPO activity was located on chromosome 2A,explaining 22.0%of the phenotypic variance,however,Ha locus accounted for the largest effect on PPO activity(55.1%).Two QTL for FPC were mapped on chromosomes 3A and 3B,accounting for 5.9%and 4.9%of the phenotypic variance.Chromosomes 1A,3B and 4A were also associated with YPC,Fa~* and KJ,among which,the QTL on chromosome 4A was stably detected across all environments.A QTL for FCI was located on chromosome 7B in homoeologous region of Psy-A1 genes locus on chromosome 7A.
     2.Glutenin loci(Glu-B1,Glu-D1 and Glu-A3),SA1B(or 1RS) and Ha locus were mainly contributed to 16 quality parameters.
     Nine QTL for TW and TKW were detected on homoeologous groups 4,5 and chromosomes 6D and 7A.Four QTL assocated FPC were mapped on chromosomes 3A,3B and 5D,with the mojor QTL associated with Ha locus explaining 27.1 of the phenotypic variance.Among the six QTL for Zeleny sedimentation value,1RS and Glu-D1 showed large effects,accounting for 19.3%and 17.6%of the phenotypic variance.
     For mixograph parameters,Glu-D1 locus contributed large effects on peak time (MPT) and weakening slope(WS),followed on peak width(MPW) and 8 min width (MTxW),explaining 41.3%,39.7%,16.2%and 24.2%of the phenotypic variance, respectively;SA1B had largest contribution to MTxW,followed on MPW and WS, accounting for 42.2%,9.8%and 11.1%of the phenotypic variance,respectively; Glu-B1 showed a large effect on MPT and relatively less on MPW,MTxW and WS, explaining 18.0%,10.3%,11.0%and 14.2%of the phenotypic variance,respectively; Glu-A3 locus showed less influence on mixograph parameters,explaining from 5.9% to 10.0%of the phenotypic variance.In addition,Ha locus had large effect on MPW, accounting for 21.8%of the phenotypic variance.
     Ha showed important effects on flour pasting parameters,RVA peak viscosity (RPV),RVA final viscosity(RFV) and RVA setback(RSb),especially on pasting temperature(RPT),explaining 71.5%of the phenotypic variance.In addition, chromosomes 1A,5B and 7D were also associated with RPV.Five QTL were detected on chromosomes 1B,3B,5B,6D and 7A,with the one on chromosome 1B explaining 21.6%of the phenotypic variance.QTL for RSb were mapped on chromosomes 1B, 3B,4B and 7A with little effects.Moreover,two QTL for RVA breakdown(RBd) were found on SA1B and chromosome 6D.
     Three QTL for Chinese white salted noodle(CWSN) adhesiveness were detected on chromosomes 1A,1B and 4D,with the one associated with Glu-B1,explaining 8.8%of the phenotypic variance.Two QTL on chromosomes 1B and 5D were detected for CWSN springiness,with the one associated with Ha locus accounting for 8.1%of the phenotypic variance.
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