发掘人工合成小麦中增加千粒重的QTL
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摘要
小麦是世界上最重要的粮食作物之一,由于瓶颈效应与长期选择,使现代栽培小麦的遗传基础狭窄。小麦野生近缘种中存在有大量可用于小麦改良的优异基因。因此,发掘并利用小麦野生近缘种中的优异基因,有利于拓宽栽培小麦的遗传基础。千粒重是小麦产量的构成因素之一,进一步提高千粒重是小麦超高产育种的一条重要途径。然而千粒重与穗粒数和穗数通常呈负相关,与株高正相关,成为小麦超高产育种的重要限制因素。本研究利用四倍体波斯小麦(2n=28,AABB)与粗山羊草Ae38(2n=14,DD)杂交人工合成的六倍体小麦Am3为供体亲本,普通小麦莱州953为轮回亲本,经5次回交的两个F_(2:3)次级分离群体(高千粒重群体85个家系和低千粒重群体75个家系)为材料,通过分子标记检测,发掘人工合成小麦中增加千粒重的QTL,并对其它农艺性状进行QTL定位,为QTL的进一步精细定位和克隆等研究奠定基础。主要结果如下:
1.采用SSR荧光标记分析技术和银染技术相结合的方法用648对不同来源的SSR引物对亲本莱州953和Am3进行多态性检测,其中348对引物在两亲本间有稳定的多态性,多态性频率为53.7%。有66对SSR引物在高千粒重导入系群体中出现多态性,占多态性标记的18.96%。有63对SSR引物在低千粒重导入系群体中出现多态性,占多态性标记的18.1%。
2.利用Map Manager软件,将高千粒重导入系群体的58个多态性SSR标记定位在小麦8条染色体上,连锁图总长为344.6cM,平均每个标记间的遗传距离为5.9cM。同样的方法将低千粒重导入系群体的52个多态性SSR标记定位在小麦9条染色体上,连锁图的总长为366.9cM,平均每个标记间的遗传距离为7.1cM。导入片段远远大于理论值。
3.利用Windows QTL Cartographer 2.0软件,采用复合区间作图法取LOD≥2.5为阈值,对两个群体进行QTL检测。在高千粒重群体中共检测到10个QTL,其中3个千粒重QTL分别位于1A、3D和4B染色体上,1个粒长QTL位于7B染色体上,2个粒宽QTL分别位于1A、和4B染色体上,2个穗数QTL分别位于4B和7B染色体上,2个株高QTL分别位于3D和4B染色体上。在低千粒重群体中共检测到17个QTL,其中3个千粒重QTL分别位于1A、4B和7B染色体上,4个粒长QTL分别位于1A和2D染色体上,1个粒宽QTL位于7B染色体上,2个穗数QTL分别位于4B和7B染色体上,2个株高QTL分别位于2D和4B染色体上,5抽穗期QTL分别位于2A、2D、6B和7B染色体上。
4.在高千粒重导入系群体中,共检测到3个来自Am3增加千粒重的新的QTL位点(QGw.caas-1A1、QGw.caas-3D、QGw.caas-4B1)。QGw.caas-3D在3个环境中都检测到,解释表型变异18.1%-31.8%,增加千粒重2.3-4.8 g,表明该QTL是一个稳定的主效QTL。QGw.caas-1A1在2个环境中检测到,解释表型变异21.4%-33.8%,增加千粒重2.7-3.8 g。QGw.caas-4B1在2个环境中检测到,解释表型变异10.9%-30.2%,增加干粒重3.9-4.8 g。所发现的增加千粒重QTL与穗数、穗粒数和株高不存在不利的相关关系,有利于千粒重、穗数、穗粒数的同步提高及株高的降低,因此是较为理想的提高千粒重的QTL。
5.虽然轮回亲本Am3的农艺性状比莱州953的差,但在导入系群体中,通过表型选择和QTL分析,选出了一些农艺性状比轮回亲本莱州953有较大改良的导入系。如高千粒重导入系IL6的平均千粒重为60.5g,平均穗粒数53.3粒,平均株高83.6cm,平均单株穗数5.8个;多粒导入系IL39的平均穗粒数83.6粒,千粒重45.9g,平均穗数5.3个,株高84.3cm。这些导入系已分发到小麦育种单位用于小麦品种的遗传改良,选育含有人工合成小麦有利基因/QTL的小麦新品种。
Common wheat is one of the most important crops in the world.During the long domestication process for common wheat,its genetic diversity has been narrowed by bottleneck effect and selection.Wild species are likely to contain favorable genetic factors that can increase the yield of modern varieties.Broadening the genetic base of common wheat by using exotic germplasm resources is an important germplasm development goal. In common wheat,thousand-gain weight(TGW) is an important component of grain yield. Increasing thousand-gain weight is one way to improve grain yield.However,it is negative correlation frequently with grain number per spike and spike number per plant, positive correlation with plant height.Am3,a synthetic wheat from crossing Triticum carthlicum(2n=28,AABB) with Aegilops tauschii(2n=14,DD),was used as donor lines and Laizhou953,a commercial Chinese cultivar used as recurrent parent in this study. After five generations of backcrossing,two BC_5F_(2:3) populations(high TGW population consisting of 85 lines and low TGW population consisting of 75 lines) were employed for genotyping and phenotyping in the present study.Our goal is to detect the favorable TGW alleles of the synthetic wheat and to fine map QTLs for TGW in the future.The results obtained were listed as below.
1.Two parents,Lalzhou953 and Am3,were screened for polymorphism with 648 SSR markers by using fluorescence and silver-staining detection systems of microsatellite markers.Among them,348 of them detect polymorphism between the parents.66 of the 348 SSR markers detected polymorphism in the high TGW population.The frequency of polymorphism was 18.96%.63of the 348 SSR markers detected polymorphism in the low population.The frequency of polymorphism was 18.1%.
2.Using Map Manager software,a map of 58 markers distributing on 8 chromosomes was constructed in the high TGW population.The total length of the linkage map was 344.6 cM with an average genetic distance of 5.9 cM.In the low population,a map of 52 markers distributing on 9 chromosomes was constructed.The total length of the linkage map was 366.9 cM with an average genetic distance of 7.1 cM.
3.The QTL analysis was performed with software Windows QTL Cartographer 2.0 based on composite interval mapping(CIM).QTL was claimed to be significant at a LOD value of 2.5.A total of 10 putative QTLs were detected in the high TGW population including 3 QTL for TGW located on chromosomes 1A,3D and 4B;1 QTL for grain length located on chromosome 7B;2 QTLs for grain width located on chromosomes 1A and 4B;2 QTLs for spike per plant located on chromosomes 4B and 7B;2 QTLs for plant height located on chromosomes 3D and 4B.A total of 17 putative QTLs were detected in the low TGW population including 3 QTL for TGW located on chromosomes 1A,4B and 7B;4 QTL for grain length located on chromosomes 1A and 2D;1 QTL for grain width located on chromosome 7B;2 QTLs for spike per plant located on chromosomes 4B and 7B;2 QTLs for plant height located on chromosomes 2D and 4B;5 QTLs for day to heading located on chromosomes 2A,2D,6B and 7B.
4.In the high TGW population,3 new QTLs(QGw.caas-1A1、QGw.caas-3D、QGw.caas-4B1) for TGW were detected.QGw.caas-1A1 was detected in two environments with phenotypic variation explained(PVE) from 21.4%to 33.8%and increasing TGW 2.7-3.8g.QGw.caas-3D was detected in three environments with PVE 18.1%-31.8%and increasing TGW 2.3-4.8 g.QGw.caas-4B1 was detected in two environments with PVE 10.9%-30.2%and increasing TGW 3.9-4.8 g.TGW was not corrected to other traits except for GINS.A significant positive correlation between TGW and GNS was found in Luoyang (r=0.232,P＜0.05),but it was not significant in Beijing.QTL mapping revealed that the QTL of TGW detected here were not located in the same regions with the other yield components and plant height except for the Qph.caas-3D,suggested that we can select these favorable traits/QTL separately and pyramid them with the other yield components, such as SP and SN.Those QTLs were not correlated with other agronomic traits.We could select the favorable traits/QTLs separately and pyramid them with the other yield components,so the QTLs were the favorable QTLs conferring increasing thousand-grain weight.
5.Despite inferior agronomic traits of donor parent Am3 to the recurrent parent Laizhou953,in the populations,some of ILs have superior agronomic traits to the recurrent parent Laizhou953,such as IL6 with:TGW 60.5g,GNS 53.3,PH 83.6 cm,SP 5.8;IL39 with:GNS 83.6,TGW 45.9g,SP 5.3,PH 84.3 cm.In this study,some of favorable lines have been distributed to breeders for improving grain yield.
1.采用SSR荧光标记分析技术和银染技术相结合的方法用648对不同来源的SSR引物对亲本莱州953和Am3进行多态性检测,其中348对引物在两亲本间有稳定的多态性,多态性频率为53.7%。有66对SSR引物在高千粒重导入系群体中出现多态性,占多态性标记的18.96%。有63对SSR引物在低千粒重导入系群体中出现多态性,占多态性标记的18.1%。
2.利用Map Manager软件,将高千粒重导入系群体的58个多态性SSR标记定位在小麦8条染色体上,连锁图总长为344.6cM,平均每个标记间的遗传距离为5.9cM。同样的方法将低千粒重导入系群体的52个多态性SSR标记定位在小麦9条染色体上,连锁图的总长为366.9cM,平均每个标记间的遗传距离为7.1cM。导入片段远远大于理论值。
3.利用Windows QTL Cartographer 2.0软件,采用复合区间作图法取LOD≥2.5为阈值,对两个群体进行QTL检测。在高千粒重群体中共检测到10个QTL,其中3个千粒重QTL分别位于1A、3D和4B染色体上,1个粒长QTL位于7B染色体上,2个粒宽QTL分别位于1A、和4B染色体上,2个穗数QTL分别位于4B和7B染色体上,2个株高QTL分别位于3D和4B染色体上。在低千粒重群体中共检测到17个QTL,其中3个千粒重QTL分别位于1A、4B和7B染色体上,4个粒长QTL分别位于1A和2D染色体上,1个粒宽QTL位于7B染色体上,2个穗数QTL分别位于4B和7B染色体上,2个株高QTL分别位于2D和4B染色体上,5抽穗期QTL分别位于2A、2D、6B和7B染色体上。
4.在高千粒重导入系群体中,共检测到3个来自Am3增加千粒重的新的QTL位点(QGw.caas-1A1、QGw.caas-3D、QGw.caas-4B1)。QGw.caas-3D在3个环境中都检测到,解释表型变异18.1%-31.8%,增加千粒重2.3-4.8 g,表明该QTL是一个稳定的主效QTL。QGw.caas-1A1在2个环境中检测到,解释表型变异21.4%-33.8%,增加千粒重2.7-3.8 g。QGw.caas-4B1在2个环境中检测到,解释表型变异10.9%-30.2%,增加干粒重3.9-4.8 g。所发现的增加千粒重QTL与穗数、穗粒数和株高不存在不利的相关关系,有利于千粒重、穗数、穗粒数的同步提高及株高的降低,因此是较为理想的提高千粒重的QTL。
5.虽然轮回亲本Am3的农艺性状比莱州953的差,但在导入系群体中,通过表型选择和QTL分析,选出了一些农艺性状比轮回亲本莱州953有较大改良的导入系。如高千粒重导入系IL6的平均千粒重为60.5g,平均穗粒数53.3粒,平均株高83.6cm,平均单株穗数5.8个;多粒导入系IL39的平均穗粒数83.6粒,千粒重45.9g,平均穗数5.3个,株高84.3cm。这些导入系已分发到小麦育种单位用于小麦品种的遗传改良,选育含有人工合成小麦有利基因/QTL的小麦新品种。
Common wheat is one of the most important crops in the world.During the long domestication process for common wheat,its genetic diversity has been narrowed by bottleneck effect and selection.Wild species are likely to contain favorable genetic factors that can increase the yield of modern varieties.Broadening the genetic base of common wheat by using exotic germplasm resources is an important germplasm development goal. In common wheat,thousand-gain weight(TGW) is an important component of grain yield. Increasing thousand-gain weight is one way to improve grain yield.However,it is negative correlation frequently with grain number per spike and spike number per plant, positive correlation with plant height.Am3,a synthetic wheat from crossing Triticum carthlicum(2n=28,AABB) with Aegilops tauschii(2n=14,DD),was used as donor lines and Laizhou953,a commercial Chinese cultivar used as recurrent parent in this study. After five generations of backcrossing,two BC_5F_(2:3) populations(high TGW population consisting of 85 lines and low TGW population consisting of 75 lines) were employed for genotyping and phenotyping in the present study.Our goal is to detect the favorable TGW alleles of the synthetic wheat and to fine map QTLs for TGW in the future.The results obtained were listed as below.
1.Two parents,Lalzhou953 and Am3,were screened for polymorphism with 648 SSR markers by using fluorescence and silver-staining detection systems of microsatellite markers.Among them,348 of them detect polymorphism between the parents.66 of the 348 SSR markers detected polymorphism in the high TGW population.The frequency of polymorphism was 18.96%.63of the 348 SSR markers detected polymorphism in the low population.The frequency of polymorphism was 18.1%.
2.Using Map Manager software,a map of 58 markers distributing on 8 chromosomes was constructed in the high TGW population.The total length of the linkage map was 344.6 cM with an average genetic distance of 5.9 cM.In the low population,a map of 52 markers distributing on 9 chromosomes was constructed.The total length of the linkage map was 366.9 cM with an average genetic distance of 7.1 cM.
3.The QTL analysis was performed with software Windows QTL Cartographer 2.0 based on composite interval mapping(CIM).QTL was claimed to be significant at a LOD value of 2.5.A total of 10 putative QTLs were detected in the high TGW population including 3 QTL for TGW located on chromosomes 1A,3D and 4B;1 QTL for grain length located on chromosome 7B;2 QTLs for grain width located on chromosomes 1A and 4B;2 QTLs for spike per plant located on chromosomes 4B and 7B;2 QTLs for plant height located on chromosomes 3D and 4B.A total of 17 putative QTLs were detected in the low TGW population including 3 QTL for TGW located on chromosomes 1A,4B and 7B;4 QTL for grain length located on chromosomes 1A and 2D;1 QTL for grain width located on chromosome 7B;2 QTLs for spike per plant located on chromosomes 4B and 7B;2 QTLs for plant height located on chromosomes 2D and 4B;5 QTLs for day to heading located on chromosomes 2A,2D,6B and 7B.
4.In the high TGW population,3 new QTLs(QGw.caas-1A1、QGw.caas-3D、QGw.caas-4B1) for TGW were detected.QGw.caas-1A1 was detected in two environments with phenotypic variation explained(PVE) from 21.4%to 33.8%and increasing TGW 2.7-3.8g.QGw.caas-3D was detected in three environments with PVE 18.1%-31.8%and increasing TGW 2.3-4.8 g.QGw.caas-4B1 was detected in two environments with PVE 10.9%-30.2%and increasing TGW 3.9-4.8 g.TGW was not corrected to other traits except for GINS.A significant positive correlation between TGW and GNS was found in Luoyang (r=0.232,P＜0.05),but it was not significant in Beijing.QTL mapping revealed that the QTL of TGW detected here were not located in the same regions with the other yield components and plant height except for the Qph.caas-3D,suggested that we can select these favorable traits/QTL separately and pyramid them with the other yield components, such as SP and SN.Those QTLs were not correlated with other agronomic traits.We could select the favorable traits/QTLs separately and pyramid them with the other yield components,so the QTLs were the favorable QTLs conferring increasing thousand-grain weight.
5.Despite inferior agronomic traits of donor parent Am3 to the recurrent parent Laizhou953,in the populations,some of ILs have superior agronomic traits to the recurrent parent Laizhou953,such as IL6 with:TGW 60.5g,GNS 53.3,PH 83.6 cm,SP 5.8;IL39 with:GNS 83.6,TGW 45.9g,SP 5.3,PH 84.3 cm.In this study,some of favorable lines have been distributed to breeders for improving grain yield.
引文
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