大麦新矮源“华矮11”主要性状的遗传分析和大麦抽穗期性状的QTL定位
摘要
大麦是我国的第四大粮食作物,对国民经济的发展和人民生活水平的提高有重要作用。社会的发展和人们物质文化生活水平的提高要求培育出产量更高、品质更优、抗性更强、适应性更广的大麦新品种,加强大麦种质资源和重要性状的遗传研究是满足这种需求的可行之道。
矮杆抗倒是大麦育种的目标性状之一,矮杆种质或基因是开展大麦矮化育种成功与否的关键所在。在大麦中已经发现了约30多个不同类型的矮杆种质,然而真正能成功应用于育种的极少。大麦育种上矮源或矮杆基因的单一化,不仅使得产量和品质的进一步提高受到限制外,而且还使新品种的遗传基础变窄,多样性降低,增强了对病、虫害的脆弱性,筛选及创造新的矮杆种质,对大麦遗传改良具有重要的理论意义与实用价值。本试验所研究的新矮源“华矮11”是一源于青藏高原地方品种的特矮秆种质,株高只有40厘米,并具有早熟和一般配合力好等特性,它的发现将拓宽大麦矮化育种的遗传基础。对“华矮11”的矮秆性进行了多年、多组合的遗传分析和等位性测定,并构建了“华矮11×华大麦6号”的DH群体。以此DH群体为定位群体,对“华矮11”的矮秆基因和部分农艺、品质性状的QTL进行了定位。
抽穗期是大麦区域和季节适应性的主要决定因素。在育种上,合适的抽穗期是大麦育种的重要目标。普遍认为种质的相互引进在相似的纬度较易成功。然而,原产于东南亚的大麦种质材料引入澳大利亚后,抽穗期提前,表现早熟,反之亦然。以Galleon×Haruna Nijo和Baudin×AC Metcalfe DH群体为定位群体,对其抽穗期基因进行定位,期望阐明在相似的纬度条件的澳大利亚和中国环境下抽穗期不同的主要原因。
本研究主要结果如下：
1.“华矮11”的矮秆性遗传：以“华矮11”为母本与9个高秆品种进行杂交,株高调查结果显示,其F1代均为高秆,而F2代则出现高秆矮秆性状分离,进一步对这9个F2分离群体的株高分离情况进行χ2检验,结果表明,所有组合的F2群体中高秆个体和矮秆个体的分离比例均符合3：1的分离规律。由此推定,华矮11的矮秆性状是受一对隐性基因控制,命名为btwd1。
2.矮秆基因btwdl的遗传等位测验：以华矮11/Monker的F1代做母本与其它4个矮秆品种杂交,后代全为高秆。表明矮秆基因btwdl与已知的矮秆基因br,uzu,sdwl和denso明显不同。
3.矮秆基因btwdl的定位：根据遗传分析结果,采用SSR标记与BSA分析方法相结合,对华矮11矮秆基因进行标记定位。结果表明来自于7H染色体的4个SSR标记(Bmac031, Bmac167, Bmag217和Bmag900)与目的基因btwdl构建成一个连锁群。该矮秆基因btwdl被定位于7H染色体长臂靠近着丝粒一侧,距离Bmac031为2.2 cM。
4.农艺性状和品质性状的QTL定位：以DH群体华矮11×华大麦6号为研究材料,在全生育期调查了农艺性状如抽穗期、有效穗、主穗长、主穗小穗数、单株小穗数、单株实粒数、穗平实粒数、单株粒重、穗平粒重、千粒重、分蘖数和品质性状蛋白质含量。选用63对在两亲本间有多态性的引物构建遗传连锁图谱。采用复合区间作图法对12个性状进行QTL定位分析,共检测到47个相关QTL,其中29个为新的QTL。
5.大麦抽穗期基因定位：以Galleon×Haruna Nijo DH群体作为研究材料,在中国(武汉,30°33'N)和澳大利亚(珀斯,31°56'S)设置3个不同的处理,3个处理包括正常大田秋播、大田晚播和温室18小时光照。通过对群体抽穗期的QTL定位分析结果表明,在中国环境下,探测到1个控制抽穗期的主效QTL(Qtl-5H)位于大麦染色体5H上,该QTL解释了抽穗期33%-50%的表型变异；在澳大利亚环境下,探测到2个控制抽穗期的主效QTL(Qtl-4H和Qtl5H),且Qtl-4H和Qtl-5H存在互作。它们的互作解释了抽穗期13%-36%的表型变异。Qtl-4H与Qtl-5H的互作可能是在澳大利亚环境下比在中国环境下群体株系更早抽穗的主要原因。
6.大麦长光周期基因的定位：以Baudin×AC Metcalfe DH群体作为研究材料,在中国(武汉,30°33'N)设置3个不同的处理,3个处理包括正常大田秋播、大田晚播和温室18小时光照,在澳大利亚(珀斯,31°56'S)设置2个不同的处理,2个处理包括正常大田秋播和温室18小时光照。通过对群体株系的抽穗期QTL定位分析结果表明,1个长光周期主效QTL(Qtl4-13)被定位在4H染色体上引物Xp12m50A199-Xp13m47B399之间。该QTL在澳大利亚和中国环境下分别解释了77.48%和37.81%的表型变异,表明它为大麦的一个新的长光周期基因。同时还定位了2个抽穗期相关QTL。
Barley is the fourth food crop in China and plays a very important role in the development of economy. To meet further social and people's living standards, new barley varieties will require high yields, better quality, high resistance and wide adaptability. Genetic characterization of barley germplasm will be crucial for the development of new barley varieties.
Dwarfism is a valuable trait in crop breeding, successful use of a dwarfing gene or dwarf source is critical for developing dwarf cultivars. In barley, more than 30 types of dwarfs or semidwarfs have been found. However, only a few of them have successfully been used in barley breeding program. Dwarf source and dwarf genes are unitary in barley breeding, not only limited further increases in yield and quality, but also narrowed the genetic basis, reduced biodiversity and increased vulnerability of disease and pest for new varieties. Therefore, discovery of new dwarf genes or dwarf sources is important for barley breeding. New dwarf source "huaai 11" has early maturity and good general combining ability. Utilization of this dwarf resource can broaden the genetic basis of barley dwarf breeding. We mapped dwarfing gene btwdl in Huaai 11 using a doubled-haploid (DH) population from a cross between Huaai 11 and Huadamai 6. In order to efficiently use this excellent germplasm for breeding program, we also analyzed the QTL underlying agronomic traits and quality traits in this line.
Heading date is a major determinant of the regional and seasonal adaptation of barley varieties. Appropriate heading date is an important goal of barley breeding. The dogma is that introduced germplasm is more likely to be adapted if it come from a similar latitude. However, barley germplasm introduced from similar latitudes of South-East Asia are extremely early heading in the Australian environments and vice versa. Therefore, we mapped heading date genes using Galleon×Haruna Nijo DH population and Baudin×AC Metcalfe DH population. The results will help us understand the main reason causing heading date difference in two similar latitude locations, Australia and China, The results of these researches are sumarized as follows:
1. Inheritance of the dwarf gene btwdl:dwarf Huaai 11 was crossed with nine tall varieties. All the F1 plants were tall. Both the tall and dwarf plants appeared in all the F2 populations with a 3:1 segregation ratio. The x2 testshowed the segregation is not significantly different from the expected 3:1 ratio, indicating that the dwarf trait is controlled by single recessive gene in the Huaai 11. This gene is designated as btwdl.
2. Allelic relationship between btwdl and other dwarfing genes:The F1 of Huaai 11 x Monker was crossed with other four dwarf varieties. All progeny in the four crosses are tall, suggesting that the gene btwdl controlling plant height in the Huaai 11 is nonallelic with br, uzu, sdwl and denso genes in these four dwarf varieties.
3. Mapping of the dwarf gene btwdl:SSR marker and BSA analysis were used to map the dwarfing gene in the Huaaill. Linkage analysis found that four SSR markers from chromosome 7H (BmacO31, Bmacl67, Bmag217 and Bmag900) were linked with the target gene btwd1. The dwarfing gene btwdl is close to the Bmac031 with a genetic distance of 2.2 cM, which is close to the centromere of 7HL.
4. QTL analysis of the agronomic and quality traits:This study using Huaai 11×Huadamai 6 DH population as research materials, Quality trait grain protein content and eleven gronomic traits such as heading date, spike numbers per plant, main spike length, spikelet number of main spike, spikelet number per plant, grain number per plant, grain number per spike, grain weight per plant, grain weight per spike,1000 grain weight, number of tillers were investigated. Sixty three pairs of SSR primers showed polymorphic between the two parents, and were used to construct a genetic linkage map. Composite interval mapping (CIM) analysis of 12 traits identifiedt a total of 47 QTL,29 of them were new QTL.
5. Mapping of heading date gene in barley:The Galleon x Haruna Nijo DH population was used to map heading date gene at two location China (Wuhan,30°33'N) and Australia (Perth,31°56' S) with three different treatments including normal autumn sowing, late sowing and pot experiment with 18h photoperiod. QTL analysis detected one major QTL (Qtl-5H) in China, which was located on chromosome 5H, and explained 33%-50% of the phenotypic variation. In Australia, two major QTL (Qtl-4H and Qtl5H) were detected for heading dates on chromosomes 4H and 5H and Qtl-4H and Qtl5H interaction. The Qtl-4H/Qtl-5H interaction explained 13%-36% of phenotypic variation. The Qtl-4H/Qtl-5H interaction may be main reason that DH lines are earlier heading in Australia than China.
6. Mapping of long photoperiod gene in barley:Bandin×AC Metcalfe DH population was used. In China (Wuhan,30°33'N), three different treatments including normal autumn sowing, late sowing and pot experiment with 18h photoperiod were conducted. In Australia (Perth,31°56' S), two different treatments including normal autumn sowing and pot experiment with 18h photoperiod were performed. QTL analysis detected a major QTL for photoperiod response and mapped onto the long arm of chromosome 4H in both Australian and Chinese locations. The QTL was flanked by the markers Xp12m50A199-Xp13m47B399 and accounted for 77.5% and 37.8% of phenotypic variation in Australia and China, respectively. Two QTL underlying heading date detected in this mapping population.
矮杆抗倒是大麦育种的目标性状之一,矮杆种质或基因是开展大麦矮化育种成功与否的关键所在。在大麦中已经发现了约30多个不同类型的矮杆种质,然而真正能成功应用于育种的极少。大麦育种上矮源或矮杆基因的单一化,不仅使得产量和品质的进一步提高受到限制外,而且还使新品种的遗传基础变窄,多样性降低,增强了对病、虫害的脆弱性,筛选及创造新的矮杆种质,对大麦遗传改良具有重要的理论意义与实用价值。本试验所研究的新矮源“华矮11”是一源于青藏高原地方品种的特矮秆种质,株高只有40厘米,并具有早熟和一般配合力好等特性,它的发现将拓宽大麦矮化育种的遗传基础。对“华矮11”的矮秆性进行了多年、多组合的遗传分析和等位性测定,并构建了“华矮11×华大麦6号”的DH群体。以此DH群体为定位群体,对“华矮11”的矮秆基因和部分农艺、品质性状的QTL进行了定位。
抽穗期是大麦区域和季节适应性的主要决定因素。在育种上,合适的抽穗期是大麦育种的重要目标。普遍认为种质的相互引进在相似的纬度较易成功。然而,原产于东南亚的大麦种质材料引入澳大利亚后,抽穗期提前,表现早熟,反之亦然。以Galleon×Haruna Nijo和Baudin×AC Metcalfe DH群体为定位群体,对其抽穗期基因进行定位,期望阐明在相似的纬度条件的澳大利亚和中国环境下抽穗期不同的主要原因。
本研究主要结果如下：
1.“华矮11”的矮秆性遗传：以“华矮11”为母本与9个高秆品种进行杂交,株高调查结果显示,其F1代均为高秆,而F2代则出现高秆矮秆性状分离,进一步对这9个F2分离群体的株高分离情况进行χ2检验,结果表明,所有组合的F2群体中高秆个体和矮秆个体的分离比例均符合3：1的分离规律。由此推定,华矮11的矮秆性状是受一对隐性基因控制,命名为btwd1。
2.矮秆基因btwdl的遗传等位测验：以华矮11/Monker的F1代做母本与其它4个矮秆品种杂交,后代全为高秆。表明矮秆基因btwdl与已知的矮秆基因br,uzu,sdwl和denso明显不同。
3.矮秆基因btwdl的定位：根据遗传分析结果,采用SSR标记与BSA分析方法相结合,对华矮11矮秆基因进行标记定位。结果表明来自于7H染色体的4个SSR标记(Bmac031, Bmac167, Bmag217和Bmag900)与目的基因btwdl构建成一个连锁群。该矮秆基因btwdl被定位于7H染色体长臂靠近着丝粒一侧,距离Bmac031为2.2 cM。
4.农艺性状和品质性状的QTL定位：以DH群体华矮11×华大麦6号为研究材料,在全生育期调查了农艺性状如抽穗期、有效穗、主穗长、主穗小穗数、单株小穗数、单株实粒数、穗平实粒数、单株粒重、穗平粒重、千粒重、分蘖数和品质性状蛋白质含量。选用63对在两亲本间有多态性的引物构建遗传连锁图谱。采用复合区间作图法对12个性状进行QTL定位分析,共检测到47个相关QTL,其中29个为新的QTL。
5.大麦抽穗期基因定位：以Galleon×Haruna Nijo DH群体作为研究材料,在中国(武汉,30°33'N)和澳大利亚(珀斯,31°56'S)设置3个不同的处理,3个处理包括正常大田秋播、大田晚播和温室18小时光照。通过对群体抽穗期的QTL定位分析结果表明,在中国环境下,探测到1个控制抽穗期的主效QTL(Qtl-5H)位于大麦染色体5H上,该QTL解释了抽穗期33%-50%的表型变异；在澳大利亚环境下,探测到2个控制抽穗期的主效QTL(Qtl-4H和Qtl5H),且Qtl-4H和Qtl-5H存在互作。它们的互作解释了抽穗期13%-36%的表型变异。Qtl-4H与Qtl-5H的互作可能是在澳大利亚环境下比在中国环境下群体株系更早抽穗的主要原因。
6.大麦长光周期基因的定位：以Baudin×AC Metcalfe DH群体作为研究材料,在中国(武汉,30°33'N)设置3个不同的处理,3个处理包括正常大田秋播、大田晚播和温室18小时光照,在澳大利亚(珀斯,31°56'S)设置2个不同的处理,2个处理包括正常大田秋播和温室18小时光照。通过对群体株系的抽穗期QTL定位分析结果表明,1个长光周期主效QTL(Qtl4-13)被定位在4H染色体上引物Xp12m50A199-Xp13m47B399之间。该QTL在澳大利亚和中国环境下分别解释了77.48%和37.81%的表型变异,表明它为大麦的一个新的长光周期基因。同时还定位了2个抽穗期相关QTL。
Barley is the fourth food crop in China and plays a very important role in the development of economy. To meet further social and people's living standards, new barley varieties will require high yields, better quality, high resistance and wide adaptability. Genetic characterization of barley germplasm will be crucial for the development of new barley varieties.
Dwarfism is a valuable trait in crop breeding, successful use of a dwarfing gene or dwarf source is critical for developing dwarf cultivars. In barley, more than 30 types of dwarfs or semidwarfs have been found. However, only a few of them have successfully been used in barley breeding program. Dwarf source and dwarf genes are unitary in barley breeding, not only limited further increases in yield and quality, but also narrowed the genetic basis, reduced biodiversity and increased vulnerability of disease and pest for new varieties. Therefore, discovery of new dwarf genes or dwarf sources is important for barley breeding. New dwarf source "huaai 11" has early maturity and good general combining ability. Utilization of this dwarf resource can broaden the genetic basis of barley dwarf breeding. We mapped dwarfing gene btwdl in Huaai 11 using a doubled-haploid (DH) population from a cross between Huaai 11 and Huadamai 6. In order to efficiently use this excellent germplasm for breeding program, we also analyzed the QTL underlying agronomic traits and quality traits in this line.
Heading date is a major determinant of the regional and seasonal adaptation of barley varieties. Appropriate heading date is an important goal of barley breeding. The dogma is that introduced germplasm is more likely to be adapted if it come from a similar latitude. However, barley germplasm introduced from similar latitudes of South-East Asia are extremely early heading in the Australian environments and vice versa. Therefore, we mapped heading date genes using Galleon×Haruna Nijo DH population and Baudin×AC Metcalfe DH population. The results will help us understand the main reason causing heading date difference in two similar latitude locations, Australia and China, The results of these researches are sumarized as follows:
1. Inheritance of the dwarf gene btwdl:dwarf Huaai 11 was crossed with nine tall varieties. All the F1 plants were tall. Both the tall and dwarf plants appeared in all the F2 populations with a 3:1 segregation ratio. The x2 testshowed the segregation is not significantly different from the expected 3:1 ratio, indicating that the dwarf trait is controlled by single recessive gene in the Huaai 11. This gene is designated as btwdl.
2. Allelic relationship between btwdl and other dwarfing genes:The F1 of Huaai 11 x Monker was crossed with other four dwarf varieties. All progeny in the four crosses are tall, suggesting that the gene btwdl controlling plant height in the Huaai 11 is nonallelic with br, uzu, sdwl and denso genes in these four dwarf varieties.
3. Mapping of the dwarf gene btwdl:SSR marker and BSA analysis were used to map the dwarfing gene in the Huaaill. Linkage analysis found that four SSR markers from chromosome 7H (BmacO31, Bmacl67, Bmag217 and Bmag900) were linked with the target gene btwd1. The dwarfing gene btwdl is close to the Bmac031 with a genetic distance of 2.2 cM, which is close to the centromere of 7HL.
4. QTL analysis of the agronomic and quality traits:This study using Huaai 11×Huadamai 6 DH population as research materials, Quality trait grain protein content and eleven gronomic traits such as heading date, spike numbers per plant, main spike length, spikelet number of main spike, spikelet number per plant, grain number per plant, grain number per spike, grain weight per plant, grain weight per spike,1000 grain weight, number of tillers were investigated. Sixty three pairs of SSR primers showed polymorphic between the two parents, and were used to construct a genetic linkage map. Composite interval mapping (CIM) analysis of 12 traits identifiedt a total of 47 QTL,29 of them were new QTL.
5. Mapping of heading date gene in barley:The Galleon x Haruna Nijo DH population was used to map heading date gene at two location China (Wuhan,30°33'N) and Australia (Perth,31°56' S) with three different treatments including normal autumn sowing, late sowing and pot experiment with 18h photoperiod. QTL analysis detected one major QTL (Qtl-5H) in China, which was located on chromosome 5H, and explained 33%-50% of the phenotypic variation. In Australia, two major QTL (Qtl-4H and Qtl5H) were detected for heading dates on chromosomes 4H and 5H and Qtl-4H and Qtl5H interaction. The Qtl-4H/Qtl-5H interaction explained 13%-36% of phenotypic variation. The Qtl-4H/Qtl-5H interaction may be main reason that DH lines are earlier heading in Australia than China.
6. Mapping of long photoperiod gene in barley:Bandin×AC Metcalfe DH population was used. In China (Wuhan,30°33'N), three different treatments including normal autumn sowing, late sowing and pot experiment with 18h photoperiod were conducted. In Australia (Perth,31°56' S), two different treatments including normal autumn sowing and pot experiment with 18h photoperiod were performed. QTL analysis detected a major QTL for photoperiod response and mapped onto the long arm of chromosome 4H in both Australian and Chinese locations. The QTL was flanked by the markers Xp12m50A199-Xp13m47B399 and accounted for 77.5% and 37.8% of phenotypic variation in Australia and China, respectively. Two QTL underlying heading date detected in this mapping population.
引文
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