中国小麦品种光周期和品质基因分子鉴定
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摘要
改良品种适应性和加工品质是我国小麦育种的重要目标。Ppd-D1a是位于普通小麦2D染色体上光周期非敏感基因,与小麦适应性密切相关;黄色素含量与面制品的外观色泽密切相关,我国的面条、馒头要求色白,需要低黄色素含量的小麦品种;麦谷蛋白亚基基因与面团强度密切相关。利用分子标记研究以上性状不仅可以了解控制相关性状的基因在不同麦区的分布规律,筛选含有优质基因的材料,而且有助于研究我国小麦在不同麦区的适应性机理和开展优质基因聚合育种。本研究利用分子标记检测了我国9个小麦种植区域926份品种(系)的光周期基因Ppd-D1a的分布规律,并结合系谱从分子水平上分析推导Ppd-D1a的来源;利用分子标记检测了我国4个冬麦区的374份改良品种(系)和197份核心地方种质控制黄色素含量的八氢番茄红素合成酶基因等位变异分布规律,并分析了改良品种的黄色素含量与八氢番茄红素合成酶基因之间的相关性,筛选了低黄色素含量材料。利用特异性分子标记研究了我国4个冬麦区的224和216份品种(系)的高、低分子量谷蛋白基因,明确了检测基因的分布频率及相关基因标记的有效性。主要研究结果如下:
1.我国小麦品种光周期非敏感基因Ppd-D1a的平均分布频率为66%,其中地方品种和改良品种分别为38.6%和90.6%;除西北高纬度地区部分春小麦和甘肃、新疆等地冬小麦外,国内1970以后推广的小麦品种(系)都携带有Ppd-D1a基因;从北到南Ppd-D1a基因的分布频率逐渐升高;结合系谱分析从分子水平上推导了中国小麦品种中Ppd-D1a基因的来源,主要是日本地方品种Akagomughi和中国地方品种蚂蚱麦和蚰子麦。Ppd-D1基因标记特异性强、重复性好,可以准确地检测小麦品种Ppd-D1a基因的存在与否。
2.控制黄色素含量的八氢番茄红素合成酶基因(Psy-A1)标记YP7A为共显性标记,在高、低黄色素含量的小麦材料中分别扩增出194 bp和231 bp片段,相应的等位基因为Psy-A1a和Psy-A1b(GenBank编号分别为EF600063和EF600064)。在374份冬小麦改良品种(系)中,含有等位基因Psy-A1a和Psy-A1b的品种(系)分别占66.8%和33.2%,二者黄色素含量平均值差异达到极显著水平(P<0.01)。其中,北方冬麦区、黄淮冬麦区、长江中下游冬麦区及西南冬麦区Psy-A1a等位基因的分布频率分别为73.8%、73.9%、25.9%和46.7%。197份核心地方种质等位基因Psy-A1a和Psy-A1b的分布频率分别为97.5%和2.5%。
3.用分子标记可以更准确地检测小麦品种(系)的HMW-GS组成。我国224份冬小麦品种(系)HMW-GS亚基的分布频率存在很大差异。其中,55份(24.6%)材料携带有Dx5,80份(35.7%)携带有By8,没有Dx5和By8等位变异的材料无扩增带产生; 5.8%的品种(系)携带有Bx20亚基。含有Bx14+By15和Bx13+By16亚基的品种分别有1和2份; 4(1.8%)和5(2.2%)份品种分别携带有Bx6和Bx17; 106(47.3%)个品种(系)含有基因By9。分子标记检测结果与SDS-PAGE结果比较,不同亚基的符合度(一致性)有一定的差异:Bx17和Bx6符合度为100%,Dx5、By8和By9的一致率分别为90.9%、88.9%和97.2%,主要原因是SDS-PAGE不能很好区分功能不同、分子量相似的亚基。
4. GLu-B3位点10个分子标记中8个特异性强、重复性好,可用于小麦品种LMW-GS的分子检测和辅助选择。10个标记检测我国216份冬小麦品种(系),发现8种等位变异,其中,34份材料(15.7%)携带Glu-B3g亚基;34份(15.7%)材料携带Glu-B3d亚基;15份材料(6.9%)含有Glu-B3b基因,4个品种含有Glu-B3e基因;13个品种(6.0%)携带Glu-B3f亚基;8个品种携带Glu-B3h亚基;4份材料含有Glu-B3i亚基;104份品种携带Glu-B3j亚基。分子标记检测结果与SDS-PAGE分析结果有一定差异,主要原因是SDS-PAGE技术对LMW-GS亚基的分辨率低所致。本研究明确了光周期非敏感基因Ppd-D1a、控制黄色素含量的八氢番茄红素合成酶基因Psy-A1、HMW-GS和Glu-B3位点等位基因在中国小麦品种(系)中的分布规律,并验证了其相应标记的有效性和实用性,为我国优质、高产和广适性小麦新品种选育提供了材料和方法。
Imcreasing the adaptability and the end processing quality of wheat are an important breeding goal in China. Ppd-D1a is photoperiod insensitive gene at Ppd-D1 locus on common wheat chromosome 2D and is closely associated with wide adaptability.Yellow pigment (YP) content is responsible for flour color. A bright white to creamy colour is preferred for white Chinese noodles and steamed bread in China, requiring a low YP content in wheat grains.Wheat HMW-GS and LMW-GS have a high correlation with dough strength that determines bread-making quality. Studying above characters using molecular makers can not only understand the distribution of genes for the related traits and screen good materials with desirable alleles; also help know adaptive mechanism in different wheat growing regions in China and converge good genes controlling different characteristic into only one genotype. Distribution of photoperiod insensitive gene Ppd-D1a was detected using allele-specific marker in a total of 926 Chinese wheat landraces and improved cultivars collected from nine wheat growing zones and origin of Ppd-D1a was deduced from molecular level in combination with pedigree analysis. Distributing frequency of phytoene synthease gene for yellow pigment content on chromosome 7A (Psy-A1) was studied with molecular marker YP7A in 374 improved cultivars from 4 winter wheat growing regions and 197 core landraces in China; the association was validated between YP content and Psy-A1 alleles and good cultivars with Psy-A1b for low yellow pigment content were screened. A total of 224 and 216 cultivars from 4 winter wheat zones were tested for HMW-GS and LMW-GS subunits at Glu-B3 locus, respectively, by allele-specific makers; frequencies of alleles detected and the availability of molecular markers were indicated. The main results in the present study were as follows:
1. The molecular marker developed from Ppd-D1 gene sequence with highly specific characteristics and repeatability could discriminate the alleles Ppd-D1a and Ppd-D1b. The average frequency of the photoperiod-insensitive Ppd-D1a allele was 66.0%, with the frequencies of 38.6% and 90.6% in landraces and improved cultivars, respectively. However, the Ppd-D1a allele was present in all improved cultivars released after 1970s except for spring wheats in high latitude northwestern China, and winter wheats in Gansu and Xinjiang. The presence of the Ppd-D1a allele in landraces and improved cultivars increased gradually from north to south, illustrating the relationship between photoperiod response and environment. Ppd-D1a in Chinese wheats was derived from three sources, Japanese landrace Akagomughi and Chinese landraces Mazhamai and Youzimai.
2. Characterization of phytoene synthease gene for yellow pigment (YP) content indicated that YP7A, a co-dominant marker of Psy-A1 gene, can amplify a 194-bp and a 231-bp fragment in the genotypes with Psy-A1a frequently for higher YP content and Psy-A1b mostly for lower YP content, respectively. Highly significant differences in YP content were detected between the genotypes with Psy-A1a and those with Psy-A1b (P<0.01). The occurrence frequencies for Psy-A1a and Psy-A1b were 66.8% and 33.2% in the 374 improved cultivars, respectively. Significant differences of allelic variations were found in improved wheat cultivars from different wheat regions of China. In the Northern Winter Wheat Region, Yellow & Huai Facultative Winter Wheat Region, Middle & Low Yangtze Valley Winter Wheat Region and Southwestern Winter Wheat Region, frequencies of the allele Psy-A1a were 73.8%、73.9%、25.9% and 46.7%, respectively. Whereas frequencies of the alleles Psy-A1a and Psy-A1b in Chinese landraces were 97.5% and 2.5%, respectively.
3. Molecular markers can more accurately evaluate HMW-GS composition in comparison with SDS-PAGE. Distribution of detected subunits were quite different in 224 Chinese winter wheat cultivars and advanced lines tested. Of them, 55 cultivars carried Dx5 subunit with a frequency of 24.6%; whereas 80 cultivars(35.7%)produced specific band (527bp) for By8. 5.8% for Bx20 based on markers Mar and ZSBy9aF1/R3, and Bx14+By15 was present in only one genotype, followed by two cultivars with Bx13+By16 according to marker ZSBy9F2/R2. Four Genotypes (1.8%) had Bx6 and five cultivars (2.2%) contained Bx17. One hundred and six (47.3%) varieties had By9, detected by ZSBy9aF1/R3, yielding 663-bp and 708-bp PCR fragments. Compared with the results of SDS-PAGE, 5 genotypes for Dx5 subunit were different in the test with allele specific molecular marker, with a 90.9% of consistent results, followed by 88.9% for By8, 100% for Bx17 and Bx6, 97.2% for By9. The difference may be mainly due to incorrect identifications of proteins that are functionally distinct, but have the similar molecular weights and relative mobilities in the test with SDS-PAGE.
4. 8 gene-specific markers with good repeatability could be used for marker-assisted selection in wheat breeding programs targeting for LMW-GS at Glu-B3 locus. A total of 216 Chinese winter wheat cultivars produced 8 allelic variations, out of them, 34 (15.7%) genotypes contained Glu-B3g, followed by 34 (15.7%) cultivars with Glu-B3d, 15 (6.9%) genotypes with Glu-B3b, 4 with Glu-B3e, 13 (6.0%) with Glu-B3f; 8 and 4 materials with Glu-B3h and Glu-B3i, respectively. Some differences were found between the results of SDS-PAGE and those of allele specific marker test. This was due to lower differentiation for LMW-GS subunits using SDS-PAGE .
In this study, the distribution were clarified for photoperiod insensitive gene Ppd-D1a, phytone synthease gene for yellow pigment content and high- and low-molecular-weight subunits allelic variation. Meanwhile, the characteristics of high efficiency and reliability of tested markers were validated in evaluating related gene alleles. The research was extremely useful for new cultivars selection with high production, good quality and wide adaptability in wheat breeding programs.
1.我国小麦品种光周期非敏感基因Ppd-D1a的平均分布频率为66%,其中地方品种和改良品种分别为38.6%和90.6%;除西北高纬度地区部分春小麦和甘肃、新疆等地冬小麦外,国内1970以后推广的小麦品种(系)都携带有Ppd-D1a基因;从北到南Ppd-D1a基因的分布频率逐渐升高;结合系谱分析从分子水平上推导了中国小麦品种中Ppd-D1a基因的来源,主要是日本地方品种Akagomughi和中国地方品种蚂蚱麦和蚰子麦。Ppd-D1基因标记特异性强、重复性好,可以准确地检测小麦品种Ppd-D1a基因的存在与否。
2.控制黄色素含量的八氢番茄红素合成酶基因(Psy-A1)标记YP7A为共显性标记,在高、低黄色素含量的小麦材料中分别扩增出194 bp和231 bp片段,相应的等位基因为Psy-A1a和Psy-A1b(GenBank编号分别为EF600063和EF600064)。在374份冬小麦改良品种(系)中,含有等位基因Psy-A1a和Psy-A1b的品种(系)分别占66.8%和33.2%,二者黄色素含量平均值差异达到极显著水平(P<0.01)。其中,北方冬麦区、黄淮冬麦区、长江中下游冬麦区及西南冬麦区Psy-A1a等位基因的分布频率分别为73.8%、73.9%、25.9%和46.7%。197份核心地方种质等位基因Psy-A1a和Psy-A1b的分布频率分别为97.5%和2.5%。
3.用分子标记可以更准确地检测小麦品种(系)的HMW-GS组成。我国224份冬小麦品种(系)HMW-GS亚基的分布频率存在很大差异。其中,55份(24.6%)材料携带有Dx5,80份(35.7%)携带有By8,没有Dx5和By8等位变异的材料无扩增带产生; 5.8%的品种(系)携带有Bx20亚基。含有Bx14+By15和Bx13+By16亚基的品种分别有1和2份; 4(1.8%)和5(2.2%)份品种分别携带有Bx6和Bx17; 106(47.3%)个品种(系)含有基因By9。分子标记检测结果与SDS-PAGE结果比较,不同亚基的符合度(一致性)有一定的差异:Bx17和Bx6符合度为100%,Dx5、By8和By9的一致率分别为90.9%、88.9%和97.2%,主要原因是SDS-PAGE不能很好区分功能不同、分子量相似的亚基。
4. GLu-B3位点10个分子标记中8个特异性强、重复性好,可用于小麦品种LMW-GS的分子检测和辅助选择。10个标记检测我国216份冬小麦品种(系),发现8种等位变异,其中,34份材料(15.7%)携带Glu-B3g亚基;34份(15.7%)材料携带Glu-B3d亚基;15份材料(6.9%)含有Glu-B3b基因,4个品种含有Glu-B3e基因;13个品种(6.0%)携带Glu-B3f亚基;8个品种携带Glu-B3h亚基;4份材料含有Glu-B3i亚基;104份品种携带Glu-B3j亚基。分子标记检测结果与SDS-PAGE分析结果有一定差异,主要原因是SDS-PAGE技术对LMW-GS亚基的分辨率低所致。本研究明确了光周期非敏感基因Ppd-D1a、控制黄色素含量的八氢番茄红素合成酶基因Psy-A1、HMW-GS和Glu-B3位点等位基因在中国小麦品种(系)中的分布规律,并验证了其相应标记的有效性和实用性,为我国优质、高产和广适性小麦新品种选育提供了材料和方法。
Imcreasing the adaptability and the end processing quality of wheat are an important breeding goal in China. Ppd-D1a is photoperiod insensitive gene at Ppd-D1 locus on common wheat chromosome 2D and is closely associated with wide adaptability.Yellow pigment (YP) content is responsible for flour color. A bright white to creamy colour is preferred for white Chinese noodles and steamed bread in China, requiring a low YP content in wheat grains.Wheat HMW-GS and LMW-GS have a high correlation with dough strength that determines bread-making quality. Studying above characters using molecular makers can not only understand the distribution of genes for the related traits and screen good materials with desirable alleles; also help know adaptive mechanism in different wheat growing regions in China and converge good genes controlling different characteristic into only one genotype. Distribution of photoperiod insensitive gene Ppd-D1a was detected using allele-specific marker in a total of 926 Chinese wheat landraces and improved cultivars collected from nine wheat growing zones and origin of Ppd-D1a was deduced from molecular level in combination with pedigree analysis. Distributing frequency of phytoene synthease gene for yellow pigment content on chromosome 7A (Psy-A1) was studied with molecular marker YP7A in 374 improved cultivars from 4 winter wheat growing regions and 197 core landraces in China; the association was validated between YP content and Psy-A1 alleles and good cultivars with Psy-A1b for low yellow pigment content were screened. A total of 224 and 216 cultivars from 4 winter wheat zones were tested for HMW-GS and LMW-GS subunits at Glu-B3 locus, respectively, by allele-specific makers; frequencies of alleles detected and the availability of molecular markers were indicated. The main results in the present study were as follows:
1. The molecular marker developed from Ppd-D1 gene sequence with highly specific characteristics and repeatability could discriminate the alleles Ppd-D1a and Ppd-D1b. The average frequency of the photoperiod-insensitive Ppd-D1a allele was 66.0%, with the frequencies of 38.6% and 90.6% in landraces and improved cultivars, respectively. However, the Ppd-D1a allele was present in all improved cultivars released after 1970s except for spring wheats in high latitude northwestern China, and winter wheats in Gansu and Xinjiang. The presence of the Ppd-D1a allele in landraces and improved cultivars increased gradually from north to south, illustrating the relationship between photoperiod response and environment. Ppd-D1a in Chinese wheats was derived from three sources, Japanese landrace Akagomughi and Chinese landraces Mazhamai and Youzimai.
2. Characterization of phytoene synthease gene for yellow pigment (YP) content indicated that YP7A, a co-dominant marker of Psy-A1 gene, can amplify a 194-bp and a 231-bp fragment in the genotypes with Psy-A1a frequently for higher YP content and Psy-A1b mostly for lower YP content, respectively. Highly significant differences in YP content were detected between the genotypes with Psy-A1a and those with Psy-A1b (P<0.01). The occurrence frequencies for Psy-A1a and Psy-A1b were 66.8% and 33.2% in the 374 improved cultivars, respectively. Significant differences of allelic variations were found in improved wheat cultivars from different wheat regions of China. In the Northern Winter Wheat Region, Yellow & Huai Facultative Winter Wheat Region, Middle & Low Yangtze Valley Winter Wheat Region and Southwestern Winter Wheat Region, frequencies of the allele Psy-A1a were 73.8%、73.9%、25.9% and 46.7%, respectively. Whereas frequencies of the alleles Psy-A1a and Psy-A1b in Chinese landraces were 97.5% and 2.5%, respectively.
3. Molecular markers can more accurately evaluate HMW-GS composition in comparison with SDS-PAGE. Distribution of detected subunits were quite different in 224 Chinese winter wheat cultivars and advanced lines tested. Of them, 55 cultivars carried Dx5 subunit with a frequency of 24.6%; whereas 80 cultivars(35.7%)produced specific band (527bp) for By8. 5.8% for Bx20 based on markers Mar and ZSBy9aF1/R3, and Bx14+By15 was present in only one genotype, followed by two cultivars with Bx13+By16 according to marker ZSBy9F2/R2. Four Genotypes (1.8%) had Bx6 and five cultivars (2.2%) contained Bx17. One hundred and six (47.3%) varieties had By9, detected by ZSBy9aF1/R3, yielding 663-bp and 708-bp PCR fragments. Compared with the results of SDS-PAGE, 5 genotypes for Dx5 subunit were different in the test with allele specific molecular marker, with a 90.9% of consistent results, followed by 88.9% for By8, 100% for Bx17 and Bx6, 97.2% for By9. The difference may be mainly due to incorrect identifications of proteins that are functionally distinct, but have the similar molecular weights and relative mobilities in the test with SDS-PAGE.
4. 8 gene-specific markers with good repeatability could be used for marker-assisted selection in wheat breeding programs targeting for LMW-GS at Glu-B3 locus. A total of 216 Chinese winter wheat cultivars produced 8 allelic variations, out of them, 34 (15.7%) genotypes contained Glu-B3g, followed by 34 (15.7%) cultivars with Glu-B3d, 15 (6.9%) genotypes with Glu-B3b, 4 with Glu-B3e, 13 (6.0%) with Glu-B3f; 8 and 4 materials with Glu-B3h and Glu-B3i, respectively. Some differences were found between the results of SDS-PAGE and those of allele specific marker test. This was due to lower differentiation for LMW-GS subunits using SDS-PAGE .
In this study, the distribution were clarified for photoperiod insensitive gene Ppd-D1a, phytone synthease gene for yellow pigment content and high- and low-molecular-weight subunits allelic variation. Meanwhile, the characteristics of high efficiency and reliability of tested markers were validated in evaluating related gene alleles. The research was extremely useful for new cultivars selection with high production, good quality and wide adaptability in wheat breeding programs.
引文
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