玉米耐荫性的QTL分析
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摘要
玉米是重要的粮食、饲料和工业原料作物,为我国第二大粮食作物,常年种植面积约4亿亩左右,总产量约1. 5亿吨,在国民经济中占有重要地位。由于荫雨寡照造成的弱光胁迫是玉米生育期间经常遇到的非生物逆境,也是当今黄淮海区玉米生产面临的重要问题。每年因弱光胁迫造成玉米减产10%-15%,重者达20%-30%。弱光胁迫已成为我国玉米生产可持续发展的主要制约因素。选育优良的耐荫玉米杂交种是当前玉米遗传育种的重要目标之一。玉米耐荫性是受多基因控制的数量性状,采用传统的育种方法改善其耐荫性效率较低。本研究以耐荫性强的自交系中72和耐荫性弱的自交系502为实验材料,配制F2作图群体,在弱光胁迫与非胁迫条件下,于郑州和浚县两地,调查F3家系田间耐荫性相关性状的表型,对抽雄天数、吐丝天数、雌雄间隔、比叶重、茎粗、株高、穗位高与耐荫性相关的QTL进行了定位和遗传分析,讨论了不同遗传背景的作图群体所定位玉米耐荫QTL的一致性和QTL之间的互作对耐荫性的影响。主要试验结论如下:
1用716对SSR引物对玉米自交系中72×502的亲本进行多态性筛选,共筛选出210对多态性引物,去除严重偏离的标记,用Mapmaker3.0作图软件进行构图,得到197个多态性标记的玉米遗传连锁图谱,图谱总长度2693.6CM,全基因组平均图距13.67 cM ,最小图距0.5 cM。
2在郑州和浚县两个试验点,F2:3家系的抽雄天数、吐丝天数、雌雄间隔、比叶重、茎粗、株高、穗位高遮荫敏感性相关性状均表现出数量性状遗传特点,F2:3家系的平均值接近双亲平均值,有一定数量的双向超亲家系。
3 F2:3家系在遮荫条件下,抽雄期、吐丝期延长,雌雄间隔期拉大,比叶重降低,株高、穗位高下降,变异范围扩大,整个群体的各个性状均符合正态分布,因此,F2:3家系群体可以用于进行QTL分析。
4 F2:3家系在郑州和浚县遮荫条件下两点各性状的QTLs定位。抽雄天数:在郑州点检测到一个QTL,在浚县点共检测到5个QTL,郑州和浚县共同检测到1个QTL,位于第1条染色体上umc1395—umc2083标记之间,贡献率分别为12.71%、6.76%;吐丝天数:郑州点检测到4个QTL,浚县点检测到3个QTL,郑州和浚县在第1条染色体上共同检测到一个QTL,位于umc1403和phi001之间,贡献率分别为10.93%、9.85%;雌雄间隔天数:郑州点检测到4个QTL,在浚县点检测到2个QTL,郑州和浚县在第一条染色体上共同检测到一个QTL,贡献率分别为10.01%和10.12%,位于umc1403和phi001之间。比叶重:郑州点检测到4个QTL,在浚县点检测到2个QTL;茎粗:在郑州没有检测到QTL,在浚县点检测到4个QTL;株高:郑州点检测到4个QTL,浚县点检测到4个QTL,郑州和浚县共同检测到3个QTL,分别位于1、4、10染色体上,分别位于umc1082和bnlg1671、bnlg2244和umc1051、umc1432和umc1962标记之间,贡献率为8.45%、8.53%,7.28%、9.92%,10.13%、8.77%。穗位高:郑州点检测到3个QTL,浚县点检测到5个QTL,郑州和浚县共同检测到3个相同的QTL,位于1、4、5染色体上,分别位于umc1082—bnlg1671、bnlg2244—umc1051、umc2036—bnlg1879标记之间,其贡献率分别为12.93%、13.79%,8.45%、7.13%,8.17%、6.36%。
5 F2:3家系郑州和浚县两点光照下各性状的QTLs定位。抽雄天数:在郑州点检测到3个QTL,浚县点共检测到4个QTL,郑州和浚县共同检测到1个QTL,位于第一条染色体上umc2083—bnlg1057标记之间,贡献率为10.25%、14.39%。吐丝天数:郑州点检测到3个QTL,浚县点检测到2个QTL,郑州和浚县在第1条染色体上共同检测到一个QTL,位于第一条染色体上umc2083—bnlg1057标记之间,贡献率分别为10.95%、10.23%。雌雄间隔天数:郑州点检测到5个QTL,浚县点检测到6个QTL,郑州和浚县共同检测到2个QTL,分别位于第1、8条染色体上umc1976—umc1403、umc1960—mmc0181标记之间,贡献率分别为10.23%、6.88%,4.66%、7.02%。比叶重:郑州点检测到4个QTL,浚县点检测到3个QTL,郑州和浚县共同检测到一个QTL,位于第10条染色体上bnlg594—bnlg2190标记之间,贡献率分别为6.75%、5.14%。茎粗:郑州点检测到2个QTL,浚县检测到2个QTL,两点没有检测到共同的QTL。株高:郑州点检测到4个QTL,浚县点检测到5个QTL,郑州和浚县共同检测到2个QTL,分别位于4、8染色体上,分别位于bnlg2244和umc1051、umc1202和umc1562标记之间,贡献率分别为11.81%、13.35%,8.2%、6.72%。穗位高:郑州点检测到5个QTL,分别位于1、3、4、5、7染色体上,贡献率变幅为4.23%—9.86%,累计贡献率为33.98%;浚县点检测到2个QTL,分别位于1、5染色体上,贡献率分别为4.97%、6.6%,累计贡献率为11.57%,两点没有检测到共同的QTL。
Maize is an important food, feed and industrial crops, as China's second largest food crops, perennial planting around an area of about 4 million mu, the total output of about 150 million tons, in the national economy occupies an important position.The stress of rain and poor light during the maize growing is often encountered in non-biological adversity,is also the important issue of maize production of the Huang-Huai-Hai area in today’s facing. Each year due to stress caused by low light, corn production is lowed by 10 -15%, in serious cases up to 20% -30%. Low-light stress has become a sustainable development of China's maize production of the major constraints. Breeding excellent tolerant maize hybrids of maize genetics and breeding is one of the important goals. Tolerant corn is controlled by multiple genes of quantitative traits using traditional breeding methods to improve their tolerant and less efficient. In this study, inbred lines of strong tolerant of the中72 and inbred lines of weak tolerant of the 502 were for experimental materials,and the F2 mapping groups were prepared.In low-light stress and non-stress conditions, in Zhengzhou and xunxian ,investigate the F3 families tolerant field-related phenotypic traits, the number of days to heading, silking days, male and female interval, specific leaf weight, stem diameter, plant height, ear height. Tolerant QTL associated to the location and genetic analysis are aslo done. Tolerant maize QTL consistency and interoper ability between the effect against tolerant are discussed in the genetic background of different groups of mapping QTL positioning . The main trial findings were as follows:
1 716 pairs of SSR primers for maize inbred lines in 72×502 were for the parental polymorphism screening, a total of 210 pairs of polymorphisms were selected primers, and the tags deviated seriously were removed.Using mapping software Mapmaker3.0 , genetic linkage map of maize was obtained with 197 polymorphism, the total length of map 2693.6CM, the average genome-wide map 13.67 cM, a minimum map distance 0.5 cM.
2 At xunxian and Zhengzhou the two pilot sites, F2:3 families tasselling days, silking days, ASI, specific leaf weight, stem diameter, plant height, ear height ,which are shade-sensitive traits related , showed genetic characteristics of quantitative traits.F2:3 family close to the average of the parents,and there are a certain number of two-line super-in-laws.
3 F2:3 families in shade conditions, tasselling stage and silking period were prolonged, male and female gap intervals were extended, specific leaf weight was lowed, plant height and ear height were decreased ,variability were expanded .The various characters of the whole group are state distribution, so that, F2: 3 family groups can be used for QTL analysis.
4 The characters of the F2: 3 families ,in xunxian and Zhengzhou ,under the shade conditions had been QTLs positioned. The number of days tasselling: in Zhengzhou ,a QTL was detected, in xunxian five QTL were detected. A common QTL was detected in Zhengzhou and xunxian, which was located on chromosome 1 , between umc1395-umc2083 and the contribution rates were 12.71%, 6.76%; silking days: four QTL were detected in Zhengzhou point, three QTL were detected in xunxian point.A common QTL was detected in Zhengzhou and xunxian, which was located on chromosome 1 , between umc1403 and phi001, and the contribution rates were 10.93%, 9.85%; ASI: four QTL were detected in Zhengzhou points, two QTL were detected in xunxian point. A common QTL was detected in Zhengzhou and xunxian, which was located on chromosome 1, and the contribution rate of 10.12 percent and 10.01 percent, between umc1403 and phi001.Specific leaf weight: four QTL were detected in Zhengzhou point, two QTL were detected in xunxian point; diameter: no QTL was detected in Zhengzhou, 4 QTL were detected in xunxian point; height: 4 QTL were detected in Zhengzhou, four QTL were detected in xunxian points, three common QTL were detected in Zhengzhou and xunxian, which were located on chromosome 1,4,10, and were located in between umc1082 and bnlg1671, bnlg2244 and umc1051, umc1432 and umc1962 tags, and the contribution rate to 8.45 percent, 8.53%, 7.28%, 9.92%, 10.13%, 8.77%. Ear height: three QTL were detected in Zhengzhou points, five QTL were detected in xunxian points, three common QTL were detected in Zhengzhou and xunxian, which were located on chromosomes 1,4,5, and were located in umc1082-bnlg1671,bnlg2244-umc1051, umc2036-bnlg1879 tags,and the contribution rate of 12.93 percent, respectively, 13.79%, 8.45%, 7.13%, 8.17%, 6.36%.
5 The characters of the F2: 3 families ,in xunxian and Zhengzhou ,under the light conditions had been QTLs positioned. The number of days tasselling: three QTL were detected in Zhengzhou points, four QTL were detected in xunxian, a common QTL was detected in Zhengzhou and xunxian, which was located on the first chromosome,between umc2083-bnlg1057 ,and the contribution rate of 10.25% , 14.39%. Silking days: three QTL were detectedin Zhengzhou points, two QTL were detected in xunxian points, a common QTL was detected in Zhengzhou and xunxian, which was located on the first chromosome, between markers umc2083-bnlg1057,and contribution rates were 10.95%, 10.23%.The number of days between the male and female: five QTL were detected in Zhengzhou points, six QTL were detected in xunxian point, two common QTL were detected in Zhengzhou and xunxian, which were located on chromosome 1and 8 ,between umc1976-umc1403, umc1960-mmc0181 ,and respectively the contribution rate of 10.23%, 6.88%, 4.66%, 7.02%. Specific leaf weight: four QTL were detected in Zhengzhou points, three QTL were detected in xunxian points, a common QTL was detected in Zhengzhou and xunxian, which was located on chromosome 10 between marker bnlg594-bnlg2190 ,and respectively the contribution rate of 6.75%, 5.14%. Diameter: two QTL were detected in Zhengzhou points, two QTL were detected in xunxian, no common QTL was detected. Height: four QTLwere detected in Zhengzhou points, five QTL were detected in xunxian points, two common QTL were detected in Zhengzhou and xunxian, which were located on chromosome 4,8, in bnlg2244 and umc1051, umc1202 and umc1562 ,and respectively the contribution rate of 11.81%, 13.35%, 8.2%, 6.72%. Ear height: five QTL were detected in Zhengzhou points, which are located on chromosome 1,3,4,5,7, the contribution rate ranged from 4.23% -9.86%, the cumulative contribution rate of 33.98%; 2 QTL were detected in xunxian, which were located on chromosome 1,5, respectively the contribution rate of 4.97%, 6.6%, the cumulative contribution rate of 11.57%, no common QTL had been detected .
1用716对SSR引物对玉米自交系中72×502的亲本进行多态性筛选,共筛选出210对多态性引物,去除严重偏离的标记,用Mapmaker3.0作图软件进行构图,得到197个多态性标记的玉米遗传连锁图谱,图谱总长度2693.6CM,全基因组平均图距13.67 cM ,最小图距0.5 cM。
2在郑州和浚县两个试验点,F2:3家系的抽雄天数、吐丝天数、雌雄间隔、比叶重、茎粗、株高、穗位高遮荫敏感性相关性状均表现出数量性状遗传特点,F2:3家系的平均值接近双亲平均值,有一定数量的双向超亲家系。
3 F2:3家系在遮荫条件下,抽雄期、吐丝期延长,雌雄间隔期拉大,比叶重降低,株高、穗位高下降,变异范围扩大,整个群体的各个性状均符合正态分布,因此,F2:3家系群体可以用于进行QTL分析。
4 F2:3家系在郑州和浚县遮荫条件下两点各性状的QTLs定位。抽雄天数:在郑州点检测到一个QTL,在浚县点共检测到5个QTL,郑州和浚县共同检测到1个QTL,位于第1条染色体上umc1395—umc2083标记之间,贡献率分别为12.71%、6.76%;吐丝天数:郑州点检测到4个QTL,浚县点检测到3个QTL,郑州和浚县在第1条染色体上共同检测到一个QTL,位于umc1403和phi001之间,贡献率分别为10.93%、9.85%;雌雄间隔天数:郑州点检测到4个QTL,在浚县点检测到2个QTL,郑州和浚县在第一条染色体上共同检测到一个QTL,贡献率分别为10.01%和10.12%,位于umc1403和phi001之间。比叶重:郑州点检测到4个QTL,在浚县点检测到2个QTL;茎粗:在郑州没有检测到QTL,在浚县点检测到4个QTL;株高:郑州点检测到4个QTL,浚县点检测到4个QTL,郑州和浚县共同检测到3个QTL,分别位于1、4、10染色体上,分别位于umc1082和bnlg1671、bnlg2244和umc1051、umc1432和umc1962标记之间,贡献率为8.45%、8.53%,7.28%、9.92%,10.13%、8.77%。穗位高:郑州点检测到3个QTL,浚县点检测到5个QTL,郑州和浚县共同检测到3个相同的QTL,位于1、4、5染色体上,分别位于umc1082—bnlg1671、bnlg2244—umc1051、umc2036—bnlg1879标记之间,其贡献率分别为12.93%、13.79%,8.45%、7.13%,8.17%、6.36%。
5 F2:3家系郑州和浚县两点光照下各性状的QTLs定位。抽雄天数:在郑州点检测到3个QTL,浚县点共检测到4个QTL,郑州和浚县共同检测到1个QTL,位于第一条染色体上umc2083—bnlg1057标记之间,贡献率为10.25%、14.39%。吐丝天数:郑州点检测到3个QTL,浚县点检测到2个QTL,郑州和浚县在第1条染色体上共同检测到一个QTL,位于第一条染色体上umc2083—bnlg1057标记之间,贡献率分别为10.95%、10.23%。雌雄间隔天数:郑州点检测到5个QTL,浚县点检测到6个QTL,郑州和浚县共同检测到2个QTL,分别位于第1、8条染色体上umc1976—umc1403、umc1960—mmc0181标记之间,贡献率分别为10.23%、6.88%,4.66%、7.02%。比叶重:郑州点检测到4个QTL,浚县点检测到3个QTL,郑州和浚县共同检测到一个QTL,位于第10条染色体上bnlg594—bnlg2190标记之间,贡献率分别为6.75%、5.14%。茎粗:郑州点检测到2个QTL,浚县检测到2个QTL,两点没有检测到共同的QTL。株高:郑州点检测到4个QTL,浚县点检测到5个QTL,郑州和浚县共同检测到2个QTL,分别位于4、8染色体上,分别位于bnlg2244和umc1051、umc1202和umc1562标记之间,贡献率分别为11.81%、13.35%,8.2%、6.72%。穗位高:郑州点检测到5个QTL,分别位于1、3、4、5、7染色体上,贡献率变幅为4.23%—9.86%,累计贡献率为33.98%;浚县点检测到2个QTL,分别位于1、5染色体上,贡献率分别为4.97%、6.6%,累计贡献率为11.57%,两点没有检测到共同的QTL。
Maize is an important food, feed and industrial crops, as China's second largest food crops, perennial planting around an area of about 4 million mu, the total output of about 150 million tons, in the national economy occupies an important position.The stress of rain and poor light during the maize growing is often encountered in non-biological adversity,is also the important issue of maize production of the Huang-Huai-Hai area in today’s facing. Each year due to stress caused by low light, corn production is lowed by 10 -15%, in serious cases up to 20% -30%. Low-light stress has become a sustainable development of China's maize production of the major constraints. Breeding excellent tolerant maize hybrids of maize genetics and breeding is one of the important goals. Tolerant corn is controlled by multiple genes of quantitative traits using traditional breeding methods to improve their tolerant and less efficient. In this study, inbred lines of strong tolerant of the中72 and inbred lines of weak tolerant of the 502 were for experimental materials,and the F2 mapping groups were prepared.In low-light stress and non-stress conditions, in Zhengzhou and xunxian ,investigate the F3 families tolerant field-related phenotypic traits, the number of days to heading, silking days, male and female interval, specific leaf weight, stem diameter, plant height, ear height. Tolerant QTL associated to the location and genetic analysis are aslo done. Tolerant maize QTL consistency and interoper ability between the effect against tolerant are discussed in the genetic background of different groups of mapping QTL positioning . The main trial findings were as follows:
1 716 pairs of SSR primers for maize inbred lines in 72×502 were for the parental polymorphism screening, a total of 210 pairs of polymorphisms were selected primers, and the tags deviated seriously were removed.Using mapping software Mapmaker3.0 , genetic linkage map of maize was obtained with 197 polymorphism, the total length of map 2693.6CM, the average genome-wide map 13.67 cM, a minimum map distance 0.5 cM.
2 At xunxian and Zhengzhou the two pilot sites, F2:3 families tasselling days, silking days, ASI, specific leaf weight, stem diameter, plant height, ear height ,which are shade-sensitive traits related , showed genetic characteristics of quantitative traits.F2:3 family close to the average of the parents,and there are a certain number of two-line super-in-laws.
3 F2:3 families in shade conditions, tasselling stage and silking period were prolonged, male and female gap intervals were extended, specific leaf weight was lowed, plant height and ear height were decreased ,variability were expanded .The various characters of the whole group are state distribution, so that, F2: 3 family groups can be used for QTL analysis.
4 The characters of the F2: 3 families ,in xunxian and Zhengzhou ,under the shade conditions had been QTLs positioned. The number of days tasselling: in Zhengzhou ,a QTL was detected, in xunxian five QTL were detected. A common QTL was detected in Zhengzhou and xunxian, which was located on chromosome 1 , between umc1395-umc2083 and the contribution rates were 12.71%, 6.76%; silking days: four QTL were detected in Zhengzhou point, three QTL were detected in xunxian point.A common QTL was detected in Zhengzhou and xunxian, which was located on chromosome 1 , between umc1403 and phi001, and the contribution rates were 10.93%, 9.85%; ASI: four QTL were detected in Zhengzhou points, two QTL were detected in xunxian point. A common QTL was detected in Zhengzhou and xunxian, which was located on chromosome 1, and the contribution rate of 10.12 percent and 10.01 percent, between umc1403 and phi001.Specific leaf weight: four QTL were detected in Zhengzhou point, two QTL were detected in xunxian point; diameter: no QTL was detected in Zhengzhou, 4 QTL were detected in xunxian point; height: 4 QTL were detected in Zhengzhou, four QTL were detected in xunxian points, three common QTL were detected in Zhengzhou and xunxian, which were located on chromosome 1,4,10, and were located in between umc1082 and bnlg1671, bnlg2244 and umc1051, umc1432 and umc1962 tags, and the contribution rate to 8.45 percent, 8.53%, 7.28%, 9.92%, 10.13%, 8.77%. Ear height: three QTL were detected in Zhengzhou points, five QTL were detected in xunxian points, three common QTL were detected in Zhengzhou and xunxian, which were located on chromosomes 1,4,5, and were located in umc1082-bnlg1671,bnlg2244-umc1051, umc2036-bnlg1879 tags,and the contribution rate of 12.93 percent, respectively, 13.79%, 8.45%, 7.13%, 8.17%, 6.36%.
5 The characters of the F2: 3 families ,in xunxian and Zhengzhou ,under the light conditions had been QTLs positioned. The number of days tasselling: three QTL were detected in Zhengzhou points, four QTL were detected in xunxian, a common QTL was detected in Zhengzhou and xunxian, which was located on the first chromosome,between umc2083-bnlg1057 ,and the contribution rate of 10.25% , 14.39%. Silking days: three QTL were detectedin Zhengzhou points, two QTL were detected in xunxian points, a common QTL was detected in Zhengzhou and xunxian, which was located on the first chromosome, between markers umc2083-bnlg1057,and contribution rates were 10.95%, 10.23%.The number of days between the male and female: five QTL were detected in Zhengzhou points, six QTL were detected in xunxian point, two common QTL were detected in Zhengzhou and xunxian, which were located on chromosome 1and 8 ,between umc1976-umc1403, umc1960-mmc0181 ,and respectively the contribution rate of 10.23%, 6.88%, 4.66%, 7.02%. Specific leaf weight: four QTL were detected in Zhengzhou points, three QTL were detected in xunxian points, a common QTL was detected in Zhengzhou and xunxian, which was located on chromosome 10 between marker bnlg594-bnlg2190 ,and respectively the contribution rate of 6.75%, 5.14%. Diameter: two QTL were detected in Zhengzhou points, two QTL were detected in xunxian, no common QTL was detected. Height: four QTLwere detected in Zhengzhou points, five QTL were detected in xunxian points, two common QTL were detected in Zhengzhou and xunxian, which were located on chromosome 4,8, in bnlg2244 and umc1051, umc1202 and umc1562 ,and respectively the contribution rate of 11.81%, 13.35%, 8.2%, 6.72%. Ear height: five QTL were detected in Zhengzhou points, which are located on chromosome 1,3,4,5,7, the contribution rate ranged from 4.23% -9.86%, the cumulative contribution rate of 33.98%; 2 QTL were detected in xunxian, which were located on chromosome 1,5, respectively the contribution rate of 4.97%, 6.6%, the cumulative contribution rate of 11.57%, no common QTL had been detected .
引文
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