玉米粗缩病相关性状的遗传学分析和QTL定位
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摘要
玉米(Zea mays L.)是世界范围内重要的粮食和饲料作物,在农业生产中占有举足轻重的地位。玉米粗缩病是世界范围内严重危害玉米生产的一种病毒病。经过我国科研工作者的长期研究,确定了我国玉米粗缩病的病原是水稻黑条矮缩病毒(RBSDV)。RBSDV主要通过传播介体灰飞虱(Laodelphax sreiatellus Fallen.)进行传播,当玉米生育期和灰飞虱迁飞的高峰期重合时,引起玉米粗缩病的爆发和流行。由于我国生产上所用的骨干自交系和杂交种的粗缩病抗性差,近年来玉米粗缩病导致玉米大面积减产,严重制约了我国玉米产量的提高。培育抗粗缩病玉米品种已成为我国玉米育种高度关注的课题和难题。玉米粗缩病抗病性鉴定试验结果表明,玉米对粗缩病的抗性呈现数量性状的特征,是受到多基因控制的。经过多年努力,我国玉米育种工作者已培育出一批抗玉米粗缩病的自交系,如90110、齐319、X178等。为了控制该病害的发生和流行、减少生产损失,开发并利用抗病玉米品种本身的抗病QTL进行遗传改良是减轻玉米粗缩病危害的关键。
近年来,国内外对玉米粗缩病研究逐步深入,但并没有取得突破性的进展,主要原因在于玉米粗缩病发病的方式较为复杂,涉及到病毒、传播介体灰飞虱和玉米植株这三方面的相互关系,导致玉米植株抗病性鉴定难度大,鉴定的准确性较差和抗病性QTL定位困难。因此,建立科学合理的粗缩病抗性鉴定方法和准确鉴定分离群体中不同基因型的粗缩病抗性是准确定位和克隆粗缩病抗病QTL的前提。本工作开展了玉米粗缩病抗性鉴定方法的改进和粗缩病抗病QTL定位研究。
1.建立田间自然发病结合人工接毒的联合鉴定方法
本研究利用灰飞虱作为传毒介体,以山东济南地区自然发病的玉米粗缩病植株叶片饲喂灰飞虱。而后从饲喂灰飞虱的饲养桶中随机挑选5头灰飞虱成虫作为检测的样本,利用我国玉米粗缩病病原水稻黑条矮缩病毒(RBSDV)基因组的特异性引物检测灰飞虱带毒情况。在来自14个灰飞虱饲养桶的样本中检测出有11个样本中的灰飞虱携带RBSDV.2010年5月在网棚中利用塑料萌种盘萌发玉米实验材料,待植株处于一叶一心期时,将携带RBSDV的灰飞虱按照500头/m2的密度放入网棚中,连续传毒5天。5天后利用虱蚜净将剩余灰飞虱全部杀灭。待植株长至三叶期时,将它们移栽到济南地区吕家村试验田中生长,由于此时适逢田间灰飞虱的迁飞高峰期,而玉米植株又处在对灰飞虱侵染敏感阶段,因而在田间又经历一次田间自然感染。通过两种接毒方法的联合使用,提高了试验材料的粗缩病发病率,提高了鉴定结果的可靠性和重复性。该方法可以在较短时期内检测大量的玉米植株,为鉴定大量分离群体材料的粗缩病抗性和相关的QTL定位奠定了基础。
2.玉米粗缩病的遗传分析
本研究利用课题组创制的“掖478×90110”的RILs的F7：9群体为实验材料,从2008年到2010年,在山东省济南和莱州两个试验点连续种植RIL群体和掖478、90110以及其F1植株,通过田间自然发病的方法和联合接毒鉴定的方法进行粗缩病的抗病性评判。在实验材料开花期对玉米粗缩病相关性状,包括上部节间性状、叶片背面蜡质性状、雄穗发育情况以及粗缩病发病指数等进行观测和统计。按照植株粗缩病的病级和各个性状的等级整理数据,然后对三年两地的实验数据进行遗传分析,结果表明玉米粗缩病在“掖478×90110”的后代中,抗病特性主要由基因型决定,受到环境效应、基因型与环境互作效应的影响较小。粗缩病抗性的广义遗传力范围为0.71～0.94。即在该RILs群体中粗缩病的抗性主要是由遗传因素决定的。分析粗缩病相关性状参数的分布可得出,玉米植株的粗缩病抗性决定于来自90110的主效QTL。
3.玉米粗缩病抗病QTL定位
利用覆盖整个玉米基因组的512对SSR引物在RILs群体中检测标记多态性,选多态性良好且扩增条带清晰的SSR引物用于QTL的定位检测,共计检测到5个控制粗缩病抗性及相关性状的加性QTL,分别位于玉米基因组的2号、6号、7号、8号以及10号染色体上,分别命名为qMRD2、qMRD6、qMRD7、qMRD8以及qMRD10。其中QTL qMRD8几乎在所有的粗缩病相关性状中被检测到,而且qMRD8单独能够解释粗缩病表型变异的12-28.9%,是一个主效QTL。QTL qMRD2、qMRD6和UqMRD7在整个QTL的定位实验中至少被检测到3次,也是重复性良好的粗缩病抗性相关QTL。QTL qMRD10仅仅在上部节间性状中被检测到了一次,且效应值较小,该QTL需要后续的工作继续进行验证。检测到的全部QTL总共可以解释粗缩病表型变异值的41.43-50.84%。本实验中qMRD6、 qMRD7和qMRD83个QTL与之前本实验室王飞等利用SSR-BSA去检测到的粗缩病抗性位点相重合,这提示在这些QTL区及其邻近位置上存在着玉米粗缩病抗性基因。本研究检测到的主效QTL可以通过MAS的方法导入抗病性较弱的骨干自交系中,用于加快玉米粗缩病抗病性育种的进程。
本工作的意义有：1.首次利用田间自然发病结合人工接毒的联合检测方法进行了玉米粗缩病抗性鉴定。该方法可有效提高玉米粗缩病抗性鉴定的准确性和稳定性。2.通过粗缩病抗病性的遗传分析确定了来自“掖478×90110”的RILs群体中抗病性主要由遗传因素决定,且存在抗粗缩病的主效QTL。3.通过粗缩病抗性QTL检测,定位了5个粗缩病抗性相关的QTL,其中3个QTL与SSR-BSA法鉴定出的相关位点重合,可用于采用MAS方法培育抗粗缩病玉米材料。
Maize (Zea mays L.) is the worldwide most important food and forage crop as well as a model plant for genetics and development biology, and plays a decisive role in agriculture. Maize rough dwarf disease (MRDD) is a viral disease that is widely distributed in the worldwide and causes great yield reduction of maize. In China, The pathogen of MRDD has been identified as rice black-streaked dwarf virus (RBSDV). RBSDV is mainly transmitted by a kind of planthopper (Laodelphax sreiatellus Fallen.) in a persistent manner among individuals. When the maize seedlings suffer the large scale migration of planthopper from wheat or other grasses in the field, it may cause the outbreak of MRDD. The MRDD has caused a great and large-scale yield reduction of maize in China for the poor resistance to MRDD of elite inbred lines and hybrids in recent years. The MRDD has become one of the hardest problems in maize production, and obstructs our grain production. The results of the assays of maize resistance to MRDD show that the maize resistance to MRDD take on the features of quantitative trait and is controlled by multiple genes. Meanwhile, Chinese breeders have selected a bulk of maize inbred lines with resistantance to MRDD, such as90110, Qi319, X178et al. To control the outbreak and transmission of MRDD and reduce the loss of maize yield, it is the key step to identify the resistant QTL in maize and improve the elite maize inbred and hybrid using these QTL.
Recently, the researches of maize resistance to MRDD have been accelerated, but the breakthrough progresses are not made. The major reason is the conditions of MRDD are complicated that involve the correlations of the virus, vector-planthopper and maize. It is hard to accurately identify the individual phenotype and to map the QTL conferring resistance to MRDD. But, the phenotype identification is the foundation for mapping and cloning the resistant QTL that develop the appropriate scientific methods to identify maize resistance to MRDD accurately in the segregation populations. In this study, the improved method to identify the maize resistance to MRDD is developed and the QTL conferring resistance to MRDD are identified.
1. The development of combined identification method based on the combination of natural infection and artificial inoculation
In this study, the planthoppers were used as the medium to transmit RBSDV and were cultured in the feeding buckets and fed on the diseased maize leaves from naturally infected field-grown plants with typical symptoms of MRDD in the Jinan area of Shandong province. Subsequently, five adult-planthoppers were randomly selected as a sample to detect RBSDV, and then the RBSDV specific primers were used to screen the samples from different feeding buckets.11out of14samples were positive. May1st in2010, the RILs and their parental lines (90110and Ye478) were germinated at plastic plates in the net house. When the seedlings grew to the1-leaf period, the planthoppers with RBSDV were dispersed into the net house. The average density of planthoppers was about500/m2. After a5-day inoculation, the insecticide was used to kill all of the planthoppers, and the seedlings were maintained to grow continually in the net house and then were transplanted to the experiment fields at the3-leaf stage in Jinan area. The seedlings were infected again by the planthoppers at the migratory flight period of planthoppers in field. The incidence of MRDD and the stability and repetitiveness of identification results were increased by the combination of the artificial inoculation and natural infection. And plenty of maize plants could be screened in a short time by using this method. The combined method has laid the foundation of identifying abundant examples in the segregation population.
2. The genetic analysis of maize rough dwarf disease
The analysis was performed in common maize using a set of recombinant inbred lines (RILs) F7;9derived from 'Ye478×90110'.The RILs population, Ye478,90110and F1were grown continually at the area of Jinan and Laizhou respectively from2008to2010, and then were identified the resistance to MRDD through the method combining the artificial inoculation and natural infection. The phenotypes related to MRDD including the shortened superior internodes, enation, tassel type and disease severity index were evaluated at the flowering stage. Subsequently, the genetic analyses were performed using the3-year experimental data at two plots according to the disease grade and the level of traits related to MRDD. The broad sense heritability of resistance to MRDD is from0.71to0.94, that is to say, the resistance to MRDD was mainly determined by the genotype in the population of Ye478and90110, with minor influence of the environment and the interaction effect between the genotype and environment. The distribution analysis of traits related to MRDD showed that there were major QTL from90110conferring resistance to MRDD.
3. Mapping of QTL conferring resistance to MRDD
The512SSR markers covering the whole maize genome were screened polymorphisms between the two parents Ye478and90110. And the polymorphism SSR markers with clear band were used for QTL mapping by using the RILs. Five additive QTL related to resistance were detected and were mapped on chromosome2,6,7,8and10respectively. The QTL were named as qMRD2, qMRD6, qMRD7, qMRD8and qMRD10, respectively. The qMRD8was a major QTL, that could be detected in all traits mapping, and explain the phenotypic variation of12-28.9%. QTL qMRD2, qMRD6and qMRD7also had good repetitiveness, which were detected3times at least through the whole mapping experiment. The qMRDIO was only detected once at the trait of shortened superior intemodes with relative small effect and need to be verified in subsequent experiment. All of these QTL could explain the phonotypical variance of41.43-50.84%. In this study, the QTL qMRD6, qMRD7and qMRD8were coincidence with the loci that were detected by Dr. Wang using SSR-BSA method, previously. The results suggested that these QTL and the loci nearby these QTL may contain the genes conferring resistance to MRDD. The major QTL detected in this study can be introduced into the elite inbred lines with little resistance to MRDD by MAS to accelerate the proceeding of maize breeding.
In this research, the combined inoculation method including natural infection and artificial inoculation was used to study the resistance to MRDD for the first time, which could enhance the stability and validity of resistance to MRDD, and could increase the detecting amount of segregation population. Moreover, this study proved that the resistance to MRDD was mainly due to the genetic factors through the genetic analysis. There were5QTL conferring resistance to MRDD detected through the QTL mapping, and3of them were coincidence with the loci detected through SSR-BSA method. Finally, this study laid the foundation of MRDD resistance breeding in maize by the method of MAS.
近年来,国内外对玉米粗缩病研究逐步深入,但并没有取得突破性的进展,主要原因在于玉米粗缩病发病的方式较为复杂,涉及到病毒、传播介体灰飞虱和玉米植株这三方面的相互关系,导致玉米植株抗病性鉴定难度大,鉴定的准确性较差和抗病性QTL定位困难。因此,建立科学合理的粗缩病抗性鉴定方法和准确鉴定分离群体中不同基因型的粗缩病抗性是准确定位和克隆粗缩病抗病QTL的前提。本工作开展了玉米粗缩病抗性鉴定方法的改进和粗缩病抗病QTL定位研究。
1.建立田间自然发病结合人工接毒的联合鉴定方法
本研究利用灰飞虱作为传毒介体,以山东济南地区自然发病的玉米粗缩病植株叶片饲喂灰飞虱。而后从饲喂灰飞虱的饲养桶中随机挑选5头灰飞虱成虫作为检测的样本,利用我国玉米粗缩病病原水稻黑条矮缩病毒(RBSDV)基因组的特异性引物检测灰飞虱带毒情况。在来自14个灰飞虱饲养桶的样本中检测出有11个样本中的灰飞虱携带RBSDV.2010年5月在网棚中利用塑料萌种盘萌发玉米实验材料,待植株处于一叶一心期时,将携带RBSDV的灰飞虱按照500头/m2的密度放入网棚中,连续传毒5天。5天后利用虱蚜净将剩余灰飞虱全部杀灭。待植株长至三叶期时,将它们移栽到济南地区吕家村试验田中生长,由于此时适逢田间灰飞虱的迁飞高峰期,而玉米植株又处在对灰飞虱侵染敏感阶段,因而在田间又经历一次田间自然感染。通过两种接毒方法的联合使用,提高了试验材料的粗缩病发病率,提高了鉴定结果的可靠性和重复性。该方法可以在较短时期内检测大量的玉米植株,为鉴定大量分离群体材料的粗缩病抗性和相关的QTL定位奠定了基础。
2.玉米粗缩病的遗传分析
本研究利用课题组创制的“掖478×90110”的RILs的F7：9群体为实验材料,从2008年到2010年,在山东省济南和莱州两个试验点连续种植RIL群体和掖478、90110以及其F1植株,通过田间自然发病的方法和联合接毒鉴定的方法进行粗缩病的抗病性评判。在实验材料开花期对玉米粗缩病相关性状,包括上部节间性状、叶片背面蜡质性状、雄穗发育情况以及粗缩病发病指数等进行观测和统计。按照植株粗缩病的病级和各个性状的等级整理数据,然后对三年两地的实验数据进行遗传分析,结果表明玉米粗缩病在“掖478×90110”的后代中,抗病特性主要由基因型决定,受到环境效应、基因型与环境互作效应的影响较小。粗缩病抗性的广义遗传力范围为0.71～0.94。即在该RILs群体中粗缩病的抗性主要是由遗传因素决定的。分析粗缩病相关性状参数的分布可得出,玉米植株的粗缩病抗性决定于来自90110的主效QTL。
3.玉米粗缩病抗病QTL定位
利用覆盖整个玉米基因组的512对SSR引物在RILs群体中检测标记多态性,选多态性良好且扩增条带清晰的SSR引物用于QTL的定位检测,共计检测到5个控制粗缩病抗性及相关性状的加性QTL,分别位于玉米基因组的2号、6号、7号、8号以及10号染色体上,分别命名为qMRD2、qMRD6、qMRD7、qMRD8以及qMRD10。其中QTL qMRD8几乎在所有的粗缩病相关性状中被检测到,而且qMRD8单独能够解释粗缩病表型变异的12-28.9%,是一个主效QTL。QTL qMRD2、qMRD6和UqMRD7在整个QTL的定位实验中至少被检测到3次,也是重复性良好的粗缩病抗性相关QTL。QTL qMRD10仅仅在上部节间性状中被检测到了一次,且效应值较小,该QTL需要后续的工作继续进行验证。检测到的全部QTL总共可以解释粗缩病表型变异值的41.43-50.84%。本实验中qMRD6、 qMRD7和qMRD83个QTL与之前本实验室王飞等利用SSR-BSA去检测到的粗缩病抗性位点相重合,这提示在这些QTL区及其邻近位置上存在着玉米粗缩病抗性基因。本研究检测到的主效QTL可以通过MAS的方法导入抗病性较弱的骨干自交系中,用于加快玉米粗缩病抗病性育种的进程。
本工作的意义有：1.首次利用田间自然发病结合人工接毒的联合检测方法进行了玉米粗缩病抗性鉴定。该方法可有效提高玉米粗缩病抗性鉴定的准确性和稳定性。2.通过粗缩病抗病性的遗传分析确定了来自“掖478×90110”的RILs群体中抗病性主要由遗传因素决定,且存在抗粗缩病的主效QTL。3.通过粗缩病抗性QTL检测,定位了5个粗缩病抗性相关的QTL,其中3个QTL与SSR-BSA法鉴定出的相关位点重合,可用于采用MAS方法培育抗粗缩病玉米材料。
Maize (Zea mays L.) is the worldwide most important food and forage crop as well as a model plant for genetics and development biology, and plays a decisive role in agriculture. Maize rough dwarf disease (MRDD) is a viral disease that is widely distributed in the worldwide and causes great yield reduction of maize. In China, The pathogen of MRDD has been identified as rice black-streaked dwarf virus (RBSDV). RBSDV is mainly transmitted by a kind of planthopper (Laodelphax sreiatellus Fallen.) in a persistent manner among individuals. When the maize seedlings suffer the large scale migration of planthopper from wheat or other grasses in the field, it may cause the outbreak of MRDD. The MRDD has caused a great and large-scale yield reduction of maize in China for the poor resistance to MRDD of elite inbred lines and hybrids in recent years. The MRDD has become one of the hardest problems in maize production, and obstructs our grain production. The results of the assays of maize resistance to MRDD show that the maize resistance to MRDD take on the features of quantitative trait and is controlled by multiple genes. Meanwhile, Chinese breeders have selected a bulk of maize inbred lines with resistantance to MRDD, such as90110, Qi319, X178et al. To control the outbreak and transmission of MRDD and reduce the loss of maize yield, it is the key step to identify the resistant QTL in maize and improve the elite maize inbred and hybrid using these QTL.
Recently, the researches of maize resistance to MRDD have been accelerated, but the breakthrough progresses are not made. The major reason is the conditions of MRDD are complicated that involve the correlations of the virus, vector-planthopper and maize. It is hard to accurately identify the individual phenotype and to map the QTL conferring resistance to MRDD. But, the phenotype identification is the foundation for mapping and cloning the resistant QTL that develop the appropriate scientific methods to identify maize resistance to MRDD accurately in the segregation populations. In this study, the improved method to identify the maize resistance to MRDD is developed and the QTL conferring resistance to MRDD are identified.
1. The development of combined identification method based on the combination of natural infection and artificial inoculation
In this study, the planthoppers were used as the medium to transmit RBSDV and were cultured in the feeding buckets and fed on the diseased maize leaves from naturally infected field-grown plants with typical symptoms of MRDD in the Jinan area of Shandong province. Subsequently, five adult-planthoppers were randomly selected as a sample to detect RBSDV, and then the RBSDV specific primers were used to screen the samples from different feeding buckets.11out of14samples were positive. May1st in2010, the RILs and their parental lines (90110and Ye478) were germinated at plastic plates in the net house. When the seedlings grew to the1-leaf period, the planthoppers with RBSDV were dispersed into the net house. The average density of planthoppers was about500/m2. After a5-day inoculation, the insecticide was used to kill all of the planthoppers, and the seedlings were maintained to grow continually in the net house and then were transplanted to the experiment fields at the3-leaf stage in Jinan area. The seedlings were infected again by the planthoppers at the migratory flight period of planthoppers in field. The incidence of MRDD and the stability and repetitiveness of identification results were increased by the combination of the artificial inoculation and natural infection. And plenty of maize plants could be screened in a short time by using this method. The combined method has laid the foundation of identifying abundant examples in the segregation population.
2. The genetic analysis of maize rough dwarf disease
The analysis was performed in common maize using a set of recombinant inbred lines (RILs) F7;9derived from 'Ye478×90110'.The RILs population, Ye478,90110and F1were grown continually at the area of Jinan and Laizhou respectively from2008to2010, and then were identified the resistance to MRDD through the method combining the artificial inoculation and natural infection. The phenotypes related to MRDD including the shortened superior internodes, enation, tassel type and disease severity index were evaluated at the flowering stage. Subsequently, the genetic analyses were performed using the3-year experimental data at two plots according to the disease grade and the level of traits related to MRDD. The broad sense heritability of resistance to MRDD is from0.71to0.94, that is to say, the resistance to MRDD was mainly determined by the genotype in the population of Ye478and90110, with minor influence of the environment and the interaction effect between the genotype and environment. The distribution analysis of traits related to MRDD showed that there were major QTL from90110conferring resistance to MRDD.
3. Mapping of QTL conferring resistance to MRDD
The512SSR markers covering the whole maize genome were screened polymorphisms between the two parents Ye478and90110. And the polymorphism SSR markers with clear band were used for QTL mapping by using the RILs. Five additive QTL related to resistance were detected and were mapped on chromosome2,6,7,8and10respectively. The QTL were named as qMRD2, qMRD6, qMRD7, qMRD8and qMRD10, respectively. The qMRD8was a major QTL, that could be detected in all traits mapping, and explain the phenotypic variation of12-28.9%. QTL qMRD2, qMRD6and qMRD7also had good repetitiveness, which were detected3times at least through the whole mapping experiment. The qMRDIO was only detected once at the trait of shortened superior intemodes with relative small effect and need to be verified in subsequent experiment. All of these QTL could explain the phonotypical variance of41.43-50.84%. In this study, the QTL qMRD6, qMRD7and qMRD8were coincidence with the loci that were detected by Dr. Wang using SSR-BSA method, previously. The results suggested that these QTL and the loci nearby these QTL may contain the genes conferring resistance to MRDD. The major QTL detected in this study can be introduced into the elite inbred lines with little resistance to MRDD by MAS to accelerate the proceeding of maize breeding.
In this research, the combined inoculation method including natural infection and artificial inoculation was used to study the resistance to MRDD for the first time, which could enhance the stability and validity of resistance to MRDD, and could increase the detecting amount of segregation population. Moreover, this study proved that the resistance to MRDD was mainly due to the genetic factors through the genetic analysis. There were5QTL conferring resistance to MRDD detected through the QTL mapping, and3of them were coincidence with the loci detected through SSR-BSA method. Finally, this study laid the foundation of MRDD resistance breeding in maize by the method of MAS.
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