栽培花生产量、品质和抗病性的遗传分析与QTL定位研究
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摘要
花生是重要的油料作物和经济作物。我国是世界上主要的花生生产、消费和出口国。因而,开展花生产量、品质、抗病性等重要性状的遗传分析、遗传图谱的构建及QTL定位具有重要的理论意义和应用价值。产量、品质和抗病性都属于典型的复杂数量性状,传统的育种方法很难实现花生诸多性状的的同步提高。现代遗传学和分子生物学的发展可将控制复杂性状的多个因子分解成简单的孟德尔遗传因子进行研究,实现对复杂数量性状的操控,实现产量、品质和抗病性等诸多性状的分子标记辅助选择和改良。本研究以多个花生重组近交系群体和F2群体为材料,采用以PCR为基础的分子标记技术,构建了花生分子遗传图谱,并结合多年份多环境的农艺性状的调查结果对花生产量、品质和抗病性等性状进行QTL定位和遗传效应研究,主要研究结果如下：
1、构建了栽培花生品种间杂交后代的重组自交系群体(郑8903×豫花4号)。对该群体产量、品质和抗病性相关的20个性状的调查表明,该群体除抗网斑病性状外,其余性状在群体中均符合正态分布,达到群体构建的预期目标,适合于进行数量性状的遗传和QTL定位等研究。
2、以本实验室构建的重组自交系群体为研究对象,对其进行多年份的重复试验,采用P1、P2及RIL家系联合分析方法对花生的产量及其相关性状、品质性状和抗病性进行主基因+多基因遗传模型分析,结果表明：主茎高、结果枝数、单株饱果数、出仁率符合3对主基因遗传模型；百果重、百仁重符合2对主基因加多基因遗传模型；蛋白质含量的遗传符合无主基因遗传模型；脂肪含量的遗传符合2对主基因加多基因遗传模型；油酸(18：1Δ9c)、亚油酸(18：2Δ9c,12c)和棕榈酸(16：0)的遗传符合2对主基因遗传模型；硬脂酸(18：0)和花生酸(20：0)的遗传符合2对主基因加多基因遗传模型；油亚比和山嵛酸(22：0)的遗传符合3对主基因遗传模型；网斑病抗性符合3对主基因加多基因遗传模型。遗传率分析结果表明,油酸、亚油酸、油亚比和棕榈酸的主基因遗传率均在60%以上,网斑病抗性的主基因遗传率在80%以上,属干主基因控制的遗传性状。
3、以重组自交系群体(郑8903×豫花4号)构建的框架图为基础,结合“郑9001×郑8903”、“白籽x豫花4号”、“开农白2号×豫花4号”等三个作图群体,以共有标记作为桥梁,构建了包含17个连锁群,共有101个标记的栽培花生复合遗传图谱,总图距为953.88 cM。
4、使用WinQTLCart 2.5和QTLNetwork 2.0两种分析软件进行QTL的检测,获得与形态、产量、品质(脂肪、蛋白、脂肪酸)、网斑病抗性相关的加性QTL位点62个筛选出与18个性状QTL位点紧密连锁(遗传距离小于3 cM)的SSR标记22个,向分子标记辅助选择育种技术的建立迈出了坚实的一步。
Peanut is an important oil crops and economic crops. China is the biggest country on production, consumption, and export of peanut in the world. Yield, quality and disease resistance are typical complex quantitative traits, traditional breeding methods is difficult to simultaneously improve many traits in peanut. With the development of modern genetics and molecular biology, complex traits can be decomposed into simple Mendelian genetic factor, so it is possible to manipulate the complex quantitative traits in crop improvement for yield, quality and disease resistance, using the method of marker-assisted selection (MAS). Therefore it is necessary and very important theoretically and practically to study the inheritance of yield, quality, disease resistance and the other important traits in peanut, to construct the linkage map, as well as the QTL mapping. In present research, molecular genetic map was constructed in peanuts, using several genetic populations included recombinant inbred lines (RILs) and F2 population as materials, based on SSR molecular markers and integrate experimental data of two year's environments, the genetic effects of agronomic traits on yield, quality, and disease resistance, as well as their QTL are studied, and the main results are as follows:
1. A recombinant inbred lines (RIL) derived from Zheng8903×Yuhua 4 in Arachis hypogaea L was developed. Investigation of 20 traits related to yield, quality and disease resistance showed that all the traits except of resistance disease trait in groups were conformed to the normal distribution, suitable in studying for inheritance of quantitative traits and QTL mapping.
2. Using the developed RILs population derived from a cross of "Zheng8903×Yuhua 4" as the material, and joint segregation analysis of major gene with minor genes mixed inheritance model as method, the inheritance of yield, quality and the resistance to web blotch disease in peanut were analyzed. As the results showed that main stem length, total bearing branches, mature pods per plant and kernel percent were controlled by three major genes, and those of 100-pod weight and 100-kernel weight followed two major genes with several minor genes. The inheritance of protein content in peanut was complied with the non-major gene model, and the fat was controlled by two major gene with several minor genes, while those of oleic acid content, linoleic acid content and palmitic acid content were controlled by two major genes only, and O/L and Behenic acid content were controlled by three major genes. The disease resistance to web blotch was controlled by three major genes with several minor genes as well. For the heritability, the results showed that those of oleic acid content, linoleic acid content and palmitic acid content were over 60%, and that of the disease resistance to web blotch was over 80%, which belong to the quality traits controlled by major gene(s).
3. Using a RIL population derived from the cross of "Zheng8903×Yuhua 4" as the basic population, combined with other the RIL population derived from "Zheng9001×Zheng8903" and two F2 population, "Baizi×Yuhua 4", "Kainongbai 2×Yuhua 4", with the help of the Join Map 3.0 and Map Chart 2.1, a composite linkage map of Arachis hypogaea L. including 17 linkage groups and 101 marker loci was constructed. The linkage map was 953.88 cM in total length with the longest linkage group of 14 SSR markers and was 158.8 cM.
4. The 62 additive QTLs related to shape, yield, quality(fat, protein, fatty acids) and the disease resistance to web blotch in peanut were mapped through the jointed linkage map based on a RILs population derived from Zheng8903×Yuhua 4, and field investigation data from two years and two locations, using the software of WinQTLCart 2.5 and QTLNetwork 2.0. And selected 18 loc trait closely linked QTL (genetic distance of less than 3 cM) of the 22 SSR markers, molecular marker assisted breeding to the establishment of technology has taken a solid step forward.
1、构建了栽培花生品种间杂交后代的重组自交系群体(郑8903×豫花4号)。对该群体产量、品质和抗病性相关的20个性状的调查表明,该群体除抗网斑病性状外,其余性状在群体中均符合正态分布,达到群体构建的预期目标,适合于进行数量性状的遗传和QTL定位等研究。
2、以本实验室构建的重组自交系群体为研究对象,对其进行多年份的重复试验,采用P1、P2及RIL家系联合分析方法对花生的产量及其相关性状、品质性状和抗病性进行主基因+多基因遗传模型分析,结果表明：主茎高、结果枝数、单株饱果数、出仁率符合3对主基因遗传模型；百果重、百仁重符合2对主基因加多基因遗传模型；蛋白质含量的遗传符合无主基因遗传模型；脂肪含量的遗传符合2对主基因加多基因遗传模型；油酸(18：1Δ9c)、亚油酸(18：2Δ9c,12c)和棕榈酸(16：0)的遗传符合2对主基因遗传模型；硬脂酸(18：0)和花生酸(20：0)的遗传符合2对主基因加多基因遗传模型；油亚比和山嵛酸(22：0)的遗传符合3对主基因遗传模型；网斑病抗性符合3对主基因加多基因遗传模型。遗传率分析结果表明,油酸、亚油酸、油亚比和棕榈酸的主基因遗传率均在60%以上,网斑病抗性的主基因遗传率在80%以上,属干主基因控制的遗传性状。
3、以重组自交系群体(郑8903×豫花4号)构建的框架图为基础,结合“郑9001×郑8903”、“白籽x豫花4号”、“开农白2号×豫花4号”等三个作图群体,以共有标记作为桥梁,构建了包含17个连锁群,共有101个标记的栽培花生复合遗传图谱,总图距为953.88 cM。
4、使用WinQTLCart 2.5和QTLNetwork 2.0两种分析软件进行QTL的检测,获得与形态、产量、品质(脂肪、蛋白、脂肪酸)、网斑病抗性相关的加性QTL位点62个筛选出与18个性状QTL位点紧密连锁(遗传距离小于3 cM)的SSR标记22个,向分子标记辅助选择育种技术的建立迈出了坚实的一步。
Peanut is an important oil crops and economic crops. China is the biggest country on production, consumption, and export of peanut in the world. Yield, quality and disease resistance are typical complex quantitative traits, traditional breeding methods is difficult to simultaneously improve many traits in peanut. With the development of modern genetics and molecular biology, complex traits can be decomposed into simple Mendelian genetic factor, so it is possible to manipulate the complex quantitative traits in crop improvement for yield, quality and disease resistance, using the method of marker-assisted selection (MAS). Therefore it is necessary and very important theoretically and practically to study the inheritance of yield, quality, disease resistance and the other important traits in peanut, to construct the linkage map, as well as the QTL mapping. In present research, molecular genetic map was constructed in peanuts, using several genetic populations included recombinant inbred lines (RILs) and F2 population as materials, based on SSR molecular markers and integrate experimental data of two year's environments, the genetic effects of agronomic traits on yield, quality, and disease resistance, as well as their QTL are studied, and the main results are as follows:
1. A recombinant inbred lines (RIL) derived from Zheng8903×Yuhua 4 in Arachis hypogaea L was developed. Investigation of 20 traits related to yield, quality and disease resistance showed that all the traits except of resistance disease trait in groups were conformed to the normal distribution, suitable in studying for inheritance of quantitative traits and QTL mapping.
2. Using the developed RILs population derived from a cross of "Zheng8903×Yuhua 4" as the material, and joint segregation analysis of major gene with minor genes mixed inheritance model as method, the inheritance of yield, quality and the resistance to web blotch disease in peanut were analyzed. As the results showed that main stem length, total bearing branches, mature pods per plant and kernel percent were controlled by three major genes, and those of 100-pod weight and 100-kernel weight followed two major genes with several minor genes. The inheritance of protein content in peanut was complied with the non-major gene model, and the fat was controlled by two major gene with several minor genes, while those of oleic acid content, linoleic acid content and palmitic acid content were controlled by two major genes only, and O/L and Behenic acid content were controlled by three major genes. The disease resistance to web blotch was controlled by three major genes with several minor genes as well. For the heritability, the results showed that those of oleic acid content, linoleic acid content and palmitic acid content were over 60%, and that of the disease resistance to web blotch was over 80%, which belong to the quality traits controlled by major gene(s).
3. Using a RIL population derived from the cross of "Zheng8903×Yuhua 4" as the basic population, combined with other the RIL population derived from "Zheng9001×Zheng8903" and two F2 population, "Baizi×Yuhua 4", "Kainongbai 2×Yuhua 4", with the help of the Join Map 3.0 and Map Chart 2.1, a composite linkage map of Arachis hypogaea L. including 17 linkage groups and 101 marker loci was constructed. The linkage map was 953.88 cM in total length with the longest linkage group of 14 SSR markers and was 158.8 cM.
4. The 62 additive QTLs related to shape, yield, quality(fat, protein, fatty acids) and the disease resistance to web blotch in peanut were mapped through the jointed linkage map based on a RILs population derived from Zheng8903×Yuhua 4, and field investigation data from two years and two locations, using the software of WinQTLCart 2.5 and QTLNetwork 2.0. And selected 18 loc trait closely linked QTL (genetic distance of less than 3 cM) of the 22 SSR markers, molecular marker assisted breeding to the establishment of technology has taken a solid step forward.
引文
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