基于微卫星DNA标记的马铃薯桃蚜不同种群遗传分化研究
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摘要
马铃薯(Solanum tuberosum L.)是世界上仅次于水稻、小麦、玉米的第四大粮食作物,也是重要的饲料作物和工业原料。我国是世界上最大的马铃薯生产国,马铃薯生产关系到我国甚至世界的粮食安全问题。蚜虫(Aphids)是马铃薯生产中重要的传毒介体昆虫和迁飞性害虫,具有寄主杂、分布广、种类多的特点,不仅直接刺吸马铃薯汁液,而且传播多种病毒病,造成植株生长发育不良、产量和品质下降、品种退化等诸多问题,有效防治蚜虫是马铃薯生产和脱毒种薯繁育的关键环节。受长期的特定环境条件胁迫和寄主选择压力的双重影响,导致蚜虫不同地理种群、不同寄主种群间产生遗传分化,从而形成新的适应性种群,然而蚜虫的迁飞行为又在一定程度上增强了种群间的基因交流,降低种群间的遗传分化。深入研究马铃薯蚜虫优势种群遗传结构和种群间生殖隔离、基因交流等问题,有利于从根本上了解马铃薯蚜虫的适应机制和成灾机理,从而为准确把握其发生规律并进行有效防治提供依据。本研究应用微卫星分子标记技术对不同寄主、不同地理马铃薯优势种群—桃蚜的遗传多样性进行了研究,旨在对不同种群桃蚜相似性进行比较,了解不同寄主间、地区间的基因交流,分析可能的迁飞路线,判断可能的虫源地和迁入地,为马铃薯蚜虫区域性成灾调控和治理提供理论基础。取得的主要研究结果如下:
1.对甘肃天水、定西、张掖三地马铃薯蚜虫形态学鉴定表明:甘肃省马铃薯田间迁飞蚜虫的主要种类有桃蚜、棉蚜、甘蓝蚜、萝卜蚜、豆蚜和长管蚜6种蚜虫,桃蚜是甘肃省马铃薯蚜虫优势蚜虫种类,发生量占到田间总蚜量的53.3%,是马铃薯病毒病的主要传毒介体。
2.针对蚜虫体型微小、体表有外骨骼的特点,采用改进的醋酸钾法(KAC)提取单头蚜虫基因组DNA,并进行了优化:在常温下直接用灭菌的一次性牙签破碎细胞,不易造成交叉污染;提取匀浆液时加入蛋白酶K、延长水浴时间可以促进蚜虫组织彻底消化和促进RNA分解;取消了平衡酚和氯仿抽提,延长DNA沉淀的时间,提高了DNA得率。本方法操作简单,对试剂和设备要求不高,解决了单头蚜虫DNA提取难、不稳定的问题,是一种较为理想的提取蚜虫DNA方法。
3.通过对不同浓度mg2+、引物、dNTP、Taq聚合酶、模板DNA对SSR-PCR扩增影响的实验研究,筛选得到各组分的适宜浓度,建立了桃蚜SSR反应体系:在20μL的反应体系中,Mg2+浓度为1.5mM,dNTP为0.25mM,Taq酶为1.0U/μL,模板DNA为40ng/μL,引物为25ng/μL,为进行桃蚜种群遗传分化的研究奠定基础。
4.从46个微卫星引物中筛选出14对多态性较高的微卫星引物,对桃蚜5种不同寄主植物桃、烟草、甘蓝、辣椒和马铃薯上的桃蚜种群的遗传关系进行了多态性分析,结果表明:14对SSR引物共扩增出70条多态性条带,多态位点比率达到98.59%,DNA片段分子量大小在70～300bp之间。马铃薯桃蚜种群的多态位点最多,为29个,甘蓝的多态位点最少,为6个。各寄主桃蚜种群的Shannon多样性指数0.5382,Nei’s指数0.3631,基因流Nm为0.1343,多样性的变化趋势为:马铃薯桃蚜种群>桃树桃蚜种群>烟草桃蚜种群>辣椒桃蚜种群>甘蓝桃蚜种群。桃蚜寄主种群遗传变异主要存在于种群内。UPGMA聚类结果表明,桃蚜在茄科的烟草和辣椒上的种群亲缘关系最近,与马铃薯桃蚜种群的亲缘关系最远,预示着桃蚜正朝着远离烟草和辣椒种群的方向进化。
5.应用7对具有高度多态性微卫星引物,对来自甘肃省张掖、武威、酒泉、兰州、临洮、临夏、定西、漳县、渭源、岷县、天水、武都、庆阳13个桃蚜地理种群进行了遗传多样性分析,结果表明,7对SSR引物在468个桃蚜个体基因组中共检测到43个多态性位点,多态性条带百分率为95.56%。13个马铃薯桃蚜地理种群观察等位基因数为1.3333,有效等位基因数为1.2293,Nei's(1973)基因多样性指数为0.1311,Shannon指数为0.1898,多态位点百分率为33.33%,基因流Nm为0.3027。临夏种群和临洮种群的遗传多样性较高,漳县种群最低。张掖和武威种群间的遗传距离最小;张掖和庆阳种群间的遗传距离最大。分子方差分析表明,遗传变异的81.03%来自于种群内部。Mantel检测结果表明,种群间的遗传距离和地理距离、海拔差距无显著相关性。
6 .聚类分析表明,马铃薯桃蚜在甘肃省分为河西和河东(以黄河为界)两个大的类群,河西类群中主要包括张掖、酒泉、武威和兰州种群,河东种群中定西、庆阳、会川种群聚为位于甘肃中东部的一支;天水、武都、岷县聚为位于甘肃东南部的一支;漳县种群、临洮种群和临夏种群则各自成为一支。进一步的分析表明,甘肃中部马铃薯优势区桃蚜来源主要有两个自然扩散途径,一是由位于甘肃东边的庆阳等地传入到定西,然后向西南部扩散;另一个则是由东南部的天水等地传入,逐渐向西南部扩散,前者的相关性较后者明显,据此推断:陕北、陇东是陇中马铃薯桃蚜主要虫源地,而陇南是次要虫源地,应加强对上述地区桃蚜的监测和治理,降低虫源基数和迁入量。
本研究将微观的分子遗传学和宏观的昆虫形态学、生态学有机的联系在一起,这对于从理论上进一步揭示马铃薯桃蚜灾变机理和迁飞规律,在实践中有效开展综合防治和甘肃省马铃薯优质种薯繁殖基地建设都具有重要的参考价值。
Potato is not only the the fourth largest grain crop of the world, after rice, wheat and maize, but also an important forage crop and raw material for chemical industry. China is the largest potato producer in the world, its production related to food security of the country and even the world. Aphids is one of the most important virus-vector insect and migration pest attacking potato, with the characteristic of large numbers and dispersed distribution. Colonies of the aphid not only feed upon the potato leaves, but also transmit a variety of plant virus, influencing the normal growth of plant, leading to decrease of yield and quality, resulting in variety degradation of potato, therefore, effective control of aphids is the key step of potato production. Influenced by long-term environmental stress and host selection pressure, different geographical populations and host populations of aphids emerged genetic differentiation, which has formed the new adaptability popuation of aphids. However, the migration of aphids enhances the genetic exchange among the populations to a certain extent, decreased the level of genetic differentiation. To make in-depth research on genetic structure of aphids dominant species, on reproductive isolation and genetic exchange among of the populations, which is beneficial to reveal the mechanism of adaptability and disaster, and provide a basis for correct understanding of occurrance and effective controlling on aphids of potato. In this paper, genetic diversities of different host-population and geo-population of dominant species of potato aphids Myzus persicae were studied by SSR-PCR, the purpose of this study was to compare genetic similarity, understand gene flow, analyse the possible migration routes, estimate the possible emigratory areas and ingoing areas of different population of Myzus persicae aphids of potato, and provide theoretical basis for regional control of aphids. The important results are shown as follows:
1. The results of morphological identification in Tianshui, Dingxi and Zhangye of Gansu province showed that the main species of aphids on potato were Myzus persicae, Aphisgossypii, Brevicoryne brassicae, Lipaphis erysimi, Acyrthasiphon pisum, and Macrosiphum sp, the dominant specie is Myzus persicae, which is the most important mediator about transmitting virus of potato.
2. The method of“improved KAC”were adopted to extract the genomic DNA of single-aphid, this method was optimised as follows:disposable and sterilized toothpicks were used to break up the cells, which can prevents cross-contamination risk. proteinase K were used, prolong the time of water bath was used to improve digest aphids and break down of RNA when homogenate of single-aphid were extracted. To increase DNA yield rate by the steps of balanced phenol and chloroform were canceled,and the DNA precipitation time was prolonged. This method is ideal way to extract the genomic DNA of aphid,because of simple operation and undemanding equipment, it also could solve the problem that extracting of genomic DNA of single-aphid was difficult and unstable.
3. The reaction conditions of SSR experiment of Myzus persicae were discussed, and the results showed the components of optimal concentration SSR reaction system of 20μL as follows: Mg2+ 1.5mM,Deoxyribonucleotide triphosphate 0.25mM,Taq DNA Polymerase 1.0U/μL,DNA template 40ng/μL,primers 25ng/μL, which was a basis for the study on genetic divergency of Myzus persicae populations.
4. Screened 14 pairs of highly polymorphic primers from 46 pairs of primers for aphids and analyzed the genetic relationship of peach,tobacco,cabbage,pepper and potato,which were the host-plants of Myzus persicae. The result of this analysis showed that 70 polymorphic bands were detected using 14 pairs of SSR primers in the five populations,percentage of polymorphic bands were 98.59%,the molecular weight of DNA fragments were from 70bp to 300bp. 29 polymorphic loci of Myzus persicae detected from the population of potato was mostly, 6 polymorphic loci from cabbage was leastly. The Shannon-Wiener diversity and Nei’s genetic diversity indexes were respectively 0.5382 and 0.3631, the gene flow was 0.1343 amomg population, there is an tendency of Myzus persicae population diversity in 5 host plants, the popato>the peach>the tobacco>the pepper>the cabbage, and the genetic variation was mainly the inner of population. Using UPGMA method, the relationship between two host-populations of Myzus persicae on tobacco and pepper are the closest, and the farthest is potato, which indicated that the evolution of Myzus persicae is towards the direction of keeping away from tobacco and pepper.
5. Seven highly polymorphic SSR primers were employed to investigate the genetic variation and structure of thirteen populations of potato aphids Myzus persicae from Zhangye, Wuwei, Jiuquan, Lanzhou, Lintao, Linxia, Dingxi, Zhangxian, Weiyuan, Minxian, Tianshui, Wudu and Qingyang in Gansu province. The result showed that 43 polymorphic locus were detected using 7 pairs of SSR primers in the 468 individuals of 13 populations,percentage of polymorphic bands were 95.56%. The number of observed alleies was 1.3333, and that of effective alleles was 1.2293, the Nei’s genetic diversity and Shannon-Wiener diversity indexes were respectively 0.1311 and 0.1898, percentage of polymorphic locus were 33.33%. The gene flow was 0.3027 amomg population. The population genetic diversity of Linxia and Lintao were higher, Zhangxian was lower. The genetic distance is the nearest between Zhangye and Wuwei, it is the farthest between Zhangye and Qingyang. Analysis of molecular variance showed that 81.03% genetic variation existed within the populations of Myzus persicae on potato. Mantel-test showed that genetic distances had no significant correlation with geographic distance and gap of elevation among popuations.
6. The cluster analysis showed that the potato aphids Myzus persicae collected from Gansu province could divide into two groups of Hexi and Hedong which of the boundary was the Huanghe River. Hexi group included with Zhangye, Jiuquan, Wuwei and Lanzhou population, in the Hedong group, Dingxi, Qingyang and Huichuan were classified one cluster that located east-central of Gansu province; Tianshui, Wudu and Minxian were southeast; Zhangxian, Lintao and Linxia were individually. A further analysis showed that there were two natural ingoing paths of the potato aphids Myzus persicae located midland of Gansu province where potato is the main crop, one maybe migrate from Qingyang located east of Gansu,the other maybe from Tianshui located southeast of Gansu,then Myzus persicae dispersed from midland to southwest of Gansu province. The relevance of the former was significant than latter. From this, we could infer that main native area of potato aphids Myzus persicae maybe north of Shanxi and east of Gansu, secondary area maybe southeast of Gansu province. We suggest to strengthen monitoring and controlling of the potato aphids Myzus persicae in these area to reduced the quantity of overwinter and ingoing population.
This study realized an organic combo of the microscopic molecular genetic and insect macroscopic morphology and ecology,which have important referential value for further to reveal the disaster mechanism and migration rule in theory,and in practice,to carry out efficiently integrated aphids control and to build reproduction base of the fine-quality potato tuber seeds.
1.对甘肃天水、定西、张掖三地马铃薯蚜虫形态学鉴定表明:甘肃省马铃薯田间迁飞蚜虫的主要种类有桃蚜、棉蚜、甘蓝蚜、萝卜蚜、豆蚜和长管蚜6种蚜虫,桃蚜是甘肃省马铃薯蚜虫优势蚜虫种类,发生量占到田间总蚜量的53.3%,是马铃薯病毒病的主要传毒介体。
2.针对蚜虫体型微小、体表有外骨骼的特点,采用改进的醋酸钾法(KAC)提取单头蚜虫基因组DNA,并进行了优化:在常温下直接用灭菌的一次性牙签破碎细胞,不易造成交叉污染;提取匀浆液时加入蛋白酶K、延长水浴时间可以促进蚜虫组织彻底消化和促进RNA分解;取消了平衡酚和氯仿抽提,延长DNA沉淀的时间,提高了DNA得率。本方法操作简单,对试剂和设备要求不高,解决了单头蚜虫DNA提取难、不稳定的问题,是一种较为理想的提取蚜虫DNA方法。
3.通过对不同浓度mg2+、引物、dNTP、Taq聚合酶、模板DNA对SSR-PCR扩增影响的实验研究,筛选得到各组分的适宜浓度,建立了桃蚜SSR反应体系:在20μL的反应体系中,Mg2+浓度为1.5mM,dNTP为0.25mM,Taq酶为1.0U/μL,模板DNA为40ng/μL,引物为25ng/μL,为进行桃蚜种群遗传分化的研究奠定基础。
4.从46个微卫星引物中筛选出14对多态性较高的微卫星引物,对桃蚜5种不同寄主植物桃、烟草、甘蓝、辣椒和马铃薯上的桃蚜种群的遗传关系进行了多态性分析,结果表明:14对SSR引物共扩增出70条多态性条带,多态位点比率达到98.59%,DNA片段分子量大小在70～300bp之间。马铃薯桃蚜种群的多态位点最多,为29个,甘蓝的多态位点最少,为6个。各寄主桃蚜种群的Shannon多样性指数0.5382,Nei’s指数0.3631,基因流Nm为0.1343,多样性的变化趋势为:马铃薯桃蚜种群>桃树桃蚜种群>烟草桃蚜种群>辣椒桃蚜种群>甘蓝桃蚜种群。桃蚜寄主种群遗传变异主要存在于种群内。UPGMA聚类结果表明,桃蚜在茄科的烟草和辣椒上的种群亲缘关系最近,与马铃薯桃蚜种群的亲缘关系最远,预示着桃蚜正朝着远离烟草和辣椒种群的方向进化。
5.应用7对具有高度多态性微卫星引物,对来自甘肃省张掖、武威、酒泉、兰州、临洮、临夏、定西、漳县、渭源、岷县、天水、武都、庆阳13个桃蚜地理种群进行了遗传多样性分析,结果表明,7对SSR引物在468个桃蚜个体基因组中共检测到43个多态性位点,多态性条带百分率为95.56%。13个马铃薯桃蚜地理种群观察等位基因数为1.3333,有效等位基因数为1.2293,Nei's(1973)基因多样性指数为0.1311,Shannon指数为0.1898,多态位点百分率为33.33%,基因流Nm为0.3027。临夏种群和临洮种群的遗传多样性较高,漳县种群最低。张掖和武威种群间的遗传距离最小;张掖和庆阳种群间的遗传距离最大。分子方差分析表明,遗传变异的81.03%来自于种群内部。Mantel检测结果表明,种群间的遗传距离和地理距离、海拔差距无显著相关性。
6 .聚类分析表明,马铃薯桃蚜在甘肃省分为河西和河东(以黄河为界)两个大的类群,河西类群中主要包括张掖、酒泉、武威和兰州种群,河东种群中定西、庆阳、会川种群聚为位于甘肃中东部的一支;天水、武都、岷县聚为位于甘肃东南部的一支;漳县种群、临洮种群和临夏种群则各自成为一支。进一步的分析表明,甘肃中部马铃薯优势区桃蚜来源主要有两个自然扩散途径,一是由位于甘肃东边的庆阳等地传入到定西,然后向西南部扩散;另一个则是由东南部的天水等地传入,逐渐向西南部扩散,前者的相关性较后者明显,据此推断:陕北、陇东是陇中马铃薯桃蚜主要虫源地,而陇南是次要虫源地,应加强对上述地区桃蚜的监测和治理,降低虫源基数和迁入量。
本研究将微观的分子遗传学和宏观的昆虫形态学、生态学有机的联系在一起,这对于从理论上进一步揭示马铃薯桃蚜灾变机理和迁飞规律,在实践中有效开展综合防治和甘肃省马铃薯优质种薯繁殖基地建设都具有重要的参考价值。
Potato is not only the the fourth largest grain crop of the world, after rice, wheat and maize, but also an important forage crop and raw material for chemical industry. China is the largest potato producer in the world, its production related to food security of the country and even the world. Aphids is one of the most important virus-vector insect and migration pest attacking potato, with the characteristic of large numbers and dispersed distribution. Colonies of the aphid not only feed upon the potato leaves, but also transmit a variety of plant virus, influencing the normal growth of plant, leading to decrease of yield and quality, resulting in variety degradation of potato, therefore, effective control of aphids is the key step of potato production. Influenced by long-term environmental stress and host selection pressure, different geographical populations and host populations of aphids emerged genetic differentiation, which has formed the new adaptability popuation of aphids. However, the migration of aphids enhances the genetic exchange among the populations to a certain extent, decreased the level of genetic differentiation. To make in-depth research on genetic structure of aphids dominant species, on reproductive isolation and genetic exchange among of the populations, which is beneficial to reveal the mechanism of adaptability and disaster, and provide a basis for correct understanding of occurrance and effective controlling on aphids of potato. In this paper, genetic diversities of different host-population and geo-population of dominant species of potato aphids Myzus persicae were studied by SSR-PCR, the purpose of this study was to compare genetic similarity, understand gene flow, analyse the possible migration routes, estimate the possible emigratory areas and ingoing areas of different population of Myzus persicae aphids of potato, and provide theoretical basis for regional control of aphids. The important results are shown as follows:
1. The results of morphological identification in Tianshui, Dingxi and Zhangye of Gansu province showed that the main species of aphids on potato were Myzus persicae, Aphisgossypii, Brevicoryne brassicae, Lipaphis erysimi, Acyrthasiphon pisum, and Macrosiphum sp, the dominant specie is Myzus persicae, which is the most important mediator about transmitting virus of potato.
2. The method of“improved KAC”were adopted to extract the genomic DNA of single-aphid, this method was optimised as follows:disposable and sterilized toothpicks were used to break up the cells, which can prevents cross-contamination risk. proteinase K were used, prolong the time of water bath was used to improve digest aphids and break down of RNA when homogenate of single-aphid were extracted. To increase DNA yield rate by the steps of balanced phenol and chloroform were canceled,and the DNA precipitation time was prolonged. This method is ideal way to extract the genomic DNA of aphid,because of simple operation and undemanding equipment, it also could solve the problem that extracting of genomic DNA of single-aphid was difficult and unstable.
3. The reaction conditions of SSR experiment of Myzus persicae were discussed, and the results showed the components of optimal concentration SSR reaction system of 20μL as follows: Mg2+ 1.5mM,Deoxyribonucleotide triphosphate 0.25mM,Taq DNA Polymerase 1.0U/μL,DNA template 40ng/μL,primers 25ng/μL, which was a basis for the study on genetic divergency of Myzus persicae populations.
4. Screened 14 pairs of highly polymorphic primers from 46 pairs of primers for aphids and analyzed the genetic relationship of peach,tobacco,cabbage,pepper and potato,which were the host-plants of Myzus persicae. The result of this analysis showed that 70 polymorphic bands were detected using 14 pairs of SSR primers in the five populations,percentage of polymorphic bands were 98.59%,the molecular weight of DNA fragments were from 70bp to 300bp. 29 polymorphic loci of Myzus persicae detected from the population of potato was mostly, 6 polymorphic loci from cabbage was leastly. The Shannon-Wiener diversity and Nei’s genetic diversity indexes were respectively 0.5382 and 0.3631, the gene flow was 0.1343 amomg population, there is an tendency of Myzus persicae population diversity in 5 host plants, the popato>the peach>the tobacco>the pepper>the cabbage, and the genetic variation was mainly the inner of population. Using UPGMA method, the relationship between two host-populations of Myzus persicae on tobacco and pepper are the closest, and the farthest is potato, which indicated that the evolution of Myzus persicae is towards the direction of keeping away from tobacco and pepper.
5. Seven highly polymorphic SSR primers were employed to investigate the genetic variation and structure of thirteen populations of potato aphids Myzus persicae from Zhangye, Wuwei, Jiuquan, Lanzhou, Lintao, Linxia, Dingxi, Zhangxian, Weiyuan, Minxian, Tianshui, Wudu and Qingyang in Gansu province. The result showed that 43 polymorphic locus were detected using 7 pairs of SSR primers in the 468 individuals of 13 populations,percentage of polymorphic bands were 95.56%. The number of observed alleies was 1.3333, and that of effective alleles was 1.2293, the Nei’s genetic diversity and Shannon-Wiener diversity indexes were respectively 0.1311 and 0.1898, percentage of polymorphic locus were 33.33%. The gene flow was 0.3027 amomg population. The population genetic diversity of Linxia and Lintao were higher, Zhangxian was lower. The genetic distance is the nearest between Zhangye and Wuwei, it is the farthest between Zhangye and Qingyang. Analysis of molecular variance showed that 81.03% genetic variation existed within the populations of Myzus persicae on potato. Mantel-test showed that genetic distances had no significant correlation with geographic distance and gap of elevation among popuations.
6. The cluster analysis showed that the potato aphids Myzus persicae collected from Gansu province could divide into two groups of Hexi and Hedong which of the boundary was the Huanghe River. Hexi group included with Zhangye, Jiuquan, Wuwei and Lanzhou population, in the Hedong group, Dingxi, Qingyang and Huichuan were classified one cluster that located east-central of Gansu province; Tianshui, Wudu and Minxian were southeast; Zhangxian, Lintao and Linxia were individually. A further analysis showed that there were two natural ingoing paths of the potato aphids Myzus persicae located midland of Gansu province where potato is the main crop, one maybe migrate from Qingyang located east of Gansu,the other maybe from Tianshui located southeast of Gansu,then Myzus persicae dispersed from midland to southwest of Gansu province. The relevance of the former was significant than latter. From this, we could infer that main native area of potato aphids Myzus persicae maybe north of Shanxi and east of Gansu, secondary area maybe southeast of Gansu province. We suggest to strengthen monitoring and controlling of the potato aphids Myzus persicae in these area to reduced the quantity of overwinter and ingoing population.
This study realized an organic combo of the microscopic molecular genetic and insect macroscopic morphology and ecology,which have important referential value for further to reveal the disaster mechanism and migration rule in theory,and in practice,to carry out efficiently integrated aphids control and to build reproduction base of the fine-quality potato tuber seeds.
引文
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