入侵植物加拿大一枝黄花（菊科）的遗传多样性和居群遗传结构研究
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摘要
生物入侵(Biological invasion)是指生物从其原来的分布区通过自然或者人为方式扩散到比较遥远的新的区域,并在新的区域繁衍,对当地的自然生态系统、生物多样性以及人类健康等产生负面影响。随着全球经济的发展和贸易活动的增加,生物入侵现象逐渐增多。生物入侵已经和全球气候变化,生境破坏一起被列为世界最棘手的三大环境难题。对入侵种的遗传多样性和居群遗传结构进行研究,不但便于掌握入侵种的入侵进化过程,揭示它的起源方式,还能够为探索其治理与根除的措施提供必要的理论依据。
加拿大一枝黄花(Solidago canadensis L.)属于菊科(Asteraceae),一枝黄花属(Solidago L.)。自交不亲和,但繁殖能力特别强,能够通过异交的方式进行有性繁殖,也通过根系进行克隆繁殖。加拿大一枝黄花原产于北美州,现在已入侵到世界许多地方。加拿大一枝黄花于1935年作为庭院花卉引入上海与苏南各市,20世纪80年代开始在我国的浙江,安徽,江西等地普遍种植,后来在长江三角洲的十几个省份扩散蔓延。对农业生产和生态环境形成直接的威胁和破坏,造成巨大的经济损失。
1采用双重抑制PCR法和复合微卫星引物的方法开发出12对加拿大一枝黄花的微卫星引物。选取35个加拿大一枝黄花的样本检测12对引物的效果。12对引物分别能扩增出2至14个等位基因,观察杂合度(Observed heterozygosity, Ho)从0.0732(SC10)到0.7391(SC45),期望杂合度(Expected heterozygosity, He)从0.1177(SC10)到0.8687(SC51),说明开发的引物能揭示较高的遗传多态性水平,可以用于加拿大一枝黄花的入侵遗传机制研究。
2选取8对微卫星引物分析中国15个居群的加拿大一枝黄花的遗传多样性水平和遗传结构。中国15个居群总体平均的多态性百分比(PPL)、平均等位基因数目(Na)、有效等位基因数目(Ne)、Nei's遗传多样性指数(H)、Shannon信息指数(1)、居群内近交系数(Fis)值为91.7%、3.758、2.71、0.529、0.981、-0.191。大多数居群的遗传多样性水平指数都较高而且相差不大,仅湖北宜昌(YC)居群遗传多样性水平偏低。AMOVA分析显示大多数的遗传变异存在于居群内部,总的基因流(Nm)值为1.207。采用UPGMA法对15个居群进行聚类分析显示没有明确的地理分布结构,PCoA分析和聚类结果相符合。IBD检测发现15个居群的地理距离和遗传距离之间没有显著性相关。加拿大一枝黄花的种子和花粉都很容易传播,而且生长于经济发达的大城市和道路两旁,所以基因流很频繁,能够保持较高的遗传多样性水平,而且居群间的差异不大。
3选取3对AFLP引物分析中国15个居群的加拿大一枝黄花的遗传多样性水平和遗传结构。15个居群总体平均的PPL、Na、Ne、H、I值分别为67.87%、1.524、1.46、0.259、0.381。AMOVA分析也显示出大多数的遗传变异存在于居群内部,总的基因流(Nm)值为1.0797。采用UPGMA法对15个居群进行聚类分析显示没有明确的地理分布结构,PCoA分析和聚类结果相符合。IBD检测发现15个居群的地理距离和遗传距离之间没有显著性相关。AFLP标记所检测到的遗传多样性水平虽低于SSR标记检测结果,但仍然显示出较高水平。对比显示两种标记都能揭示较高的遗传多样性差异,但SSR分子标记更加方便快捷,而AFLP标记则能检测整个基因组的变异且更加经济节约成本。
4使用3对AFLP引物分析美国17个居群的加拿大一枝黄花的遗传多样性水平和遗传结构。17个居群总体平均的PPL、Na、Ne、H、I值分别为70.76%、1.572、1.487、0.279、0.400。AMOVA分析显示出大多数的遗传变异存在于居群内部,总的基因流(Nm)值为1.581。采用UPGMA法对17个居群进行聚类分析显示也没有明确的地理分布结构,PCoA分析和聚类结果相符合。IBD检测发现17个居群的地理距离和遗传距离之间没有显著性相关。将美国和中国居群对比分析,发现中国的加拿大一枝黄花遗传多样性水平没有降低,仅基因流水平略低一些。UPGMA法对两个国家的32个居群进行聚类分析显示能够明显聚成中国和美国两类；而苏州(SZ)居群和美国的居群聚类在一起,这可能是因为苏州(SZ)居群位于首次引进加拿大一枝黄花的区域范围内,所以存在和美国较多的相似的遗传基因。中国的加拿大一枝黄花的遗传多样性水平较高,一方面由于其多起源,另一方面由于其特殊的繁育系统。
Biological invasion is the species disperses from the native distribution area to remote new areas by natural or human activities, and causes damage for the local natural ecosystems, biodiversity and human health. Associated with the increase of global economy and trade activities, the biological invasion is become more and more common. Now, biological invasions as well as global climate change and habitat destruction are considered as the world's three most intractable environmental problems. Exploring the genetic diversity of invasive plant is not only helpful for understanding the evolution history of invasive species, but also providing necessary theory for control invasive species. In addition, the possible origin areas for the invasive species could be revealed by analyzing the genetic structure.
Solidago canadensis belongs to the Asteraceae. It is a self-incompatible, but could propagate through outcrossing and asexual reproduction. S. canadensis is native to North America but now have invaded many parts of the world. S. canadensis was introduced to Shanghai and southern Jiangsu (China) in1935as an ornamental plant, but escaping from gardens and spreading throughout the southeastern China in80s of the20th century. It has occupied the farmland and forests and caused huge economic losses. S. canadensis has caused damage to agriculture and environment, resulted in huge great economic losses.
112microsatellite loci were isolated for S. canadensis using dual-suppression polymerase chain reaction (PCR) technique and improved technique for isolating compound SSR markers. The quality of the12pairs of primers was analyzed using35samples of S. canadensis:12pairs of primers can amplify2-14alleles, Observed Heterozygosity (Ho) ranged from0.0732(SC10) to0.7391(SC45), while Expected Heterozygosity (He) ranged from0.1177(SC10) to0.8687(SC51). These primers were able to get high level of polymorphism, could be used for the genetic diversity and genetic structure analysis of S. canadensis.
2The genetic diversity and genetic structure of15Chinese populations was analyzed by8microsatellite primers. The average percentage of polymorphism (PPL) for the15populations is91.7%, the average number of alleles (Na) is3.758, effective number of alleles (Ne) is2.71, Nei's genetic diversity index (H) is0.529, Shannon information index (Ⅰ) is0.981, fixation index (Fis) within populations is-0.191. The genetic diversity for most populations is high except the Yichang (YC) populations, which has a relative low level of genetic diversity. AMOVA analysis showed that most genetic variation occurred within populations and the total gene flow (Nm) is1.207. UPGMA and PCoA analyses resulted similar results that no clear geographical distribution structure was shown. IBD analyses suggested the geographical distance and genetic distance of the15populations was not significantly correlated. S.canadensis usually grow in large cities and along the both sides of road, the gene flow is very frequently. Therefore, it has a high level of genetic diversity, and the interpopulation variation is slight.
3The genetic diversity and genetic structure of15Chinese populations were analyzed by3AFLP primers. The overall average of15populations of PPL、Na、Ne、H、 I were shown to be67.87%、1.524、1.46、0.259、0.381respectively. Most of genetic variation were shown to be existed within populations by AMOVA analysis, and the total gene flow (Nm) was shown to be1.0797. UPGMA and PCoA analyses resulted similar results that no clear geographical distribution structure was shown. Different from the results recovered from SSR primers, IBD analyses suggested the geographical distance and genetic distance of the15populations was significantly negative. Although the genetic diversity recovered by AFLP primers is relatively lower than that recovered by SSR, the genetic diversity is also high. SSR marker is faster and convenient, while AFLP markers more economical and could be able to detect the variation throughout the whole genome.
4The genetic diversity and genetic structure of17USA populations were analyzed by3AFLP primers. The overall average of17populations of PPL、Na、Ne、H、Iwere shown to be70.76%、1.572、1.487、0.279、0.400respectively. Most of genetic variation were shown to be existed within populations by AMOVA analysis, and the total gene flow (Nm) was shown to be1.581. UPGMA and PCoA analyses resulted similar results that no clear geographical distribution structure was shown. IBD analyses suggested the geographical distance and genetic distance of the17populations was significantly negative. The genetic diversity of USA populations is similar to that of Chinese populations. The populations from the two countries were divided into two groups in the UPGMA analyses except the SZ (China) population, which was clustered together with USA populations. The position of SZ population is possibly because SZ population lies in the first introduction areas of S. canadensis, so it is similar to USA population. The study suggested the S. canadensis from Chinese has a high genetic diversity, it is possibly related to the multiple origins, also related to the distinctively reproduction system of this species.
加拿大一枝黄花(Solidago canadensis L.)属于菊科(Asteraceae),一枝黄花属(Solidago L.)。自交不亲和,但繁殖能力特别强,能够通过异交的方式进行有性繁殖,也通过根系进行克隆繁殖。加拿大一枝黄花原产于北美州,现在已入侵到世界许多地方。加拿大一枝黄花于1935年作为庭院花卉引入上海与苏南各市,20世纪80年代开始在我国的浙江,安徽,江西等地普遍种植,后来在长江三角洲的十几个省份扩散蔓延。对农业生产和生态环境形成直接的威胁和破坏,造成巨大的经济损失。
1采用双重抑制PCR法和复合微卫星引物的方法开发出12对加拿大一枝黄花的微卫星引物。选取35个加拿大一枝黄花的样本检测12对引物的效果。12对引物分别能扩增出2至14个等位基因,观察杂合度(Observed heterozygosity, Ho)从0.0732(SC10)到0.7391(SC45),期望杂合度(Expected heterozygosity, He)从0.1177(SC10)到0.8687(SC51),说明开发的引物能揭示较高的遗传多态性水平,可以用于加拿大一枝黄花的入侵遗传机制研究。
2选取8对微卫星引物分析中国15个居群的加拿大一枝黄花的遗传多样性水平和遗传结构。中国15个居群总体平均的多态性百分比(PPL)、平均等位基因数目(Na)、有效等位基因数目(Ne)、Nei's遗传多样性指数(H)、Shannon信息指数(1)、居群内近交系数(Fis)值为91.7%、3.758、2.71、0.529、0.981、-0.191。大多数居群的遗传多样性水平指数都较高而且相差不大,仅湖北宜昌(YC)居群遗传多样性水平偏低。AMOVA分析显示大多数的遗传变异存在于居群内部,总的基因流(Nm)值为1.207。采用UPGMA法对15个居群进行聚类分析显示没有明确的地理分布结构,PCoA分析和聚类结果相符合。IBD检测发现15个居群的地理距离和遗传距离之间没有显著性相关。加拿大一枝黄花的种子和花粉都很容易传播,而且生长于经济发达的大城市和道路两旁,所以基因流很频繁,能够保持较高的遗传多样性水平,而且居群间的差异不大。
3选取3对AFLP引物分析中国15个居群的加拿大一枝黄花的遗传多样性水平和遗传结构。15个居群总体平均的PPL、Na、Ne、H、I值分别为67.87%、1.524、1.46、0.259、0.381。AMOVA分析也显示出大多数的遗传变异存在于居群内部,总的基因流(Nm)值为1.0797。采用UPGMA法对15个居群进行聚类分析显示没有明确的地理分布结构,PCoA分析和聚类结果相符合。IBD检测发现15个居群的地理距离和遗传距离之间没有显著性相关。AFLP标记所检测到的遗传多样性水平虽低于SSR标记检测结果,但仍然显示出较高水平。对比显示两种标记都能揭示较高的遗传多样性差异,但SSR分子标记更加方便快捷,而AFLP标记则能检测整个基因组的变异且更加经济节约成本。
4使用3对AFLP引物分析美国17个居群的加拿大一枝黄花的遗传多样性水平和遗传结构。17个居群总体平均的PPL、Na、Ne、H、I值分别为70.76%、1.572、1.487、0.279、0.400。AMOVA分析显示出大多数的遗传变异存在于居群内部,总的基因流(Nm)值为1.581。采用UPGMA法对17个居群进行聚类分析显示也没有明确的地理分布结构,PCoA分析和聚类结果相符合。IBD检测发现17个居群的地理距离和遗传距离之间没有显著性相关。将美国和中国居群对比分析,发现中国的加拿大一枝黄花遗传多样性水平没有降低,仅基因流水平略低一些。UPGMA法对两个国家的32个居群进行聚类分析显示能够明显聚成中国和美国两类；而苏州(SZ)居群和美国的居群聚类在一起,这可能是因为苏州(SZ)居群位于首次引进加拿大一枝黄花的区域范围内,所以存在和美国较多的相似的遗传基因。中国的加拿大一枝黄花的遗传多样性水平较高,一方面由于其多起源,另一方面由于其特殊的繁育系统。
Biological invasion is the species disperses from the native distribution area to remote new areas by natural or human activities, and causes damage for the local natural ecosystems, biodiversity and human health. Associated with the increase of global economy and trade activities, the biological invasion is become more and more common. Now, biological invasions as well as global climate change and habitat destruction are considered as the world's three most intractable environmental problems. Exploring the genetic diversity of invasive plant is not only helpful for understanding the evolution history of invasive species, but also providing necessary theory for control invasive species. In addition, the possible origin areas for the invasive species could be revealed by analyzing the genetic structure.
Solidago canadensis belongs to the Asteraceae. It is a self-incompatible, but could propagate through outcrossing and asexual reproduction. S. canadensis is native to North America but now have invaded many parts of the world. S. canadensis was introduced to Shanghai and southern Jiangsu (China) in1935as an ornamental plant, but escaping from gardens and spreading throughout the southeastern China in80s of the20th century. It has occupied the farmland and forests and caused huge economic losses. S. canadensis has caused damage to agriculture and environment, resulted in huge great economic losses.
112microsatellite loci were isolated for S. canadensis using dual-suppression polymerase chain reaction (PCR) technique and improved technique for isolating compound SSR markers. The quality of the12pairs of primers was analyzed using35samples of S. canadensis:12pairs of primers can amplify2-14alleles, Observed Heterozygosity (Ho) ranged from0.0732(SC10) to0.7391(SC45), while Expected Heterozygosity (He) ranged from0.1177(SC10) to0.8687(SC51). These primers were able to get high level of polymorphism, could be used for the genetic diversity and genetic structure analysis of S. canadensis.
2The genetic diversity and genetic structure of15Chinese populations was analyzed by8microsatellite primers. The average percentage of polymorphism (PPL) for the15populations is91.7%, the average number of alleles (Na) is3.758, effective number of alleles (Ne) is2.71, Nei's genetic diversity index (H) is0.529, Shannon information index (Ⅰ) is0.981, fixation index (Fis) within populations is-0.191. The genetic diversity for most populations is high except the Yichang (YC) populations, which has a relative low level of genetic diversity. AMOVA analysis showed that most genetic variation occurred within populations and the total gene flow (Nm) is1.207. UPGMA and PCoA analyses resulted similar results that no clear geographical distribution structure was shown. IBD analyses suggested the geographical distance and genetic distance of the15populations was not significantly correlated. S.canadensis usually grow in large cities and along the both sides of road, the gene flow is very frequently. Therefore, it has a high level of genetic diversity, and the interpopulation variation is slight.
3The genetic diversity and genetic structure of15Chinese populations were analyzed by3AFLP primers. The overall average of15populations of PPL、Na、Ne、H、 I were shown to be67.87%、1.524、1.46、0.259、0.381respectively. Most of genetic variation were shown to be existed within populations by AMOVA analysis, and the total gene flow (Nm) was shown to be1.0797. UPGMA and PCoA analyses resulted similar results that no clear geographical distribution structure was shown. Different from the results recovered from SSR primers, IBD analyses suggested the geographical distance and genetic distance of the15populations was significantly negative. Although the genetic diversity recovered by AFLP primers is relatively lower than that recovered by SSR, the genetic diversity is also high. SSR marker is faster and convenient, while AFLP markers more economical and could be able to detect the variation throughout the whole genome.
4The genetic diversity and genetic structure of17USA populations were analyzed by3AFLP primers. The overall average of17populations of PPL、Na、Ne、H、Iwere shown to be70.76%、1.572、1.487、0.279、0.400respectively. Most of genetic variation were shown to be existed within populations by AMOVA analysis, and the total gene flow (Nm) was shown to be1.581. UPGMA and PCoA analyses resulted similar results that no clear geographical distribution structure was shown. IBD analyses suggested the geographical distance and genetic distance of the17populations was significantly negative. The genetic diversity of USA populations is similar to that of Chinese populations. The populations from the two countries were divided into two groups in the UPGMA analyses except the SZ (China) population, which was clustered together with USA populations. The position of SZ population is possibly because SZ population lies in the first introduction areas of S. canadensis, so it is similar to USA population. The study suggested the S. canadensis from Chinese has a high genetic diversity, it is possibly related to the multiple origins, also related to the distinctively reproduction system of this species.
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