沙棘木蠹蛾种群遗传结构与灾害发生机制的分子生态学研究
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摘要
沙棘木蠹蛾(Holcocerus hippophaecolus Hua, Chou, Fang et Chen)属鳞翅目(Lepidoptera)木蠹蛾科(Cossidae)线角木蠹蛾属(Holcocerus)。它主要危害沙棘(Hippophae rhamnoides L.),是我国近15年来新出现的暴发性害虫。木研究利用分子生态学的研究方法分析了不同地区、不同寄主种群沙棘木蠹蛾的遗传结构,以及影响因子。结合系统发育的分析方法,探讨沙棘木蠹蛾与其近缘种的遗传关系,揭示了寄主适应性在沙棘木蠹蛾进化历史中的重要作用。主要研究结果如下：
1.建立了沙棘木蠹蛾AFLP多态性荧光标记体系。选取三对引物组合,采用AFLP荧光标记和银染法对相同的沙棘木蠹蛾全基因组DNA样品的多态性进行了分析。结果显示荧光标记法在精确度和检测效果方面都较为理想。因此,木研究建立了一套以银染法AFLP技术筛选引物,荧光标记法进行种群多样性分析的沙棘木蠹蛾AFLP研究体系。从100对AFLP引物组合中筛选出9对条带清晰、多态性较高的引物组合。
2.利用AFLP分子标记技术揭示了我国沙棘木蠹蛾不同地区种群间分化明显的遗传特性,明确了沙棘木蠹蛾种群遗传结构受到多种因素的综合影响。较高的遗传分化可能与不同地区长时间的地理隔离、不同的环境条件以及沙棘木蠹蛾扩散能力较弱等影响有关。进一步分析沙棘木蠹蛾种群遗传结构和多样性参数与地理距离、干旱因子以及暴发因子的相关性。结果表明,种群遗传结构并不符合地理隔离的模式,而暴发因子和干旱因子对其遗传结构的影响也不显著。沙棘木蠹蛾种群平均杂合度与干旱、历史暴发事件均无显著相关。说明沙棘木蠹蛾的种群遗传结构是受到了多种因素综合影响的结果。
3.明确了在种群遗传变异水平不存在与暴发相关的显著分化,但少数位点可能与沙棘木蠹蛾的暴发相关。根据沙棘木蠹蛾的历史发生情况,将十个不同地区种群划分为“曾暴发地区种群”和“未曾暴发地区种群”两组。遗传距离的聚类和分子方差分析结果表明,两组间混合聚类,组间遗传变异百分比很小且不显著。但是,我们在组间检测到了少数(4个)可能与暴发相关的受选择位点,说明沙棘木蠹蛾的基因组中可能存在某些特异位点与种群数量的波动相关。
4.明确了沙棘木蠹蛾的暴发源于本地种群的数量增长。AFLP分子标记分析十个不同地区人工林的沙棘木蠹蛾种群遗传关系的结果表明,该害虫的灾害发生并非源于同一致害能力强的种群的扩散。基于线粒体DNA COⅠ序列的遗传变异分析结果证实,代表性地区沙棘人工林中危害的沙棘木蠹蛾样本的基因型与当地天然林中样本较为一致,证实了沙棘木蠹蛾人工林中的种群来源于本地。
5.利用AFLP分子标记分析建平地区不同寄主种群沙棘木蠹蛾的遗传结构,推测其在沙棘上大面积暴发的内因基础,明确了沙棘木蠹蛾对沙棘的适应性分化。通过检测不同寄主种群的遗传多样性水平和种群间的遗传分化,揭示出沙棘上的沙棘木蠹蛾种群比其它寄主上种群更高的遗传多样性水平,较丰富的遗传变异对应着更强的适应能力,有利于该害虫的生存与繁殖。另外,寄主选择对沙棘木蠹蛾种群的分化作用大于地理隔离,沙棘木蠹蛾的“寄主族”现象很可能存在于沙棘与其他寄主之间,而这种对寄主的适应性分化很可能早在沙棘引进到建平地区之前就已形成。
6.初步明确了沙棘木蠹蛾与部分近缘种的遗传关系。利用线粒体DNA CO I基因的部分序列对沙棘木蠹蛾同属的4个近缘种以及3个外群种进行遗传关系的研究。系统发育树的结果显示沙棘木蠹蛾与榆木蠹蛾以中度的置信度聚为一支。AMOVA分析的结果表明也这两个物种在COⅠ基因序列上没有显著的分化。结合AFLP标记对沙棘木蠹蛾与榆木蠹蛾混合集群基因组DNA的扫描,在242个AFLP位点中发现了5个可能与寄主相关的选择性位点,这一结果为沙棘木蠹蛾进化过程中寄主适应性的重要作用提供了线索。
木研究揭示了沙棘木蠹蛾暴发成灾的虫源、内在遗传因素以及寄主选择性的分子机理,同时为它区域成灾调控和制定行之有效的防治策略等方面提供理论基础。
The seabuckthorn carpenter moth (Holcocerus hippophaecolus Hua. Chou, Fang et Chen) is a native species throughout the north of China and is considered the main threat to seabuckthorn, Hippophae rhamnoides L. in last 15 years. The influence of outbreaks, environmental factors and host species in shaping the genetic variation and structure of H. hippophaecolus were assessed by using Amplified Fragment Length Polymorphism (AFLP). The phylogenetic relationships among H. hippophaecolus and its closely related species were analyzed, the results uncovered important role of host selection in the evolutionary history of H. hippophaecolus.
1. The fluorescence labeling technique of AFLP was established. Three primers combinations were used for amplification of genomic DNA by both fluorescence labeling technique and silver staining technique. The results showed that, compared with silver staining technique, using fluorescence labeling work efficiency and could get much more bands. Nine selective primer combinations which revealed sufficient amplified bands with high resolution and polymorphism were selected from 100 pairs of primer and clear fingerprints were obtained. The fluorescence labeling technique AFLP analyze system can be used to study population genetic diversity and structure.
2. The population genetic variation and structure of H. hippophaecolus from 10 different areas across its range with contrasting historical patterns of outbreak events were described by using AFLP markers. The evidence of limited gene flow among samples collected from 10 locations was detected. Independent population structure of H. hippophaecolus can be explained by geographical isolation, environmental heterogeneity and low dispersal capacity. H. hippophaecolus did not follow an isolation-by-distance pattern. The results also rejected the hypothesis that outbreak and drought events were driving the genetic structure of H. hippophaecolus. Rather, the genetic structure appears to be influenced by various confounding bio-geographical factors.
3. The hypothesis that outbreak-associated genetic divergence exist among populations was rejected, as evidenced by genetic clusters containing a combination of populations from historical outbreak areas, as well as non-outbreak areas. AMOVA results confirmed the presence of mild and nonsignificant genetic differentiation total variability was due to the variation between outbreak and non-outbreak groups. A small number of markers (4 of 933 loci) were identified as candidates under selection in response to population densities.
4. Our results support the notion that outbreak events were likely to be endemic population changes from latent to epidemic rather than being due to insects with an outbreak-associated genotype spreading to outbreak areas. H. hippophaecolus samples were collected from both plantation and natural forestry in two different locations. The genetic relationship among samples based on mtDNA CO I sequences indicated that H. hippophaecolus in plantation are indigenous to those two locations respectively.
5. Genetic diversity and differentiation of H. hippophaecolus sympatric populations from four different host trees were detected in Jianping. Higher Nei's heterozygosity was found in the population feeding on seabuckthorn than other hosts populations. There were detectable genetic differences between H. hippophaecolus populations occupying different host trees. Host races might exist in H. hippophaecolus used seabuckthorn and other host plant. In Jianping, genetic divergence pre-dating the introduction of seabuckthorn, the possibilities of an ancestral host shift in seabuckthorn carpenter moth are suggested.
6. Based on MP and Bayes tree reconstructed from the mtDNA CO I sequences. The H. hippophaecolus and H. vicarious were interspersed throughout the tree indicated they are the most closely related. Variation within and between species from population genetic perspectives were detected. Divergence between H. hippophaecolus and H. Vicariou has been relatively recent. There is little phylogenetic resolution and no significant genetic variance between species. Genome scan approach was further used to detect positive loci under divergent selection among populations of H. hippophaecolus / H. vicariou complex feeding on different host plant. Based on 242 Amplified Fragment Length Polymorphism (AFLP) markers, four loci (1.6%) were identified as candidates for adaptation to host plants. The evidence for positive selection suggests an important role of host selection in the evolutionary history of H. hippophaecolus.
The results provide scientific evidence in revealing the molecular mechanism of outbreak of H. hippophaecolus disaster. Furthermore, they are also useful in establishing control measures to H. hippophaecolus.
1.建立了沙棘木蠹蛾AFLP多态性荧光标记体系。选取三对引物组合,采用AFLP荧光标记和银染法对相同的沙棘木蠹蛾全基因组DNA样品的多态性进行了分析。结果显示荧光标记法在精确度和检测效果方面都较为理想。因此,木研究建立了一套以银染法AFLP技术筛选引物,荧光标记法进行种群多样性分析的沙棘木蠹蛾AFLP研究体系。从100对AFLP引物组合中筛选出9对条带清晰、多态性较高的引物组合。
2.利用AFLP分子标记技术揭示了我国沙棘木蠹蛾不同地区种群间分化明显的遗传特性,明确了沙棘木蠹蛾种群遗传结构受到多种因素的综合影响。较高的遗传分化可能与不同地区长时间的地理隔离、不同的环境条件以及沙棘木蠹蛾扩散能力较弱等影响有关。进一步分析沙棘木蠹蛾种群遗传结构和多样性参数与地理距离、干旱因子以及暴发因子的相关性。结果表明,种群遗传结构并不符合地理隔离的模式,而暴发因子和干旱因子对其遗传结构的影响也不显著。沙棘木蠹蛾种群平均杂合度与干旱、历史暴发事件均无显著相关。说明沙棘木蠹蛾的种群遗传结构是受到了多种因素综合影响的结果。
3.明确了在种群遗传变异水平不存在与暴发相关的显著分化,但少数位点可能与沙棘木蠹蛾的暴发相关。根据沙棘木蠹蛾的历史发生情况,将十个不同地区种群划分为“曾暴发地区种群”和“未曾暴发地区种群”两组。遗传距离的聚类和分子方差分析结果表明,两组间混合聚类,组间遗传变异百分比很小且不显著。但是,我们在组间检测到了少数(4个)可能与暴发相关的受选择位点,说明沙棘木蠹蛾的基因组中可能存在某些特异位点与种群数量的波动相关。
4.明确了沙棘木蠹蛾的暴发源于本地种群的数量增长。AFLP分子标记分析十个不同地区人工林的沙棘木蠹蛾种群遗传关系的结果表明,该害虫的灾害发生并非源于同一致害能力强的种群的扩散。基于线粒体DNA COⅠ序列的遗传变异分析结果证实,代表性地区沙棘人工林中危害的沙棘木蠹蛾样本的基因型与当地天然林中样本较为一致,证实了沙棘木蠹蛾人工林中的种群来源于本地。
5.利用AFLP分子标记分析建平地区不同寄主种群沙棘木蠹蛾的遗传结构,推测其在沙棘上大面积暴发的内因基础,明确了沙棘木蠹蛾对沙棘的适应性分化。通过检测不同寄主种群的遗传多样性水平和种群间的遗传分化,揭示出沙棘上的沙棘木蠹蛾种群比其它寄主上种群更高的遗传多样性水平,较丰富的遗传变异对应着更强的适应能力,有利于该害虫的生存与繁殖。另外,寄主选择对沙棘木蠹蛾种群的分化作用大于地理隔离,沙棘木蠹蛾的“寄主族”现象很可能存在于沙棘与其他寄主之间,而这种对寄主的适应性分化很可能早在沙棘引进到建平地区之前就已形成。
6.初步明确了沙棘木蠹蛾与部分近缘种的遗传关系。利用线粒体DNA CO I基因的部分序列对沙棘木蠹蛾同属的4个近缘种以及3个外群种进行遗传关系的研究。系统发育树的结果显示沙棘木蠹蛾与榆木蠹蛾以中度的置信度聚为一支。AMOVA分析的结果表明也这两个物种在COⅠ基因序列上没有显著的分化。结合AFLP标记对沙棘木蠹蛾与榆木蠹蛾混合集群基因组DNA的扫描,在242个AFLP位点中发现了5个可能与寄主相关的选择性位点,这一结果为沙棘木蠹蛾进化过程中寄主适应性的重要作用提供了线索。
木研究揭示了沙棘木蠹蛾暴发成灾的虫源、内在遗传因素以及寄主选择性的分子机理,同时为它区域成灾调控和制定行之有效的防治策略等方面提供理论基础。
The seabuckthorn carpenter moth (Holcocerus hippophaecolus Hua. Chou, Fang et Chen) is a native species throughout the north of China and is considered the main threat to seabuckthorn, Hippophae rhamnoides L. in last 15 years. The influence of outbreaks, environmental factors and host species in shaping the genetic variation and structure of H. hippophaecolus were assessed by using Amplified Fragment Length Polymorphism (AFLP). The phylogenetic relationships among H. hippophaecolus and its closely related species were analyzed, the results uncovered important role of host selection in the evolutionary history of H. hippophaecolus.
1. The fluorescence labeling technique of AFLP was established. Three primers combinations were used for amplification of genomic DNA by both fluorescence labeling technique and silver staining technique. The results showed that, compared with silver staining technique, using fluorescence labeling work efficiency and could get much more bands. Nine selective primer combinations which revealed sufficient amplified bands with high resolution and polymorphism were selected from 100 pairs of primer and clear fingerprints were obtained. The fluorescence labeling technique AFLP analyze system can be used to study population genetic diversity and structure.
2. The population genetic variation and structure of H. hippophaecolus from 10 different areas across its range with contrasting historical patterns of outbreak events were described by using AFLP markers. The evidence of limited gene flow among samples collected from 10 locations was detected. Independent population structure of H. hippophaecolus can be explained by geographical isolation, environmental heterogeneity and low dispersal capacity. H. hippophaecolus did not follow an isolation-by-distance pattern. The results also rejected the hypothesis that outbreak and drought events were driving the genetic structure of H. hippophaecolus. Rather, the genetic structure appears to be influenced by various confounding bio-geographical factors.
3. The hypothesis that outbreak-associated genetic divergence exist among populations was rejected, as evidenced by genetic clusters containing a combination of populations from historical outbreak areas, as well as non-outbreak areas. AMOVA results confirmed the presence of mild and nonsignificant genetic differentiation total variability was due to the variation between outbreak and non-outbreak groups. A small number of markers (4 of 933 loci) were identified as candidates under selection in response to population densities.
4. Our results support the notion that outbreak events were likely to be endemic population changes from latent to epidemic rather than being due to insects with an outbreak-associated genotype spreading to outbreak areas. H. hippophaecolus samples were collected from both plantation and natural forestry in two different locations. The genetic relationship among samples based on mtDNA CO I sequences indicated that H. hippophaecolus in plantation are indigenous to those two locations respectively.
5. Genetic diversity and differentiation of H. hippophaecolus sympatric populations from four different host trees were detected in Jianping. Higher Nei's heterozygosity was found in the population feeding on seabuckthorn than other hosts populations. There were detectable genetic differences between H. hippophaecolus populations occupying different host trees. Host races might exist in H. hippophaecolus used seabuckthorn and other host plant. In Jianping, genetic divergence pre-dating the introduction of seabuckthorn, the possibilities of an ancestral host shift in seabuckthorn carpenter moth are suggested.
6. Based on MP and Bayes tree reconstructed from the mtDNA CO I sequences. The H. hippophaecolus and H. vicarious were interspersed throughout the tree indicated they are the most closely related. Variation within and between species from population genetic perspectives were detected. Divergence between H. hippophaecolus and H. Vicariou has been relatively recent. There is little phylogenetic resolution and no significant genetic variance between species. Genome scan approach was further used to detect positive loci under divergent selection among populations of H. hippophaecolus / H. vicariou complex feeding on different host plant. Based on 242 Amplified Fragment Length Polymorphism (AFLP) markers, four loci (1.6%) were identified as candidates for adaptation to host plants. The evidence for positive selection suggests an important role of host selection in the evolutionary history of H. hippophaecolus.
The results provide scientific evidence in revealing the molecular mechanism of outbreak of H. hippophaecolus disaster. Furthermore, they are also useful in establishing control measures to H. hippophaecolus.
引文
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