日本伏翼（Pipistrellus abramus）种群遗传学研究
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摘要
相对于欧洲和北美洲,第四纪冰川时期的气候变化对亚洲生物群落的分布和遗传结构形成的影响的研究报导至今还很少。在冰川后期的种群扩张期间,地理屏障对某个物种的扩散能力的促进或限制的作用同样对其种群遗传结构有重要的影响。日本伏翼(Pipistrellus abramus)在中国大陆、海南岛和舟山群岛等很多地理区域都有分布。近些年来,由于生态环境的严重破坏以及城乡建设的快速发展,对于主要栖息于居民老房里的翼手目动物类群,如日本伏翼,在某些地区其种群数量急剧下降。遗传多样性是种群长期生存繁衍和抵御疾病等外界胁迫的基础,种群数量的演变历史对种群的保护有很重要的指导作用。我们对日本伏翼的遗传多样性以及种群间的基因流动等方面进行研究,可以使我们能更全面了解日本伏翼的生存现状及其种群历史演替情况,并制定正确的、合理的动物保护措施。
本文运用线粒体DNA和微卫星DNA两种不同分子遗传标记对日本伏翼分布于国内的17个地理种群的种群遗传结构及其种群历史情况进行研究。其结果如下：
1、以富集法首次构建了日本伏翼微卫星DNA文库,共筛选出微卫星引物10对。以这10对引物分别扩增广西桂林的日本伏翼种群37份个体DNA样品,结果显示,每对引物扩增条带均呈现出多态性,所有微卫星位点的平均等位基因数达到10个,范围在7-13个不等。杂合度观测值和期望值范围分别为0.486-0.971和0.752-0.876。分析发现,3个微卫星位点(WW6、PA133和1-20)偏离了哈迪-温伯格平衡,但没发现任何2个位点存在连锁不平衡。成功筛选出这些微卫星位点对促进日本伏翼及本属其他物种的种群遗传结构深入研究有重要的作用。
2、线粒体DNA数据和微卫星DNA(8对位点引物)数据共同表明,日本伏翼的不同地理种群均具有较高的遗传多样性。种群遗传多样性主要指标的变化范围较大,线粒体DNA单倍型多样性(h)为0.286-0.9,核苷酸多样性(П)为0.00025-0.01878；微卫星DNA的种群杂合度观测值(Ho)为0.58-0.89,杂合度期望值(HE)为0.67-0.82,种群的平均等位基因丰富度为4.13-10.36。
3、以两种分子遗传标记对17个种群的研究结果共同说明,自然地理环境对日本伏翼种群结构的连续性产生明显的影响。更重要的是,这两种标记的结果都揭示,横亘在海南岛和中国大陆间的琼州海峡对日本伏翼的基因流产生显著的阻隔作用。同时,在两种遗传标记的结果中,所有种群的遗传距离与地理距离均呈显著相关(线粒体DNA, R2=0.27, P<0.001;微卫星DNA, R2=0.31, P<0.001)。
4、线粒体DNA序列单倍型的错配分布分析(Mismatch distribution)说明,自第四纪冰期以来,大陆/舟山群岛日本伏翼种群和海南岛的日本伏翼种群都经历了种群扩张事件,且前者在TCS的基因网络分析中呈现星状拓扑结构。线粒体DNA分析结果揭示,在相互隔离7000-9000年后,大陆与舟山群岛日本伏翼种群间仍能检测到共同祖先单倍型,不过,海南岛与大陆相互隔离的时间虽然也在10000年左右,但海南岛日本伏翼种群内没能检测到共享单倍型,说明没有发现共同的祖先或通过近期种群扩张而来。
5、微卫星DNA数据分析结果并没有完全支持线粒体DNA数据的分析结果,反之,发现在大陆/舟山群岛日本伏翼种群和海南岛日本伏翼种群间存在一些差异(见结果部分),进一步说明了只有综合运用不同分子标记对种群遗传空间尺度进行分析,才能更为准确和有效地评价物种种群遗传结构及其种群历史演化情况。
本文运用了线粒体DNA和微卫星DNA两种不同的分子标记比较分析了日本伏翼不同种群的遗传特征与地理分布的关系,为进一步研究该物种的谱系地理学特征奠定了基础。
Compared to Europe and North America, the influence of climatic oscillations during the Pleistocene in shaping the distribution and genetic structure of biota in Asia is not well known. The influence of geographical barriers in promoting or restricting the dispersal ability of a species during post-glacial expansions will also have a large impact on its population genetic structure. The Japanese pipistrelle bat, Pipistrellus abramus, is found over a wide range in China, including Hainan Island and the Zhoushan Archipelago. Recently, in some areas, the number individuals of some bat species have dramatically been in decline due to seriously destructive ecological environment and fast development of towns. Genetic diversity is the basis of population long-term survival and resisting the diseases. Demographic history analysis is vital to population protection. In order to comprehensively understand survival status and demographic history and establish accurate protection steps for Pipistrellus abramus, it plays an important role in estimation of genetic diversity and gene flow between populations.
We applied both mitochondrial (cytochrome b) and microsatellite markers to examine population genetic structure and demographic history in this species. The main results are as follows:
1. We describe the first set of ten microsatellite markers isolated in Pipistrellus abramus. The number of alleles per locus ranged from 7 to 13. The observed and expected heterozygosities values ranged from 0.486 to 0.971 and from 0.752 to 0.876, respectively. Three loci (WW6、PA133 andl-20) revealed significant departure from Hardy-Weinberg equilibrium and no linkage disequilibrium was found between loci pairs. These informative microsatellite markers will be a powerful molecular tool for studying the population genetic structure of P. abramus, as well as other species of this genus.
2. Both results of mtDNA and microsatellite DNA (8 loci) revealed Pipistrellus abramus contain relatively high genetic diversities among different geographic populations. For the mitochondrial DNA (mtDNA) data, the haplotype diversity ranged from 0.286 to 0.9, and nucleotide diversity ranged from 0.00025 to 0.01878. For the microsatellite data, the observed heterozygosity (HO) and expected (HE) heterozygosity ranged from 0.58 to 0.89 and from 0.67 to 0.82, respectively. The allelic richness (RS) ranged from 4.13 to 10.36 among sampled populations.
3. Our results suggest that geographical features have had a clear impact on population connectivity in P. abramus. Specifically, both markers indicate that the Qiongzhou Strait, which separates Hainan Island from the China mainland, acts as a significant barrier to gene flow. Likewise, both genetic distance of mtDNA and microsatellite DNA were significant with natural geography distance (mtDNA, R2=0.27, P<0.001; microsatellite DNA, R2=0.31, P<0.001).
4. Mismatch distributions show that populations from China mainland/Zhoushan Archipelago and Hainan Island appear to have undergone expansions since the Pleistocene, with evidence of a star-like topology in the former. Mt DNA reveals evidence of common ancestry between the colonies sampled on the Zhoushan Archipelago and those on the mainland, which are thought to have been isolated approximately 7000-9000 years ago. Although the Qiongzhou Strait seperates the Hainan Island and mainland for over 10000 years, no haplotype was found to share among individuals sampled from the island and mainland, and it revealed that the bats sampled from the Hainan Island may colony into the island via recent population expansion since the Pleistocene.
5. Microsatellite data did not fully support the mitochondrial data and instead detected some differentiation between the two groups, highlighting the importance of using different markers at different spatial scales.
In summary, we were explicit that the thesis data were combined with mitochondrial (cytochrome b) and microsatellite molecular markers for comparison of different geographical populations and geography distribution. Further more, the results were the basic study of phylogeography of the Japanese pipistrelle bat, Pipistrellus abramus.
本文运用线粒体DNA和微卫星DNA两种不同分子遗传标记对日本伏翼分布于国内的17个地理种群的种群遗传结构及其种群历史情况进行研究。其结果如下：
1、以富集法首次构建了日本伏翼微卫星DNA文库,共筛选出微卫星引物10对。以这10对引物分别扩增广西桂林的日本伏翼种群37份个体DNA样品,结果显示,每对引物扩增条带均呈现出多态性,所有微卫星位点的平均等位基因数达到10个,范围在7-13个不等。杂合度观测值和期望值范围分别为0.486-0.971和0.752-0.876。分析发现,3个微卫星位点(WW6、PA133和1-20)偏离了哈迪-温伯格平衡,但没发现任何2个位点存在连锁不平衡。成功筛选出这些微卫星位点对促进日本伏翼及本属其他物种的种群遗传结构深入研究有重要的作用。
2、线粒体DNA数据和微卫星DNA(8对位点引物)数据共同表明,日本伏翼的不同地理种群均具有较高的遗传多样性。种群遗传多样性主要指标的变化范围较大,线粒体DNA单倍型多样性(h)为0.286-0.9,核苷酸多样性(П)为0.00025-0.01878；微卫星DNA的种群杂合度观测值(Ho)为0.58-0.89,杂合度期望值(HE)为0.67-0.82,种群的平均等位基因丰富度为4.13-10.36。
3、以两种分子遗传标记对17个种群的研究结果共同说明,自然地理环境对日本伏翼种群结构的连续性产生明显的影响。更重要的是,这两种标记的结果都揭示,横亘在海南岛和中国大陆间的琼州海峡对日本伏翼的基因流产生显著的阻隔作用。同时,在两种遗传标记的结果中,所有种群的遗传距离与地理距离均呈显著相关(线粒体DNA, R2=0.27, P<0.001;微卫星DNA, R2=0.31, P<0.001)。
4、线粒体DNA序列单倍型的错配分布分析(Mismatch distribution)说明,自第四纪冰期以来,大陆/舟山群岛日本伏翼种群和海南岛的日本伏翼种群都经历了种群扩张事件,且前者在TCS的基因网络分析中呈现星状拓扑结构。线粒体DNA分析结果揭示,在相互隔离7000-9000年后,大陆与舟山群岛日本伏翼种群间仍能检测到共同祖先单倍型,不过,海南岛与大陆相互隔离的时间虽然也在10000年左右,但海南岛日本伏翼种群内没能检测到共享单倍型,说明没有发现共同的祖先或通过近期种群扩张而来。
5、微卫星DNA数据分析结果并没有完全支持线粒体DNA数据的分析结果,反之,发现在大陆/舟山群岛日本伏翼种群和海南岛日本伏翼种群间存在一些差异(见结果部分),进一步说明了只有综合运用不同分子标记对种群遗传空间尺度进行分析,才能更为准确和有效地评价物种种群遗传结构及其种群历史演化情况。
本文运用了线粒体DNA和微卫星DNA两种不同的分子标记比较分析了日本伏翼不同种群的遗传特征与地理分布的关系,为进一步研究该物种的谱系地理学特征奠定了基础。
Compared to Europe and North America, the influence of climatic oscillations during the Pleistocene in shaping the distribution and genetic structure of biota in Asia is not well known. The influence of geographical barriers in promoting or restricting the dispersal ability of a species during post-glacial expansions will also have a large impact on its population genetic structure. The Japanese pipistrelle bat, Pipistrellus abramus, is found over a wide range in China, including Hainan Island and the Zhoushan Archipelago. Recently, in some areas, the number individuals of some bat species have dramatically been in decline due to seriously destructive ecological environment and fast development of towns. Genetic diversity is the basis of population long-term survival and resisting the diseases. Demographic history analysis is vital to population protection. In order to comprehensively understand survival status and demographic history and establish accurate protection steps for Pipistrellus abramus, it plays an important role in estimation of genetic diversity and gene flow between populations.
We applied both mitochondrial (cytochrome b) and microsatellite markers to examine population genetic structure and demographic history in this species. The main results are as follows:
1. We describe the first set of ten microsatellite markers isolated in Pipistrellus abramus. The number of alleles per locus ranged from 7 to 13. The observed and expected heterozygosities values ranged from 0.486 to 0.971 and from 0.752 to 0.876, respectively. Three loci (WW6、PA133 andl-20) revealed significant departure from Hardy-Weinberg equilibrium and no linkage disequilibrium was found between loci pairs. These informative microsatellite markers will be a powerful molecular tool for studying the population genetic structure of P. abramus, as well as other species of this genus.
2. Both results of mtDNA and microsatellite DNA (8 loci) revealed Pipistrellus abramus contain relatively high genetic diversities among different geographic populations. For the mitochondrial DNA (mtDNA) data, the haplotype diversity ranged from 0.286 to 0.9, and nucleotide diversity ranged from 0.00025 to 0.01878. For the microsatellite data, the observed heterozygosity (HO) and expected (HE) heterozygosity ranged from 0.58 to 0.89 and from 0.67 to 0.82, respectively. The allelic richness (RS) ranged from 4.13 to 10.36 among sampled populations.
3. Our results suggest that geographical features have had a clear impact on population connectivity in P. abramus. Specifically, both markers indicate that the Qiongzhou Strait, which separates Hainan Island from the China mainland, acts as a significant barrier to gene flow. Likewise, both genetic distance of mtDNA and microsatellite DNA were significant with natural geography distance (mtDNA, R2=0.27, P<0.001; microsatellite DNA, R2=0.31, P<0.001).
4. Mismatch distributions show that populations from China mainland/Zhoushan Archipelago and Hainan Island appear to have undergone expansions since the Pleistocene, with evidence of a star-like topology in the former. Mt DNA reveals evidence of common ancestry between the colonies sampled on the Zhoushan Archipelago and those on the mainland, which are thought to have been isolated approximately 7000-9000 years ago. Although the Qiongzhou Strait seperates the Hainan Island and mainland for over 10000 years, no haplotype was found to share among individuals sampled from the island and mainland, and it revealed that the bats sampled from the Hainan Island may colony into the island via recent population expansion since the Pleistocene.
5. Microsatellite data did not fully support the mitochondrial data and instead detected some differentiation between the two groups, highlighting the importance of using different markers at different spatial scales.
In summary, we were explicit that the thesis data were combined with mitochondrial (cytochrome b) and microsatellite molecular markers for comparison of different geographical populations and geography distribution. Further more, the results were the basic study of phylogeography of the Japanese pipistrelle bat, Pipistrellus abramus.
引文
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