我国子午沙鼠分子系统地理学研究
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摘要
子午沙鼠(Meriones meridianus Pallas,1773)(英文名：midday gerbil)隶属于啮齿目(Rodentia),仓鼠科(Cricetidae),沙鼠属(Meriones)。主要分布于我国西北部地区沙漠地带边缘,垂直分布可从海拔-150m的吐鲁番盆地,上升到昆仑山脉北坡海拔3200m的高山荒漠地区,沙漠地区的建群种和优势种,通常是环境沙漠化的指示物种。对环境的敏感性,有限的扩散能力,时代周期短以及突变速率快等特征使子午沙鼠成为系统地理学理想的研究对象。子午沙鼠国内的分布区域地形地貌复杂,包括山体,盆地,湖盆,森林草原等,这些都是潜在的地理隔离屏障,且天山等地区在第四纪时期深受冰期-间冰期的气候波动和环境变化的影响,可能会作用于子午沙鼠的系统地理结构,留下遗传印记。因此,本文基于线粒体基因(Cyt-b基因和D-loop基因)和核基因(IRBP基因和AVPR2基因)研究子午沙鼠的系统地理结构。
本文的研究目的是：(1)国内子午沙鼠分布区内潜在的地理隔离屏障对其系统地理结构的影响；(2)第四纪气候波动和环境变化带来的生境变化对子午沙鼠的遗传谱系空间分布的历史格局以及种群动态历史的影响：(3)通过分子遗传的手段来重新审视子午沙鼠亚种的分类地位；(4)比较探讨不同遗传标记位点的遗传变异以及系统发生分析；(5)在子午沙鼠的分布范围内当属青藏高原地形地貌最为复杂,海拔较高,环境恶劣,因此将与子午沙鼠同域分布于柴达木盆地地区的三趾跳鼠和五趾跳鼠进行系统地理学和种群历史动态方面的比较分析,以分子遗传学的角度来佐证青藏高原动物区系形成与演化过程。
本文的研究结果表明：
(1)基于子午沙鼠线粒体基因(Cyt-b和D-loop)、核基因(IRBP基因)以及串联合并基因构建的系统发生树显示分布于中国西北部的子午沙鼠显著分化为两个支系,大多数天山地区的样本聚为一个支系,来自于中国西北部的其它沙漠地区以及少量的天山地区的样本聚为另一个支系。两个支系的分歧时间发生在天山抬升之后的中更新世中期,说明天山抬升有效地阻止子午沙鼠基因交流而产生了明显的地理隔离作用,此外,间冰期的暖湿气候带来的吐鲁番盆地森林草原扩张,也对其具有效的隔离作用导致遗传分化。
(2)由于分布于天山地区的种群长期处于环境稳定的状态,因此近期没有发生种群扩张。而另一个支系受到末次盛冰期气候的影响,沙漠环境快速急剧扩张和湖泊萎缩消失,从而分布于中国西北部其它沙漠地区的种群经历了近期种群扩张的历史。
(3)子午沙鼠两个支系间的遗传距离小于沙鼠属姐妹种种间的遗传距离,表明在分子遗传角度上,子午沙鼠两个支系的分类地位为亚种水平。
(4)基于新疆地区的134个子午沙鼠样本的IRBP序列和AVPR2序列分别构建的系统发生树。IRBP系统发生树显示83个单倍型聚为两个支系,两个支系之间具有显著的遗传分化,呈现互为单系群关系。而AVPR2系统发生树显示13个单倍型之间没有遗传分化,呈现多系群关系。对于夜行性动物子午沙鼠来说,具有暗适应调节能力的IRBP基因功能上对生命生存制约性低,如果IRBP基因发生有害突变导致暗适应功能不完全或丧失,可以通过听觉、嗅觉等的代偿作用来弥补,因此对子午沙鼠的生存繁殖几乎没有影响,因此根据遗传结果可以推断IRBP基因很可能接近于中性选择,有效的积累了遗传漂变等产生的突变。而具有缩尿功能的AVPR2基因所编码的蛋白对于生存于缺水干旱环境的沙漠地区的子午沙鼠是致命的,如果AVPR2基因发生有害突变导致缩尿功能的丧失就会威胁到子午沙鼠的生存,因此AVPR2基因很有可能受到强烈的自然选择作用,不能够有效的积累基因突变,从而系统发生树呈现没有遗传分化的多系群关系。
(5)五趾跳鼠柴达木地理单元与甘肃内蒙地理单元具有显著的遗传分化,而子午沙鼠和三趾跳鼠这两个地理单元之间几乎没有遗传分化。五趾跳鼠生境偏好选择海拔较高的山地草原带,因此在中更新世时期青藏高原抬升带来的气候干旱化导致的干草原发育扩张,为五趾跳鼠由甘肃内蒙等地迁移扩散到柴达木盆地提供了有利条件,由于高海拔地区的地形地貌复杂,容易受到生境片段化而导致遗传分化,本文的结果还显示随着种群由西向东分布共祖时间逐渐递减,可以看出其在柴达木盆地中由西向东迁移扩散。而子午沙鼠和三趾跳鼠生境偏好选择沙漠且分布海拔较低,随着晚更新世时期气候更加干旱化导致柴达木盆地沙漠迅速形成发育,使子午沙鼠和三趾跳鼠辐射性迅速迁移扩散到柴达木盆地,没有足够的时间产生遗传分化。
The range of Meriones meridianus (Rodentia:Cricetidae:Meriones) spans several major arid and semi-arid biogeographic regions in northern China (Luo, Chen et al.2000). In northern China, this desert-dwelling rodent species is distributed along the edge of deserts, and it is an important component of the desert systems that it inhabits. It could serve as an indicator of desertification. Northern China is endowed with a complicated terrain of deserts, high mountains, rivers, lakes, and basins which vary widely in elevation from the Qaidam Basin, which is over2700m above sea level to the Turpan Basis, which is103m below sea level. These complex and varied terrains, in conjunction with climate fluctuations, have facilitated the diversification and vicariance of M. meridianus. The species provide good models for reconstructions of how environmental parameters influenced the history of biota. This is because its short generation time, rapid mtDNA substitution rate, relatively limited dispersal ability, and strong associations with particular habitats give rise to informative contemporary patterns of genetic variation. In particular, the midday gerbil M. meridianus may display especially informative phylogeographical patterns due to its strong habitat preferences in arid regions. Here, we used three separate gene regions (Cyt-b, D-loop and IRBP) and the combined dataset to analyse phylogeographical patterns and the historical demography of M. meridianus.
In this paper our aims are:(1) to determine the influence of the potential geographical isolation barrier on the phylogeographical structure within the current geographical range of M. meridianus.(2) to investigate the historical demography of this species and to determine the influence of climatic fluctuations and environmental variation on the phylogeographical structure within the current geographical range of M. meridianus.(3) solve the taxonomical issues by using phylogenetic analyses and sequence divergence values.(4) to assess whether, and to what extent, out results, obtained using a mitochondrial marker, are concordant with those achieved by nuclear markers.(5) Herein, through testing the genetic differentiation between the populations inhabiting inside and outside the regions of the Qaidam Basin, we presented a comparative phylogeographic study to determine whether there are similar in the phylogeographic patterns of the three rodents that may have resulted from a parallel response to the same historical events, or the three species display spatially and temporally distinct phylogeographic patterns as a consequence of different ecological niche modeling and altitude distributions. We then reconstructed the demographic history of each species to determine whether qualitative differences in demographic history. The main results are following:
(1) The phylogeographical patterns and demographic history of mitochondrial DNA (cytochrome b,n=327and D-loop, n=252) and nuclear DNA (IRBP gene, n=235) haplotypes were studied for M. meridianus in northern China, a desert-dwelling gerbil species. The phylogenetic analyses, which were performed on the separate and combined (mitochondrial+nuclear) datasets, revealed two divergent clades (Clade A and Clade B) corresponding to distinct geographical regions. Clade A contained the haplotypes found mostly in individuals from the Tianshan Mountains area. Clade B contained haplotypes from populations located in other deserts in northern China. The divergence times indicated that the history of M. meridianus was influenced by the uplift of the Tianshan Mountains and climate-induced habitat fluctuations. In the Pleistocene, the expansion of forests and grasslands during interglacial period led to the isolation of M. meridianus, which preferred to inhabit deserts. Hence, long geological isolation and M. meridianus adaptation to local ecological conditions led to its genetic divergence.
(2) Clade A had long-lasting demographic stability, most likely because the populations of this clade remained in a stable desert environment for a long time. However, the extension of other deserts and disappearance of palaeolakes during the last glacial period resulted in demographic expansion of Clade B.
(3) In our study, the K2P distances (for the Cyt-b, D-loop, and IRBP gene regions) between clades suggested the intraspecific value. Thus, our results indicated that two subspecies may exist within the M. meridianus.
(4)The IRBP-gene tree supports a reciprocally monophyletic relationship, whereas AVPR2-gene tree did not. Compared to the IRBP-gene tree, low genetic divergence was observed in the the AVPR2-gene tree, depicting a polyphyletic relationship. The inactivating mutations of the AVPR2gene are lethal to the desert-dwelling rodent under conditions of water restriction, because it causes an inherited NDI which is caraterized by the inability to concentrate the urine and an increased urine volume. Thus, AVPR2gene of this desert-dwelling rodent is likely suffer strong selective constraints to maintain the vital important gene function. When the gene expressed in several tissues some mutations are detrimental to be weeded out by selection. Thus, the AVPR2gene is more likely to be conserved among populations. For M. meridianus which is a nocturnal species, functional and structural abnormalities in IRBP do not play fatal role in the survive and reproduce because of an evolutionary sensory modality tradeoff, and visual impairment can be complemented by other sense, such as olfactory and auditory. The IRBP gene tend to accumulate mutations and lead to high level of genetic diversity.
(5) We conducted a comparative phylogeography of three rodent species(Allactaga sibirica, M. meridianus and Dipus sagitta) endemic on the northeast margin of the Tibetan Plateau (i.e. Qaidam Basin) and outside Qaidam Basin based on fragments of the mitochondrial Cyt-b gene in275individuals from34populations. Our analyses indicated that intraspecific genetic structures across the three rodents did not originate by the same historical events and they experienced different geological and climatic histories. A. sibirica with high-altitude distributions was subjected to strong isolation as a result of the fragmented habitats caused by complicated topology of high-altitude regions in Qaidam Basin. No clear signal of population expansion was recorded in A. sibirica, with a potential explanation that successive phases of population expansion obscured the signal during its migration process. M. meridianus and D. sagitta with low-altitude distributions experienced less isolation due to a relatively homogeneous habitat and lack of geographical barriers through Qaidam Basin. The two desert-dwelling rodent species, M. meridianus and D. sagitta, experienced a sudden demographic expansion during LGM with disappearance of the ancient lakes and the maximal extent of desert expansion.
本文的研究目的是：(1)国内子午沙鼠分布区内潜在的地理隔离屏障对其系统地理结构的影响；(2)第四纪气候波动和环境变化带来的生境变化对子午沙鼠的遗传谱系空间分布的历史格局以及种群动态历史的影响：(3)通过分子遗传的手段来重新审视子午沙鼠亚种的分类地位；(4)比较探讨不同遗传标记位点的遗传变异以及系统发生分析；(5)在子午沙鼠的分布范围内当属青藏高原地形地貌最为复杂,海拔较高,环境恶劣,因此将与子午沙鼠同域分布于柴达木盆地地区的三趾跳鼠和五趾跳鼠进行系统地理学和种群历史动态方面的比较分析,以分子遗传学的角度来佐证青藏高原动物区系形成与演化过程。
本文的研究结果表明：
(1)基于子午沙鼠线粒体基因(Cyt-b和D-loop)、核基因(IRBP基因)以及串联合并基因构建的系统发生树显示分布于中国西北部的子午沙鼠显著分化为两个支系,大多数天山地区的样本聚为一个支系,来自于中国西北部的其它沙漠地区以及少量的天山地区的样本聚为另一个支系。两个支系的分歧时间发生在天山抬升之后的中更新世中期,说明天山抬升有效地阻止子午沙鼠基因交流而产生了明显的地理隔离作用,此外,间冰期的暖湿气候带来的吐鲁番盆地森林草原扩张,也对其具有效的隔离作用导致遗传分化。
(2)由于分布于天山地区的种群长期处于环境稳定的状态,因此近期没有发生种群扩张。而另一个支系受到末次盛冰期气候的影响,沙漠环境快速急剧扩张和湖泊萎缩消失,从而分布于中国西北部其它沙漠地区的种群经历了近期种群扩张的历史。
(3)子午沙鼠两个支系间的遗传距离小于沙鼠属姐妹种种间的遗传距离,表明在分子遗传角度上,子午沙鼠两个支系的分类地位为亚种水平。
(4)基于新疆地区的134个子午沙鼠样本的IRBP序列和AVPR2序列分别构建的系统发生树。IRBP系统发生树显示83个单倍型聚为两个支系,两个支系之间具有显著的遗传分化,呈现互为单系群关系。而AVPR2系统发生树显示13个单倍型之间没有遗传分化,呈现多系群关系。对于夜行性动物子午沙鼠来说,具有暗适应调节能力的IRBP基因功能上对生命生存制约性低,如果IRBP基因发生有害突变导致暗适应功能不完全或丧失,可以通过听觉、嗅觉等的代偿作用来弥补,因此对子午沙鼠的生存繁殖几乎没有影响,因此根据遗传结果可以推断IRBP基因很可能接近于中性选择,有效的积累了遗传漂变等产生的突变。而具有缩尿功能的AVPR2基因所编码的蛋白对于生存于缺水干旱环境的沙漠地区的子午沙鼠是致命的,如果AVPR2基因发生有害突变导致缩尿功能的丧失就会威胁到子午沙鼠的生存,因此AVPR2基因很有可能受到强烈的自然选择作用,不能够有效的积累基因突变,从而系统发生树呈现没有遗传分化的多系群关系。
(5)五趾跳鼠柴达木地理单元与甘肃内蒙地理单元具有显著的遗传分化,而子午沙鼠和三趾跳鼠这两个地理单元之间几乎没有遗传分化。五趾跳鼠生境偏好选择海拔较高的山地草原带,因此在中更新世时期青藏高原抬升带来的气候干旱化导致的干草原发育扩张,为五趾跳鼠由甘肃内蒙等地迁移扩散到柴达木盆地提供了有利条件,由于高海拔地区的地形地貌复杂,容易受到生境片段化而导致遗传分化,本文的结果还显示随着种群由西向东分布共祖时间逐渐递减,可以看出其在柴达木盆地中由西向东迁移扩散。而子午沙鼠和三趾跳鼠生境偏好选择沙漠且分布海拔较低,随着晚更新世时期气候更加干旱化导致柴达木盆地沙漠迅速形成发育,使子午沙鼠和三趾跳鼠辐射性迅速迁移扩散到柴达木盆地,没有足够的时间产生遗传分化。
The range of Meriones meridianus (Rodentia:Cricetidae:Meriones) spans several major arid and semi-arid biogeographic regions in northern China (Luo, Chen et al.2000). In northern China, this desert-dwelling rodent species is distributed along the edge of deserts, and it is an important component of the desert systems that it inhabits. It could serve as an indicator of desertification. Northern China is endowed with a complicated terrain of deserts, high mountains, rivers, lakes, and basins which vary widely in elevation from the Qaidam Basin, which is over2700m above sea level to the Turpan Basis, which is103m below sea level. These complex and varied terrains, in conjunction with climate fluctuations, have facilitated the diversification and vicariance of M. meridianus. The species provide good models for reconstructions of how environmental parameters influenced the history of biota. This is because its short generation time, rapid mtDNA substitution rate, relatively limited dispersal ability, and strong associations with particular habitats give rise to informative contemporary patterns of genetic variation. In particular, the midday gerbil M. meridianus may display especially informative phylogeographical patterns due to its strong habitat preferences in arid regions. Here, we used three separate gene regions (Cyt-b, D-loop and IRBP) and the combined dataset to analyse phylogeographical patterns and the historical demography of M. meridianus.
In this paper our aims are:(1) to determine the influence of the potential geographical isolation barrier on the phylogeographical structure within the current geographical range of M. meridianus.(2) to investigate the historical demography of this species and to determine the influence of climatic fluctuations and environmental variation on the phylogeographical structure within the current geographical range of M. meridianus.(3) solve the taxonomical issues by using phylogenetic analyses and sequence divergence values.(4) to assess whether, and to what extent, out results, obtained using a mitochondrial marker, are concordant with those achieved by nuclear markers.(5) Herein, through testing the genetic differentiation between the populations inhabiting inside and outside the regions of the Qaidam Basin, we presented a comparative phylogeographic study to determine whether there are similar in the phylogeographic patterns of the three rodents that may have resulted from a parallel response to the same historical events, or the three species display spatially and temporally distinct phylogeographic patterns as a consequence of different ecological niche modeling and altitude distributions. We then reconstructed the demographic history of each species to determine whether qualitative differences in demographic history. The main results are following:
(1) The phylogeographical patterns and demographic history of mitochondrial DNA (cytochrome b,n=327and D-loop, n=252) and nuclear DNA (IRBP gene, n=235) haplotypes were studied for M. meridianus in northern China, a desert-dwelling gerbil species. The phylogenetic analyses, which were performed on the separate and combined (mitochondrial+nuclear) datasets, revealed two divergent clades (Clade A and Clade B) corresponding to distinct geographical regions. Clade A contained the haplotypes found mostly in individuals from the Tianshan Mountains area. Clade B contained haplotypes from populations located in other deserts in northern China. The divergence times indicated that the history of M. meridianus was influenced by the uplift of the Tianshan Mountains and climate-induced habitat fluctuations. In the Pleistocene, the expansion of forests and grasslands during interglacial period led to the isolation of M. meridianus, which preferred to inhabit deserts. Hence, long geological isolation and M. meridianus adaptation to local ecological conditions led to its genetic divergence.
(2) Clade A had long-lasting demographic stability, most likely because the populations of this clade remained in a stable desert environment for a long time. However, the extension of other deserts and disappearance of palaeolakes during the last glacial period resulted in demographic expansion of Clade B.
(3) In our study, the K2P distances (for the Cyt-b, D-loop, and IRBP gene regions) between clades suggested the intraspecific value. Thus, our results indicated that two subspecies may exist within the M. meridianus.
(4)The IRBP-gene tree supports a reciprocally monophyletic relationship, whereas AVPR2-gene tree did not. Compared to the IRBP-gene tree, low genetic divergence was observed in the the AVPR2-gene tree, depicting a polyphyletic relationship. The inactivating mutations of the AVPR2gene are lethal to the desert-dwelling rodent under conditions of water restriction, because it causes an inherited NDI which is caraterized by the inability to concentrate the urine and an increased urine volume. Thus, AVPR2gene of this desert-dwelling rodent is likely suffer strong selective constraints to maintain the vital important gene function. When the gene expressed in several tissues some mutations are detrimental to be weeded out by selection. Thus, the AVPR2gene is more likely to be conserved among populations. For M. meridianus which is a nocturnal species, functional and structural abnormalities in IRBP do not play fatal role in the survive and reproduce because of an evolutionary sensory modality tradeoff, and visual impairment can be complemented by other sense, such as olfactory and auditory. The IRBP gene tend to accumulate mutations and lead to high level of genetic diversity.
(5) We conducted a comparative phylogeography of three rodent species(Allactaga sibirica, M. meridianus and Dipus sagitta) endemic on the northeast margin of the Tibetan Plateau (i.e. Qaidam Basin) and outside Qaidam Basin based on fragments of the mitochondrial Cyt-b gene in275individuals from34populations. Our analyses indicated that intraspecific genetic structures across the three rodents did not originate by the same historical events and they experienced different geological and climatic histories. A. sibirica with high-altitude distributions was subjected to strong isolation as a result of the fragmented habitats caused by complicated topology of high-altitude regions in Qaidam Basin. No clear signal of population expansion was recorded in A. sibirica, with a potential explanation that successive phases of population expansion obscured the signal during its migration process. M. meridianus and D. sagitta with low-altitude distributions experienced less isolation due to a relatively homogeneous habitat and lack of geographical barriers through Qaidam Basin. The two desert-dwelling rodent species, M. meridianus and D. sagitta, experienced a sudden demographic expansion during LGM with disappearance of the ancient lakes and the maximal extent of desert expansion.
引文
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