长柄水青冈种群遗传结构和系统地理学研究
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摘要
系统地理学主要研究影响物种的基因谱系形成现有地理分布格局的历史过程,常用于系统地理学研究的基因是细胞器DNA(线粒体DNA、叶绿体DNA)和核DNA。细胞器DNA和核DNA是两种不同遗传模式的基因,两种基因片段结合起来用于系统地理学的研究,可以揭示种群的遗传结构并推测影响种群现有遗传分布格局的历史过程。欧洲和北美最早开展系统地理学方面的研究。第四纪冰期时欧洲和北美大部分地区都被冰盖覆盖,许多古老的物种只在少数避难所存活下来,存活下来的种群在冰期结束后由南向北重新扩散;而东亚大陆并没有被巨大的冰川覆盖,但气温比现在降低8-12℃。那么当时许多东亚物种的分布区是否与当时欧洲和北美生物的分布区发生了相同的变化?这些东亚物种是否也存在冰期避难所?冰期后又是沿怎样的扩散路线形成现在的分布格局?
为了解决以上的问题,我们选取了长柄水青冈作为研究材料。第三纪晚期时,水青冈属植物广泛分布于我国大部分地区,甚至北部地区也有分布。第四纪冰期气候使其退缩到南方亚热带地区,但是冰期后气候回暖时,喜马拉雅山和青藏高原的抬升,使我国东部和西南部季风性气候加强,西北华北大陆性气候加强,气候条件的改变限制了水青冈向北、向西北的扩散,水青冈继续保留在南方亚热带地区。长柄水青冈是我国水青冈属中分布最广泛的种,主要在我国秦岭以南、五岭南坡以北各地分布,是我国亚热带常绿落叶阔叶混交林的优势种之一,分布区呈东西样带分布,东部地区分布呈明显片断化。第四纪冰期对水青冈现在的分布格局有着重要的影响。那么,第四纪冰期时长柄水青冈的避难所在那里?冰期后沿怎样的路线向外扩散并形成现有分布格局?本文针对以上问题进行探讨:采用核DNA的SSR标记和叶绿体DNA标记,分析长柄水青冈的种群遗传结构,探讨种子流和花粉流在种群遗传分化中的贡献率;希望通过分析种群遗传结构和叶绿体单倍型的地理分布格局,揭示长柄水青冈的冰期避难所和冰期后的扩散路线。主要结果如下:
(1)用7对微卫星引物对27个种群576个个体分析表明,长柄水青冈种群有较低的遗传多样性。大部分的遗传变异存在于种群内。种群间遗传分化程度高于其它水青冈种。
东部种群和西部种群表现出不同的遗传多样性分布格局。东部种群的遗传多样性低于西部种群。东部分布区种群间的遗传分化比较大,表明长期的片断化阻隔了种群间花粉和种子的传播,使种群间的遗传分化加大。东部不同种群的遗传多样性变化很大:分布区北部的种群遗传多样性较低,种群规模最小;分布区中部的种群具有最高的遗传多样性。WL种群和MD种群检测到了瓶颈效应。这说明长柄水青冈种群分布有衰退的趋势,分布区的南缘和北缘逐渐变得不适合长柄水青冈的生长。
西部地区虽然有高山的阻隔,但种群为大规模连续分布,所以相对于东部种群而言,西部种群的种子流和花粉流能在短距离内扩散,种群间的基因流比较高,遗传分化程度比较低。
(2)用2对叶绿体通用引物(trnS-trnG、psbB-psbF)对22个种群201个个体测序,共得到13种单倍型,只有4个种群(JL、WL、JS、LS)存在单倍型多样性。叶绿体DNA分析表明93.197%的遗传变异存在于种群间,种群间基因分化系数G_(ST)为0.900。长柄水青冈与亮叶水青冈两个种间检测到3种共享单倍型,表明两个种的分化时间较近,二者间曾发生基因渐渗。所有种群都没有检测到种群扩张,表明冰期后气候等环境条件限制了长柄水青冈种群大规模的扩张。
叶绿体单倍型分布呈显著的地理格局,表现出东部分布区与西部分布区单倍型地理分布格局的不同。西部地区种群有较多的原始单倍型和独特的单倍型;同一单倍型在南部地区分布很近,表明冰期后的扩散发生在短距离内;北部因为没有高山的阻隔,单倍型扩散距离较远。东部地区也有较多独特的单倍型,同一单倍型的分布比较广,表明冰期后存在长距离的扩散。
(3)根据两种遗传标记估算的花粉-种子迁移率r为34.228,即在长柄水青冈种群中通过花粉的基因流要比通过种子的基因流高。通过cpDNA计算的遗传分化G_(STm)与其它壳斗科物种接近,表明片断化对长柄水青冈种群间的种子流影响不大,对花粉流的影响比较大。西部高山对花粉的传播也起了一定的阻隔作用。
(4)微卫星揭示的种群遗传结构和cpDNA单倍型地理格局表明,长柄水青冈存在东部和西部两种不同的避难所格局。
西部的避难所以广西、贵州为中心,冰期后种群从低纬度地区向北扩散,高山的阻隔使种群只能短距离扩散或者沿山体上下迁移。避难所北部的种群,向北的迁移扩散不受高山的限制,沿四川盆地周围的山系向北扩散,并越过四川盆地,沿中部低海拔地区向东部地区扩散。
东部避难所位于武夷山山脉的两侧,因武夷山的阻隔,表现出两种不同的扩散路线。武夷山脉以东、南岭东段的避难所,沿武夷山脉东侧向北扩散,到达武夷山脉北段的浙江省境内。而山脉西侧的避难所种群则是沿武夷山西侧向北扩散至浙江省南部。这两条扩散路线都没有越过武夷山脊,是单独发生的。浙江省南部武夷山系的北段单倍型比较丰富,遗传多样性比较高,是两条迁移路线的汇总处。
Phylogeography studies the principles and processes governing the geographical distributions of genealogical lineages,which emphasizes historical aspects of the contemporary spatial distributions of gene lineages.Compared with biparentally inherited nuclear markers,the cpDNA variation provides special markers for phylogeny reconstruction at the population level and the geographic structure of variation,which provide insights into the phylogeograpy of the species. The contrasted patterns of inheritance of organelle and nuclear genes can be used to unravel the complexity of gene flow in plants,as they are predicted to result in very different distribution of genetic diversity within and among populations.In the last two decades,extensive phylogeographic studies have been conducted on taxa from Europe and North America,where were covered by huge glaciers during the Quaternary.Some species only survived in a few refugia and recolonized after glaciation in Europe and North America.Contrasting to Europe and North America,East Asia had not been covered by huge glaciers during the Quaternary glaciation. However,the temperature had decreased about 8-12℃.Did distribution ranges of the species in East Asia also retract like those in European and North America? Where did they survive from the cold temperature and how did they recolonize after the Quaternary glacial ages?
In order to answer these questions,we choose Fagus longipetiolata as the focal species. Fagus(beech)is among the most abundant and economically important genera of broadleaved trees in northern hemisphere temperate forests.The genus Fagus comprises 10 to 14 species of monoecious trees that are disjunctly distributed in Europe and south-western Asia(1-5 species), eastern Northern America and Mexico(1-2 species)and eastern Asia(7-12 species).F. longipetiolata distributes widely in subtropical southern China,which distributed much further north during the late Tertiary in China.During the early Quaternary,it declined and disappeared in northern China,but expanded in subtropical southern China.And its distribution range is from Xichou County,Yunnan Province,to Tiantai Mountain,Zhejiang Province.F.longipetiolata is a monoecious,long-lived woody angiosperm.It is pollinated by wind and dispersed by gravity,but rodents may serve as the secondary dispersers.
Nuclear microsatellites and cpDNA variations among populations of F.longipetiolata were investigated in the study.The main results are as following.
(1)A total of 576 individuals were collected from 27 populations covering most distribution range of E longipetiolata.We used 7 pairs of microsatellite primers,among them 6 were screened from congeneric species,E sylvatica and E crenata,and one,locus FL05,was developed by ourself.Low genetic diversity was found within populations.The genetic differentiation among populations was high(F_(ST)=0.23).Gene flow for 21 populations(Nm)was estimated to be 0.83.
Different genetic patterns were found in populations of eastern and western range. Populations of western range had higher genetic diversity than those of eastern range.And genetic differentiation was small among populations.However,genetic diversity was much more diverse among populations of eastern range.Higher genetic diversity was found within large populations than small populations.Populations of northern and southern part had lower genetic diversity than that in central populations in eastern China.Recent bottleneck effect was found in a southern population,MD population,and a northern population,SH population.
(2)Two pairs of universal cpDNA primers,trnS-trnG and psbB-psbF,were used.A total of 201 individuals from 26 populations were sequenced successfully.15 polymorphic sites over 1664bp and 13 haplotypes were observed.However,within-population haplotype diversity was very low.Only 4 populations had more than one haplotypes.About 93.197 percentage of genetic differentiation was among populations.The genetic differentiation value was high(G_(ST)=0.900, 21 populations).There were 3 haplotypes shared with F.lucida indicating the early introgression between these two species or they shared a common ancestor.
Significant geographic structure of haplotypes was detected.Populations of western and eastern ranges had own unique haplotypes.More ancestral haplotypes were found and distributed widely in populations of western range.Dispersion of the same haplotype in western range happened in short distance.However the same haplotype in eastern China was found in a long distance,meaning that it was distributed from south refugia to north mountains in eastern China.
(3)Based on Ennos' model,the pollen-seed ratio(r)was estimated to be 34.228,indicating that pollen flow had played a more important role in the maintenance of genetic structure than seed flow.However this value was much smaller than that in other species of Fagaceae,due to the decreased pollen flow among populations by natural fragmentation,given similar cpDNA-based G_(ST)with other Fagus species.
(4)From this study,two contrasting pattern in glacial refugia and recolonization routes were found in eastern and western distribution range.
Southwestern China including Guangxi Province and Guizhou Province provided refugia for F.longipetiolata during Quaternary glaciations.When it became warm,its postglacial populations migrated from one region to the other along the lower mountains.It was short-distance disseminated for populations of south part in this refugia because of high mountains.While there were Sichuan Basin in north part of this refugia,population migrated northward along the mountains surrounding Sichuan Basin.An exceptional case of long-distance dissemination,the modern distribution of haplotype H1 was ascribed to a postglacial colonization route starting from north part of this refugia,eastward crossing the Sichuan Basin,central of China and reaching East China.
Wuyishan-Nanling served as the other refugia.This refugia was separated into two parts: south and north,by Wuyishan.After glaciations,populations in the south part migrated northeastward along Wuyishan.In western Zhejiang,E longipetiolata migrated to the north and east parts of Zhejiang Province.Populations survived in the north Wuyishan refugia migrated to the north as well as northeastward along the north ridge of Wuyishan and then mixed with the migrators of south part in western Zhejiang Province.Therefore,high and mixed haplotype diversity was found in western Zhejiang Province.
Given the closely phylogenetic relationship based on haplotype and SSRs between populations of Fujian Province and populations of western China,populations of eastern range might colonize from western China before the Quaternary glaciations.
为了解决以上的问题,我们选取了长柄水青冈作为研究材料。第三纪晚期时,水青冈属植物广泛分布于我国大部分地区,甚至北部地区也有分布。第四纪冰期气候使其退缩到南方亚热带地区,但是冰期后气候回暖时,喜马拉雅山和青藏高原的抬升,使我国东部和西南部季风性气候加强,西北华北大陆性气候加强,气候条件的改变限制了水青冈向北、向西北的扩散,水青冈继续保留在南方亚热带地区。长柄水青冈是我国水青冈属中分布最广泛的种,主要在我国秦岭以南、五岭南坡以北各地分布,是我国亚热带常绿落叶阔叶混交林的优势种之一,分布区呈东西样带分布,东部地区分布呈明显片断化。第四纪冰期对水青冈现在的分布格局有着重要的影响。那么,第四纪冰期时长柄水青冈的避难所在那里?冰期后沿怎样的路线向外扩散并形成现有分布格局?本文针对以上问题进行探讨:采用核DNA的SSR标记和叶绿体DNA标记,分析长柄水青冈的种群遗传结构,探讨种子流和花粉流在种群遗传分化中的贡献率;希望通过分析种群遗传结构和叶绿体单倍型的地理分布格局,揭示长柄水青冈的冰期避难所和冰期后的扩散路线。主要结果如下:
(1)用7对微卫星引物对27个种群576个个体分析表明,长柄水青冈种群有较低的遗传多样性。大部分的遗传变异存在于种群内。种群间遗传分化程度高于其它水青冈种。
东部种群和西部种群表现出不同的遗传多样性分布格局。东部种群的遗传多样性低于西部种群。东部分布区种群间的遗传分化比较大,表明长期的片断化阻隔了种群间花粉和种子的传播,使种群间的遗传分化加大。东部不同种群的遗传多样性变化很大:分布区北部的种群遗传多样性较低,种群规模最小;分布区中部的种群具有最高的遗传多样性。WL种群和MD种群检测到了瓶颈效应。这说明长柄水青冈种群分布有衰退的趋势,分布区的南缘和北缘逐渐变得不适合长柄水青冈的生长。
西部地区虽然有高山的阻隔,但种群为大规模连续分布,所以相对于东部种群而言,西部种群的种子流和花粉流能在短距离内扩散,种群间的基因流比较高,遗传分化程度比较低。
(2)用2对叶绿体通用引物(trnS-trnG、psbB-psbF)对22个种群201个个体测序,共得到13种单倍型,只有4个种群(JL、WL、JS、LS)存在单倍型多样性。叶绿体DNA分析表明93.197%的遗传变异存在于种群间,种群间基因分化系数G_(ST)为0.900。长柄水青冈与亮叶水青冈两个种间检测到3种共享单倍型,表明两个种的分化时间较近,二者间曾发生基因渐渗。所有种群都没有检测到种群扩张,表明冰期后气候等环境条件限制了长柄水青冈种群大规模的扩张。
叶绿体单倍型分布呈显著的地理格局,表现出东部分布区与西部分布区单倍型地理分布格局的不同。西部地区种群有较多的原始单倍型和独特的单倍型;同一单倍型在南部地区分布很近,表明冰期后的扩散发生在短距离内;北部因为没有高山的阻隔,单倍型扩散距离较远。东部地区也有较多独特的单倍型,同一单倍型的分布比较广,表明冰期后存在长距离的扩散。
(3)根据两种遗传标记估算的花粉-种子迁移率r为34.228,即在长柄水青冈种群中通过花粉的基因流要比通过种子的基因流高。通过cpDNA计算的遗传分化G_(STm)与其它壳斗科物种接近,表明片断化对长柄水青冈种群间的种子流影响不大,对花粉流的影响比较大。西部高山对花粉的传播也起了一定的阻隔作用。
(4)微卫星揭示的种群遗传结构和cpDNA单倍型地理格局表明,长柄水青冈存在东部和西部两种不同的避难所格局。
西部的避难所以广西、贵州为中心,冰期后种群从低纬度地区向北扩散,高山的阻隔使种群只能短距离扩散或者沿山体上下迁移。避难所北部的种群,向北的迁移扩散不受高山的限制,沿四川盆地周围的山系向北扩散,并越过四川盆地,沿中部低海拔地区向东部地区扩散。
东部避难所位于武夷山山脉的两侧,因武夷山的阻隔,表现出两种不同的扩散路线。武夷山脉以东、南岭东段的避难所,沿武夷山脉东侧向北扩散,到达武夷山脉北段的浙江省境内。而山脉西侧的避难所种群则是沿武夷山西侧向北扩散至浙江省南部。这两条扩散路线都没有越过武夷山脊,是单独发生的。浙江省南部武夷山系的北段单倍型比较丰富,遗传多样性比较高,是两条迁移路线的汇总处。
Phylogeography studies the principles and processes governing the geographical distributions of genealogical lineages,which emphasizes historical aspects of the contemporary spatial distributions of gene lineages.Compared with biparentally inherited nuclear markers,the cpDNA variation provides special markers for phylogeny reconstruction at the population level and the geographic structure of variation,which provide insights into the phylogeograpy of the species. The contrasted patterns of inheritance of organelle and nuclear genes can be used to unravel the complexity of gene flow in plants,as they are predicted to result in very different distribution of genetic diversity within and among populations.In the last two decades,extensive phylogeographic studies have been conducted on taxa from Europe and North America,where were covered by huge glaciers during the Quaternary.Some species only survived in a few refugia and recolonized after glaciation in Europe and North America.Contrasting to Europe and North America,East Asia had not been covered by huge glaciers during the Quaternary glaciation. However,the temperature had decreased about 8-12℃.Did distribution ranges of the species in East Asia also retract like those in European and North America? Where did they survive from the cold temperature and how did they recolonize after the Quaternary glacial ages?
In order to answer these questions,we choose Fagus longipetiolata as the focal species. Fagus(beech)is among the most abundant and economically important genera of broadleaved trees in northern hemisphere temperate forests.The genus Fagus comprises 10 to 14 species of monoecious trees that are disjunctly distributed in Europe and south-western Asia(1-5 species), eastern Northern America and Mexico(1-2 species)and eastern Asia(7-12 species).F. longipetiolata distributes widely in subtropical southern China,which distributed much further north during the late Tertiary in China.During the early Quaternary,it declined and disappeared in northern China,but expanded in subtropical southern China.And its distribution range is from Xichou County,Yunnan Province,to Tiantai Mountain,Zhejiang Province.F.longipetiolata is a monoecious,long-lived woody angiosperm.It is pollinated by wind and dispersed by gravity,but rodents may serve as the secondary dispersers.
Nuclear microsatellites and cpDNA variations among populations of F.longipetiolata were investigated in the study.The main results are as following.
(1)A total of 576 individuals were collected from 27 populations covering most distribution range of E longipetiolata.We used 7 pairs of microsatellite primers,among them 6 were screened from congeneric species,E sylvatica and E crenata,and one,locus FL05,was developed by ourself.Low genetic diversity was found within populations.The genetic differentiation among populations was high(F_(ST)=0.23).Gene flow for 21 populations(Nm)was estimated to be 0.83.
Different genetic patterns were found in populations of eastern and western range. Populations of western range had higher genetic diversity than those of eastern range.And genetic differentiation was small among populations.However,genetic diversity was much more diverse among populations of eastern range.Higher genetic diversity was found within large populations than small populations.Populations of northern and southern part had lower genetic diversity than that in central populations in eastern China.Recent bottleneck effect was found in a southern population,MD population,and a northern population,SH population.
(2)Two pairs of universal cpDNA primers,trnS-trnG and psbB-psbF,were used.A total of 201 individuals from 26 populations were sequenced successfully.15 polymorphic sites over 1664bp and 13 haplotypes were observed.However,within-population haplotype diversity was very low.Only 4 populations had more than one haplotypes.About 93.197 percentage of genetic differentiation was among populations.The genetic differentiation value was high(G_(ST)=0.900, 21 populations).There were 3 haplotypes shared with F.lucida indicating the early introgression between these two species or they shared a common ancestor.
Significant geographic structure of haplotypes was detected.Populations of western and eastern ranges had own unique haplotypes.More ancestral haplotypes were found and distributed widely in populations of western range.Dispersion of the same haplotype in western range happened in short distance.However the same haplotype in eastern China was found in a long distance,meaning that it was distributed from south refugia to north mountains in eastern China.
(3)Based on Ennos' model,the pollen-seed ratio(r)was estimated to be 34.228,indicating that pollen flow had played a more important role in the maintenance of genetic structure than seed flow.However this value was much smaller than that in other species of Fagaceae,due to the decreased pollen flow among populations by natural fragmentation,given similar cpDNA-based G_(ST)with other Fagus species.
(4)From this study,two contrasting pattern in glacial refugia and recolonization routes were found in eastern and western distribution range.
Southwestern China including Guangxi Province and Guizhou Province provided refugia for F.longipetiolata during Quaternary glaciations.When it became warm,its postglacial populations migrated from one region to the other along the lower mountains.It was short-distance disseminated for populations of south part in this refugia because of high mountains.While there were Sichuan Basin in north part of this refugia,population migrated northward along the mountains surrounding Sichuan Basin.An exceptional case of long-distance dissemination,the modern distribution of haplotype H1 was ascribed to a postglacial colonization route starting from north part of this refugia,eastward crossing the Sichuan Basin,central of China and reaching East China.
Wuyishan-Nanling served as the other refugia.This refugia was separated into two parts: south and north,by Wuyishan.After glaciations,populations in the south part migrated northeastward along Wuyishan.In western Zhejiang,E longipetiolata migrated to the north and east parts of Zhejiang Province.Populations survived in the north Wuyishan refugia migrated to the north as well as northeastward along the north ridge of Wuyishan and then mixed with the migrators of south part in western Zhejiang Province.Therefore,high and mixed haplotype diversity was found in western Zhejiang Province.
Given the closely phylogenetic relationship based on haplotype and SSRs between populations of Fujian Province and populations of western China,populations of eastern range might colonize from western China before the Quaternary glaciations.
引文
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