太平洋鲱和大头鳕的群体遗传学研究
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摘要
海洋鱼类通常展现三种遗传结构模式(群体间遗传差异较小并且是不显著的、群体间遗传差异较小但却是显著的、群体间遗传分化较大并伴有系统发育上的分支)和三种系统地理格局(系统发育上具有较大分化的类群在空间上异域分布或同域分布、系统发育上连续的类群在空间上也是连续的)。这些遗传差异模式的形成主要和更新世剧烈的气候波动、当前的海洋环境因子以及海洋鱼类自身的生态特性密切相关。太平洋鲱和大头鳕是北太平洋两个重要经济鱼种,二者都起源于大西洋,通过跨北冰洋扩散进入太平洋,更新世的冰期循环和北太平洋独特的海洋地理环境很可能在两个种种内形成具有代表性的遗传结构和系统地理格局。
我们采用形态学、线粒体DNA和微卫星三种标记来检测太平洋鲱的群体结构和系统地理格局,并探讨了太平洋鲱与大西洋鲱的分化事件。主要研究结果如下：
(1)对北太平洋沿岸8个地理群体424个个体的分节特征和量度特征开展了多变量分析,结果表明这些群体之间在这些特征变量上均存在显著差异,些特征变量在群体间存在纬度渐变趋势。
(2)应用线粒体DNA控制区对9个地理群体开展了分子系统地理学研究,在太平洋鲱种内检测到了3个高度分化的系统发育分支,一个分支占据西北太平洋,和另外两个分支在东太平洋发生二次连接,3个分支可能形成于更新世冰期的隔离。3个分支的频率分布差异定义了两个地理组群,西北太平洋组群和东太平洋组群。
(3)通过检测9个微卫星多态位点差异,在太平洋鲱两两群体间发现了小但极显著的遗传差异,9个群体被进一步细分成3个地理组群,在之前两个地理组群划分的基础上,黄海群体独立成群。地形的阻隔、海流和海水温度的限制很可能是组群间较大遗传差异产生的原因。
(4)为检测黄海群体时间上的遗传结构,比较了3个不同时期黄海群体的控制区序列差异,结果表明黄海内的鲱鱼群体为一个独立的随机交配种群。我们推测黄海群体的特殊性可能与黑潮暖流及其支流的高温限制作用有关。
(5)为探讨太平洋鲱的进化起源,以及与其姊妹种大西洋鲱之间在形态上和遗传上的关系。我们比较了太平洋鲱黄海群体和一个大西洋鲱群体的24个形态特征差异,基于11个形态特征参数建立了判别式,判别率达98.8%。分析了两个种的细胞色素b和控制区序列,基于8.1%的控制区净遗传距离推测两个种在线粒体上分化发生在距今约125万年前。基于分支间的净遗传距离估算太平洋鲱种内分支进化发生在更新世晚期。核苷酸不配对分布结果表明两个种都经历了群体扩张。
为检测大头鳕的群体遗传结构和历史动态,我们采用线粒体DNA和微卫星两种分子标记对采自北太平洋的9个地理群体开展分析,主要研究结果如下：
(1)基于线粒体DNA控制区的分子系统地理学研究结果,大头鳕种内无谱系结构发生,且具有相当低的遗传多样度水平,西北太平洋群体间存在小但显著遗传结构,东太平洋群体间存在中度的距离隔离趋势。
(2)基于8个微卫星多态位点的差异,在西北太平洋4个地理群体间检测到微弱但显著的遗传结构,黄海群体和其他3个群体存在明显分化,黑潮暖流及其支流同样可能对大头鳕的黄海群体起到限制作用。我们的微卫星结果发现大头鳕具有相当高的遗传多样度水平。
我们推测线粒体和微卫星结果的不一致主要来自于线粒体相比核基因四倍高的遗传漂变率以及微卫星相当高的突变速率。线粒体和微卫星结果都提示大头鳕群体经历了近期的瓶颈效应。
Marine fish usually exhibited three genetic structuring patterns (i. little genetic differentiations, ii. small but significant genetic differentiations, iii. large genetic divergence with phylogenetic lineages) and three phylogeographic patterns (i. deep phylogenetic break with lineages allopatric, ii. deep phylogenetic break with lineages sympatric, iii. shallow phylogenetic relationship or no genealogy structure). All these patterns of genetic variations involved historical climatical fluctuations, contemporary marine environment factors and life-history traits of marine fish. Pacific herring and Pacific cod are two commercially important fish species in North Pacific. They were considered derived from an Atlantic invader by trans-Arctic dispersal. Large glacial-interglacial circles in Pleistocene and unique oceanographic characters of North Pacific may have influenced heavily the population structuring and phylogeographic patterns in the two species.
Morphological, mitochondrial DNA sequence and microsatellite DNA markers were used to estimate the population structure and phylogeographic pattern of Pacific herring in North Pacific, the divergence between Pacific herring and Atlantic herring was also discussed.
(1) Meristic and morphometric characters were compared by multivariate analysis for eight populations of total 424 individuals, significant differences were found in means of all these characters among eight populations, and a latitudinal cline was observed in several morphological characters.
(2) Three high distinct lineages were detected in 318 individuals from nine populations across the range of the species based on sequence variations of mtDNA control region, which might be isolated and diverged during glacial periods of Pleistocene. One lineage dominated the Northwestern Pacific, while the other two lineages occurred secondary contact in Eastern Pacific. The frequency difference of the three lineages in 9 populations defined two geographic groups, a Northwestern Pacific group and an Eastern Pacific Group.
(3) High significant genetic differentiations were found among 9 populations by scanning 9 microsatellite loci variations, which subdivided populations into three groups. This grouping picked up the Yellow Sea population as an independent group. Sea water temperature barriers from Kuroshio Current and Tsushima Current might restrict the gene flows between populations in Yellow Sea and other locations.
(4) MtDNA control region sequences were used to investigate temporal population structure of Pacific herring which were collected from three different periods in Yellow Sea. No significant genetic differentiations were detected among the three populations and the result suggested populations in Yellow Sea belong to a single panmictic stock.
(5) An Atlantic herring population was also analyzed as an outgroup to discuss the origin and evolution of Pacific herring. Meristic and morphometric characters were compared between the Atlantic herring population and a Pacific herring population from Yellow Sea. A discriminant formula was created with 11 character parameters, and the accuracy was 98.8%. MtDNA variations were analyzed using sequence data from cytochrome b and control region. Based on 8.1% net genetic distance in control region, we postulated the divergence in mtDNA between the two species occurred at about 1.25 million years ago. Three lineages of Pacific herring were considered to diverge during late Pleistocene. Mismatch distribution analyses indicated population expansion in both species. MtDNA sequence and microsatellite DNA markers were used to estimate the population structure and demographic history of Pacific cod in North Pacific.
(1) MtDNA control region were analyzed for 259 individuals from nine localities over the species'range. The result showed lack of genealogical structure and remarkably low control region diversity. Small but significant genetic differentiations were detected among Northwestern Pacific populations. Four populations from coastal North America exhibited a trend of isolation by distance.
(2) Small but significant genetic structure was also found in four populations of Northwestern Pacific based on microsatellite analyses. Like Pacific herring, large genetic differentiations between the Yellow Sea population and other populations might be also attributed to sea water temperature barriers. Contrary to mtDNA result, microsatellite analyses revealed relative high genetic diversity for Pacific cod.
We interpreted such discordance between mitochondrial and microsatellites might be caused by the four-fold higher rate of drift for mtDNA than nuclear genes and fairly high mutation rate for microsatellite DNA. Both mtDNA and microsatellite results suggested a recent bottleneck in Pacific cod.
我们采用形态学、线粒体DNA和微卫星三种标记来检测太平洋鲱的群体结构和系统地理格局,并探讨了太平洋鲱与大西洋鲱的分化事件。主要研究结果如下：
(1)对北太平洋沿岸8个地理群体424个个体的分节特征和量度特征开展了多变量分析,结果表明这些群体之间在这些特征变量上均存在显著差异,些特征变量在群体间存在纬度渐变趋势。
(2)应用线粒体DNA控制区对9个地理群体开展了分子系统地理学研究,在太平洋鲱种内检测到了3个高度分化的系统发育分支,一个分支占据西北太平洋,和另外两个分支在东太平洋发生二次连接,3个分支可能形成于更新世冰期的隔离。3个分支的频率分布差异定义了两个地理组群,西北太平洋组群和东太平洋组群。
(3)通过检测9个微卫星多态位点差异,在太平洋鲱两两群体间发现了小但极显著的遗传差异,9个群体被进一步细分成3个地理组群,在之前两个地理组群划分的基础上,黄海群体独立成群。地形的阻隔、海流和海水温度的限制很可能是组群间较大遗传差异产生的原因。
(4)为检测黄海群体时间上的遗传结构,比较了3个不同时期黄海群体的控制区序列差异,结果表明黄海内的鲱鱼群体为一个独立的随机交配种群。我们推测黄海群体的特殊性可能与黑潮暖流及其支流的高温限制作用有关。
(5)为探讨太平洋鲱的进化起源,以及与其姊妹种大西洋鲱之间在形态上和遗传上的关系。我们比较了太平洋鲱黄海群体和一个大西洋鲱群体的24个形态特征差异,基于11个形态特征参数建立了判别式,判别率达98.8%。分析了两个种的细胞色素b和控制区序列,基于8.1%的控制区净遗传距离推测两个种在线粒体上分化发生在距今约125万年前。基于分支间的净遗传距离估算太平洋鲱种内分支进化发生在更新世晚期。核苷酸不配对分布结果表明两个种都经历了群体扩张。
为检测大头鳕的群体遗传结构和历史动态,我们采用线粒体DNA和微卫星两种分子标记对采自北太平洋的9个地理群体开展分析,主要研究结果如下：
(1)基于线粒体DNA控制区的分子系统地理学研究结果,大头鳕种内无谱系结构发生,且具有相当低的遗传多样度水平,西北太平洋群体间存在小但显著遗传结构,东太平洋群体间存在中度的距离隔离趋势。
(2)基于8个微卫星多态位点的差异,在西北太平洋4个地理群体间检测到微弱但显著的遗传结构,黄海群体和其他3个群体存在明显分化,黑潮暖流及其支流同样可能对大头鳕的黄海群体起到限制作用。我们的微卫星结果发现大头鳕具有相当高的遗传多样度水平。
我们推测线粒体和微卫星结果的不一致主要来自于线粒体相比核基因四倍高的遗传漂变率以及微卫星相当高的突变速率。线粒体和微卫星结果都提示大头鳕群体经历了近期的瓶颈效应。
Marine fish usually exhibited three genetic structuring patterns (i. little genetic differentiations, ii. small but significant genetic differentiations, iii. large genetic divergence with phylogenetic lineages) and three phylogeographic patterns (i. deep phylogenetic break with lineages allopatric, ii. deep phylogenetic break with lineages sympatric, iii. shallow phylogenetic relationship or no genealogy structure). All these patterns of genetic variations involved historical climatical fluctuations, contemporary marine environment factors and life-history traits of marine fish. Pacific herring and Pacific cod are two commercially important fish species in North Pacific. They were considered derived from an Atlantic invader by trans-Arctic dispersal. Large glacial-interglacial circles in Pleistocene and unique oceanographic characters of North Pacific may have influenced heavily the population structuring and phylogeographic patterns in the two species.
Morphological, mitochondrial DNA sequence and microsatellite DNA markers were used to estimate the population structure and phylogeographic pattern of Pacific herring in North Pacific, the divergence between Pacific herring and Atlantic herring was also discussed.
(1) Meristic and morphometric characters were compared by multivariate analysis for eight populations of total 424 individuals, significant differences were found in means of all these characters among eight populations, and a latitudinal cline was observed in several morphological characters.
(2) Three high distinct lineages were detected in 318 individuals from nine populations across the range of the species based on sequence variations of mtDNA control region, which might be isolated and diverged during glacial periods of Pleistocene. One lineage dominated the Northwestern Pacific, while the other two lineages occurred secondary contact in Eastern Pacific. The frequency difference of the three lineages in 9 populations defined two geographic groups, a Northwestern Pacific group and an Eastern Pacific Group.
(3) High significant genetic differentiations were found among 9 populations by scanning 9 microsatellite loci variations, which subdivided populations into three groups. This grouping picked up the Yellow Sea population as an independent group. Sea water temperature barriers from Kuroshio Current and Tsushima Current might restrict the gene flows between populations in Yellow Sea and other locations.
(4) MtDNA control region sequences were used to investigate temporal population structure of Pacific herring which were collected from three different periods in Yellow Sea. No significant genetic differentiations were detected among the three populations and the result suggested populations in Yellow Sea belong to a single panmictic stock.
(5) An Atlantic herring population was also analyzed as an outgroup to discuss the origin and evolution of Pacific herring. Meristic and morphometric characters were compared between the Atlantic herring population and a Pacific herring population from Yellow Sea. A discriminant formula was created with 11 character parameters, and the accuracy was 98.8%. MtDNA variations were analyzed using sequence data from cytochrome b and control region. Based on 8.1% net genetic distance in control region, we postulated the divergence in mtDNA between the two species occurred at about 1.25 million years ago. Three lineages of Pacific herring were considered to diverge during late Pleistocene. Mismatch distribution analyses indicated population expansion in both species. MtDNA sequence and microsatellite DNA markers were used to estimate the population structure and demographic history of Pacific cod in North Pacific.
(1) MtDNA control region were analyzed for 259 individuals from nine localities over the species'range. The result showed lack of genealogical structure and remarkably low control region diversity. Small but significant genetic differentiations were detected among Northwestern Pacific populations. Four populations from coastal North America exhibited a trend of isolation by distance.
(2) Small but significant genetic structure was also found in four populations of Northwestern Pacific based on microsatellite analyses. Like Pacific herring, large genetic differentiations between the Yellow Sea population and other populations might be also attributed to sea water temperature barriers. Contrary to mtDNA result, microsatellite analyses revealed relative high genetic diversity for Pacific cod.
We interpreted such discordance between mitochondrial and microsatellites might be caused by the four-fold higher rate of drift for mtDNA than nuclear genes and fairly high mutation rate for microsatellite DNA. Both mtDNA and microsatellite results suggested a recent bottleneck in Pacific cod.
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