基于AFLP标记技术的几种海洋经济鱼类分子系统地理学研究
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摘要
本研究采用AFLP标记技术进行花鲈、刀鲚、梭鱼、棘头梅童鱼、大泷六线鱼、太平洋鲱鱼和斑头鱼七种海洋鱼类的群体遗传结构和遗传多样性研究，并探讨了花鲈和鲈鱼、棘头梅童鱼和黑鳃梅童鱼种间分化，得出了如下结论：
（1）对采自中国沿海的花鲈3个群体和日本沿海的鲈鱼4个群体进行了AFLP分析，结果表明日本有明海鲈鱼群体的Nei遗传多样性指数和Shannon多样性指数略高于其他6个群体的遗传多样性指数，且都处于中等程度偏下水平；UPGMA树显示来自中国的3个花鲈群体和来自日本的4个鲈鱼群体分别聚类；花鲈两两群体的F_(st)值（0.084-0.140）和鲈鱼两两群体之间的F_(st)值（0.045-0.153）明显低于花鲈群体和鲈鱼群体之间的F_(st)值（0.283-0.556）；花鲈群体之间和鲈鱼群体之间的基因流均大于1，而花鲈与鲈鱼群体之间的基因流均小于1。因此，本研究的结果也支持中日鲈鱼属于不同物种这一结论。
（2）为进一步分析中国沿海与日本沿海的刀鲚群体间遗传分化水平，本研究采用AFLP标记技术对中日刀鲚群体的遗传多样性和群体遗传结构进行了研究。UPGMA树结果显示日本两个群体的所有个体在系统树上形成了一个小的分支，但是整体上仍然与中国群体形成了一个大支；中国群体之间的F_(st)值以及日本群体之间的F_(st)值显著小于中日群体之间的F_(st)值，中日刀鲚群体之间存在较为显著的遗传分化，但刀鲚中日种群间的遗传分化指数远小于刀鲚与凤鲚之间的遗传分化指数；研究结果显示中日刀鲚群体间的基因交流可能十分有限。
（3）对来自中日沿海的8个地理群体的91尾梭鱼样品进行了AFLP分析，从核基因角度探讨梭鱼不同地理群体的遗传变异，结果显示梭鱼群体的多态位点比例仅为22.73%-38.46%，相对较低；UPGMA聚类树显示梭鱼8个群体分为两个不同的地理组群，南方群体和北方群体，南方群体包括珠江和云林群体，北方群体包括其余的6个群体；两两群体之间的遗传分化指数F_(st)也显示南方群体与北方群体存在较大的遗传分化，暗示其分布范围内的南海海区群体与北方群体（包括东海、黄渤海和日本沿海）存在较大的遗传分化；研究结果认为梭鱼线粒体所揭示的3个支系不是代表了3个隐蔽种，而南北两个群体可能代表两个隐蔽种。
（4）石首鱼科是自上新世至今十分繁盛的海洋鱼类，广泛分布于热带、亚热带的近海水域。本研究以黑鳃梅童鱼为外群，对采自中国沿海的棘头梅童鱼5个地理群体进行了AFLP分析，同时探讨两种梅童鱼种间遗传差异，研究结果显示，棘头梅童鱼60个个体中共扩增出458条清晰的条带，多态位点比例为95.85%；两种梅童鱼UPGMA聚类树的结果表明两种梅童鱼明显分为两个支，在棘头梅童鱼5个群体中，舟山和上海聚为一支，温州和厦门聚为一支，最后与深圳聚为一支；棘头梅童鱼5个群体之间的遗传分化指数均较大，从0.1648（舟山和上海）到0.3074（深圳和上海）；5个群体间的基因流系数均远小于1，表明各个群体间基因流受阻；研究结果支持棘头梅童鱼与黑鳃梅童鱼为2种不同的梅童鱼的观点，并认为棘头梅童鱼不同地理群体之间的基因交流非常有限。
（5）针对线粒体DNA和微卫星研究结果的差异，采用AFLP标记技术对采自中国沿海的5个地理群体以及日本沿海1个地理群体的大泷六线鱼进行补充研究。结果显示日本青森群体的Nei遗传多样性指数和Shannon遗传多样性指数最高，丹东群体最低；UPGMA聚类树的结果显示大泷六线鱼的拓扑结构比较简单，虽然存在两个分支，但是没有检测到与采样地点相对应的分支；两两群体的Fst值较小，仅为0.016-0.122；基于Fst值计算两两群体之间的基因流，结果显示每组基因流都大于1，并且有5组群体之间的基因流都大于4；AFLP标记技术的研究结果支持大泷六线鱼是一个随机交配的群体。
（6）采用AFLP标记技术对采自整个分布范围的太平洋鲱鱼开展了分子系统地理学研究。8个群体的遗传多样性指数都比较相近，并无明显差异；UPGMA树结果显示所有个体分为两大支，其中一支由希托克岛的个体组成，另一支由其他七个群体组成，除鄂霍次克海群体存在一个小的分支外，不同的地理群体间无明显的分支，存在一定程度的交叉；托克岛群体与其他群体的遗传分化指数Fst较大，西勘察加与石狩湾群体之间的遗传分化指数最小；两两群体之间的基因流的计算结果显示希托克岛群体与其他群体的基因流均小于1，而其他两两群体之间的基因流均大于1。本研究结果支持其他学者的分支A起源于西北太平洋某处避难所、而分支B应该位于东太平洋避难所的结论。
（7）对采自中国青岛和威海俚岛近海的两个斑头鱼群体进行了AFLP比较分析。结果显示两群体的遗传多样性指数水平都比较相近，并无明显差异；UPGMA树上存在两个分支，但是两个群体的个体相互混杂，并没有与采样地点相对应的分支的存在；两群体之间的F_(st)值为0.056，且统计检验结果显著；两群体之间的基因流大于4。AFLP标记技术的研究结果认为斑头鱼与大泷六线鱼具有十分相似的遗传多样性水平和遗传结构差异，并支持斑头鱼是一个随机交配的群体。
In the present study, AFLP molecular marker was used to estimate thepopulation genetic diversity and genetic structure of Lateolabrax maculates, Coilianasus, Liza haematocheilus, Collichthys lucidus, Hexagrammos otakii, Clupeapallasii and H. agrammus. Phylogenetic relationships between L. maculates and L.japonicus, C. lucidus and C. niveatus were also discussed. The results of the presentstudy were as follow:
(1) To examine population genetic structure and phylogenetic relationship of L.maculates and L. japonicus, samples of L. maculates from Chinese Coast andsamples of L. japonicus from Japanese Coast were collected and analyzed by AFLP.The results showed that Nei gene diversity and Shannon’s information index ofAriake Sea were higher than the other6populations. Two clades were detected byUPGMA tree were, which were Chinese clade and Japanese clade, respectively.Pairwise F_(st)between L. maculates and L. japonicus was significantly higher thanthat between populations of L. maculates and between populations of L. japonicus.Gene flow values between populations of L. maculates and between populations of L.maculates were higher than1, while gene flow between these two species was lowerthan1. The results of present study supported that the L. maculates and L. japonicuswere different speices.
(2) AFLP marker was used to examine the genetic diversity and geneticstructure of Coilia nasus and to analyze the genetic differentiation between Chineseand Japanese populations. The results showed that all individuals from Japanesecoast were clustered into one clade, but this clade was still surrounded by Chineseindividuals on the whole and no obvious differentiation was detected by UPGMA.Pairwise F_(st)values between Chinese populations and Japanese populations were higher than those between Chinese populations and between Japanese populations,but significantly lower than that between C. nasus and C. mystus. The results ofpresent study indicated limited gene flow between Chinese populations and Japanesepopulations, but did not supported they were different species.
(3) Eight populations from Chinese and Japanese coast were analyzed by AFLP.The results showed that proportion of polymorphic loci were very low, only22.73%-38.46%. These8populations were clustered into two clades, namely,southern group including populations from Zhujiang and Yunlin and northern groupincluding the other6populations. AMOVA and pairwise F_(st)also supported theseparation of north and south populations of L. haematocheilus. The present studydemonstrated that L. haematocheilus was indeed composed of at least twogenetically divergent species, but not three cryptic species in the NorthwesternPacific.
(4) In the present study, the genetic diversity and population genetic structure ofC. lucidus were examined by AFLP marker. C. niveatus was collected as theoutgroup to discuss the differentiation between these two species. A total of458bands were identified from60individuals and the percentage of polymorphic bandswas95.85%. These two species clustered into two clades. Among the5populationsfor C. lucidus, populations from Zhoushan and Shanghai clustered into one clade,populations from Wenzhou and Xiamen clustered into another clade, and these twoclades subsequently clustered with Shenzhen population. The pairwise F_(st)rangedfrom0.1648(between populations from Zhoushan and Shanghai) to0.3074(betweenpopulations from Shenzhen and Shanghai). Gene flow values among differentpopulations were lower than1. The results of present study indicated the limitedgene flow among different geographic populations, in conformity with the point thatC. lucidus and C. niveatus were different species.
(5) Six geographic populations of H. otakii from Chinese and Japanese coastawere further analyzed to estimate the population genetic structure, due to thedifference of results between morphological and microsatellite DNA analysis. Theresults showed that the genetic diversity of Hachinohe population was the highest and the lowest occurred in Dandong population. Two clades were detected byUPGMA tree, but no significant genealogical branches or clusters corresponding tosampling localities were detected. Pairwise F_(st)values (0.016-0.122) showed nogenetic divergence among different geographic populations and gene flow valueswere high among5populations. The results of the present study supposed that H.otakii was a random mating population.
(6) AFLP marker was used to estimate the population structure andphylogeographic pattern of Clupea pallasii in North Pacific. The genetic diversityindices of8populations were similar to each other. Two branches were detected, oneof which consisted of all individuals from Sitka Island and another consisted of theother7populations. All individuals of Okhotsk Sea formed into one small clade andno significant genealogical branches or clusters corresponding to sampling localitieswere detected. The pairwise F_(st)values between Sitka Island and other populationswere higher and the lowest differentiation was observed between populations ofWest Kamchatka and Ishikari Bay. Gene flow values between Sitka Island and otherpopulations were less than1, contrary to the other combinations. The results ofpresent study supported the opinion that clade A should originate from some refugeof the Northwestern Pacific and clade B should originate from some refuge of theEast Pacific.
(7) Two populations of H. agrammus from Qingdao and Lidao of Weihai coastwere analyzed by AFLP marker. The results showed that genetic diversity of the twopopulations was similar to each other. Two branches were detected by UPGMA treewith no significant genealogical branches or clusters corresponded to samplinglocalities. The pairwise F_(st)value between the two populations was0.056and geneflow was more than4. The results of the present study indicated that H. agrammuswas a random mating population and showed the similar genetic diversity with H.otakii.
（1）对采自中国沿海的花鲈3个群体和日本沿海的鲈鱼4个群体进行了AFLP分析，结果表明日本有明海鲈鱼群体的Nei遗传多样性指数和Shannon多样性指数略高于其他6个群体的遗传多样性指数，且都处于中等程度偏下水平；UPGMA树显示来自中国的3个花鲈群体和来自日本的4个鲈鱼群体分别聚类；花鲈两两群体的F_(st)值（0.084-0.140）和鲈鱼两两群体之间的F_(st)值（0.045-0.153）明显低于花鲈群体和鲈鱼群体之间的F_(st)值（0.283-0.556）；花鲈群体之间和鲈鱼群体之间的基因流均大于1，而花鲈与鲈鱼群体之间的基因流均小于1。因此，本研究的结果也支持中日鲈鱼属于不同物种这一结论。
（2）为进一步分析中国沿海与日本沿海的刀鲚群体间遗传分化水平，本研究采用AFLP标记技术对中日刀鲚群体的遗传多样性和群体遗传结构进行了研究。UPGMA树结果显示日本两个群体的所有个体在系统树上形成了一个小的分支，但是整体上仍然与中国群体形成了一个大支；中国群体之间的F_(st)值以及日本群体之间的F_(st)值显著小于中日群体之间的F_(st)值，中日刀鲚群体之间存在较为显著的遗传分化，但刀鲚中日种群间的遗传分化指数远小于刀鲚与凤鲚之间的遗传分化指数；研究结果显示中日刀鲚群体间的基因交流可能十分有限。
（3）对来自中日沿海的8个地理群体的91尾梭鱼样品进行了AFLP分析，从核基因角度探讨梭鱼不同地理群体的遗传变异，结果显示梭鱼群体的多态位点比例仅为22.73%-38.46%，相对较低；UPGMA聚类树显示梭鱼8个群体分为两个不同的地理组群，南方群体和北方群体，南方群体包括珠江和云林群体，北方群体包括其余的6个群体；两两群体之间的遗传分化指数F_(st)也显示南方群体与北方群体存在较大的遗传分化，暗示其分布范围内的南海海区群体与北方群体（包括东海、黄渤海和日本沿海）存在较大的遗传分化；研究结果认为梭鱼线粒体所揭示的3个支系不是代表了3个隐蔽种，而南北两个群体可能代表两个隐蔽种。
（4）石首鱼科是自上新世至今十分繁盛的海洋鱼类，广泛分布于热带、亚热带的近海水域。本研究以黑鳃梅童鱼为外群，对采自中国沿海的棘头梅童鱼5个地理群体进行了AFLP分析，同时探讨两种梅童鱼种间遗传差异，研究结果显示，棘头梅童鱼60个个体中共扩增出458条清晰的条带，多态位点比例为95.85%；两种梅童鱼UPGMA聚类树的结果表明两种梅童鱼明显分为两个支，在棘头梅童鱼5个群体中，舟山和上海聚为一支，温州和厦门聚为一支，最后与深圳聚为一支；棘头梅童鱼5个群体之间的遗传分化指数均较大，从0.1648（舟山和上海）到0.3074（深圳和上海）；5个群体间的基因流系数均远小于1，表明各个群体间基因流受阻；研究结果支持棘头梅童鱼与黑鳃梅童鱼为2种不同的梅童鱼的观点，并认为棘头梅童鱼不同地理群体之间的基因交流非常有限。
（5）针对线粒体DNA和微卫星研究结果的差异，采用AFLP标记技术对采自中国沿海的5个地理群体以及日本沿海1个地理群体的大泷六线鱼进行补充研究。结果显示日本青森群体的Nei遗传多样性指数和Shannon遗传多样性指数最高，丹东群体最低；UPGMA聚类树的结果显示大泷六线鱼的拓扑结构比较简单，虽然存在两个分支，但是没有检测到与采样地点相对应的分支；两两群体的Fst值较小，仅为0.016-0.122；基于Fst值计算两两群体之间的基因流，结果显示每组基因流都大于1，并且有5组群体之间的基因流都大于4；AFLP标记技术的研究结果支持大泷六线鱼是一个随机交配的群体。
（6）采用AFLP标记技术对采自整个分布范围的太平洋鲱鱼开展了分子系统地理学研究。8个群体的遗传多样性指数都比较相近，并无明显差异；UPGMA树结果显示所有个体分为两大支，其中一支由希托克岛的个体组成，另一支由其他七个群体组成，除鄂霍次克海群体存在一个小的分支外，不同的地理群体间无明显的分支，存在一定程度的交叉；托克岛群体与其他群体的遗传分化指数Fst较大，西勘察加与石狩湾群体之间的遗传分化指数最小；两两群体之间的基因流的计算结果显示希托克岛群体与其他群体的基因流均小于1，而其他两两群体之间的基因流均大于1。本研究结果支持其他学者的分支A起源于西北太平洋某处避难所、而分支B应该位于东太平洋避难所的结论。
（7）对采自中国青岛和威海俚岛近海的两个斑头鱼群体进行了AFLP比较分析。结果显示两群体的遗传多样性指数水平都比较相近，并无明显差异；UPGMA树上存在两个分支，但是两个群体的个体相互混杂，并没有与采样地点相对应的分支的存在；两群体之间的F_(st)值为0.056，且统计检验结果显著；两群体之间的基因流大于4。AFLP标记技术的研究结果认为斑头鱼与大泷六线鱼具有十分相似的遗传多样性水平和遗传结构差异，并支持斑头鱼是一个随机交配的群体。
In the present study, AFLP molecular marker was used to estimate thepopulation genetic diversity and genetic structure of Lateolabrax maculates, Coilianasus, Liza haematocheilus, Collichthys lucidus, Hexagrammos otakii, Clupeapallasii and H. agrammus. Phylogenetic relationships between L. maculates and L.japonicus, C. lucidus and C. niveatus were also discussed. The results of the presentstudy were as follow:
(1) To examine population genetic structure and phylogenetic relationship of L.maculates and L. japonicus, samples of L. maculates from Chinese Coast andsamples of L. japonicus from Japanese Coast were collected and analyzed by AFLP.The results showed that Nei gene diversity and Shannon’s information index ofAriake Sea were higher than the other6populations. Two clades were detected byUPGMA tree were, which were Chinese clade and Japanese clade, respectively.Pairwise F_(st)between L. maculates and L. japonicus was significantly higher thanthat between populations of L. maculates and between populations of L. japonicus.Gene flow values between populations of L. maculates and between populations of L.maculates were higher than1, while gene flow between these two species was lowerthan1. The results of present study supported that the L. maculates and L. japonicuswere different speices.
(2) AFLP marker was used to examine the genetic diversity and geneticstructure of Coilia nasus and to analyze the genetic differentiation between Chineseand Japanese populations. The results showed that all individuals from Japanesecoast were clustered into one clade, but this clade was still surrounded by Chineseindividuals on the whole and no obvious differentiation was detected by UPGMA.Pairwise F_(st)values between Chinese populations and Japanese populations were higher than those between Chinese populations and between Japanese populations,but significantly lower than that between C. nasus and C. mystus. The results ofpresent study indicated limited gene flow between Chinese populations and Japanesepopulations, but did not supported they were different species.
(3) Eight populations from Chinese and Japanese coast were analyzed by AFLP.The results showed that proportion of polymorphic loci were very low, only22.73%-38.46%. These8populations were clustered into two clades, namely,southern group including populations from Zhujiang and Yunlin and northern groupincluding the other6populations. AMOVA and pairwise F_(st)also supported theseparation of north and south populations of L. haematocheilus. The present studydemonstrated that L. haematocheilus was indeed composed of at least twogenetically divergent species, but not three cryptic species in the NorthwesternPacific.
(4) In the present study, the genetic diversity and population genetic structure ofC. lucidus were examined by AFLP marker. C. niveatus was collected as theoutgroup to discuss the differentiation between these two species. A total of458bands were identified from60individuals and the percentage of polymorphic bandswas95.85%. These two species clustered into two clades. Among the5populationsfor C. lucidus, populations from Zhoushan and Shanghai clustered into one clade,populations from Wenzhou and Xiamen clustered into another clade, and these twoclades subsequently clustered with Shenzhen population. The pairwise F_(st)rangedfrom0.1648(between populations from Zhoushan and Shanghai) to0.3074(betweenpopulations from Shenzhen and Shanghai). Gene flow values among differentpopulations were lower than1. The results of present study indicated the limitedgene flow among different geographic populations, in conformity with the point thatC. lucidus and C. niveatus were different species.
(5) Six geographic populations of H. otakii from Chinese and Japanese coastawere further analyzed to estimate the population genetic structure, due to thedifference of results between morphological and microsatellite DNA analysis. Theresults showed that the genetic diversity of Hachinohe population was the highest and the lowest occurred in Dandong population. Two clades were detected byUPGMA tree, but no significant genealogical branches or clusters corresponding tosampling localities were detected. Pairwise F_(st)values (0.016-0.122) showed nogenetic divergence among different geographic populations and gene flow valueswere high among5populations. The results of the present study supposed that H.otakii was a random mating population.
(6) AFLP marker was used to estimate the population structure andphylogeographic pattern of Clupea pallasii in North Pacific. The genetic diversityindices of8populations were similar to each other. Two branches were detected, oneof which consisted of all individuals from Sitka Island and another consisted of theother7populations. All individuals of Okhotsk Sea formed into one small clade andno significant genealogical branches or clusters corresponding to sampling localitieswere detected. The pairwise F_(st)values between Sitka Island and other populationswere higher and the lowest differentiation was observed between populations ofWest Kamchatka and Ishikari Bay. Gene flow values between Sitka Island and otherpopulations were less than1, contrary to the other combinations. The results ofpresent study supported the opinion that clade A should originate from some refugeof the Northwestern Pacific and clade B should originate from some refuge of theEast Pacific.
(7) Two populations of H. agrammus from Qingdao and Lidao of Weihai coastwere analyzed by AFLP marker. The results showed that genetic diversity of the twopopulations was similar to each other. Two branches were detected by UPGMA treewith no significant genealogical branches or clusters corresponded to samplinglocalities. The pairwise F_(st)value between the two populations was0.056and geneflow was more than4. The results of the present study indicated that H. agrammuswas a random mating population and showed the similar genetic diversity with H.otakii.
引文
陈大刚．黄渤海渔业生态学．北京：海洋出版社，1991．
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