中国斑腿蝗科部分种类分子系统学研究
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摘要
斑腿蝗科(Catantopidae)隶属于直翅目蝗亚目蝗总科(Orthoptera:Caelifera:Acridoidea),是蝗总科中最大的科。我国已知16亚科93属,分布于东洋区的居多,古北区种类较少。其中中华稻蝗、日本黄脊蝗、棉蝗、短星翅蝗等种类为农业生产上的重要害虫,对它们的系统分类学研究不仅具有理论意义,同时在蝗虫防治的农业生产实践中也具有较为重要的应用价值。近些年来从形态学,细胞学等方面对斑腿蝗科系统学研究具有一些报道,但目前国内外对斑腿蝗科昆虫较为全面的分子系统学研究还非常少。本研究以线粒体基因16SrDNA、Cytb及核基因28SrDNA为分子标记,对斑腿蝗科部分种类的系统发育关系进行了重建,为斑腿蝗科的系统学研究提供分子证据。
本研究采用PCR产物直接测序法测定了9亚科30种斑腿蝗及1种癞蝗(外群)的线粒体Cytb(708bp)(只有其中的27种)、16SrDNA(522bp左右)以及核基因28SrDNA(512bp左右)的部分片段,对三个基因序列片段的组成成分、替换饱和性、分类单元间碱基组成的偏向性以及系统发育信号进行了检验。对三个基因片段之间的相合性进行了分析。采用MP、NJ、ML和BI分别对Cytb、16SrDNA、28SrDNA及三基因片段组成的联合数据集进行建树并比较得出以下结论:
1.30种斑腿蝗科昆虫线粒体基因Cytb及16SrDNA具有昆虫线粒体所具有的高A+T含量,二基因片段分别为72.9%与68.7%;而核基因28SrDNA只有36%。替换饱和性分析结果表明16SrDNA的环区及Cytb的第三位点存在部分饱和现象。Cytb第三位点在各分类单元间存在碱基组成偏向性。PTP及树长分布检测表明,三数据组及子集、联合数据集都具有较强的系统发育信号,同时发现各数据集子集的gl值具有叠加性。联合数据集的gl也表现为各数据集之间的叠加结果,但其gl值却小于Cytb的gl值,分析认为这一现象是由于数据集之间的部分冲突所致。
2.ILD检验结果表明16SrDNA环区与Cytb第三位点间具有显著的不相合性,相关系数检验得出相反的结论,分析认为二者之间存在潜在的相合性,不相合原因是二者所含有的进化噪音引起的,故认为可以进行联合分析。
3.采用四种方法对三数据集及联合数集分别构建MP、NJ、ML、BI树,比较三数据集与联合数据集所建树,三数据集四种方法所建树与联合树都存在一定的差异,但都支持大多数的分支关系。
Catantopidae is the biggest family in Acridoidea (Orthptera: Caelifera). It includes 16 subfamilies, 93 genera in China, mostly distributed in Oriental realm, others in Paleoarctic realm. In this family, there are some important pests in agriculture, such as Oxya chinensis, Patanga japonica, Chondracris rosea rosea, Calliptamus abbreviatus. The systematics study of them has not only theoretical but also practical applications in grasshopper control and plant protection. Recently, some cytological, morphological systematic research works on Catantopidae have been reported. However, the molecular systematic research on Catantopidae insects is still very limited. In this study, we adopted 2 mitochondrial genes (16S and Cyt b) and nuclear gene 28S rDNA sequence as molecular markers, and reconstructed the phylogenetic relationships of 30 species of Catantopidae insects which belong to 9 subfamilies, hence providing molecular evidence for the systematic study for Catantopidae.Partial sequences of Cyt b, 16S rDNA and 28S rDNA of 30 species Catantopidae grasshoppers representing 9 subfamily and 1 outgroup species (belong to Pamphagidae) have been sequenced. The base composition, substitution saturation, base composition bias among taxa, phylogenetic signal have been tested. Incongruence between different data sets and their subdata sets has been analyzed. Reconstruction of phylogeny based on Cyt b, 16S rDNA , 28S rDNA and their combined data set using MP, NJ, ML and BI and with comparison of the trees inferred from four method draw conclusion as follows.1. It is found that there are high ratio of A+T in both of Cyt b and 16S rDNA in 30 species of Catantopidae, with 72.9% and 68.7% respectively, whereas content of A+T in 28S rDNA is only 36%. Substitution saturation analysis indicates that saturation occur in loop of 16S rDNA and the 3rd code position of Cyt b, and base bias on the 3rd position of Cyt b among taxa. Strong phylogenetic signal exist in 3 subdata sets and combined data set by PTP and g1 testing. It is found that there are additive of g1 in subdata sets. So does gl of the combined data set, with value less than that of Cyt b. The phenomenon may be caused by conflict among the subdata sets.2. ILD test shows significant incongruence between loop of 16S rDNA and the 3rd position of Cyt b, the correlative co-efficiency draw the reversal conclusion. Further analyses suggest that there is potential congruence between 16S rDNA and Cyt b,
incongruence caused by evolutionary noise. Thus, both of them can be used for combined analysis.3. MPT, NJT, MLT and BIT were constructed based on 3 data sets and combined data set. Comparing these trees, there are some difference between trees from each other, and that of combined data set.4. The combined unweighted tree was tested by PBS test. The branch supports for combined trees are 73.9% for 16S rDNA, 82.6% for 28S rDNA, and 86.9% for Cyt b respectively. The value of PBS is much higher than the support value of topology. The low of support value of the topology is caused by substitution saturation, base composition bias, compensating substitution.5. The MPT obtained form down weighted on the 3rd positions of Cyt b, loop and stem regions of 16S rDNA with 0 or 0.5 value, compared with unweihted MPT show that the noise of these subdata sets does not affect the topology of combined tree, and suggest combined trees is more robust.6. Based on combined data set, MP, NJ, ML and BI method were separately used to reconstruct the phylogenetic relationships of the 26 Catantopidae insects, and KH test based on likelihood method was used to compare the tree topologies produced by different methods. The result of KH test indicates that all trees based on combined data set have no significant difference in topologies, and each of them can be reasonable in the explanation of phylogenetic relationships among taxa. Based on the trees constructed from 4 methods, we can conclude:(1) The relationship of 9 subfamilies can be described as: ((Melanoplinae, Podisminae,)(((Caryandinae,Oxyinae),Coptocrinae,((CallipIaminae,Eyprepocnemidina), Cyrtacanthacridinae)), Catantopinae)).(2) Eyprepocnemidinae is a paraphyletic group and Podisminae is a polyphyletic group. We proposed to merge them into one subfamily.(3) Sinopodisma isn't a monophyletic group, and need further studies with more data.(4) Oxyinae is allied to Caryandinae and both subfamilies need to clarify whether they are monophyletic respectively.(5) The monophyly of Coptocrinae is not supported and Traualia should be moved out of Coptocrinae.(6) Traualia and Catantops form a clade. It is uncertain that these genera should be
本研究采用PCR产物直接测序法测定了9亚科30种斑腿蝗及1种癞蝗(外群)的线粒体Cytb(708bp)(只有其中的27种)、16SrDNA(522bp左右)以及核基因28SrDNA(512bp左右)的部分片段,对三个基因序列片段的组成成分、替换饱和性、分类单元间碱基组成的偏向性以及系统发育信号进行了检验。对三个基因片段之间的相合性进行了分析。采用MP、NJ、ML和BI分别对Cytb、16SrDNA、28SrDNA及三基因片段组成的联合数据集进行建树并比较得出以下结论:
1.30种斑腿蝗科昆虫线粒体基因Cytb及16SrDNA具有昆虫线粒体所具有的高A+T含量,二基因片段分别为72.9%与68.7%;而核基因28SrDNA只有36%。替换饱和性分析结果表明16SrDNA的环区及Cytb的第三位点存在部分饱和现象。Cytb第三位点在各分类单元间存在碱基组成偏向性。PTP及树长分布检测表明,三数据组及子集、联合数据集都具有较强的系统发育信号,同时发现各数据集子集的gl值具有叠加性。联合数据集的gl也表现为各数据集之间的叠加结果,但其gl值却小于Cytb的gl值,分析认为这一现象是由于数据集之间的部分冲突所致。
2.ILD检验结果表明16SrDNA环区与Cytb第三位点间具有显著的不相合性,相关系数检验得出相反的结论,分析认为二者之间存在潜在的相合性,不相合原因是二者所含有的进化噪音引起的,故认为可以进行联合分析。
3.采用四种方法对三数据集及联合数集分别构建MP、NJ、ML、BI树,比较三数据集与联合数据集所建树,三数据集四种方法所建树与联合树都存在一定的差异,但都支持大多数的分支关系。
Catantopidae is the biggest family in Acridoidea (Orthptera: Caelifera). It includes 16 subfamilies, 93 genera in China, mostly distributed in Oriental realm, others in Paleoarctic realm. In this family, there are some important pests in agriculture, such as Oxya chinensis, Patanga japonica, Chondracris rosea rosea, Calliptamus abbreviatus. The systematics study of them has not only theoretical but also practical applications in grasshopper control and plant protection. Recently, some cytological, morphological systematic research works on Catantopidae have been reported. However, the molecular systematic research on Catantopidae insects is still very limited. In this study, we adopted 2 mitochondrial genes (16S and Cyt b) and nuclear gene 28S rDNA sequence as molecular markers, and reconstructed the phylogenetic relationships of 30 species of Catantopidae insects which belong to 9 subfamilies, hence providing molecular evidence for the systematic study for Catantopidae.Partial sequences of Cyt b, 16S rDNA and 28S rDNA of 30 species Catantopidae grasshoppers representing 9 subfamily and 1 outgroup species (belong to Pamphagidae) have been sequenced. The base composition, substitution saturation, base composition bias among taxa, phylogenetic signal have been tested. Incongruence between different data sets and their subdata sets has been analyzed. Reconstruction of phylogeny based on Cyt b, 16S rDNA , 28S rDNA and their combined data set using MP, NJ, ML and BI and with comparison of the trees inferred from four method draw conclusion as follows.1. It is found that there are high ratio of A+T in both of Cyt b and 16S rDNA in 30 species of Catantopidae, with 72.9% and 68.7% respectively, whereas content of A+T in 28S rDNA is only 36%. Substitution saturation analysis indicates that saturation occur in loop of 16S rDNA and the 3rd code position of Cyt b, and base bias on the 3rd position of Cyt b among taxa. Strong phylogenetic signal exist in 3 subdata sets and combined data set by PTP and g1 testing. It is found that there are additive of g1 in subdata sets. So does gl of the combined data set, with value less than that of Cyt b. The phenomenon may be caused by conflict among the subdata sets.2. ILD test shows significant incongruence between loop of 16S rDNA and the 3rd position of Cyt b, the correlative co-efficiency draw the reversal conclusion. Further analyses suggest that there is potential congruence between 16S rDNA and Cyt b,
incongruence caused by evolutionary noise. Thus, both of them can be used for combined analysis.3. MPT, NJT, MLT and BIT were constructed based on 3 data sets and combined data set. Comparing these trees, there are some difference between trees from each other, and that of combined data set.4. The combined unweighted tree was tested by PBS test. The branch supports for combined trees are 73.9% for 16S rDNA, 82.6% for 28S rDNA, and 86.9% for Cyt b respectively. The value of PBS is much higher than the support value of topology. The low of support value of the topology is caused by substitution saturation, base composition bias, compensating substitution.5. The MPT obtained form down weighted on the 3rd positions of Cyt b, loop and stem regions of 16S rDNA with 0 or 0.5 value, compared with unweihted MPT show that the noise of these subdata sets does not affect the topology of combined tree, and suggest combined trees is more robust.6. Based on combined data set, MP, NJ, ML and BI method were separately used to reconstruct the phylogenetic relationships of the 26 Catantopidae insects, and KH test based on likelihood method was used to compare the tree topologies produced by different methods. The result of KH test indicates that all trees based on combined data set have no significant difference in topologies, and each of them can be reasonable in the explanation of phylogenetic relationships among taxa. Based on the trees constructed from 4 methods, we can conclude:(1) The relationship of 9 subfamilies can be described as: ((Melanoplinae, Podisminae,)(((Caryandinae,Oxyinae),Coptocrinae,((CallipIaminae,Eyprepocnemidina), Cyrtacanthacridinae)), Catantopinae)).(2) Eyprepocnemidinae is a paraphyletic group and Podisminae is a polyphyletic group. We proposed to merge them into one subfamily.(3) Sinopodisma isn't a monophyletic group, and need further studies with more data.(4) Oxyinae is allied to Caryandinae and both subfamilies need to clarify whether they are monophyletic respectively.(5) The monophyly of Coptocrinae is not supported and Traualia should be moved out of Coptocrinae.(6) Traualia and Catantops form a clade. It is uncertain that these genera should be
引文
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