缘蝽科昆虫线粒体COI基因的分子系统学研究（半翅目：异翅亚目）
摘要
缘蝽科隶属于半翅目Hemiptera，异翅亚目Heteroptera，缘蝽次目Infraorder Pentatomomorpha，缘蝽总科Coreoidea。属世界性分布，已知250属约1800种。我国已知约63属近200种，全部为植食性。其吸取寄主的营养器官和繁殖器官的汁液，为害作物。国内外已从分类学、分子系统学、生物地理学、生物防治等各个方面对缘蝽科进行了广泛的研究。在系统学方面，缘蝽科内族间、属种间的系统发育关系一直存在着较大的争论，目前已从外部形态特征、分子生物学特征、酯酶同工酶等方面进行过探讨。
线粒体编码细胞色素C氧化酶亚基I基因(COI)已成为当前动物分子系统学中应用最广泛的分子标记之一，特别是在昆虫中。本研究采用PCR产物直接测序法测定(13种)及从NCBI上下载(3种)共计缘蝽科11属16种昆虫的线粒体COI 726 bp的序列，借助系统发育分析软件，采用邻接法(NJ)、最小进化法(ME)、最大简约法(MP)、最大似然法(ML)及贝叶斯推论法(BI)五种方法探讨了缘蝽科部分种类的系统发育关系。通过分析讨论，得出如下结论：
1．缘蝽科16种昆虫COI基因T、C、A、G的平均含量分别为37.0％、14.9％、33.4％和14.8％。A+T平均含量达到70.4％，G+C平均含量达到29.6％，A+T的含量明显高于G+C的含量，表现出非常强的A+T含量偏向性，同多数昆虫的核苷酸频率相一致。缘蝽科16种昆虫COI基因726 bp序列共编码242个氨基酸，由20种氨基酸组成，其中丝氨酸(Ser)、异亮氨酸(Ile)含量最高，反映出COI基因在氨基酸组成上具有一定偏向性。
2．缘蝽科16种昆虫COI转换频率和颠换相同；转换主要发生在T与C之间，颠换主要发生在A与T之间。密码子第二位点的转换与颠换数目都比较小，占转换与颠换总数的5.4％和1.9％；密码子第三位的转换和颠换的频率都较高，转换频率占总数的71.4％，颠换占总数的80.4％。
3．从GenBank上下载红蝽总科的Physopelta quadriguttata和猎蝽总科的Triatoma maculate作为外群，建立缘蝽科11属16种间的系统发育关系。结果显示狭义巨缘蝽族两属为一个单系群，竹缘蝽族的两个属为一单系群，缘蝽属与棘缘蝽属亲缘关系非常近。
4．五种建树方法显示出了几个问题：同缘蝽族是否为一单系群，普缘蝽属为何与同缘蝽族聚在一起；瘤缘蝽属与怪缘蝽属是否应为一单系群并同归于菲缘蝽族；怪缘蝽属与黑缘蝽属(沟缘蝽亚科)关系似乎更为接近。
5．运用COI基因的核苷酸序列对缘蝽科昆虫进行分子系统研究时，各属种间的分类关系都比较明确，得到的树的拓扑结构大都一致。因此，COI基因在分析种间和属间的系统发育关系时，是一个较为有效的分子标记。
Coreidae are a big family of Heteroptera. There are 250 genera, 1800species known in the world, and about 63 genera, 200 species known in China. Coreidaewhich is a kind of phytophagous insects and important pests of agriculture and foresthave been studied widely in many fields such as taxonomy, molecular phylogeny,biogeography, etc. However, there are still lots of debates on the phylogeneticrelationships among tribes and genera levels. Phylogenetic relationships of Coreidaehave been studied by external morphological characters, molecular characters,Isoenzymes, etc.
The mitochondrial cytochrome oxidase subunit I (COI) gene is one of the mostwidely used genetic marker for molecular systematics of animals, especially in insects.In this research, the COI gene sequences were amplified by PCR, and then sequenceddirectly through cycling sequencing method. 726 bp sequences of 16 species ofCoreidae were obtained. Based on the analyses of COI gene sequences, wereconstructed the phylogenetic relationships of these species by using Neighbor-Joining(NJ), Minimum Evolution (ME), Maximum Parsimony (MP), Maximum Likelihood(ML) and Bayesian Inference (BI) methods. The conclusions are showed as follows:
1) The average contents of T, C, A andG to COI gene of 16 species in Coreidae are37.0%, 14.9%, 33.4% and 14.8%, respectively, A+T content is 70.4%, while G+Ccontent is 29.6%, which shows a strong A+T bias. These sequences code 242 aminoacids and contain 20 kinds of amino acids, respectively. The contents of Set and Ile arethe highest.
2) To these 16 species of Coreidae, ratio of transition to transversion (R) of COIgene sequence is 1; Transitions are the same with transversions; most transitions occurbetween T and C, while most transversions occur between A and T. Substitution of thesecond position of codons are the lowest: transition are 5.4% of all transition, whiletransversion are 1.9%. And Substitutions of the third position of codons are the highest:transitions are 71.4%, while transversions are 80.4%.
3) The phylogenetic relationships among 16 species of Coreidae and outgroups which include Physopelta quadriguttata (Hemiptera: Pyrrhocoroidea) and Triatomamaculate (Hemiptera: Reduviidae) had been reconstructed. The results show that: towspecies of the Mictini s. str is a monophyletic group; two genera of Cloresmini is amonophyletic group; Coreus has a high relative relationship with Cletus.
4) The five tree-building methods provide some questions: Homoeocerini is amonophyletic group or not? And why is the Plinachtus clustered with theHomoeocerini? Acanthocoris and Cordysceles belong to a monophyletic group or not?And are they belong to Physomerini? The relative relationships between Cordyscelesand Hygia seem to be high.
5) The research on phylogenetic tree shows that COI gene is an efficient symbol instudy of the phylogenetic evolution of interspecies and intragenus, so it is a usefulmarker.
线粒体编码细胞色素C氧化酶亚基I基因(COI)已成为当前动物分子系统学中应用最广泛的分子标记之一，特别是在昆虫中。本研究采用PCR产物直接测序法测定(13种)及从NCBI上下载(3种)共计缘蝽科11属16种昆虫的线粒体COI 726 bp的序列，借助系统发育分析软件，采用邻接法(NJ)、最小进化法(ME)、最大简约法(MP)、最大似然法(ML)及贝叶斯推论法(BI)五种方法探讨了缘蝽科部分种类的系统发育关系。通过分析讨论，得出如下结论：
1．缘蝽科16种昆虫COI基因T、C、A、G的平均含量分别为37.0％、14.9％、33.4％和14.8％。A+T平均含量达到70.4％，G+C平均含量达到29.6％，A+T的含量明显高于G+C的含量，表现出非常强的A+T含量偏向性，同多数昆虫的核苷酸频率相一致。缘蝽科16种昆虫COI基因726 bp序列共编码242个氨基酸，由20种氨基酸组成，其中丝氨酸(Ser)、异亮氨酸(Ile)含量最高，反映出COI基因在氨基酸组成上具有一定偏向性。
2．缘蝽科16种昆虫COI转换频率和颠换相同；转换主要发生在T与C之间，颠换主要发生在A与T之间。密码子第二位点的转换与颠换数目都比较小，占转换与颠换总数的5.4％和1.9％；密码子第三位的转换和颠换的频率都较高，转换频率占总数的71.4％，颠换占总数的80.4％。
3．从GenBank上下载红蝽总科的Physopelta quadriguttata和猎蝽总科的Triatoma maculate作为外群，建立缘蝽科11属16种间的系统发育关系。结果显示狭义巨缘蝽族两属为一个单系群，竹缘蝽族的两个属为一单系群，缘蝽属与棘缘蝽属亲缘关系非常近。
4．五种建树方法显示出了几个问题：同缘蝽族是否为一单系群，普缘蝽属为何与同缘蝽族聚在一起；瘤缘蝽属与怪缘蝽属是否应为一单系群并同归于菲缘蝽族；怪缘蝽属与黑缘蝽属(沟缘蝽亚科)关系似乎更为接近。
5．运用COI基因的核苷酸序列对缘蝽科昆虫进行分子系统研究时，各属种间的分类关系都比较明确，得到的树的拓扑结构大都一致。因此，COI基因在分析种间和属间的系统发育关系时，是一个较为有效的分子标记。
Coreidae are a big family of Heteroptera. There are 250 genera, 1800species known in the world, and about 63 genera, 200 species known in China. Coreidaewhich is a kind of phytophagous insects and important pests of agriculture and foresthave been studied widely in many fields such as taxonomy, molecular phylogeny,biogeography, etc. However, there are still lots of debates on the phylogeneticrelationships among tribes and genera levels. Phylogenetic relationships of Coreidaehave been studied by external morphological characters, molecular characters,Isoenzymes, etc.
The mitochondrial cytochrome oxidase subunit I (COI) gene is one of the mostwidely used genetic marker for molecular systematics of animals, especially in insects.In this research, the COI gene sequences were amplified by PCR, and then sequenceddirectly through cycling sequencing method. 726 bp sequences of 16 species ofCoreidae were obtained. Based on the analyses of COI gene sequences, wereconstructed the phylogenetic relationships of these species by using Neighbor-Joining(NJ), Minimum Evolution (ME), Maximum Parsimony (MP), Maximum Likelihood(ML) and Bayesian Inference (BI) methods. The conclusions are showed as follows:
1) The average contents of T, C, A andG to COI gene of 16 species in Coreidae are37.0%, 14.9%, 33.4% and 14.8%, respectively, A+T content is 70.4%, while G+Ccontent is 29.6%, which shows a strong A+T bias. These sequences code 242 aminoacids and contain 20 kinds of amino acids, respectively. The contents of Set and Ile arethe highest.
2) To these 16 species of Coreidae, ratio of transition to transversion (R) of COIgene sequence is 1; Transitions are the same with transversions; most transitions occurbetween T and C, while most transversions occur between A and T. Substitution of thesecond position of codons are the lowest: transition are 5.4% of all transition, whiletransversion are 1.9%. And Substitutions of the third position of codons are the highest:transitions are 71.4%, while transversions are 80.4%.
3) The phylogenetic relationships among 16 species of Coreidae and outgroups which include Physopelta quadriguttata (Hemiptera: Pyrrhocoroidea) and Triatomamaculate (Hemiptera: Reduviidae) had been reconstructed. The results show that: towspecies of the Mictini s. str is a monophyletic group; two genera of Cloresmini is amonophyletic group; Coreus has a high relative relationship with Cletus.
4) The five tree-building methods provide some questions: Homoeocerini is amonophyletic group or not? And why is the Plinachtus clustered with theHomoeocerini? Acanthocoris and Cordysceles belong to a monophyletic group or not?And are they belong to Physomerini? The relative relationships between Cordyscelesand Hygia seem to be high.
5) The research on phylogenetic tree shows that COI gene is an efficient symbol instudy of the phylogenetic evolution of interspecies and intragenus, so it is a usefulmarker.
引文
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