栎瘿蜂族和客瘿蜂族部分瘿蜂共生菌Wolbachia的感染、传播途径及其MLST分析
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摘要
Wolbachia是一类细胞质共生菌,感染了20-76%的昆虫,是丰度最高、分布最广的共生细菌。Wolbachia能调控宿主的生殖方式,如诱导胞质不亲和、诱导孤雌生殖、雄性个体的雌性化和杀雄。Wolbachia传播方式主要是垂直传播,但也存在水平传播现象。瘿蜂亚科(Cynipinae)隶属于膜翅目的瘿蜂科(Cynipidae),已知约1400多种,绝大多数为植食性种类。我国(包括台湾省)已记录的瘿蜂种类共11属,29种。
本研究主要在中国南方地区采集瘿蜂标本,应用PCR法,检测获得的瘿蜂种类的Wolbachia感染；基于wsp基因序列及多位点序列分型(Multi-locus Sequence Typing, MLST)分析瘿蜂Wolbachia的系统发育关系；探讨瘿蜂Wolbachia的传播途径及其对宿主生殖模式的影响。
主要研究结果如下：
(1)从河南林州,安徽安庆,湖南长沙、岳阳、娄底、常德、邵阳、益阳,江西吉安,广东韶关和福建宁德等地共采集瘿蜂虫瘿14种,其寄主植物分别隶属于壳斗科(Fagaceae)的栗属Castanea、栎属Quercus、石栎属Lithocarpus、栲属Castanopsis和蔷薇科的蔷薇属Rosa。获得瘿蜂羽化成虫共15种。其中,新种10种(含待发表新种9种),分别为锥栗瘿蜂Dryocosmus zhuili Liu, ms、白球瘿蜂Cynips crystaloides Liu,ms、光顶并腹客瘿蜂Synergus intricatus Liu, ms、尖盾阔头瘿蜂Trigonaspis acuminatus Liu, ms、苔形瘿安瘿蜂Andricus elodeoides Liu, ms、河南并腹客瘿蜂Synergus henanensis Liu, ms、脊顶并腹客瘿蜂Synergus carinatus Liu, ms、平滑客瘿蜂Saphonecrus glaberus Liu,ms、重纹并腹客瘿蜂Synergus brutus Liu, ms和壶瓶山客瘿蜂Saphonecrus hupingshanensis Liu;中国新纪录种3种,分别为向川安瘿蜂Andricus mukaigawae、日本并腹客瘿蜂Synergus japonicus和日本蔷薇瘿蜂Diplolepis japonica;中国已记录种2种,分别是麦氏安瘿蜂Andricus mairei和栗瘿蜂Dryocosmus kuriphilus。
(2)对3种群麦氏安瘿蜂的Wolbachia感染进行了PCR检测,发现岳阳、长沙和邵阳种群的麦氏安瘿蜂雌、雄成虫均存在Wolbachia的感染,除邵阳雌成虫Wolbachia的感染率为80%之外,其它均为100%,且3种群所测得Wolbachia的wsp基因序列完全一致。岳阳、长沙及邵阳种群麦氏安瘿蜂均获得了雌、雄成虫,雌性率分别为15.3%、12.1%和19.8%,雄性比显著偏高,因此,可以认为Wolbachia的共生并未诱导其营产雌孤雌生殖。
(3)采集了8地理种群的向川安瘿蜂虫瘿,并获得了造瘿瘿蜂(向川安瘿蜂)、客居瘿蜂(日本并腹客瘿蜂)及寄生性长尾小蜂Torymus sp的羽化成虫。三者均具Wolbachia的感染,感染率分别为60%、37.1%和100%。除寄生蜂Torymussp.仅感染1个Wolbachia株系之外,向川安瘿蜂和日本并腹客瘿蜂分别感染了5个和8个株系,证实了这2种瘿蜂均存在Wolbachia高水平的多重感染。检测到向川安瘿蜂群落的3组重组株系,发现其Wolbachia的wsp基因存在明显的遗传重组现象。在向川安瘿蜂群落中的9个Wolbachia株系中,有5个株系为向川安瘿蜂和日本并腹客瘿蜂的共有株系,说明造瘿瘿蜂和客居瘿蜂之间存在Wolbachia的水平传播现象。可以推测,向川安瘿蜂和日本并腹客瘿蜂Wolbachia高水平多重感染的原因有二,即基因的遗传重组现象和水平传播。
(4)对获得的15种瘿蜂进行了Wolbachia感染的PCR检测,发现8种瘿蜂具Wolbachia感染,分别是向川安瘿蜂、麦氏安瘿蜂、锥栗瘿蜂、白球瘿蜂、日本并腹客瘿蜂、光顶并腹客瘿蜂、平滑客瘿蜂和重纹并腹客瘿蜂,感染的种类达53.3%。其中,麦氏安瘿蜂的感染率最高,接近100%；重纹并腹客瘿蜂的感染率最低,为26.7%。
(5)应用gatB、coxA、hcpA、ftsZ和fbpA5个保守基因序列,采用多位点序列分型(MLST)分析,从9种瘿蜂中获得了11个Wolbachia株系,且均为新株系。在11个株系中,具MLST等位基因共39个,其中有18个新等位基因,分别为gatB等位基因4个、coxA等位基因4个、hcpA等位基因5个、ftsZ等位基因2个和(?)fbpA等位基因3个。基于MLST分析,构建了瘿蜂Wolbachia的系统发育树；基于线粒体CO I基因序列,构建了宿主瘿蜂的系统发育树。瘿蜂感染Wolbachia的系统发生树与宿主的并不一致,且一些株系间Wolbachia的wsp基因序列一致,但保守的MLST序列却差异较大,显示本研究的瘿蜂种类间存在Wolbachia的水平传播。
Wolbachia are maternally inherited endosymbiotic bacteria. Recent study of Wolbachia incidence estimated that20-76%of insect species are infected with Wolbachia, and Wolbachia may be the most abundant and widespread endosymbiotic bacteria on earth. They could manipulate the reproduction of their hosts by causing cytoplasmic incompatibility, parthenogenesis induction, feminization of genetic males and male-killing. Cynipinae (Hymenoptera:Cynipoidea), which comprises about1400described species, are phytophagous species.29gall wasp species belonging to11genera had been recorded in China (including Taiwan).
In this paper, I collected plant galls of gall wasps in southern China and used PCR to detect the infection of Wolbachia in gall wasps. The phylogeny of Wolbachia in gall wasps was analysed by wsp gene and Multi Locus Sequence Typing (MLST). The transmission mode of Wolbachia and its effect on the reproduction mode of gall wasps were discussed.
The primary achievements of this dissertation are listed as follows:
(1)14plant galls of gall wasps were collected from Changsha, Yueyang, Loudi, Changde, Shaoyang, Yiyang in Hunan, Jian in Jiangxi, Shaoguang in Guangdong, Anqing in Anhui, Ningde in Fujian and LinZhou in Henan of China. Their host plants belonged to Castanea, Quercus, Lithocarpus and Castanopsis of Fagaceae and Rosa of Rosaceae. The15gall wasps were reared from these plant galls, and10of15gall wasps were new species:Dryocosmus zhuili Liu, ms; Cynips crystaloides Liu, ms; Synergus intricatus Liu, ms; Trigonaspis acuminatus Liu, ms; Andricus elodeoides Liu, ms; Synergus henanensis Liu, ms; Synergus carinatus Liu, ms; Saphonecrus glaberus Liu, ms; Synergus brutus Liu, ms; Saphonecrus hupingshanensis Liu, ms. A. mukaigawae, Synergus japonicus and Diplolepis japonica were recorded for the first time in China. Andricus mairei and Dryocosmus kuriphilus had been recorded in China.
(2) Our survey results showed Wolbachia infection rate Andricus mairei in Yueyang, Changsha, and Shaoyang populations:100%for the male wasps from all populations, while,100%,100%and80%for the Yueyang, Changsha and Shaoyang populations of the female wasps, respectively. The sequencing results showed that the wsp gene from all the three populations has the same sequence. Our study discovered strikingly male-biased sex ratios in3populations of A. mairei. Females only accounted for15.3%,12.1%and19.8%of the Yueyang, Changsha and Shaoyang populations, respectively. Thus, these results suggested that the Wolbachia did not induce parthenogenesis in A. mairei.
(3) Plant galls of Andricus mukaigawae were collected from eight locations in China. A. mukaigawae (gall marker), Synergus japonicus (inquiline) and Torymus sp.(parasitic wasp) emerged from plant galls of A. mukaigawae. The infection rate of Wolbachia in A. mukaigawae, S. japonicus and Torymus sp. was60%,37.1%and100%, respectively. The high levels of multiple Wolbachia infection was found in A. mukaigawae and its associated inquiline S. japonicus, with5and8Wolbachia strains, respectively. The Torymus sp. was single infection with Wolbachia. The results showed obvious recombination of wsp gene and revealed3recombination groups for Wolbachia in A. mukaigawae communities. The results suggested the horizontal transmission of Wolbachia between A. mukaigawae and S. japonicus according to the fact that A. mukaigawae and S. japonicus shared5of9Wolbachia strains in A. mukaigawae communities. The results suggested that horizontal transmission and recombination played an important role in the high levels of multiple Wolbachia infection in A. mukaigawae and S. japonicus.
(4) The results revealed Wolbachia infection in A. mukaigawae, A. mairei, Dryocosmus zhuili Liu, ms; Cynips crystaloides Liu, ms; Synergus japonicus, Synergus intricatus Liu, ms; Saphonecrus glaberus Liu, ms and Synergus brutus Liu, ms by PCR detection. The Wolbachia incidence of15gall wasps was53.3%. The infection ratio of A. mairei was the highest (100%), and that of S. brutus was lowest (26.7%).
(5)11new Wolbachia strains were found in9gall wasps using conserved gatB-. coxA、hcpA、ftsL and fbpA genes according to MLST analysis. The11strains had39MLST alleles, and18of39MLST alleles were novel alleles. The gatB, coxA, hep A, ftsZ and fbpA genes of MLST genes had4,4,5,2and3novel alleles, respectively. The MLST phylogenetic tree of Wolbachia and CO I phylogenetic tree of gall wasps were constructed, and the MLST tree and CO I tree were incongruent. Some Wolbachia strains sharing same wsp gene had a high sequence differences of conserved MLST genes. Thus, the results suggested horizontal transmission of Wolbachia between gall wasps.
本研究主要在中国南方地区采集瘿蜂标本,应用PCR法,检测获得的瘿蜂种类的Wolbachia感染；基于wsp基因序列及多位点序列分型(Multi-locus Sequence Typing, MLST)分析瘿蜂Wolbachia的系统发育关系；探讨瘿蜂Wolbachia的传播途径及其对宿主生殖模式的影响。
主要研究结果如下：
(1)从河南林州,安徽安庆,湖南长沙、岳阳、娄底、常德、邵阳、益阳,江西吉安,广东韶关和福建宁德等地共采集瘿蜂虫瘿14种,其寄主植物分别隶属于壳斗科(Fagaceae)的栗属Castanea、栎属Quercus、石栎属Lithocarpus、栲属Castanopsis和蔷薇科的蔷薇属Rosa。获得瘿蜂羽化成虫共15种。其中,新种10种(含待发表新种9种),分别为锥栗瘿蜂Dryocosmus zhuili Liu, ms、白球瘿蜂Cynips crystaloides Liu,ms、光顶并腹客瘿蜂Synergus intricatus Liu, ms、尖盾阔头瘿蜂Trigonaspis acuminatus Liu, ms、苔形瘿安瘿蜂Andricus elodeoides Liu, ms、河南并腹客瘿蜂Synergus henanensis Liu, ms、脊顶并腹客瘿蜂Synergus carinatus Liu, ms、平滑客瘿蜂Saphonecrus glaberus Liu,ms、重纹并腹客瘿蜂Synergus brutus Liu, ms和壶瓶山客瘿蜂Saphonecrus hupingshanensis Liu;中国新纪录种3种,分别为向川安瘿蜂Andricus mukaigawae、日本并腹客瘿蜂Synergus japonicus和日本蔷薇瘿蜂Diplolepis japonica;中国已记录种2种,分别是麦氏安瘿蜂Andricus mairei和栗瘿蜂Dryocosmus kuriphilus。
(2)对3种群麦氏安瘿蜂的Wolbachia感染进行了PCR检测,发现岳阳、长沙和邵阳种群的麦氏安瘿蜂雌、雄成虫均存在Wolbachia的感染,除邵阳雌成虫Wolbachia的感染率为80%之外,其它均为100%,且3种群所测得Wolbachia的wsp基因序列完全一致。岳阳、长沙及邵阳种群麦氏安瘿蜂均获得了雌、雄成虫,雌性率分别为15.3%、12.1%和19.8%,雄性比显著偏高,因此,可以认为Wolbachia的共生并未诱导其营产雌孤雌生殖。
(3)采集了8地理种群的向川安瘿蜂虫瘿,并获得了造瘿瘿蜂(向川安瘿蜂)、客居瘿蜂(日本并腹客瘿蜂)及寄生性长尾小蜂Torymus sp的羽化成虫。三者均具Wolbachia的感染,感染率分别为60%、37.1%和100%。除寄生蜂Torymussp.仅感染1个Wolbachia株系之外,向川安瘿蜂和日本并腹客瘿蜂分别感染了5个和8个株系,证实了这2种瘿蜂均存在Wolbachia高水平的多重感染。检测到向川安瘿蜂群落的3组重组株系,发现其Wolbachia的wsp基因存在明显的遗传重组现象。在向川安瘿蜂群落中的9个Wolbachia株系中,有5个株系为向川安瘿蜂和日本并腹客瘿蜂的共有株系,说明造瘿瘿蜂和客居瘿蜂之间存在Wolbachia的水平传播现象。可以推测,向川安瘿蜂和日本并腹客瘿蜂Wolbachia高水平多重感染的原因有二,即基因的遗传重组现象和水平传播。
(4)对获得的15种瘿蜂进行了Wolbachia感染的PCR检测,发现8种瘿蜂具Wolbachia感染,分别是向川安瘿蜂、麦氏安瘿蜂、锥栗瘿蜂、白球瘿蜂、日本并腹客瘿蜂、光顶并腹客瘿蜂、平滑客瘿蜂和重纹并腹客瘿蜂,感染的种类达53.3%。其中,麦氏安瘿蜂的感染率最高,接近100%；重纹并腹客瘿蜂的感染率最低,为26.7%。
(5)应用gatB、coxA、hcpA、ftsZ和fbpA5个保守基因序列,采用多位点序列分型(MLST)分析,从9种瘿蜂中获得了11个Wolbachia株系,且均为新株系。在11个株系中,具MLST等位基因共39个,其中有18个新等位基因,分别为gatB等位基因4个、coxA等位基因4个、hcpA等位基因5个、ftsZ等位基因2个和(?)fbpA等位基因3个。基于MLST分析,构建了瘿蜂Wolbachia的系统发育树；基于线粒体CO I基因序列,构建了宿主瘿蜂的系统发育树。瘿蜂感染Wolbachia的系统发生树与宿主的并不一致,且一些株系间Wolbachia的wsp基因序列一致,但保守的MLST序列却差异较大,显示本研究的瘿蜂种类间存在Wolbachia的水平传播。
Wolbachia are maternally inherited endosymbiotic bacteria. Recent study of Wolbachia incidence estimated that20-76%of insect species are infected with Wolbachia, and Wolbachia may be the most abundant and widespread endosymbiotic bacteria on earth. They could manipulate the reproduction of their hosts by causing cytoplasmic incompatibility, parthenogenesis induction, feminization of genetic males and male-killing. Cynipinae (Hymenoptera:Cynipoidea), which comprises about1400described species, are phytophagous species.29gall wasp species belonging to11genera had been recorded in China (including Taiwan).
In this paper, I collected plant galls of gall wasps in southern China and used PCR to detect the infection of Wolbachia in gall wasps. The phylogeny of Wolbachia in gall wasps was analysed by wsp gene and Multi Locus Sequence Typing (MLST). The transmission mode of Wolbachia and its effect on the reproduction mode of gall wasps were discussed.
The primary achievements of this dissertation are listed as follows:
(1)14plant galls of gall wasps were collected from Changsha, Yueyang, Loudi, Changde, Shaoyang, Yiyang in Hunan, Jian in Jiangxi, Shaoguang in Guangdong, Anqing in Anhui, Ningde in Fujian and LinZhou in Henan of China. Their host plants belonged to Castanea, Quercus, Lithocarpus and Castanopsis of Fagaceae and Rosa of Rosaceae. The15gall wasps were reared from these plant galls, and10of15gall wasps were new species:Dryocosmus zhuili Liu, ms; Cynips crystaloides Liu, ms; Synergus intricatus Liu, ms; Trigonaspis acuminatus Liu, ms; Andricus elodeoides Liu, ms; Synergus henanensis Liu, ms; Synergus carinatus Liu, ms; Saphonecrus glaberus Liu, ms; Synergus brutus Liu, ms; Saphonecrus hupingshanensis Liu, ms. A. mukaigawae, Synergus japonicus and Diplolepis japonica were recorded for the first time in China. Andricus mairei and Dryocosmus kuriphilus had been recorded in China.
(2) Our survey results showed Wolbachia infection rate Andricus mairei in Yueyang, Changsha, and Shaoyang populations:100%for the male wasps from all populations, while,100%,100%and80%for the Yueyang, Changsha and Shaoyang populations of the female wasps, respectively. The sequencing results showed that the wsp gene from all the three populations has the same sequence. Our study discovered strikingly male-biased sex ratios in3populations of A. mairei. Females only accounted for15.3%,12.1%and19.8%of the Yueyang, Changsha and Shaoyang populations, respectively. Thus, these results suggested that the Wolbachia did not induce parthenogenesis in A. mairei.
(3) Plant galls of Andricus mukaigawae were collected from eight locations in China. A. mukaigawae (gall marker), Synergus japonicus (inquiline) and Torymus sp.(parasitic wasp) emerged from plant galls of A. mukaigawae. The infection rate of Wolbachia in A. mukaigawae, S. japonicus and Torymus sp. was60%,37.1%and100%, respectively. The high levels of multiple Wolbachia infection was found in A. mukaigawae and its associated inquiline S. japonicus, with5and8Wolbachia strains, respectively. The Torymus sp. was single infection with Wolbachia. The results showed obvious recombination of wsp gene and revealed3recombination groups for Wolbachia in A. mukaigawae communities. The results suggested the horizontal transmission of Wolbachia between A. mukaigawae and S. japonicus according to the fact that A. mukaigawae and S. japonicus shared5of9Wolbachia strains in A. mukaigawae communities. The results suggested that horizontal transmission and recombination played an important role in the high levels of multiple Wolbachia infection in A. mukaigawae and S. japonicus.
(4) The results revealed Wolbachia infection in A. mukaigawae, A. mairei, Dryocosmus zhuili Liu, ms; Cynips crystaloides Liu, ms; Synergus japonicus, Synergus intricatus Liu, ms; Saphonecrus glaberus Liu, ms and Synergus brutus Liu, ms by PCR detection. The Wolbachia incidence of15gall wasps was53.3%. The infection ratio of A. mairei was the highest (100%), and that of S. brutus was lowest (26.7%).
(5)11new Wolbachia strains were found in9gall wasps using conserved gatB-. coxA、hcpA、ftsL and fbpA genes according to MLST analysis. The11strains had39MLST alleles, and18of39MLST alleles were novel alleles. The gatB, coxA, hep A, ftsZ and fbpA genes of MLST genes had4,4,5,2and3novel alleles, respectively. The MLST phylogenetic tree of Wolbachia and CO I phylogenetic tree of gall wasps were constructed, and the MLST tree and CO I tree were incongruent. Some Wolbachia strains sharing same wsp gene had a high sequence differences of conserved MLST genes. Thus, the results suggested horizontal transmission of Wolbachia between gall wasps.
引文
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