洋葱伯克氏群细菌的分子致病基础及其对生境适应性研究
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摘要
洋葱伯克氏菌群(Bcc)由17个相近的亚种组成,它们对植物均有有利和有害的两个方面。有一些Bcc菌株能够降解自然环境中和人工合成的污染物,有一些能促进植物生长和防治植物病害。同时,Bcc的一些菌株被发现是植物病原菌而另一些是人体条件致病菌。然而,在过去的几十年里,相关研究表明人体致病Bcc菌出现了多重抗药性。因此,对Bcc菌进行风险评估,对来自于土壤、水、医院环境、植物病原菌等不同来源的Bcc菌进行分子检测及环境适应性的研究势在必行。
本研究建立了一个简单易行并且结果稳定的生菜致病模型来检测Bcc菌的致病性。同时和Bcc菌致病性的其他评估模型如苜蓿模型、腊蛾模型和大鼠琼脂珠模型进行了比较。此外,本研究通过寄主模型培养基培养、1D蛋白质电泳、液相层析串联质朴分析和MASCOT数据库检索,以及硅肺功能分析,研究了Bcc菌的外膜蛋白。同时,为了找到一种抗菌制剂,促进医学和工业环境的卫生,本研究首次评估了金属铜表面对Bcc菌的抑菌作用。以不锈钢为对照研究了金属铜表面对Bcc菌的作用机制。
主要的研究结果如下：
1.利用生菜模型评价了265个菌株的致病性,细菌生长24小时后接种,观察5天。在184株被调查的B. pyrrocinia菌株中,有50株是弱致病力,5株致病力一般,有2株表现很强的致病症状,另外127株没有致病作用。然而,B. anthina, B. stabilis和B. arboris的所有菌株对生菜都有致病型；在大鼠致病模型上也得到了同样的结果。致病基因BCAM0218, BCAM0224和]BCAM0219在所有265个菌株均没有被检测到,BCESM在B. arbroris中被检测含有。
2.测定了来源于医院、水、土壤和植物病原菌的70多株Bcc菌株在苜蓿、腊蛾和大鼠琼脂珠上的毒力。来源于医院和水中的菌株比土壤菌和植物病原菌的致病力强,但是也有少数的医院菌株表现出致病力减弱的症状。此外,来源于水的Bcc菌株致病力比土壤菌和植物病原菌强,但是比医院菌弱。在显微镜下观察生菜中脉组织,可以明显的看出细菌的入侵、定殖和复制。对生物膜的检测研究也得到相似的结果：来源于水和医院的菌株形成生物膜的能力比土壤菌和植物病原菌强。在致病模型试验中,B. cepacia LMG1222被用作参考菌株。
3.在外膜蛋白研究中,通过LC-MS/MS方法进行了外膜蛋白的蛋白质组分析,在水、囊胞性纤维症、植物和土壤模拟生境下总共鉴定了400多个蛋白。在整个鉴定蛋白目录中,已检测发现了一些共同蛋白及一些在每个培养基条件下差异表达的蛋白。一些共同蛋白如ABC转运蛋白、外排泵蛋白、OmpW等是Bcc菌生存和生长的基础。一些蛋白如3型、4型和6型分泌系统(T3SS、T4SS、 T6SS), ZmpA、鞭毛蛋白、孔蛋白、LysR等在不同寄主模拟生境下表达差异有所不同,可能是Bcc菌在进化过程中对寄主及不同生存条件的反应,这也是未来研究工作的努力方向。
4.铜表面对Bcc菌表现很强的抑菌活性。大量的铜离子进入细菌细胞,细菌在接触铜表面之后细胞裂解,但是并未发现DNA的损坏。可能的机制是,接触杀菌过程通常破坏细胞膜、铜大量的进入细胞、细胞死亡,然后DNA降解。
5.我们正在开展B. seminalis的4个不同菌株的基因组和转录组的研究,目的是为了开启Bcc的生态学大门.
Burkholderia cepacia complex (Bcc) is composed of17closely related species and found ubiquitously, where they can have both beneficial and detrimental effects on plants. Some can degrade natural and man-made pollutants; some can promote plant growth or can control plant diseases. Some members are also recognized as a plant pathogen and some opportunistic human pathogens. However, over the past decade, there have been several reports of emergence of multidrug resistant Bcc infecting the vulnerable people. This has increased the need for the risk assessment, molecular basis and niche adaptation of Bcc isolates resourced from soil, water, clinical as well as plant pathogen.
To test the pathogenicity, an alternative, potential, easy accessible and high throughput lettuce infection model was developed in this study. Alfalfa, Galleria mellonella (wax moth) and rat agar bead model was also used to evaluate the virulence intensity among Bcc isolates comparing with lettuce. Moreover, the outer membrane (OM) proteins of Bcc were investigated for molecular basis by growing Bcc bacteria in host mimic media.1D-SDS-PAGE and Lipid chromatography tandem mass spectrometry (LC-MS/MS) peptides profiles and MASCOT search database as well as in silico functional analysis were performed. Furthermore, to find possible antibacterial agent to keep environment particularly hospital and industries hygienic, copper surfaces as antibacterial agent, first time, was evaluated against Bcc. Mode of action of dry metallic copper surfaces against Bcc was also evaluated at copper exposed Bcc cells, while stainless steel was used as a control
The main results of these studies were as follows:
1. The lettuce model takes～24h to launch experiment including bacterial growth using store-bought lettuce and monitor for up to5d post-infection. Total,265Bcc isolates were used to investigate the virulence intensity and efficacy of lettuce. All isolates belonging to B. anthina, B. stabilis and B. arboris were determined avirulent in lettuce except B. pyrrocinia, where out of184,50isolates showed weak,5 moderate and2isolates showed severe disease symptoms and127found totally avirulent. Selected isolates in rat infection model showed same level of infection as in lettuce infection model. The transmissibility marker BCAM0218, BCAM0224, BCAM0219and BCESM were exclusively absent in265isolates except in B. arboris in which BCAM0219were determined.
2. In the further study,70more Bcc isolates resourced from different places e.g clinical, water, soil and plant pathogens were tested in lettuce. Following lettuce pathogenicity was also tested in alfalfa, Galleria mellonella and rat agar bead alternative infection model. The biofilm formation ability was also tested. Severe to moderate pathogenicity was observed for isolates of clinical and water origin compared to soil and plant pathogen, with the exception of a few clinical isolates exhibiting reduced pathogenicity. Moreover, among water isolates, pathogenicity was also higher than soil and plant resource isolates but less than clinical. The lettuce midrib tissues show remarkable invasion, localization, and replication of bacteria when observed by transmission electron microscopic study. Similarly, biofilm formation ability was also higher among water and clinical virulent isolates compared to soil and plant pathogen isolates. B. cepacia LMG1222was used as a reference strain in these infection models.
3. In the OM study, shotgun proteomic analyses of OM by LC-MS/MS, overall identified>400proteins under water, Cystic Fibrosis (CF) mimic, water mimic, plant mimic and soil mimic media. Among the catalog of entire identified proteins, shared proteins as well as several proteins remarkably differentially expressed under each growth medium were detected. Shared proteins like ABC transporter proteins, efflux proteins, OmpW etc could be fundamental for growth and survival. While differentially expressed proteins e.g type3, type4, type6secretion system (T3SS, T4SS, T6SS), ZmpA, flagellin, porin, LysR etc under different host mimic medium, perhaps reflecting the evolution due to different environmental survival toward host-adaptation and may represent prime for further studies.
4. Copper surfaces were tested against Bcc and it showed the strong antibacterial activity. Higher influx of copper ions into the bacterial cells and cells disintegration among the cells exposed to copper surfaces were noted, while no DNA damage was detected at specified time. At the existing level of knowledge, it looks that contact killing proceeds by successive membrane damage, copper influx into the cells, cell death, and DNA degradation.
5. The genomics and transcriptomics studies of four B. seminalis strains for the purpose to unlock the ecology of Bcc are under process.
本研究建立了一个简单易行并且结果稳定的生菜致病模型来检测Bcc菌的致病性。同时和Bcc菌致病性的其他评估模型如苜蓿模型、腊蛾模型和大鼠琼脂珠模型进行了比较。此外,本研究通过寄主模型培养基培养、1D蛋白质电泳、液相层析串联质朴分析和MASCOT数据库检索,以及硅肺功能分析,研究了Bcc菌的外膜蛋白。同时,为了找到一种抗菌制剂,促进医学和工业环境的卫生,本研究首次评估了金属铜表面对Bcc菌的抑菌作用。以不锈钢为对照研究了金属铜表面对Bcc菌的作用机制。
主要的研究结果如下：
1.利用生菜模型评价了265个菌株的致病性,细菌生长24小时后接种,观察5天。在184株被调查的B. pyrrocinia菌株中,有50株是弱致病力,5株致病力一般,有2株表现很强的致病症状,另外127株没有致病作用。然而,B. anthina, B. stabilis和B. arboris的所有菌株对生菜都有致病型；在大鼠致病模型上也得到了同样的结果。致病基因BCAM0218, BCAM0224和]BCAM0219在所有265个菌株均没有被检测到,BCESM在B. arbroris中被检测含有。
2.测定了来源于医院、水、土壤和植物病原菌的70多株Bcc菌株在苜蓿、腊蛾和大鼠琼脂珠上的毒力。来源于医院和水中的菌株比土壤菌和植物病原菌的致病力强,但是也有少数的医院菌株表现出致病力减弱的症状。此外,来源于水的Bcc菌株致病力比土壤菌和植物病原菌强,但是比医院菌弱。在显微镜下观察生菜中脉组织,可以明显的看出细菌的入侵、定殖和复制。对生物膜的检测研究也得到相似的结果：来源于水和医院的菌株形成生物膜的能力比土壤菌和植物病原菌强。在致病模型试验中,B. cepacia LMG1222被用作参考菌株。
3.在外膜蛋白研究中,通过LC-MS/MS方法进行了外膜蛋白的蛋白质组分析,在水、囊胞性纤维症、植物和土壤模拟生境下总共鉴定了400多个蛋白。在整个鉴定蛋白目录中,已检测发现了一些共同蛋白及一些在每个培养基条件下差异表达的蛋白。一些共同蛋白如ABC转运蛋白、外排泵蛋白、OmpW等是Bcc菌生存和生长的基础。一些蛋白如3型、4型和6型分泌系统(T3SS、T4SS、 T6SS), ZmpA、鞭毛蛋白、孔蛋白、LysR等在不同寄主模拟生境下表达差异有所不同,可能是Bcc菌在进化过程中对寄主及不同生存条件的反应,这也是未来研究工作的努力方向。
4.铜表面对Bcc菌表现很强的抑菌活性。大量的铜离子进入细菌细胞,细菌在接触铜表面之后细胞裂解,但是并未发现DNA的损坏。可能的机制是,接触杀菌过程通常破坏细胞膜、铜大量的进入细胞、细胞死亡,然后DNA降解。
5.我们正在开展B. seminalis的4个不同菌株的基因组和转录组的研究,目的是为了开启Bcc的生态学大门.
Burkholderia cepacia complex (Bcc) is composed of17closely related species and found ubiquitously, where they can have both beneficial and detrimental effects on plants. Some can degrade natural and man-made pollutants; some can promote plant growth or can control plant diseases. Some members are also recognized as a plant pathogen and some opportunistic human pathogens. However, over the past decade, there have been several reports of emergence of multidrug resistant Bcc infecting the vulnerable people. This has increased the need for the risk assessment, molecular basis and niche adaptation of Bcc isolates resourced from soil, water, clinical as well as plant pathogen.
To test the pathogenicity, an alternative, potential, easy accessible and high throughput lettuce infection model was developed in this study. Alfalfa, Galleria mellonella (wax moth) and rat agar bead model was also used to evaluate the virulence intensity among Bcc isolates comparing with lettuce. Moreover, the outer membrane (OM) proteins of Bcc were investigated for molecular basis by growing Bcc bacteria in host mimic media.1D-SDS-PAGE and Lipid chromatography tandem mass spectrometry (LC-MS/MS) peptides profiles and MASCOT search database as well as in silico functional analysis were performed. Furthermore, to find possible antibacterial agent to keep environment particularly hospital and industries hygienic, copper surfaces as antibacterial agent, first time, was evaluated against Bcc. Mode of action of dry metallic copper surfaces against Bcc was also evaluated at copper exposed Bcc cells, while stainless steel was used as a control
The main results of these studies were as follows:
1. The lettuce model takes～24h to launch experiment including bacterial growth using store-bought lettuce and monitor for up to5d post-infection. Total,265Bcc isolates were used to investigate the virulence intensity and efficacy of lettuce. All isolates belonging to B. anthina, B. stabilis and B. arboris were determined avirulent in lettuce except B. pyrrocinia, where out of184,50isolates showed weak,5 moderate and2isolates showed severe disease symptoms and127found totally avirulent. Selected isolates in rat infection model showed same level of infection as in lettuce infection model. The transmissibility marker BCAM0218, BCAM0224, BCAM0219and BCESM were exclusively absent in265isolates except in B. arboris in which BCAM0219were determined.
2. In the further study,70more Bcc isolates resourced from different places e.g clinical, water, soil and plant pathogens were tested in lettuce. Following lettuce pathogenicity was also tested in alfalfa, Galleria mellonella and rat agar bead alternative infection model. The biofilm formation ability was also tested. Severe to moderate pathogenicity was observed for isolates of clinical and water origin compared to soil and plant pathogen, with the exception of a few clinical isolates exhibiting reduced pathogenicity. Moreover, among water isolates, pathogenicity was also higher than soil and plant resource isolates but less than clinical. The lettuce midrib tissues show remarkable invasion, localization, and replication of bacteria when observed by transmission electron microscopic study. Similarly, biofilm formation ability was also higher among water and clinical virulent isolates compared to soil and plant pathogen isolates. B. cepacia LMG1222was used as a reference strain in these infection models.
3. In the OM study, shotgun proteomic analyses of OM by LC-MS/MS, overall identified>400proteins under water, Cystic Fibrosis (CF) mimic, water mimic, plant mimic and soil mimic media. Among the catalog of entire identified proteins, shared proteins as well as several proteins remarkably differentially expressed under each growth medium were detected. Shared proteins like ABC transporter proteins, efflux proteins, OmpW etc could be fundamental for growth and survival. While differentially expressed proteins e.g type3, type4, type6secretion system (T3SS, T4SS, T6SS), ZmpA, flagellin, porin, LysR etc under different host mimic medium, perhaps reflecting the evolution due to different environmental survival toward host-adaptation and may represent prime for further studies.
4. Copper surfaces were tested against Bcc and it showed the strong antibacterial activity. Higher influx of copper ions into the bacterial cells and cells disintegration among the cells exposed to copper surfaces were noted, while no DNA damage was detected at specified time. At the existing level of knowledge, it looks that contact killing proceeds by successive membrane damage, copper influx into the cells, cell death, and DNA degradation.
5. The genomics and transcriptomics studies of four B. seminalis strains for the purpose to unlock the ecology of Bcc are under process.
引文
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