鼠多形螺旋线虫感染对宿主抗细菌天然免疫反应的调控研究
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摘要
蠕虫在世界范围内每年大约有30亿人口受到感染，并且发现蠕虫感染高发地区往往与原虫性、细菌性、病毒性疾病高发地区一致或重叠，比如疟疾、肺结核、艾滋病等。这种情况一般由蠕虫与其他病原共感染后蠕虫引起强烈的免疫调节有关。在发展中国家蠕虫感染与其他病原比如细菌等共感染经常发生。如鼠伤寒沙门氏菌，可以通过食物经口传播，可感染多种动物及人类，其感染人类可导致急性肠炎，该病原的感染依赖于宿主的免疫状况及共感染情况，因此鼠伤寒沙门氏菌容易感染婴幼儿及免疫力低下人群易感染。肠道蠕虫感染会对宿主进行免疫调节和改变宿主免疫状况，蠕虫感染诱导产生的Th2免疫反应会对细菌诱导的Th1免疫反应有抑制作用，从而导致宿主对细菌的控制与清除能力减弱。肠道蠕虫则主要诱导宿主Th2免疫反应，而应对鼠伤寒沙门氏菌则主要依赖宿主天然免疫系统，在肠道蠕虫感染情况下共感染鼠伤寒沙门氏菌可能会导致更严重的病症。目前蠕虫感染对宿主抗细菌免疫调节研究很多，但是对宿主细菌感染天然免疫反应的影响尚不清楚。本课题首先对小鼠进行寄生虫检测，排除鼠自身携带寄生虫的可能。为研究蠕虫感染对宿主抗细菌天然免疫反应的影响，本课题建立了肠道蠕虫多形螺旋线虫与鼠伤寒沙门氏菌或者鼠柠檬酸杆菌共感染C57BL/6小鼠作为动物感染模型，并对共感染肠道蠕虫与细菌小鼠天然免疫反应进行分析。
本研究利用鼠伤寒沙门氏菌感染造成急性感染模型，对共感染肠道蠕虫与细菌组及各对照组小鼠肠道细菌增殖进行涂板计数分析，并利用红色荧光鼠伤寒沙门氏菌对细菌在盲肠组织中的侵入情况进行观察分析；通过冰冻切片对各组中急性肠炎病理变化观察打分并分析；利用流式细胞分筛技术、免疫荧光染色技术、ELISA技术和荧光定量PCR技术对各组小鼠天然免疫反应水平进行比较分析；通过Th2细胞因子体外刺激并感染巨噬细胞对肠道蠕虫影响宿主抗细菌天然免疫机制进行研究。研究结果发现共感染肠道蠕虫与鼠伤寒沙门氏菌组小鼠比单独感染鼠伤寒沙门氏菌小鼠出现更严重的肠道病理变化和更低的生存率，包括粪便中细菌排出量增加、盲肠和结肠组织炎症加重、细菌侵入增多等。肠道蠕虫感染导致宿主在抗鼠伤寒沙门氏菌感染中抗菌肽分泌不足、细胞因子分泌受到抑制以及抗鼠伤寒沙门氏菌关键单核细胞--中性粒细胞数量不足。通过对其影响机制进一步分析研究发现，蠕虫感染导致宿主在抗鼠伤寒沙门氏菌免疫反应时中性粒细胞召集相关的趋化因子KC和MIP2以及细胞因子TNFα表达水平。体外巨噬细胞刺激及鼠伤寒沙门氏菌感染实验证明，蠕虫感染诱导的Th2细胞因子IL-4/IL-13刺激提高巨噬细胞中调节性细胞因子IL-10表达水平，进而导致巨噬细胞在感染鼠伤寒沙门氏菌后表达趋化因子KC、MIP2及细胞因子TNFα水平较未处理组巨噬细胞下降显著。本研究还利用鼠柠檬酸杆菌引起肠道慢性细菌感染模型进行验证，同样发现肠道蠕虫多形螺旋线虫感染导致鼠柠檬酸杆菌诱导的慢性细菌性结肠炎发病严重，且体重下降迅速及生存率下降，肠道组织中抗菌肽Reg3γ与趋化因子KC的分泌表达受到抑制，以及中性粒细胞召集受阻。
本课题通过研究寄生性蠕虫对宿主抗细菌天然免疫反应的影响，确定了肠道蠕虫感染抑制宿主抗细菌感染时抗菌肽、趋化因子及细胞因子的分泌表达，阻碍了中性粒细胞的召集。为进一步了解蠕虫感染对细菌性疾病的影响提供依据，也为预防和治疗寄生虫和细菌混合感染这一重要公共卫生问题的提供新方法和参考。
Helminths are estimated to infect3billion people worldwide. The distribution ofseveral helminth pathogens coincides geographically with many devastating microbialdiseases including HIV, malaria, and tuberculosis, and it is possible that the strongimmunomodulatory effects of helminthes on host responses may have a significantimpact on such coincident infections. In many developing countries, exposure tohelminth infections and simultaneous infection with other pathogens such as bacteriais very common.. Such as Salmonella enterica serovar Typhimurium (S.Typhimurium), it is a Gram-negative food-borne pathogen that is a major cause ofacute gastroenteritis in humans. The ability of the host to control such bacterialpathogens may be influenced by host immune status and by concurrent infections. Across-regulatory suppression of Th1responses by a helminth-driven strong Th2response has been suggested as a contributing factor to the alteration of the hostresponse to concurrent bacterial infections.Although much is known about thepotential role of helminth stimulated T cells, typically Th2and Treg, in altering hostprotection against the bacterial infection, the impact of helminth infection on theinnate immune response to enteric bacterial pathogens is less well understood. To testthe hypothesis that helminth infection may negatively regulate host mucosal innateimmunity against bacterial enteropathogens, a murine co-infection model wasestablished by using the intestinal nematode Heligmosomoides polygyrus and S.Typhimurium.
In this study, Salmonella-induced acute colitis was used, and the bacterialevasion and output was calculated. Also the pathology and innate immune reponsesinclude monocyte recruitment, cytokine expressions and chemokine expressions weredetermined through the HE staining, FACs and Real-time PCR. The in vitrostimulation of macrophage by Th2cytokines and infection with Salmonella wasprocessed for understanding the mechanism of the way by which the helminthinfection suppresses the bacteria induced innate immune responses. We found that mice co-infected with S. Typhimurium and H. polygyrus developed more severeintestinal inflammation than animals infected with S. Typhimurium alone. Theenhanced susceptibility to Salmonella-induced intestinal injury in co-infected micewas found to be associated with diminished neutrophil recruitment to the site ofbacterial infection that correlated with decreased expression of the chemoattractantsMIP-2and KC, poor control of bacterial replication and exacerbated intestinalinflammation. The mechanism of helminth-induced inhibition of MIP-2and KCexpression involved IL-10and, to a less extent, IL-4and IL-13. Ly6G antibody-mediated depletion of neutrophils reproduced the adverse effects of H. polygyrus onSalmonella infection. Our results suggest that impaired neutrophil recruitment is animportant contributor to the enhanced severity of Salmonella enterocolitis associatedwith helminth co-infection. The Citrobacter rodentium (C. rodentium) infected micewith co-infected helminth H. polygyrus were used for verifying the impairedrecruitment of nrutrophil by the helminth infection. The results show the sameinbition of KC and Reg3γ in the conlon tissue from the co-infected mice withhelminth and bacteria. And the neutrophil recruitment was also suppressed by thehelminth infection.
In the current study, we determined that the helminth infection may suppressesthe bacteria induced innate immune responses, and the helminth infection decreasesthe anti-microbial peptides, chemokine and cytokine expressions in the co-infectedmice with bacteria, failed the neutrophil recruitment during the innate immunityagainst bacteria. The better understanding of the regulation of helminth infection onthe innate immune responses induced by bacteria provides the novel insight and toolto prevent and control the co-current infection with helminth and bacteria.
本研究利用鼠伤寒沙门氏菌感染造成急性感染模型，对共感染肠道蠕虫与细菌组及各对照组小鼠肠道细菌增殖进行涂板计数分析，并利用红色荧光鼠伤寒沙门氏菌对细菌在盲肠组织中的侵入情况进行观察分析；通过冰冻切片对各组中急性肠炎病理变化观察打分并分析；利用流式细胞分筛技术、免疫荧光染色技术、ELISA技术和荧光定量PCR技术对各组小鼠天然免疫反应水平进行比较分析；通过Th2细胞因子体外刺激并感染巨噬细胞对肠道蠕虫影响宿主抗细菌天然免疫机制进行研究。研究结果发现共感染肠道蠕虫与鼠伤寒沙门氏菌组小鼠比单独感染鼠伤寒沙门氏菌小鼠出现更严重的肠道病理变化和更低的生存率，包括粪便中细菌排出量增加、盲肠和结肠组织炎症加重、细菌侵入增多等。肠道蠕虫感染导致宿主在抗鼠伤寒沙门氏菌感染中抗菌肽分泌不足、细胞因子分泌受到抑制以及抗鼠伤寒沙门氏菌关键单核细胞--中性粒细胞数量不足。通过对其影响机制进一步分析研究发现，蠕虫感染导致宿主在抗鼠伤寒沙门氏菌免疫反应时中性粒细胞召集相关的趋化因子KC和MIP2以及细胞因子TNFα表达水平。体外巨噬细胞刺激及鼠伤寒沙门氏菌感染实验证明，蠕虫感染诱导的Th2细胞因子IL-4/IL-13刺激提高巨噬细胞中调节性细胞因子IL-10表达水平，进而导致巨噬细胞在感染鼠伤寒沙门氏菌后表达趋化因子KC、MIP2及细胞因子TNFα水平较未处理组巨噬细胞下降显著。本研究还利用鼠柠檬酸杆菌引起肠道慢性细菌感染模型进行验证，同样发现肠道蠕虫多形螺旋线虫感染导致鼠柠檬酸杆菌诱导的慢性细菌性结肠炎发病严重，且体重下降迅速及生存率下降，肠道组织中抗菌肽Reg3γ与趋化因子KC的分泌表达受到抑制，以及中性粒细胞召集受阻。
本课题通过研究寄生性蠕虫对宿主抗细菌天然免疫反应的影响，确定了肠道蠕虫感染抑制宿主抗细菌感染时抗菌肽、趋化因子及细胞因子的分泌表达，阻碍了中性粒细胞的召集。为进一步了解蠕虫感染对细菌性疾病的影响提供依据，也为预防和治疗寄生虫和细菌混合感染这一重要公共卫生问题的提供新方法和参考。
Helminths are estimated to infect3billion people worldwide. The distribution ofseveral helminth pathogens coincides geographically with many devastating microbialdiseases including HIV, malaria, and tuberculosis, and it is possible that the strongimmunomodulatory effects of helminthes on host responses may have a significantimpact on such coincident infections. In many developing countries, exposure tohelminth infections and simultaneous infection with other pathogens such as bacteriais very common.. Such as Salmonella enterica serovar Typhimurium (S.Typhimurium), it is a Gram-negative food-borne pathogen that is a major cause ofacute gastroenteritis in humans. The ability of the host to control such bacterialpathogens may be influenced by host immune status and by concurrent infections. Across-regulatory suppression of Th1responses by a helminth-driven strong Th2response has been suggested as a contributing factor to the alteration of the hostresponse to concurrent bacterial infections.Although much is known about thepotential role of helminth stimulated T cells, typically Th2and Treg, in altering hostprotection against the bacterial infection, the impact of helminth infection on theinnate immune response to enteric bacterial pathogens is less well understood. To testthe hypothesis that helminth infection may negatively regulate host mucosal innateimmunity against bacterial enteropathogens, a murine co-infection model wasestablished by using the intestinal nematode Heligmosomoides polygyrus and S.Typhimurium.
In this study, Salmonella-induced acute colitis was used, and the bacterialevasion and output was calculated. Also the pathology and innate immune reponsesinclude monocyte recruitment, cytokine expressions and chemokine expressions weredetermined through the HE staining, FACs and Real-time PCR. The in vitrostimulation of macrophage by Th2cytokines and infection with Salmonella wasprocessed for understanding the mechanism of the way by which the helminthinfection suppresses the bacteria induced innate immune responses. We found that mice co-infected with S. Typhimurium and H. polygyrus developed more severeintestinal inflammation than animals infected with S. Typhimurium alone. Theenhanced susceptibility to Salmonella-induced intestinal injury in co-infected micewas found to be associated with diminished neutrophil recruitment to the site ofbacterial infection that correlated with decreased expression of the chemoattractantsMIP-2and KC, poor control of bacterial replication and exacerbated intestinalinflammation. The mechanism of helminth-induced inhibition of MIP-2and KCexpression involved IL-10and, to a less extent, IL-4and IL-13. Ly6G antibody-mediated depletion of neutrophils reproduced the adverse effects of H. polygyrus onSalmonella infection. Our results suggest that impaired neutrophil recruitment is animportant contributor to the enhanced severity of Salmonella enterocolitis associatedwith helminth co-infection. The Citrobacter rodentium (C. rodentium) infected micewith co-infected helminth H. polygyrus were used for verifying the impairedrecruitment of nrutrophil by the helminth infection. The results show the sameinbition of KC and Reg3γ in the conlon tissue from the co-infected mice withhelminth and bacteria. And the neutrophil recruitment was also suppressed by thehelminth infection.
In the current study, we determined that the helminth infection may suppressesthe bacteria induced innate immune responses, and the helminth infection decreasesthe anti-microbial peptides, chemokine and cytokine expressions in the co-infectedmice with bacteria, failed the neutrophil recruitment during the innate immunityagainst bacteria. The better understanding of the regulation of helminth infection onthe innate immune responses induced by bacteria provides the novel insight and toolto prevent and control the co-current infection with helminth and bacteria.
引文
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