Wilson-Welder Jennifer H, Frank Ami T, Hornsby Richard L, Olsen Steven C, Alt David P
Infectious Bacterial Diseases Research Unit, National Animal Disease Center Ames, IA, USA.
Front Microbiol. 2016 Jul 19;7:1110. doi: 10.3389/fmicb.2016.01110. eCollection 2016.
Cattle are the reservoir hosts of Leptospira borgpetersenii serovar Hardjo, and can also be reservoir hosts of other Leptospira species such as L. kirschneri, and Leptospira interrogans. As a reservoir host, cattle shed Leptospira, infecting other animals, including humans. Previous studies with human and murine neutrophils have shown activation of neutrophil extracellular trap or NET formation, and upregulation of inflammatory mediators by neutrophils in the presence of Leptospira. Humans, companion animals and most widely studied models of Leptospirosis are of acute infection, hallmarked by systemic inflammatory response, neutrophilia, and septicemia. In contrast, cattle exhibit chronic infection with few outward clinical signs aside from reproductive failure. Taking into consideration that there is host species variation in innate immunity, especially in pathogen recognition and response, the interaction of bovine peripheral blood polymorphonuclear cells (PMNs) and several Leptospira strains was evaluated. Studies including bovine-adapted strains, human pathogen strains, a saprophyte and inactivated organisms. Incubation of PMNs with Leptospira did induce slight activation of neutrophil NETs, greater than unstimulated cells but less than the quantity from E. coli P4 stimulated PMNs. Very low but significant from non-stimulated, levels of reactive oxygen peroxides were produced in the presence of all Leptospira strains and E. coli P4. Similarly, significant levels of reactive nitrogen intermediaries (NO2) was produced from PMNs when incubated with the Leptospira strains and greater quantities in the presence of E. coli P4. PMNs incubated with Leptospira induced RNA transcripts of IL-1β, MIP-1α, and TNF-α, with greater amounts induced by live organisms when compared to heat-inactivated leptospires. Transcript for inflammatory cytokine IL-8 was also induced, at similar levels regardless of Leptospira strain or viability. However, incubation of Leptospira strains with bovine PMNs did not affect Leptospira viability as measured by limiting dilution culture. This is in contrast to previously reported results of innate inflammatory activation by Leptospira in human and other animal models, or the activation and interaction of bovine PMNs with Escherichia coli and other bacterial pathogens. While it could be hypothesized that variations in innate receptor recognition, specifically variance in toll-like receptor 2, could underlie the observed reduction of activation in bovine PMNs, additional studies would be needed to explore this possibility. Reduction in neutrophil responses may help to establish nearly asymptomatic chronic Leptospira infection of cattle. This study emphasizes the importance of studying host-pathogen relationships in the appropriate species as extrapolation from other animal models may be incorrect and confounded by differences in the host responses.
牛是波氏疏螺旋体哈焦血清型的储存宿主,也可以是其他疏螺旋体物种的储存宿主,如克氏疏螺旋体和问号状疏螺旋体。作为储存宿主,牛排出疏螺旋体,感染包括人类在内的其他动物。先前对人类和小鼠中性粒细胞的研究表明,在存在疏螺旋体的情况下,中性粒细胞胞外陷阱或NET形成会被激活,并且中性粒细胞会上调炎症介质。人类、伴侣动物以及研究最为广泛的钩端螺旋体病模型均为急性感染,其特征为全身炎症反应、中性粒细胞增多和败血症。相比之下,牛表现为慢性感染,除繁殖失败外几乎没有明显的临床症状。考虑到先天免疫存在宿主物种差异,尤其是在病原体识别和反应方面,对牛外周血多形核细胞(PMN)与几种钩端螺旋体菌株的相互作用进行了评估。研究包括适应牛的菌株、人类病原体菌株、腐生菌和灭活生物体。PMN与钩端螺旋体的孵育确实诱导了中性粒细胞NETs的轻微激活,比未刺激的细胞更强,但比大肠杆菌P4刺激的PMN产生的量少。在所有钩端螺旋体菌株和大肠杆菌P4存在的情况下,产生了非常低但与未刺激状态相比有显著差异的活性氧过氧化物水平。同样,当与钩端螺旋体菌株孵育时,PMN产生了显著水平但数量低于大肠杆菌P4的活性氮中间体(NO2)。与钩端螺旋体孵育的PMN诱导了IL-1β、MIP-1α和TNF-α的RNA转录本,与热灭活的钩端螺旋体相比,活生物体诱导的量更多。炎症细胞因子IL-8的转录本也被诱导,无论钩端螺旋体菌株或活力如何,其水平相似。然而,通过有限稀释培养测量,钩端螺旋体菌株与牛PMN的孵育并未影响钩端螺旋体的活力。这与先前报道的钩端螺旋体在人类和其他动物模型中引起的先天炎症激活结果,或牛PMN与大肠杆菌和其他细菌病原体的激活及相互作用结果形成对比。虽然可以假设先天受体识别的差异,特别是Toll样受体2的差异,可能是观察到的牛PMN激活减少的基础,但还需要进一步的研究来探索这种可能性。中性粒细胞反应的减少可能有助于牛建立几乎无症状的慢性钩端螺旋体感染。这项研究强调了在合适的物种中研究宿主-病原体关系的重要性,因为从其他动物模型推断可能是不正确的,并且会因宿主反应的差异而混淆。