Rickettsiae are small, Gram-negative bacilli that have evolved in such close association with arthropod hosts that they are adapted to survive within the host cells. They represent a rather diverse collection of bacteria, and therefore listing characteristics that apply to the entire group is difficult. The common threads that hold the rickettsiae into a group are their epidemiology, their obligate intracellular lifestyle, and the laboratory technology required to work with them. In the laboratory, rickettsiae cannot be cultivated on agar plates or in broth, but only in viable eukaryotic host cells (e.g., in cell culture, embryonated eggs, or susceptible animals). The exception, which shows the artificial nature of using obligate intracellular parasitism as a defining phenotypic characteristic, is , which is cultivable axenically, but was traditionally considered as a rickettsia. The diversity of rickettsiae is demonstrated in the variety of specific intracellular locations where they live and the remarkable differences in their major outer membrane proteins and genetic relatedness (Table 38-1). An example of extreme adaptation is that the metabolic activity of is greatly increased in the acidic environment of the phagolysosome, which is a harsh location for survival for most other organisms. Obligate intracellular parasitism among bacteria is not unique to rickettsiae. Chlamydiae also have evolved to occupy an intracellular niche, and numerous bacteria (e.g., , , , , , and ) are facultative intracellular parasites. In contrast with chlamydiae, all rickettsiae can synthesize ATP. is the only rickettsia that appears to have a developmental cycle. Some organisms in the family are closely related genetically (e.g., , , , and ); others are related less closely to species (e.g., and ); and others not related to (e.g., ). The phenotypic traits of the medically important organism () suggest that the species may be an example of convergent evolution in a similar ecologic niche. Rickettsioses are zoonoses that, except for Q fever, are usually transmitted to humans by arthropods (tick, mite, flea, louse, or chigger) (Table 38-2). Therefore, their geographic distribution is determined by that of the infected arthropod, which for most rickettsial species is the reservoir host. Rickettsiae are important causes of human diseases in the United States (Rocky Mountain spotted fever, Q fever, murine typhus, sylvatic typhus, human monocytic ehrlichiosis, human granulocytic ehrlichiosis, and rickettsialpox) and around the world (Q fever, murine typhus, scrub typhus, epidemic typhus, boutonneuse fever, and other spotted fevers) (Table 38-2).
立克次氏体是小型革兰氏阴性杆菌,它们与节肢动物宿主紧密共生,已适应在宿主细胞内生存。它们代表了一类相当多样化的细菌,因此很难列出适用于整个群体的特征。将立克次氏体归为一类的共同主线是它们的流行病学、专性胞内生活方式以及处理它们所需的实验室技术。在实验室中,立克次氏体不能在琼脂平板或肉汤中培养,而只能在有活力的真核宿主细胞中培养(例如在细胞培养物、胚胎卵或易感动物中)。例外情况是五日热巴通体,它可以在无细胞环境中培养,但传统上被认为是一种立克次氏体,这表明将专性胞内寄生作为定义性表型特征具有人为性。立克次氏体的多样性体现在它们生存的特定细胞内位置的多样性、主要外膜蛋白的显著差异以及遗传相关性上(表38 - 1)。一个极端适应的例子是五日热巴通体的代谢活性在吞噬溶酶体的酸性环境中大大增加,而吞噬溶酶体对大多数其他生物来说是一个恶劣的生存场所。细菌中的专性胞内寄生并非立克次氏体所特有。衣原体也已进化到占据细胞内生态位,并且许多细菌(例如大肠杆菌、沙门氏菌、布鲁氏菌、李斯特菌、军团菌和分枝杆菌)是兼性胞内寄生虫。与衣原体不同,所有立克次氏体都能合成ATP。五日热巴通体是唯一似乎具有发育周期的立克次氏体。立克次氏体科中的一些生物体在遗传上密切相关(例如立氏立克次体、普氏立克次体、恙虫病东方体和五日热巴通体);其他一些与立克次体属物种的关系不太密切(例如斑点热立克次体和加拿大立克次体);还有一些与立克次体属无关(例如汉赛巴尔通体)。具有医学重要性的病原体五日热巴通体的表型特征表明,该物种可能是在相似生态位中趋同进化的一个例子。立克次体病是人畜共患病,除Q热外,通常通过节肢动物(蜱、螨、跳蚤、虱子或恙螨)传播给人类(表38 - 2)。因此,它们的地理分布由受感染节肢动物的分布决定,对大多数立克次体物种来说,受感染节肢动物是储存宿主。立克次氏体是美国(落基山斑疹热、Q热、鼠型斑疹伤寒、丛林斑疹伤寒、人单核细胞埃立克体病、人粒细胞埃立克体病和立克次体痘)以及世界各地(Q热、鼠型斑疹伤寒、恙虫病、流行性斑疹伤寒、纽扣热和其他斑点热)人类疾病的重要病因(表38 - 2)。