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Introduction

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 Bartonella (Rochalimaea) quintana, 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 Coxiella burnetii 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., Mycobacteria, Legionella, Salmonella, Shigella, Francisella, and Brucella) are facultative intracellular parasites. In contrast with chlamydiae, all rickettsiae can synthesize ATP. Coxiella burnetii is the only rickettsia that appears to have a developmental cycle.

Some organisms in the family Rickettsiaceae are closely related genetically (e.g., Rickettsia rickettsii, R akari, R prowazekii, and R typhi); others are related less closely to Rickettsia species (e.g., Ehrlichia and Bartonella); and others not related to Rickettsia species (e.g., C burnetii). The phenotypic traits of the medically important organism Orientia (Rickettsia) tsutsugamushi suggest that the species may be an example of convergent evolution in a similar ecologic niche.

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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).

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