Rickettsia

 Rickettsia is a genus of nonmotile, Gram-negative, nonspore-forming, highly pleomorphic bacteria that may occur in the forms of cocci (0.1 μm in diameter), bacilli (1–4 μm long), or threads (up to about 10 μm long). The term "rickettsia" has nothing to do with rickets (which is a deficiency disease resulting from lack of vitamin D); the bacterial genus Rickettsia was named after Howard Taylor Ricketts, in honor of his pioneering work on tick-borne spotted fever.

Rickettsia
Red-stained Rickettsia rickettsii visible in the cell of an Ixodid vector tick
Red-stained Rickettsia rickettsii visible in cells of an Ixodid vector tick
Scientific classificatione
Domain:Bacteria
Phylum:Proteobacteria
Class:Alphaproteobacteria
Order:Rickettsiales
Family:Rickettsiaceae
Tribe:Rickettsieae
Genus:Rickettsia
da Rocha-Lima, 1916
Species groups and Species[8]
  • belli group
    • Candidatus Rickettsia angustus
    • Candidatus Rickettsia kingi
    • Candidatus Rickettsia mendelii
    • Candidatus Rickettsia tarasevichiae
    • Rickettsia bellii
    • Rickettsia canadensis [1][2]
    • Rickettsia monteiroi
  • spotted fever group
    • Candidatus Rickettsia andeanae
    • Candidatus Rickettsia antechini
    • Candidatus Rickettsia barbariae
    • Candidatus Rickettsia goldwasserii
    • Candidatus Rickettsia hungarica
    • Candidatus Rickettsia jingxinensis
    • Candidatus Rickettsia kellyi
    • Candidatus Rickettsia kotlanii
    • Candidatus Rickettsia kulagini
    • Candidatus Rickettsia leptotrombidium
    • Candidatus Rickettsia longicornii
    • Candidatus Rickettsia nicoyana
    • Candidatus Rickettsia principis
    • Candidatus Rickettsia rara
    • Candidatus Rickettsia rioja
    • Candidatus Rickettsia senegalensis
    • Candidatus Rickettsia siciliensis
    • Candidatus Rickettsia tasmanensis
    • Candidatus Rickettsia uralica
    • Candidatus Rickettsia wissemanii
    • Rickettsia aeschlimannii [3]
    • Rickettsia africae [4]
    • Rickettsia akari [1]
    • Rickettsia amblyommatis
    • Rickettsia argasii
    • Rickettsia asembonensis
    • Rickettsia asiatica [5]
    • Rickettsia australis [1]
    • Rickettsia buchneri
    • Rickettsia conorii
      • R. conorii subsp. caspia
      • R. conorii subsp. conorii
      • R. conorii subsp. indica
      • R. conorii subsp. israelensis
    • Rickettsia cooleyi [6]
    • Rickettsia felis [7]
    • Rickettsia fournieri
    • Rickettsia gravesii
    • Rickettsia heilongjiangensis
    • Rickettsia helvetica
    • Rickettsia honei
      • R. honei subsp. marmionii
    • Rickettsia hoogstraalii
    • Rickettsia hulinensis
    • Rickettsia hulinii
    • Rickettsia japonica
    • Rickettsia lusitaniae
    • Rickettsia marmionii
    • Rickettsia martinet
    • Rickettsia massiliae
    • Rickettsia monacensis
    • Rickettsia montanensis
    • Rickettsia moreli
    • Rickettsia parkeri [1]
    • Rickettsia peacockii
    • Rickettsia philipii
    • Rickettsia raoultii
    • Rickettsia rhipicephali
    • Rickettsia rickettsii [1]
    • Rickettsia sibirica subgroup
      • Rickettsia sibirica [1]
    • Rickettsia slovaca
    • Rickettsia tamurae
    • Rickettsia vini
  • typhus group
    • Rickettsia prowazekii [1]
    • Rickettsia typhi [1]
  • Rickettsia incertae sedis
    • Rickettsia limoniae
    • Candidatus Rickettsia colombianensi
    • Candidatus Rickettsia laoensis
    • Candidatus Rickettsia liberiensis
    • Candidatus Rickettsia mahosotii

Properly, Rickettsia is the name of a single genus, but the informal term "rickettsia", plural "rickettsias", usually not capitalised, commonly applies to any members of the order Rickettsiales. Being obligate intracellular parasites, rickettsias depend on entry, growth, and replication within the cytoplasm of living eukaryotic host cells (typically endothelial cells).[9] Accordingly, Rickettsia species cannot grow in artificial nutrient culture; they must be grown either in tissue or embryo cultures; typically, chicken embryos are used, following a method developed by Ernest William Goodpasture and his colleagues at Vanderbilt University in the early 1930s.

Rickettsia species are transmitted by numerous types of arthropod, including chiggerticksfleas, and lice, and are associated with both human and plant diseases.[10] Most notably, Rickettsia species are the pathogens responsible for typhusrickettsialpoxboutonneuse feverAfrican tick-bite feverRocky Mountain spotted feverFlinders Island spotted fever, and Queensland tick typhus (Australian tick typhus).[11] The majority of Rickettsia bacteria are susceptible to antibiotics of the tetracycline group.

ClassificationEdit

The classification of Rickettsia into three groups (spotted fever, typhus, and scrub typhus) was initially based on serology. This grouping has since been confirmed by DNA sequencing. All three of these groups include human pathogens. The scrub typhus group has been reclassified as a related new genus, Orientia, but they still are in the order Rickettsiales and accordingly still are grouped with the rest of the rickettsial diseases.

Rickettsias are more widespread than previously believed and are known to be associated with arthropodsleeches, and protists. Divisions have also been identified in the spotted fever group and this group likely should be divided into two clades.[12] Arthropod-inhabiting rickettsiae are generally associated with reproductive manipulation (such as parthenogenesis) to persist in host lineage[10]

In March 2010, Swedish researchers reported a case of bacterial meningitis in a woman caused by Rickettsia helvetica previously thought to be harmless.[13]

Spotted fever groupEdit

  • Rickettsia rickettsii (Western Hemisphere)
Rocky Mountain spotted fever
  • Rickettsia akari (USA, former Soviet Union)
Rickettsialpox
  • Rickettsia conorii (Mediterranean countries, Africa, Southwest Asia, India)
Boutonneuse fever
  • Rickettsia sibirica (Siberia, Mongolia, northern China)
Siberian tick typhus or North Asian tick typhus
  • Rickettsia australis (Australia)
Australian tick typhus
  • Rickettsia felis (North and South America, Southern Europe, Australia)
Flea-borne spotted fever
  • Rickettsia japonica (Japan)
Oriental spotted fever
  • Rickettsia africae (South Africa)
African tick bite fever
  • Rickettsia hoogstraalii (Croatia, Spain and Georgia USA)[14]
Unknown pathogenicity

Typhus groupEdit

  • Rickettsia prowazekii (worldwide)
Epidemic typhus, recrudescent typhus, and sporadic typhus
  • Rickettsia typhi (worldwide)
Murine typhus (endemic typhus)

Scrub typhus groupEdit

  • The causative agent of scrub typhus formerly known as R. tsutsugamushi has been reclassified into the genus Orientia.
Schematic ribosomal RNA phylogeny of Alphaproteobacteria
  Magnetococcidae  

  Magnetococcus marinus

  Caulobacteridae  

  RhodospirillalesSphingomonadales,
  RhodobacteraceaeRhizobialesetc.

  Holosporales

  Rickettsidae  
  Pelagibacterales  
  Pelagibacteraceae  

  Pelagibacter

  Subgroups Ib, II, IIIa, IIIb, IV and V

  Rickettsiales  

  Proto-mitochondria

  Anaplasmataceae  

  Ehrlichia

  Anaplasma

  Wolbachia

  Neorickettsia

  Midichloriaceae  

  Midichloria

  Rickettsiaceae  

  Rickettsia

  Orientia

The cladogram of Rickettsidae has been inferred by Ferla et al. [15] from the comparison of 16S + 23S ribosomal RNA sequences.

Flora and fauna pathogenesisEdit

Plant diseases have been associated with these Rickettsia-like organisms (RLOs):[16]

  • Beet latent rosette RLO
  • Citrus greening bacterium possibly this citrus greening disease
  • Clover leaf RLO
  • Grapevine infectious necrosis RLO
  • Grapevine Pierce's RLO
  • Grapevine yellows RLO
  • Witch's broom disease on Larix spp.
  • Peach phony RLO
  • Papaya Bunchy Top Disease [17]

Infection occurs in nonhuman mammals; for example, species of Rickettsia have been found to afflict the South American guanacoLama guanacoe.[18]

PathophysiologyEdit

Rickettsial organisms are obligate intracellular parasites and invade vascular endothelial cells in target organs, damaging them and producing increased vascular permeability with consequent oedemahypotension, and hypoalbuminaemia.[19]

GenomicsEdit

Certain segments of rickettsial genomes resemble those of mitochondria.[20] The deciphered genome of R. prowazekii is 1,111,523 bp long and contains 834 genes.[21] Unlike free-living bacteria, it contains no genes for anaerobic glycolysis or genes involved in the biosynthesis and regulation of amino acids and nucleosides. In this regard, it is similar to mitochondrial genomes; in both cases, nuclear (host) resources are used.

ATP production in Rickettsia is the same as that in mitochondria. In fact, of all the microbes known, the Rickettsia is probably the closest relative (in a phylogenetic sense) to the mitochondria. Unlike the latter, the genome of R. prowazekii, however, contains a complete set of genes encoding for the tricarboxylic acid cycle and the respiratory chain complex. Still, the genomes of the Rickettsia, as well as the mitochondria, are frequently said to be "small, highly derived products of several types of reductive evolution".

The recent discovery of another parallel between Rickettsia and viruses may become a basis for fighting HIV infection.[22] Human immune response to the scrub typhus pathogen, Orientia tsutsugamushi, appears to provide a beneficial effect against HIV infection progress, negatively influencing the virus replication process. A probable reason for this actively studied phenomenon is a certain degree of homology between the rickettsiae and the virus, namely, common epitope(s) due to common genome fragment(s) in both pathogens. Surprisingly, the other infection reported to be likely to provide the same effect (decrease in viral load) is the virus-caused illness dengue fever.

Comparative analysis of genomic sequences have also identified five conserved signature indels in important proteins, which are uniquely found in members of the genus Rickettsia. These indels consist of a four-amino-acid insertion in transcription repair coupling factor Mfd, a 10-amino-acid insertion in ribosomal protein L19, a one-amino-acid insertion in FtsZ, a one-amino-acid insertion in major sigma factor 70, and a one-amino-acid deletion in exonuclease VII. These indels are all characteristic of the genus and serve as molecular markers for Rickettsia.[23]

Bacterial small RNAs play critical roles in virulence and stress/adaptation responses. Although their specific functions have not been discovered in Rickettsia, few studies showed the expression of novel sRNA in human microvascular endothelial cells (HMEC) infected with Rickettsia.[24][25]

NamingEdit

The genus Rickettsia is named after Howard Taylor Ricketts (1871–1910), who studied Rocky Mountain spotted fever in the Bitterroot Valley of Montana, and eventually died of typhus after studying that disease in Mexico City.


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