Rhodococcus equi

Last updated

Rhodococcus equi
Rhodococcus species.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Bacteria
Phylum: Actinomycetota
Class: Actinomycetia
Order: Mycobacteriales
Family: Nocardiaceae
Genus: Rhodococcus
Species:
R. equi
Binomial name
Rhodococcus equi
(Magnusson 1923) Goodfellow and Alderson 1977 (Approved Lists 1980)

Rhodococcus equi is a Gram-positive coccobacillus bacterium. The organism is commonly found in dry and dusty soil and can be important for diseases of domesticated animals (horses and goats). The frequency of infection can reach near 60%. [1] R. equi is an important pathogen causing pneumonia in foals. Since 2008, R. equi has been known to infect wild boar and domestic pigs. [2] R. equi can infect immunocompromised people, such as HIV-AIDS patients or transplant recipients. Rhodococcus infection in these patients resemble clinical and pathological signs of pulmonary tuberculosis. It is a facultative intracellular mycobacterial pathogen. [3]

Contents

Hosts

Virulence

The most common route of infection in horses is likely via inhalation of contaminated dust particles. Inhaled virulent strains of R. equi are phagocytosed by alveolar macrophages. During normal phagocytosis, bacteria are enclosed by the phagosome, which fuses with the lysosome to become a phagolysosome. The internal environment of the phagolysosome contains nucleases and proteases, which are activated by the low pH of the compartment. The macrophage produces bacteriocidal compounds (e.g., oxygen radicals) following the respiratory burst. However, like its close relative Mycobacterium tuberculosis , R. equi prevents the fusion of the phagosome with the lysosome and acidification of the phagosome. [4] [5] [6] Additionally, the respiratory burst is inhibited. This allows R. equi to multiply within the phagosome where it is shielded from the immune system by the very cell that was supposed to kill it. [7] After about 48 hours, the macrophage is killed by necrosis, not apoptosis. [8] Necrosis is pro-inflammatory, attracting additional phagocytic cells to the site of infection, eventually resulting in massive tissue damage.[ citation needed ]

Virulence plasmid

All strains isolated from foals and the majority of human, cattle, and pig isolates contain a large plasmid. This plasmid has been shown to be essential for infection of foals, and presumably plays a similar role for infection of other hosts, although this has not been established yet. Strains that lack the virulence plasmid are unable to proliferate in macrophages. This virulence plasmid has been characterised in detail from equine and porcine strains, although only the former has been functionally characterised. [9] [10] These circular plasmids consist of a conserved backbone responsible for replication and bacterial conjugation of the plasmid. This portion of the plasmid is highly conserved and found in nonpathogenic Rhodococci plasmids. In addition to the conserved region, the virulence plasmids contain a highly variable region that has undergone substantial genetic rearrangements, including inversion and deletions. This region has a different GC-content from the rest of the plasmid, and is flanked by genes associated with mobile genetic elements. It is therefore assumed to be derived from a different bacterial species than the backbone of the plasmid via lateral gene transfer.[ citation needed ]

Pathogenicity island

The variable region of the virulence plasmid contain genes that are highly expressed following phagocytosis of R. equi by macrophages. [11] This variable region is believed to be a pathogenicity island that contains genes essential for virulence.

A hallmark of the pathogenicity island (PAI) is that many genes within it do not have homologues in other species. The most notable of these are the virulence-associated protein (vap) genes. All foals infected with R. equi produce high levels of antibodies specific for vapA, the first vap gene to be characterised. Deletion of vapA renders the resulting strain avirulent. [12] In addition to vapA, the PAI encodes a further five full-length vap homologues, one truncated vap gene, and two vap pseudogenes. The porcine PAI contains five full-length vap genes, including the vapA homologue, vapB. In addition to these unique genes, the PAI contains genes that have a known function, in particular two regulatory genes encoding the LysR-type regulator VirR and the response regulator Orf8. These two proteins have been shown to control expression of a number of PAI genes including vapA. [13] Other genes have homology to transport proteins and enzymes. However, the functionality of these genes or how the proteins encoded within PAI subvert the macrophage has not yet been established.[ citation needed ]

Taxonomic debate

While this organism is generally known as Rhodococcus equi, there has been taxonomic debate since the 1980s [14] about whether this name is the valid name, with Rhodococcus hoagii and Prescottella equi both proposed as official alternative names. [15] Other names used include Nocardia restricta, [14] Corynebacterium equi, [16] Bacillus hoagii, [16] Corynebacterium purulentus, [16] Mycobacterium equi, [16] Mycobacterium restrictum, [16] and Proactinomyces restrictus. [16]

Related Research Articles

<i>Candida albicans</i> Species of fungus

Candida albicans is an opportunistic pathogenic yeast that is a common member of the human gut flora. It can also survive outside the human body. It is detected in the gastrointestinal tract and mouth in 40–60% of healthy adults. It is usually a commensal organism, but it can become pathogenic in immunocompromised individuals under a variety of conditions. It is one of the few species of the genus Candida that cause the human infection candidiasis, which results from an overgrowth of the fungus. Candidiasis is, for example, often observed in HIV-infected patients. C. albicans is the most common fungal species isolated from biofilms either formed on (permanent) implanted medical devices or on human tissue. C. albicans, C. tropicalis, C. parapsilosis, and C. glabrata are together responsible for 50–90% of all cases of candidiasis in humans. A mortality rate of 40% has been reported for patients with systemic candidiasis due to C. albicans. By one estimate, invasive candidiasis contracted in a hospital causes 2,800 to 11,200 deaths yearly in the US. Nevertheless, these numbers may not truly reflect the true extent of damage this organism causes, given new studies indicating that C. albicans can cross the blood–brain barrier in mice.

<i>Cryptococcus neoformans</i> Species of yeast

Cryptococcus neoformans is an encapsulated yeast belonging to the class Tremellomycetes and an obligate aerobe that can live in both plants and animals. Its teleomorph is a filamentous fungus, formerly referred to Filobasidiella neoformans. In its yeast state, it is often found in bird excrement. Cryptococcus neoformans can cause disease in apparently immunocompetent, as well as immunocompromised, hosts.

<i>Rickettsia rickettsii</i> Species of bacterium

Rickettsia rickettsii is a Gram-negative, intracellular, coccobacillus bacterium that was first discovered in 1902. Having a reduced genome, the bacterium harvests nutrients from its host cell to carry out respiration, making it an organoheterotroph. Maintenance of its genome is carried out through vertical gene transfer where specialization of the bacterium allows it to shuttle host sugars directly into its TCA cycle.

<i>Coxiella burnetii</i> Species of bacterium

Coxiella burnetii is an obligate intracellular bacterial pathogen, and is the causative agent of Q fever. The genus Coxiella is morphologically similar to Rickettsia, but with a variety of genetic and physiological differences. C. burnetii is a small Gram-negative, coccobacillary bacterium that is highly resistant to environmental stresses such as high temperature, osmotic pressure, and ultraviolet light. These characteristics are attributed to a small cell variant form of the organism that is part of a biphasic developmental cycle, including a more metabolically and replicatively active large cell variant form. It can survive standard disinfectants, and is resistant to many other environmental changes like those presented in the phagolysosome.

Pathogenicity islands (PAIs), as termed in 1990, are a distinct class of genomic islands acquired by microorganisms through horizontal gene transfer. Pathogenicity islands are found in both animal and plant pathogens. Additionally, PAIs are found in both gram-positive and gram-negative bacteria. They are transferred through horizontal gene transfer events such as transfer by a plasmid, phage, or conjugative transposon. Therefore, PAIs contribute to microorganisms' ability to evolve.

<i>Corynebacterium diphtheriae</i> Species of prokaryote

Corynebacterium diphtheriae is a gram-positive pathogenic bacterium that causes diphtheria. It is also known as the Klebs–Löffler bacillus because it was discovered in 1884 by German bacteriologists Edwin Klebs (1834–1912) and Friedrich Löffler (1852–1915). The bacteria are usually harmless unless they are infected by a bacteriophage that carries a gene that gives rise to a toxin. This toxin causes the disease. Diphtheria is caused by the adhesion and infiltration of the bacteria into the mucosal layers of the body, primarily affecting the respiratory tract and the subsequent release of an exotoxin. The toxin has a localized effect on skin lesions, as well as a metastatic, proteolytic effects on other organ systems in severe infections. Originally a major cause of childhood mortality, diphtheria has been almost entirely eradicated due to the vigorous administration of the diphtheria vaccination in the 1910s.

<i>Aspergillus fumigatus</i> Species of fungus

Aspergillus fumigatus is a species of fungus in the genus Aspergillus, and is one of the most common Aspergillus species to cause disease in individuals with an immunodeficiency.

<i>Francisella tularensis</i> Species of bacterium

Francisella tularensis is a pathogenic species of Gram-negative coccobacillus, an aerobic bacterium. It is nonspore-forming, nonmotile, and the causative agent of tularemia, the pneumonic form of which is often lethal without treatment. It is a fastidious, facultative intracellular bacterium, which requires cysteine for growth. Due to its low infectious dose, ease of spread by aerosol, and high virulence, F. tularensis is classified as a Tier 1 Select Agent by the U.S. government, along with other potential agents of bioterrorism such as Yersinia pestis, Bacillus anthracis, and Ebola virus. When found in nature, Francisella tularensis can survive for several weeks at low temperatures in animal carcasses, soil, and water. In the laboratory, F. tularensis appears as small rods, and is grown best at 35–37 °C.

<i>Vibrio parahaemolyticus</i> Species of bacterium

Vibrio parahaemolyticus (V. parahaemolyticus) is a curved, rod-shaped, Gram-negative bacterial species found in the sea and in estuaries which, when ingested, may cause gastrointestinal illness in humans. V. parahaemolyticus is oxidase positive, facultatively aerobic, and does not form spores. Like other members of the genus Vibrio, this species is motile, with a single, polar flagellum.

<i>Brucella suis</i> Bacterium that causes swine brucellosis

Brucella suis is a bacterium that causes swine brucellosis, a zoonosis that affects pigs. The disease typically causes chronic inflammatory lesions in the reproductive organs of susceptible animals or orchitis, and may even affect joints and other organs. The most common symptom is abortion in pregnant susceptible sows at any stage of gestation. Other manifestations are temporary or permanent sterility, lameness, posterior paralysis, spondylitis, and abscess formation. It is transmitted mainly by ingestion of infected tissues or fluids, semen during breeding, and suckling infected animals.

<i>Shigella flexneri</i> Species of bacterium

Shigella flexneri is a species of Gram-negative bacteria in the genus Shigella that can cause diarrhea in humans. Several different serogroups of Shigella are described; S. flexneri belongs to group B. S. flexneri infections can usually be treated with antibiotics, although some strains have become resistant. Less severe cases are not usually treated because they become more resistant in the future. Shigella are closely related to Escherichia coli, but can be differentiated from E.coli based on pathogenicity, physiology and serology.

<i>Yersinia pseudotuberculosis</i> Species of bacterium

Yersinia pseudotuberculosis is a Gram-negative bacterium that causes Far East scarlet-like fever in humans, who occasionally get infected zoonotically, most often through the food-borne route. Animals are also infected by Y. pseudotuberculosis. The bacterium is urease positive.

<span class="mw-page-title-main">Phagolysosome</span> Cytoplasmic body

In biology, a phagolysosome, or endolysosome, is a cytoplasmic body formed by the fusion of a phagosome with a lysosome in a process that occurs during phagocytosis. Formation of phagolysosomes is essential for the intracellular destruction of microorganisms and pathogens. It takes place when the phagosome's and lysosome's membranes 'collide', at which point the lysosomal contents—including hydrolytic enzymes—are discharged into the phagosome in an explosive manner and digest the particles that the phagosome had ingested. Some products of the digestion are useful materials and are moved into the cytoplasm; others are exported by exocytosis.

<i>Rhodococcus</i> Genus of bacteria

Rhodococcus is a genus of aerobic, nonsporulating, nonmotile Gram-positive bacteria closely related to Mycobacterium and Corynebacterium. While a few species are pathogenic, most are benign, and have been found to thrive in a broad range of environments, including soil, water, and eukaryotic cells. Some species have large genomes, including the 9.7 megabasepair genome of Rhodococcus sp. RHA1.

<i>Rhodococcus fascians</i> Species of bacterium

Rhodococcus fascians is a Gram positive bacterial phytopathogen that causes leafy gall disease. R. fascians is the only phytopathogenic member of the genus Rhodococcus; its host range includes both dicotyledonous and monocotyledonous hosts. Because it commonly afflicts tobacco (Nicotiana) plants, it is an agriculturally significant pathogen.

Listeriolysin O (LLO) is a hemolysin produced by the bacterium Listeria monocytogenes, the pathogen responsible for causing listeriosis. The toxin may be considered a virulence factor, since it is crucial for the virulence of L. monocytogenes.

<i>Streptococcus zooepidemicus</i> Species of bacterium

Streptococcus zooepidemicus is a Lancefield group C streptococcus that was first isolated in 1934 by P. R. Edwards, and named Animal pyogens A. It is a mucosal commensal and opportunistic pathogen that infects several animals and humans, but most commonly isolated from the uterus of mares. It is a subspecies of Streptococcus equi, a contagious upper respiratory tract infection of horses, and shares greater than 98% DNA homology, as well as many of the same virulence factors.

<i>Salmonella enterica <span style="font-style:normal;">subsp.</span> enterica</i> Subspecies of bacterium

Salmonella enterica subsp. enterica is a subspecies of Salmonella enterica, the rod-shaped, flagellated, aerobic, Gram-negative bacterium. Many of the pathogenic serovars of the S. enterica species are in this subspecies, including that responsible for typhoid.

<i>Vibrio anguillarum</i> Species of bacterium

Vibrio anguillarum is a species of prokaryote that belongs to the family Vibrionaceae, genus Vibrio. V. anguillarum is typically 0.5 - 1 µm in diameter and 1 - 3 µm in length. It is a gram-negative, comma-shaped rod bacterium that is commonly found in seawater and brackish waters. It is polarly flagellated, non-spore-forming, halophilic, and facultatively anaerobic. V. anguillarum has the ability to form biofilms. V. anguillarum is pathogenic to various fish species, crustaceans, and mollusks.

Listeria ivanovii is a species of bacteria in the genus Listeria. The listeria are rod-shaped bacteria, do not produce spores, and become positively stained when subjected to Gram staining. Of the six bacteria species within the genus, L. ivanovii is one of the two pathogenic species. In 1955 Bulgaria, the first known isolation of this species was found from sheep. It behaves like L. monocytogenes, but is found almost exclusively in ruminants. The species is named in honor of Bulgarian microbiologist Ivan Ivanov. This species is facultatively anaerobic, which makes it possible for it to go through fermentation when there is oxygen depletion.

References

  1. Muscatello, G; Leadon, DP; Klayt, M; Ocampo-Sosa, A; Lewis, DA; Fogarty, U; Buckley, T; Gilkerson, JR; Meijer, WG; Vazquez-Boland, JA (September 2007). "Rhodococcus equi infection in foals: the science of 'rattles'". Equine Veterinary Journal. 39 (5): 470–8. doi:10.2746/042516407X209217. PMID   17910275.
  2. Makrai, L; Kobayashi, A; Matsuoka, M; Sasaki, Y; Kakuda, T; Dénes, B; Hajtós, I; Révész, I; Jánosi, K; Fodor, L; Varga, J; Takai, S (15 October 2008). "Isolation and characterisation of Rhodococcus equi from submaxillary lymph nodes of wild boars (Sus scrofa)". Veterinary Microbiology. 131 (3–4): 318–23. doi:10.1016/j.vetmic.2008.04.009. PMID   18499361.
  3. Kelly, B. G.; Wall, D. M.; Boland, C. A.; Meijer, W. G. (2002). "Isocitrate lyase of the facultative intracellular pathogen Rhodococcus equi". Microbiology. 148 (Pt 3): 793–798. doi: 10.1099/00221287-148-3-793 . PMID   11882714.
  4. von Bargen, K; Polidori, M; Becken, U; Huth, G; Prescott, JF; Haas, A (December 2009). "Rhodococcus equi virulence-associated protein A is required for diversion of phagosome biogenesis but not for cytotoxicity". Infection and Immunity. 77 (12): 5676–81. doi:10.1128/IAI.00856-09. PMC   2786453 . PMID   19797071.
  5. Fernandez-Mora, E; Polidori, M; Lührmann, A; Schaible, UE; Haas, A (August 2005). "Maturation of Rhodococcus equi-containing vacuoles is arrested after completion of the early endosome stage". Traffic. 6 (8): 635–53. doi: 10.1111/j.1600-0854.2005.00304.x . PMID   15998320. S2CID   30122137.
  6. Sydor, T; von Bargen, K; Hsu, FF; Huth, G; Holst, O; Wohlmann, J; Becken, U; Dykstra, T; Söhl, K; Lindner, B; Prescott, JF; Schaible, UE; Utermöhlen, O; Haas, A (March 2013). "Diversion of phagosome trafficking by pathogenic Rhodococcus equi depends on mycolic acid chain length". Cellular Microbiology. 15 (3): 458–73. doi:10.1111/cmi.12050. PMC   3864644 . PMID   23078612.
  7. Hondalus, MK; Mosser, DM (October 1994). "Survival and replication of Rhodococcus equi in macrophages". Infection and Immunity. 62 (10): 4167–75. doi:10.1128/IAI.62.10.4167-4175.1994. PMC   303092 . PMID   7927672.
  8. Lührmann, A; Mauder, N; Sydor, T; Fernandez-Mora, E; Schulze-Luehrmann, J; Takai, S; Haas, A (February 2004). "Necrotic death of Rhodococcus equi-infected macrophages is regulated by virulence-associated plasmids". Infection and Immunity. 72 (2): 853–62. doi:10.1128/iai.72.2.853-862.2004. PMC   321572 . PMID   14742529.
  9. Letek, M; Ocampo-Sosa, AA; Sanders, M; Fogarty, U; Buckley, T; Leadon, DP; González, P; Scortti, M; Meijer, WG; Parkhill, J; Bentley, S; Vázquez-Boland, JA (September 2008). "Evolution of the Rhodococcus equi vap pathogenicity island seen through comparison of host-associated vapA and vapB virulence plasmids". Journal of Bacteriology. 190 (17): 5797–805. doi:10.1128/JB.00468-08. PMC   2519538 . PMID   18606735.
  10. Takai, S; Hines, SA; Sekizaki, T; Nicholson, VM; Alperin, DA; Osaki, M; Takamatsu, D; Nakamura, M; Suzuki, K; Ogino, N; Kakuda, T; Dan, H; Prescott, JF (December 2000). "DNA sequence and comparison of virulence plasmids from Rhodococcus equi ATCC 33701 and 103". Infection and Immunity. 68 (12): 6840–7. doi:10.1128/iai.68.12.6840-6847.2000. PMC   97788 . PMID   11083803.
  11. Ren, J; Prescott, JF (1 July 2003). "Analysis of virulence plasmid gene expression of intra-macrophage and in vitro grown Rhodococcus equi ATCC 33701". Veterinary Microbiology. 94 (2): 167–82. doi:10.1016/S0378-1135(03)00099-3. PMID   12781484.
  12. Jain, S; Bloom, BR; Hondalus, MK (October 2003). "Deletion of vapA encoding Virulence Associated Protein A attenuates the intracellular actinomycete Rhodococcus equi". Molecular Microbiology. 50 (1): 115–28. doi: 10.1046/j.1365-2958.2003.03689.x . PMID   14507368. S2CID   42313934.
  13. Russell, DA; Byrne, GA; O'Connell, EP; Boland, CA; Meijer, WG (September 2004). "The LysR-type transcriptional regulator VirR is required for expression of the virulence gene vapA of Rhodococcus equi ATCC 33701". Journal of Bacteriology. 186 (17): 5576–84. doi:10.1128/JB.186.17.5576-5584.2004. PMC   516814 . PMID   15317761.
  14. 1 2 Garrity, GM (January 2014). "Conservation of Rhodococcus equi (Magnusson 1923) Goodfellow and Alderson 1977 and rejection of Corynebacterium hoagii (Morse 1912) Eberson 1918". International Journal of Systematic and Evolutionary Microbiology. 64 (Pt 1): 311–2. doi:10.1099/ijs.0.059741-0. PMID   24408953. Open Access logo PLoS transparent.svg
  15. Goodfellow, M; Sangal, V; Jones, AL; Sutcliffe, IC (September 2015). "Charting stormy waters: A commentary on the nomenclature of the equine pathogen variously named Prescottella equi, Rhodococcus equi and Rhodococcus hoagii". Equine Veterinary Journal. 47 (5): 508–509. doi: 10.1111/evj.12399 . PMID   25912143.
  16. 1 2 3 4 5 6 Berman, Jules J. (2012). Taxonomic guide to infectious diseases : understanding the biologic classes of pathogenic organisms. London: Elsevier/Academic Press. p. 266. ISBN   978-0-12-415895-5.

Further reading