Biology:Burkholderia pseudomallei

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Short description: Species of bacterium

Burkholderia pseudomallei
Bps close.JPG
Burkholderia pseudomallei colonies on Ashdown's agar showing the characteristic cornflower head morphology
Scientific classification edit
Domain: Bacteria
Phylum: Pseudomonadota
Class: Betaproteobacteria
Order: Burkholderiales
Family: Burkholderiaceae
Genus: Burkholderia
Species:
B. pseudomallei
Binomial name
Burkholderia pseudomallei
(Whitmore 1913)
Yabuuchi et al. 1993[1]
Synonyms

Bacillus pseudomallei Whitmore 1913
Bacterium whitmori Stanton and Fletcher 1921
Malleomyces pseudomallei Breed 1939
Loefflerella pseudomallei Brindle and Cowan 1951
Pfeiferella pseudomallei
Pseudomonas pseudomallei (Whitmore 1913) Haynes 1957

Burkholderia pseudomallei (also known as Pseudomonas pseudomallei) is a Gram-negative, bipolar, aerobic, motile rod-shaped bacterium.[2] It is a soil-dwelling bacterium endemic in tropical and subtropical regions worldwide, particularly in Thailand and northern Australia.[3] It was reported in 2008 that there had been an expansion of the affected regions due to significant natural disasters, and it could be found in Southern China, Hong Kong, and countries in America.[4] B. pseudomallei, amongst other pathogens, has been found in monkeys imported into the United States from Asia for laboratory use, posing a risk that the pathogen could be introduced into the country.[5]

Although it is mainly a soil-dwelling bacteria, a study performed by Apinya Pumpuang and others showed that Burkholderia pseudomallei survived in distilled water for 16 years, demonstrating that it is capable of living in water if a specific environment is provided.[6] It is resistant to a variety of harsh conditions including nutrient deficiency, extreme temperature or pH.[7] It infects humans, causing the disease melioidosis;[8] mortality is 20–50% even with treatment. The CDC classifies it as a "Tier 1 select agent" with potential as a bioterrorism agent.[5] It infects other animals, most commonly livestock such as goats, pigs, and sheep, less frequently.[9] It is also capable of infecting plants in a laboratory setting.[10]

Burkholderia pseudomallei measures 2–5 μm in length and 0.4–0.8 μm in diameter and is capable of self-propulsion using flagella. The bacteria can grow in a number of artificial nutrient environments, especially betaine- and arginine-containing ones.

In vitro, optimal proliferation temperature is reported around 40 °C in neutral or slightly acidic environments (pH 6.8–7.0). The majority of strains are capable of oxidation, not fermentation, of sugars without gas formation (most importantly, glucose and galactose; older cultures are reported to also metabolize maltose and starch). Bacteria produce both exo- and endotoxins. The role of the toxins identified in the process of melioidosis symptom development has not been fully elucidated.[11]

Identification

Burkholderia pseudomallei is not fastidious and grows on a large variety of culture media (blood agar, MacConkey agar, EMB, etc.). Ashdown's medium (or Burkholderia cepacia medium) may be used for selective isolation.[12] Cultures typically become positive in 24 to 48 hours (this rapid growth rate differentiates the organism from B. mallei, which typically takes a minimum of 72 hours to grow). Colonies are wrinkled, have a metallic appearance, and possess an earthy odor. On Gram staining, the organism is a Gram-negative rod with a characteristic "safety pin" appearance (bipolar staining). On sensitivity testing, the organism appears highly resistant (it is innately resistant to many antibiotics including colistin and gentamicin) and that again differentiates it from B. mallei, which is in contrast, exquisitely sensitive to many antibiotics. For environmental specimens only, differentiation from the nonpathogenic B. thailandensis using an arabinose test is necessary (B. thailandensis is never isolated from clinical specimens).[13] The laboratory identification of B. pseudomallei has been described in the literature.[14]

The classic textbook description of B. pseudomallei in clinical samples is of an intracellular, bipolar-staining, Gram-negative rod, but this is of little value in identifying the organism from clinical samples.[14] Some[15] suggest the Wayson stain is useful for this purpose, but this has been shown not to be the case.[16]

Laboratory identification of B. pseudomallei can be difficult, especially in Western countries where it is rarely seen. The large, wrinkled colonies look like environmental contaminants, so are often discarded as being of no clinical significance. Colony morphology is very variable and a single strain may display multiple colony types,[17][18] so inexperienced laboratory staff may mistakenly believe the growth is not pure. The organism grows more slowly than other bacteria that may be present in clinical specimens, and in specimens from nonsterile sites, is easily overgrown. Nonsterile specimens should, therefore, be cultured in selective media (e.g., Ashdown's[19][20] or B. cepacia medium).[12] For heavily contaminated samples, such as feces, a modified version of Ashdown's that includes norfloxacin, amoxicillin, and polymyxin B has been proposed.[21] In blood culture, the BacT/ALERT MB system (normally used for culturing mycobacteria) by bioMérieux has been shown to have superior yields compared to conventional blood culture media.[22]

Even when the isolate is recognized to be significant, commonly used identification systems may misidentify the organism as Chromobacterium violaceum or other nonfermenting, Gram-negative bacilli such as Burkholderia cepacia or Pseudomonas aeruginosa.[23][24] Again, because the disease is rarely seen in Western countries, identification of B. pseudomallei in cultures may not actually trigger alarms in physicians unfamiliar with the disease.[25] Routine biochemical methods for identification of bacteria vary widely in their identification of this organism: the API 20NE system accurately identifies B. pseudomallei in 99% of cases,[26] as does the automated Vitek 1 system, but the automated Vitek 2 system only identifies 19% of isolates.[24]

The pattern of resistance to antimicrobials is distinctive, and helps to differentiate the organism from P. aeruginosa. The majority of B. pseudomallei isolates are intrinsically resistant to all aminoglycosides (via an efflux pump mechanism),[27] but sensitive to co-amoxiclav:[28] this pattern of resistance almost never occurs in P. aeruginosa and is helpful in identification.[29] Unfortunately, the majority of strains in Sarawak, Borneo, are susceptible to aminoglycosides and macrolides, which means the conventional recommendations for isolation and identification do not apply there.[30]

Molecular methods (PCR) of diagnosis are possible, but not routinely available for clinical diagnosis.[31][32] Fluorescence in situ hybridisation has also been described, but has not been clinically validated, and it is not commercially available.[33] In Thailand, a latex agglutination assay is widely used,[26] while a rapid immunofluorescence technique is also available in a small number of centres.[34]

Characteristics

Morphological, physiological, and biochemical characteristics of Burkholderia pseudomallei are shown in the Table below.

Test type Test Characteristics
Colony characters Size 2–5 μm in length and 0.4–0.8 μm in diameter
Type Round
Color Whitish
Shape Multiple
Morphological characters Shape Rod (Variable)
Physiological characters Motility +
Growth at 6.5% NaCl +
Biochemical characters Gram staining -
Oxidase +
Catalase +[35]
Oxidative-Fermentative
Motility +
Methyl Red
Voges-Proskauer
Indole -[36]
H2S Production -
Urease
Nitrate reductase +
β-Galactosidase
Hydrolysis of Gelatin +
Casein
Utilization of Glycerol +
Galactose +
D-Glucose +
D-Fructose +
D-Mannose +
Mannitol Variable

Note: + = Positive, – =Negative

Disinfection

Burkholderia pseudomallei is susceptible to numerous disinfectants, including benzalkonium chloride, iodine, mercuric chloride, potassium permanganate, 1% sodium hypochlorite, 70% ethanol, 2% glutaraldehyde, and to a lesser extent, phenolic preparations.[37] B. pseudomallei is effectively killed by the commercial disinfectants, Perasafe and Virkon.[38] The microorganism can also be destroyed by heating to above 74 °C for 10 min or by ultraviolet irradiation.[39]

Medical importance

Main page: Medicine:Melioidosis

Burkholderia pseudomallei infection in humans is called melioidosis or Whitmore’s disease. It is spread though direct contact with water or soil that holds the bacteria. There have been few cases of transmission of the bacteria perinatally.[40] Its mortality is 20 to 50% even with treatment.[28]

Antibiotic treatment and sensitivity testing

The antibiotic of choice is ceftazidime.[28] While various antibiotics are active in vitro (e.g., chloramphenicol, doxycycline, co-trimoxazole), they have been proven to be inferior in vivo for the treatment of acute melioidosis.[41] Disc diffusion tests are unreliable when looking for co-trimoxazole resistance in B. pseudomallei (they greatly overestimate resistance) and Etests or agar dilution tests should be used in preference.[42][43] The actions of co-trimoxazole and doxycycline are antagonistic, which suggests these two drugs ought not to be used together.[44]

The organism is intrinsically resistant to gentamicin[45] and colistin, and this fact is helpful in the identification of the organism.[46] Kanamycin is used to kill B. pseudomallei in the laboratory, but the concentrations used are much higher than those achievable in humans.[47]

Pathogenicity mechanisms and virulence factors

Burkholderia pseudomallei is an opportunistic pathogen. An environmental organism, it has no requirement to pass through an animal host to replicate. From the point of view of the bacterium, human infection is a developmental "dead end".[48]

Strains which cause disease in humans differ from those causing disease in other animals, by possessing certain genomic islands.[49] It may have the ability to cause disease in humans because of DNA it has acquired from other microorganisms.[49] Its mutation rate is also high, and the organism continues to evolve even after infecting a host.[50]

Burkholderia pseudomallei is able to invade cells (it is an intracellular pathogen).[51] It is able to polymerise actin, and to spread from cell to cell, causing cell fusion and the formation of multinucleated giant cells.[52] It possesses a uniquely fusogenic type VI secretion system that is required for cell-cell spread and virulence in mammalian hosts.[53] The bacterium also expresses a toxin called lethal factor 1.[54] B. pseudomallei is one of the first Proteobacteria to be identified as containing an active type VI secretion system. It is also the only organism identified that contains up to six different type VI secretion systems.[55]

B. pseudomallei is intrinsically resistant to many antimicrobial agents by virtue of its efflux pump mechanism. This mediates resistance to aminoglycosides (AmrAB-OprA), tetracyclines, fluoroquinolones, and macrolides (BpeAB-OprB).[56]

Vaccine candidates

(As of 2023) no vaccine had been licensed, although many had been evaluated in pre-clinical studies.[57][58]

Vaccine candidates have been suggested. Aspartate-β-semialdehyde dehydrogenase (asd) gene deletion mutants are auxotrophic for diaminopimelate (DAP) in rich media and auxotrophic for DAP, lysine, methionine and threonine in minimal media.[59] The Δasd bacterium (bacterium with the asd gene removed) protects against inhalational melioidosis in mice.[60]

Transformation

Burkholderia pseudomoallei can go through transformation. The bacteria is able to uptake a free plasmid using electroporation and the plasmid material will integrate into the host DNA when they are electrocompetent. An example is given by an experiment performed by Mack K. and Titball R. W., which showed that a Burkholderia pseudomallei 4845 was capable of transformation by electroporation, utilizing the incQ plasmid pKT230.[61]

References

  1. ↑ Yabuuchi, E; Kosako, Y; Oyaizu, H; Yano, I; Hotta, H; Hashimoto, Y; Ezaki, T; Arakawa, M (1992). "Proposal of Burkholderia gen. nov. and transfer of seven species of the genus Pseudomonas homology group II to the new genus, with the type species Burkholderia cepacia (Palleroni and Holmes 1981) comb. nov.". Microbiol Immunol 36 (12): 1251–75. doi:10.1111/j.1348-0421.1992.tb02129.x. PMID 1283774. 
  2. ↑ "Burkholderia pseudomallei". VirginiaTech Pathogen Database. http://pathport.vbi.vt.edu/pathinfo/pathogens/Burkholderia_pseudomallei.html. 
  3. ↑ Limmathurotsakul, Direk; Golding, Nick; Dance, David A. B.; Messina, Jane P.; Pigott, David M.; Moyes, Catherine L.; Rolim, Dionne B.; Bertherat, Eric et al. (2016-01-11). "Predicted global distribution of Burkholderia pseudomallei and burden of melioidosis" (in en). Nature Microbiology 1 (1): 15008. doi:10.1038/nmicrobiol.2015.8. ISSN 2058-5276. PMID 26877885. 
  4. ↑ Currie, Bart J.; Dance, David A.B.; Cheng, Allen C. (December 2008). "The global distribution of Burkholderia pseudomallei and melioidosis: an update" (in en). Transactions of the Royal Society of Tropical Medicine and Hygiene 102: S1–S4. doi:10.1016/S0035-9203(08)70002-6. PMID 19121666. https://academic.oup.com/trstmh/article-lookup/doi/10.1016/S0035-9203(08)70002-6. 
  5. ↑ 5.0 5.1 Colley, Claire (18 December 2022). "US public not warned that monkeys imported from Cambodia carried deadly pathogens". The Guardian. https://www.theguardian.com/us-news/2022/dec/18/monkeys-imported-us-from-cambodia-carried-deadly-pathogens. 
  6. ↑ Pumpuang, Apinya; Chantratita, Narisara; Wikraiphat, Chanthiwa; Saiprom, Natnaree; Day, Nicholas P.J.; Peacock, Sharon J.; Wuthiekanun, Vanaporn (October 2011). "Survival of Burkholderia pseudomallei in distilled water for 16 years". Transactions of the Royal Society of Tropical Medicine and Hygiene 105 (10): 598–600. doi:10.1016/j.trstmh.2011.06.004. ISSN 0035-9203. PMID 21764093. PMC 3183224. http://dx.doi.org/10.1016/j.trstmh.2011.06.004. 
  7. ↑ Cheng, Allen C.; Currie, Bart J. (April 2005). "Melioidosis: Epidemiology, Pathophysiology, and Management". Clinical Microbiology Reviews 18 (2): 383–416. doi:10.1128/CMR.18.2.383-416.2005. ISSN 0893-8512. PMID 15831829. 
  8. ↑ "Melioidosis" (in en-us). 2022-07-27. https://www.cdc.gov/melioidosis/index.html. 
  9. ↑ "Melioidosis". New Jersey Department of Agriculture. 2003. https://www.nj.gov/agriculture/divisions/ah/diseases/melioidosis.html#:~:text=Infection%20with%20B.,be%20infected%20in%20the%20laboratory.. 
  10. ↑ "Identification of tomato plant as a novel host model for Burkholderia pseudomallei". BMC Microbiol 10: 28. 2010. doi:10.1186/1471-2180-10-28. PMID 20109238. 
  11. ↑ "Toxin production by Burkholderia pseudomallei strains and correlation with severity of melioidosis". Journal of Medical Microbiology 46 (7): 557–63. July 1997. doi:10.1099/00222615-46-7-557. PMID 9236739. 
  12. ↑ 12.0 12.1 "Comparison of Ashdown's medium, Burkholderia cepacia medium, and Burkholderia pseudomallei selective agar for clinical isolation of Burkholderia pseudomallei". Journal of Clinical Microbiology 43 (10): 5359–61. October 2005. doi:10.1128/JCM.43.10.5359-5361.2005. PMID 16208018. 
  13. ↑ "Differences in genomic macrorestriction patterns of arabinose-positive (Burkholderia thailandensis) and arabinose-negative Burkholderia pseudomallei". Microbiology and Immunology 43 (7): 625–30. 1999. doi:10.1111/j.1348-0421.1999.tb02449.x. PMID 10529102. 
  14. ↑ 14.0 14.1 "The laboratory diagnosis of melioidosis.". Br J Biomed Sci 53 (4): 249–53. 1996. PMID 9069100. 
  15. ↑ "Four fatal cases of melioidosis in U. S. soldiers in Vietnam. Bacteriologic and pathologic characteristics". The American Journal of Tropical Medicine and Hygiene 17 (2): 183–91. March 1968. doi:10.4269/ajtmh.1968.17.183. PMID 4869109. 
  16. ↑ "Evaluation of the Wayson stain for the rapid diagnosis of melioidosis". Journal of Clinical Microbiology 45 (5): 1669–70. May 2007. doi:10.1128/JCM.00396-07. PMID 17360835. 
  17. ↑ "Biological relevance of colony morphology and phenotypic switching by Burkholderia pseudomallei.". J Bacteriol 189 (3): 807–17. 2007. doi:10.1128/JB.01258-06. PMID 17114252. 
  18. ↑ "Survival of Burkholderia pseudomallei in distilled water for 16 years". Trans R Soc Trop Med Hyg 105 (10–2): 598–600. 2011. doi:10.1016/j.trstmh.2011.06.004. PMID 21764093. 
  19. ↑ Ashdown LR (1979). "An improved screening technique for isolation of Pseudomonas pseudomallei from clinical specimens". Pathology 11 (2): 293–7. doi:10.3109/00313027909061954. PMID 460953. 
  20. ↑ Roesnita B; Tay ST; Puthucheary SD; Sam IC. (2012). "Diagnostic use of Burkholderia pseudomallei selective media in a low prevalence setting.". Trans R Soc Trop Med Hyg 106 (2): 131–3. doi:10.1016/j.trstmh.2011.10.007. PMID 22112687. 
  21. ↑ "An improved selective culture medium enhances the isolation of Burkholderia pseudomallei from contaminated specimens.". Am J Trop Med Hyg 89 (5): 973–82. 2013. doi:10.4269/ajtmh.13-0119. PMID 24062483. PMC 3820346. http://www.ajtmh.org/cgi/pmidlookup?view=long&pmid=24062483. 
  22. ↑ "Contribution of the BacT/ALERT MB Mycobacteria Bottle to bloodstream infection surveillance in Thailand: added yield for Burkholderia pseudomallei.". J Clin Microbiol 53 (3): 910–4. 2015. doi:10.1128/JCM.02008-14. PMID 25588650. 
  23. ↑ "Potential misidentification of Burkholderia pseudomallei by API 20NE". Pathology 30 (1): 62–4. February 1998. doi:10.1080/00313029800169685. PMID 9534210. 
  24. ↑ 24.0 24.1 "Comparison of automated and nonautomated systems for identification of Burkholderia pseudomallei". Journal of Clinical Microbiology 40 (12): 4625–7. December 2002. doi:10.1128/JCM.40.12.4625-4627.2002. PMID 12454163. 
  25. ↑ "Imported Melioidosis—South Florida, 2005". Morb Mortal Wkly Rep 55 (32): 873–6. 2006. PMID 16915220. https://www.cdc.gov/mmwr/preview/mmwrhtml/mm5532a1.htm. 
  26. ↑ 26.0 26.1 "Accuracy of Burkholderia pseudomallei identification using the API 20NE system and a latex agglutination test". Journal of Clinical Microbiology 45 (11): 3774–6. November 2007. doi:10.1128/JCM.00935-07. PMID 17804660. 
  27. ↑ "Efflux-mediated aminoglycoside and macrolide resistance in Burkholderia pseudomallei". Antimicrobial Agents and Chemotherapy 43 (3): 465–70. March 1999. doi:10.1128/AAC.43.3.465. PMID 10049252. 
  28. ↑ 28.0 28.1 28.2 "Management of melioidosis". Expert Review of Anti-infective Therapy 4 (3): 445–55. June 2006. doi:10.1586/14787210.4.3.445. PMID 16771621. 
  29. ↑ "A comparison of routine bench and molecular diagnostic methods in the identification of Burkholderia pseudomallei". J Clin Microbiol 47 (5): 1578–80. 2009. doi:10.1128/JCM.02507-08. PMID 19279182. 
  30. ↑ "Burkholderia pseudomallei from Sarawak, Malaysian Borneo are predominantly susceptible to aminoglycosides and macrolides". Antimicrob Agents Chemother 58 (1): 162–6. 2013. doi:10.1128/AAC.01842-13. PMID 24145517. 
  31. ↑ "Suitability of partial 16S ribosomal RNA gene sequence analysis for the identification of dangerous bacterial pathogens". Journal of Applied Microbiology 102 (3): 852–9. March 2007. doi:10.1111/j.1365-2672.2006.03107.x. PMID 17309636. 
  32. ↑ "Identification of Burkholderia pseudomallei and related bacteria by multiple-locus sequence typing-derived PCR and real-time PCR". Journal of Clinical Microbiology 45 (3): 1045–8. March 2007. doi:10.1128/JCM.02350-06. PMID 17251403. 
  33. ↑ "Rapid identification of Burkholderia pseudomallei and Burkholderia mallei by fluorescence in situ hybridization (FISH) from culture and paraffin-embedded tissue samples". International Journal of Medical Microbiology 301 (7): 585–90. 2011. doi:10.1016/j.ijmm.2011.04.017. PMID 21658996. 
  34. ↑ "Rapid immunofluorescence microscopy for diagnosis of melioidosis". Clinical and Diagnostic Laboratory Immunology 12 (4): 555–6. April 2005. doi:10.1128/CDLI.12.4.555-556.2005. PMID 15817767. 
  35. ↑ "Burkholderia Pseudomallei". http://www.antimicrobe.org/ClinicMicro/Burkholderia_pseudomallei1.htm. 
  36. ↑ Hemarajata, Peera; Baghdadi, Jonathan D.; Hoffman, Risa; Humphries, Romney M. (December 2016). "Burkholderia pseudomallei: Challenges for the Clinical Microbiology Laboratory". Journal of Clinical Microbiology 54 (12): 2866–2873. doi:10.1128/JCM.01636-16. ISSN 0095-1137. PMID 27654336. 
  37. ↑ Miller, WR; Pannell, L; Cravitz, L; Tanner, WA; Ingalls, MS (1948). "Studies on certain biological characteristics of Malleomyces mallei and Malleomyces pseudomallei: I. Morphology, cultivation, viability, and isolation from contaminated specimens". J Bacteriol 55 (1): 115–126. doi:10.1128/JB.55.1.115-126.1948. PMID 16561426. 
  38. ↑ "Perasafe, Virkon and bleach are bactericidal for Burkholderia pseudomallei, a select agent and the cause of melioidosis". J Hosp Infect 77 (2): 183–4. 2010. doi:10.1016/j.jhin.2010.06.026. PMID 20832143. 
  39. ↑ Rose, L. J.; O'Connell, H. (2009-05-01). "UV Light Inactivation of Bacterial Biothreat Agents". Applied and Environmental Microbiology 75 (9): 2987–2990. doi:10.1128/AEM.02180-08. ISSN 0099-2240. PMID 19270145. Bibcode: 2009ApEnM..75.2987R. 
  40. ↑ Rodríguez, José Y.; Huertas, Mónica G.; Rodríguez, Gerson J.; Vargas-Otalora, Sandra; Benıtez-Peñuela, Miguel A.; Esquea, Kelin; Rios, Rafael; Mendoza, Laura R. et al. (2020-11-04). "Case Report: Gestational Melioidosis through Perinatal Transmission". The American Journal of Tropical Medicine and Hygiene 103 (5): 1838–1840. doi:10.4269/ajtmh.20-0223. ISSN 0002-9637. PMID 32748772. PMC 7646751. http://dx.doi.org/10.4269/ajtmh.20-0223. 
  41. ↑ "Halving of mortality of severe melioidosis by ceftazidime". Lancet 2 (8665): 697–701. September 1989. doi:10.1016/S0140-6736(89)90768-X. PMID 2570956. 
  42. ↑ "Comparison between the antimicrobial susceptibility of Burkholderia pseudomallei to trimethoprim-sulfamethoxazole by standard disk diffusion method and by minimal inhibitory concentration determination". J Med Assoc Thai 83 (8): 856–60. 2000. PMID 10998837. 
  43. ↑ "Trimethoprim/sulfamethoxazole resistance in clinical isolates of Burkholderia pseudomallei". J Antimicrob Chemother 55 (6): 1029–31. 2005. doi:10.1093/jac/dki151. PMID 15886263. 
  44. ↑ "In vitro interactions between cotrimoxazole and doxycycline in Burkholderia pseudomallei: how important is this combination in maintenance therapy of melioidosis?". Indian J Med Microbiol 27 (1): 88–9. 2009. doi:10.1016/S0255-0857(21)01773-4. PMID 19172079. 
  45. ↑ Trunck LA; Propst, KL; Wuthiekanun, V; Tuanyok, A; Beckstrom-Sternberg, SM; Beckstrom-Sternberg, JS; Peacock, SJ; Keim, P et al. (2009). Picardeau, Mathieu. ed. "Molecular basis of rare aminoglycoside susceptibility and pathogenesis of Burkholderia pseudomallei clinical isolates from Thailand". PLOS Negl Trop Dis 3 (9): e519. doi:10.1371/journal.pntd.0000519. PMID 19771149. 
  46. ↑ Ashdown, LR (1979). "Identification of Pseudomonas pseudomallei in the clinical laboratory". J Clin Pathol 32 (5): 500–4. doi:10.1136/jcp.32.5.500. PMID 381328. 
  47. ↑ "Virulent Burkholderia pseudomallei is more efficient than avirulent Burkholderia thailandensis in invasion of and adherence to cultured human epithelial cells". Microbial Pathogenesis 36 (5): 287–9. 2004. doi:10.1016/j.micpath.2004.01.001. PMID 15043863. 
  48. ↑ Guttman, David S., ed (2010). "A genomic survey of positive selection in Burkholderia pseudomallei provides insights into the evolution of accidental virulence". PLOS Pathog. 6 (4): e1000845. doi:10.1371/journal.ppat.1000845. PMID 20368977. 
  49. ↑ 49.0 49.1 Achtman, Mark, ed (October 2008). "The core and accessory genomes of Burkholderia pseudomallei: implications for human melioidosis". PLOS Pathog. 4 (10): e1000178. doi:10.1371/journal.ppat.1000178. PMID 18927621. 
  50. ↑ Guttman, David S., ed (2010). "Within-host evolution of Burkholderia pseudomallei in four cases of acute melioidosis". PLOS Pathog. 6 (1): e1000725. doi:10.1371/journal.ppat.1000725. PMID 20090837. 
  51. ↑ "Melioidosis: insights into the pathogenicity of Burkholderia pseudomallei.". Nature Reviews Microbiology 4 (4): 272–82. 2006. doi:10.1038/nrmicro1385. PMID 16541135. 
  52. ↑ "Burkholderia pseudomallei induces cell fusion and actin-associated membrane protrusion: a possible mechanism for cell-to-cell spreading". Infect. Immun. 68 (9): 5377–84. 2000. doi:10.1128/IAI.68.9.5377-5384.2000. PMID 10948167. 
  53. ↑ Toesca, Isabelle J.; French, Christopher T.; Miller, Jeff F. (2014-04-01). "The Type VI secretion system spike protein VgrG5 mediates membrane fusion during intercellular spread by pseudomallei group Burkholderia species". Infection and Immunity 82 (4): 1436–1444. doi:10.1128/IAI.01367-13. ISSN 1098-5522. PMID 24421040. 
  54. ↑ "A Burkholderia pseudomallei toxin inhibits helicase activity of translation factor eIF4A.". Science 334 (6057): 821–4. 2011. doi:10.1126/science.1211915. PMID 22076380. Bibcode: 2011Sci...334..821C. 
  55. ↑ "In vivo expression technology identifies a type VI secretion system locus in Burkholderia pseudomallei that is induced upon invasion of macrophages". Microbiology 153 (Pt 8): 2689–99. August 2007. doi:10.1099/mic.0.2007/006585-0. PMID 17660433. http://mic.sgmjournals.org/cgi/pmidlookup?view=long&pmid=17660433. 
  56. ↑ "The BpeAB-OprB efflux pump of Burkholderia pseudomallei 1026b does not play a role in quorum sensing, virulence factor production, or extrusion of aminoglycosides, but is a broad-spectrum drug efflux system". Antimicrob. Agents Chemother. 54 (8): 3113–20. 2010. doi:10.1128/AAC.01803-09. PMID 20498323. 
  57. ↑ Baker, Sarah M.; Settles, Erik W.; Davitt, Christopher; Gellings, Patrick; Kikendall, Nicole; Hoffmann, Joseph; Wang, Yihui; Bitoun, Jacob et al. (29 January 2021). "Burkholderia pseudomallei OMVs derived from infection mimicking conditions elicit similar protection to a live-attenuated vaccine". npj Vaccines 6 (1): 18. doi:10.1038/s41541-021-00281-z. ISSN 2059-0105. PMID 33514749. 
  58. ↑ (in en) Vaccines for Neglected Pathogens: Strategies, Achievements and Challenges. 2023. p. 329. doi:10.1007/978-3-031-24355-4. ISBN 978-3-031-24354-7. https://link.springer.com/book/10.1007/978-3-031-24355-4. Retrieved 6 October 2023. 
  59. ↑ "Glyphosate resistance as a novel select-agent-compliant, non-antibiotic-selectable marker in chromosomal mutagenesis of the essential genes asd and dapB of Burkholderia pseudomallei.". Appl Environ Microbiol 75 (19): 6062–75. 2009. doi:10.1128/AEM.00820-09. PMID 19648360. Bibcode: 2009ApEnM..75.6062N. 
  60. ↑ "The Burkholderia pseudomallei Δasd mutant exhibits attenuated intracellular infectivity and imparts protection against acute inhalation melioidosis in mice". Infect Immun 79 (10): 4010–8. 2011. doi:10.1128/IAI.05044-11. PMID 21807903. 
  61. ↑ Mack, K.; Titball, R. W. (1996-11-01). "Transformation of Burkholderia pseudomallei by electroporation". Analytical Biochemistry 242 (1): 73–76. doi:10.1006/abio.1996.0430. ISSN 0003-2697. PMID 8923967. https://pubmed.ncbi.nlm.nih.gov/8923967/. 

External links

Wikidata ☰ Q140475 entry



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