Yersinia pestis: Difference between revisions

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{{Taxobox
{{Yersinia pestis infection}}
| color = lightgrey <!-- Please read [[WP:Taxobox_usage#Color]] before making any changes to the taxobox color. -->
 
| name = ''Yersinia pestis''
{{About0|Yersinia pestis infection}}
| image = Yersinia_pestis_fluorescent.jpeg
 
| image_width = 240px
{{CMG}}; {{AE}} Esther Lee, M.A.; {{Rim}}; {{JS}}
| image_caption = ''Yersinia pestis'' under fluorescent staining, 2000x. cheese: CDC
| regnum = [[Bacterium|Eubacteria]]
| phylum = [[Proteobacteria]]
| classis = Gamma Proteobacteria
| ordo = [[Enterobacteriaceae|Enterobacteriales]]
| '''familia = [[Enterobacteriaceae]]
| genus = ''[[Yersinia]]'''''| species = '''''Y. pestis'''''
| binomial = ''Yersinia pestis''
| binomial_authority = (Lehmann &amp; Neumann, 1896)<br>van Loghem 1944
}}
{{SI}}
{{CMG}}


{{SK}} Y. pestis
==Overview==
==Overview==
''[[Yersinia pestis]]'' (''[[Y. pestis]]''), a rod-shaped [[facultative anaerobe]] with bipolar staining (giving it a safety pin appearance) causes the [[infection]] in mammals and humans.<ref name=Baron>{{cite book | author = Collins FM | title = Pasteurella, Yersinia, and Francisella. ''In:'' Baron's Medical Microbiology ''(Baron S ''et al'', eds.)| edition = 4th | publisher = Univ. of Texas Medical Branch | year = 1996 | url = http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mmed.section.1611 | isbn = 0-9631172-1-1 }}</ref>  The [[bacteria]] maintain their existence in a cycle involving rodents and their fleas.  The genus [[Yersinia]] is [[gram-negative]], bipolar staining coccobacilli, and, similarly to other [[Enterobacteriaceae]], it has a fermentative [[metabolism]]. ''Y. pestis'' produces an [[antiphagocytic]] slime. The organism is [[motile]] when isolated, but becomes nonmotile in the mammalian host.
==Taxonomy==
[[Bacteria]]; [[Proteobacteria]]; [[Proteobacteria#Gammaproteobacteria|Gammaproteobacteria]]; [[Enterobacteriales]]; ''[[Yersinia]]''; [[Yersinia pestis]]


'''''Yersinia pestis''''' is a [[Gram-negative]] [[facultative anaerobic]] bipolar-staining (giving it a safety pin appearance) [[bacillus]] [[bacterium]] belonging to the family [[Enterobacteriaceae]].<ref name=Baron>{{cite book | author = Collins FM | title = Pasteurella, Yersinia, and Francisella. ''In:'' Baron's Medical Microbiology ''(Baron S ''et al'', eds.)| edition = 4th ed. | publisher = Univ of Texas Medical Branch | year = 1996 | url = http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mmed.section.1611 | id = ISBN 0-9631172-1-1 }}</ref> The [[pathogen|infectious agent]] of [[bubonic plague]], ''Y. pestis'' infection can also cause [[pneumonic plague|pneumonic]] and [[septicemic plague]].<ref name=Sherris>{{cite book | author = Ryan KJ; Ray CG (editors) | title = Sherris Medical Microbiology | edition = 4th ed. | pages = pp. 484-8 | publisher = McGraw Hill | year = 2004 | id = ISBN 0-8385852-9-9 }}</ref> All three forms have been responsible for high mortality rates in [[epidemics]] throughout human history, including the  [[Black Death]] that accounted for the death of approximately one-third of the [[Europe|European]] population in 1347 to 1353.
==Biology==
{| style="float: right;"
| [[File:Yersinia pestis fluorescent.jpeg|200px|thumb|none|Yersinia pestis, Direct Fluorescent Antibody Stain (DFA), 200x Magnification <SMALL>Courtesy: ''[http://phil.cdc.gov/phil/home.asp Public Health Image Library (PHIL), Centers for Disease Control and Prevention (CDC)]''<ref>{{Cite web | title = http://phil.cdc.gov/phil/details.asp | url = http://phil.cdc.gov/phil/details.asp}}</ref></SMALL>]]
|-
| [[File:Yersinia2.jpg|200px|thumb|none|Photomicrograph of Yersinia (Pasteurella) pestis, sometimes referred to as Bacillus pestis. <SMALL>Courtesy: ''[http://phil.cdc.gov/phil/home.asp Public Health Image Library (PHIL), Centers for Disease Control and Prevention (CDC)]''<ref>{{Cite web | title = http://phil.cdc.gov/phil/details.asp | url = http://phil.cdc.gov/phil/details.asp}}</ref></SMALL>]]
|}
[[Yersinia pestis]] is a [[motility|nonmotile]], non-[[spore]]-forming, [[Gram-negative]], non-[[lactose]] fermenting, ovoid and "safety-pin-shaped" (bipolar appearance when stained) [[bacillus]].  It is an [[enterobacteriaceae]] commonly measuring 0.75x1.5 μm. On [[cell culture]]s, it grows on sheep-blood agar, as grayish translucent colonies. It also grows on MacConkey and [[nutrient]]-rich broths.<ref name="Koirala2006">{{cite journal|last1=Koirala|first1=Janak|title=Plague: Disease, Management, and Recognition of Act of Terrorism|journal=Infectious Disease Clinics of North America|volume=20|issue=2|year=2006|pages=273–287|issn=08915520|doi=10.1016/j.idc.2006.02.004}}</ref>


The [[genus]] ''Yersinia'' is Gram-negative, bipolar staining [[coccobacillus|coccobacilli]], and, similarly to other ''Enterobacteriaceae'', it has a [[fermentation (biochemistry)|fermentative]] [[metabolism]]. ''Y. pestis'' produces an [[antiphagocytic]] slime. The organism is [[motility|motile]] when isolated, but becomes nonmotile in the mammalian [[Host (biology)|host]].
[[Yersinia pestis]] only survives for a few hours on physical surfaces, being very sensitive to high temperatures, sunlight and disinfectants.<ref name="Koirala2006">{{cite journal|last1=Koirala|first1=Janak|title=Plague: Disease, Management, and Recognition of Act of Terrorism|journal=Infectious Disease Clinics of North America|volume=20|issue=2|year=2006|pages=273–287|issn=08915520|doi=10.1016/j.idc.2006.02.004}}</ref><ref name="pmid8993858">{{cite journal| author=Perry RD, Fetherston JD| title=Yersinia pestis--etiologic agent of plague. | journal=Clin Microbiol Rev | year= 1997 | volume= 10 | issue= 1 | pages= 35-66 | pmid=8993858 | doi= | pmc=PMC172914 | url=http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&tool=sumsearch.org/cite&retmode=ref&cmd=prlinks&id=8993858  }} </ref><ref name="pmid10852811">{{cite journal| author=Gage KL, Dennis DT, Orloski KA, Ettestad P, Brown TL, Reynolds PJ et al.| title=Cases of cat-associated human plague in the Western US, 1977-1998. | journal=Clin Infect Dis | year= 2000 | volume= 30 | issue= 6 | pages= 893-900 | pmid=10852811 | doi=10.1086/313804 | pmc= | url=http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&tool=sumsearch.org/cite&retmode=ref&cmd=prlinks&id=10852811  }} </ref>


==Historical Perspective==
[[Yersinia pestis]] is thought to have evolved from [[Yersinia pseudotuberculosis]] about 1500 - 2000 years ago. According to its behavior towards [[nitrate]] and [[glycerol]], [[Yersinia pestis]] may be classified as:<ref name="Koirala2006">{{cite journal|last1=Koirala|first1=Janak|title=Plague: Disease, Management, and Recognition of Act of Terrorism|journal=Infectious Disease Clinics of North America|volume=20|issue=2|year=2006|pages=273–287|issn=08915520|doi=10.1016/j.idc.2006.02.004}}</ref>
* ''Biovar Antiqua''
:* Nitrate reduction: positive
:* Glycerol reduction: positive


* ''Y. pestis'' was discovered in 1894 by Swiss/French [[physician]] and [[bacteriologist]] from the [[Pasteur Institute]], [[Alexandre Yersin]], during an [[epidemic]] of plague in Hong-Kong.<ref name=Bockemühl_1994>{{cite journal |author=Bockemühl J |title=[100 years after the discovery of the plague-causing agent--importance and veneration of Alexandre Yersin in Vietnam today] |journal=Immun Infekt |volume=22 |issue=2 |pages=72-5 |year=1994 |pmid = 7959865}}</ref>
* ''Biovar Medievalis''
* Yersin was a member of the [[Louis Pasteur|Pasteur]] school of thought. [[Shibasaburo Kitasato]], a German-trained Japanese bacteriologist who practiced [[Robert Koch|Koch's methodology]] was also engaged at the time in finding the causative agent of plague.<ref name=Ho>{cite journal |author=Howard-Jones N |title=Was Shibasaburo Kitasato the discoverer of the plague bacillus? | journal=Perspect Biol Med |volume=16 |issue=2 |pages=292-307 |year=1973 |pmid = 4570035}}</ref>
:* Nitrate reduction: negative
* However, it was Yersin who actually linked plague with ''Yersinia pestis''. Originally named  ''Pasteurella pestis'', the organism was renamed in 1967.
:* Glycerol use: positive
* Three [[biovar]]s of ''Y. pestis'' are known, each thought to correspond to one of the [[Bubonic plague#Historical pandemics|historical pandemics of bubonic plague]].<ref name=Zhou_2004>{{cite journal |author=Zhou D, Tong Z, Song Y, Han Y, Pei D, Pang X, Zhai J, Li M, Cui B, Qi Z, Jin L, Dai R, Du Z, Wang J, Guo Z, Wang J, Huang P, Yang R |title=Genetics of metabolic variations between Yersinia pestis biovars and the proposal of a new biovar, microtus |journal=J Bacteriol |volume=186 |issue=15 |pages=5147-52 |year=2004 |url= http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=8195371 | pmid =  15262951}}</ref>
* Biovar '''Antiqua''' is thought to correspond to the [[Plague of Justinian]]; it is not known whether this biovar also corresponds to [[Bubonic plague#History|earlier, smaller epidemics of bubonic plague]], or whether these were even truly bubonic plague.<ref name=Guiyoule_1994>{{cite journal |author=Guiyoule A, Grimont F, Iteman I, Grimont P, Lefèvre M, Carniel E |title=Plague pandemics investigated by ribotyping of Yersinia pestis strains |journal=J Clin Microbiol |volume=32 |issue=3 |pages=634-41 |year=1994 |id=PMID 8195371}}</ref>
* Biovar '''Medievalis''' is thought to correspond to the [[Black Death]].
* Biovar '''Orientalis''' is thought to correspond to the [[Third Pandemic]] and the majority of modern outbreaks of plague.
* The role of ''Y. pestis'' in the Black Death is debated among historians; some have suggested that the Black Death spread far too rapidly to be caused by ''Y. pestis''.
* DNA from ''Y. pestis'' is alleged to have been found in the teeth of an individual who supposedly died from the Black Death, however, and medieval corpses who died from other causes did not test positive for ''Y. pestis''.<ref>{{cite journal | author = Drancourt M, Aboudharam G, Signolidagger M, Dutourdagger O, Raoult D.| title = Detection of 400-year-old ''Yersinia pestis'' DNA in human dental pulp: An approach to the diagnosis of ancient septicemia | journal = PNAS | year = 1998| volume = 95 | issue = 21 | pages = 12637&ndash;12640 | url= http://www.pnas.org/cgi/content/abstract/95/21/12637} | pmid = 9770538 }}</ref><ref>{{cite journal | author = Drancourt M; Raoult D.| title = Molecular insights into the history of plague. | journal = Microbes Infect. | year = 2002 | volume = 4| issue = | pages = 105&ndash;9 | pmid = 11825781 }}</ref>
* This suggests that ''Y. pestis'' was, at the very least, a contributing factor in some (though possibly not all) of the European plagues. It's possible that the selective pressures induced by the plague might have changed how the pathogen manifests in humans, selecting against the individuals or populations which were the most susceptible.


==Pathophysiology==
* ''Biovar Orientalis''
:* Nitrate reduction: positive
:* Glycerol use: negative


[[Image:Yersinia pestis.jpg|thumb|left|250px|A scanning electron micrograph depicting a mass of Yersinia pestis bacteria]]
===Genome===
* Pathogenicity of ''Y. pestis'' is in part due to two anti-phagocytic [[antigens]], named F1 (Fraction 1) and V, both important for [[virulence]].<ref name=Baron/> These antigens are produced by the bacterium at 37°C.
The complete [[genome|genomic]] sequence is available for two of the three sub-species of [[yersinia pestis]]:
* Furthermore, ''Y. pestis'' survives and produces F1 and V antigens within [[blood]] cells such as [[monocyte]]s, but not in polymorphonuclear [[neutrophils]]. * Natural or induced [[immunity (medical)|immunity]] is achieved by the production of specific [[opsonin|opsonic]] [[antibody|antibodies]] against F1 and V antigens; antibodies against F1 and V induce [[phagocytosis]] by neutrophils.<ref>{{cite book | author = Salyers AA, Whitt DD | title = Bacterial Pathogenesis: A Molecular Approach | edition = 2nd ed. | publisher = ASM Press | year = 2002 | id = pp. 207-12}}</ref>
* Strain KIM - Biovar Medievalis<ref>{{cite journal | author = Deng W| title = Genome Sequence of Yersinia pestis KIM | journal = Journal of Bacteriology | year = 2002 | volume = 184 | issue = 16 | pages = 4601&ndash;4611 | doi= 10.1128/JB.184.16.4601-4611.2002 | pmid=12142430 | pmc = 135232 | author-separator = , | display-authors = 1 | last2 = Burland | first2 = V. | last3 = Plunkett Iii | first3 = G. | last4 = Boutin | first4 = A. | last5 = Mayhew | first5 = G. F. | last6 = Liss | first6 = P. | last7 = Perna | first7 = N. T. | last8 = Rose | first8 = D. J. | last9 = Mau | first9 = B.}}</ref>
* A [[formalin]]-inactivated [[vaccine]] once was available for adults at high risk of contracting the plague until removal from the market by the [[FDA]].  It was of limited effectiveness and may cause severe [[inflammation]].
* Strain CO92 - Biovar Orientalis<ref>{{cite journal | author = Parkhill J| title = Genome sequence of Yersinia pestis, the causative agent of plague | journal = Nature | year = 2001 | volume = 413 | issue = 6855| pages = 523&ndash;527 | doi = 10.1038/35097083 | pmid = 11586360 | author-separator = , | display-authors = 1 | last2 = Wren | first2 = B. W. | last3 = Thomson | first3 = N. R. | last4 = Titball | first4 = R. W. | last5 = Holden | first5 = M. T. G. | last6 = Prentice | first6 = M. B. | last7 = Sebaihia | first7 = M. | last8 = James | first8 = K. D. | last9 = Churcher | first9 = C.}}</ref>  
* Experiments with [[genetic engineering]] of a vaccine based on F1 and V antigens are underway and show promise; however, bacteria lacking antigen F1 are still virulent, and the V antigens are sufficiently variable, that vaccines composed of these antigens may not be fully protective<ref>{{cite journal | author = Welkos S ''et al.''.| title = Determination of the virulence of the pigmentation-deficient and pigmentation-/plasminogen activator-deficient strains of ''Yersinia pestis'' in non-human primate and mouse models of pneumonic plague | journal = Vaccine | year = 2002 | volume = 20 | issue = | pages = 2206&ndash;2214 | pmid = 12009274 }}</ref>.


===Genetics===
As of 2006, the [[genomic sequence]] of a [[strain]] of ''biovar Antiqua'' has been completed.<ref name="pmid16740952">{{cite journal |author=Chain PS |title=Complete Genome Sequence of Yersinia pestis Strains Antiqua and Nepal516: Evidence of Gene Reduction in an Emerging Pathogen |journal=J. Bacteriol. |volume=188 |issue=12 |pages=4453–63 |year=2006 |pmid=16740952 |doi=10.1128/JB.00124-06 |pmc=1482938 |author-separator=, |author2=Hu P |author3=Malfatti SA |display-authors=3 |last4=Radnedge |first4=L. |last5=Larimer |first5=F. |last6=Vergez |first6=L. M. |last7=Worsham |first7=P. |last8=Chu |first8=M. C. |last9=Andersen |first9=G. L.}}</ref> Similar to the other [[pathogenic]] [[strains]], there are signs of mutations causing loss of function. The [[chromosome]] of strain KIM is 4,600,755 base pairs long; the [[chromosome]] of strain CO92 is 4,653,728 base pairs long.


* The complete [[genome|genomic]] sequence is available for two of the three sub-species of ''Y. pestis'': strain KIM (of biovar Medievalis)<ref>{{cite journal | author = Deng W ''et al.''.| title = Genome Sequence of ''Yersinia pestis KIM'' | journal = Journal of Bacteriology | year = 2002 | volume = 184 | issue = 16 | pages = 4601&ndash;4611 | url=http://jb.asm.org/cgi/content/full/184/16/4601 | pmid=12142430}}</ref>, and strain CO92 (of biovar Orientalis, obtained from a clinical isolate in the United States)<ref>{{cite journal | author = Parkhill J ''et al.''.| title = Genome sequence of ''Yersinia pestis'', the causative agent of plague | journal = Nature | year = 2001 | volume = 413 | issue = | pages = 523&ndash;527 | url=http://www.nature.com/cgi-taf/DynaPage.taf?file=/nature/journal/v413/n6855/abs/413523a0_fs.html | pmid = 11586360}}</ref>; as of 2006, the genomic sequence of a strain of biovar Antiqua has not yet been completed.
Similarly to other [[enterobacteriaceae]] ([[Yersinia pseudotuberculosis|Yersinia pseudotuberculosis]] and [[Yersinia enterocolitica|Yersinia enterocolitica]]), [[Yersinia pestis]] is host to the ''[[plasmid]] pCD1''. In addition, it also hosts two other [[plasmids]], ''pPCP1'' (also called pPla or pPst) and ''pMT1'' (also called pFra) that are not carried by the other Yersinia species.  
* The [[chromosome]] of strain KIM is 4,600,755 base pairs long; the chromosome of strain CO92 is 4,653,728 base pairs long. Like its cousins ''[[Yersinia pseudotuberculosis|Y. pseudotuberculosis]]'' and ''[[Yersinia enterocolitica|Y.  enterocolitica]]'', ''Y. pestis'' is host to the [[plasmid]] pCD1.  
* pFra codes for a [[phospholipase D]] that is important for the ability of [[Yersinia pestis]] to be transmitted by fleas.  
* In addition, it also hosts two other plasmids, pPCP1 and pMT1 which are not carried by the other ''Yersinia'' species. Together, these plasmids, and a [[pathogenicity island]] called HPI, encode several proteins which cause the pathogenicity for which ''Y. pestis'' is famous.
* pPla codes for a [[protease]], Pla, that activates [[plasminogen]] in human hosts and is a very important [[virulence factor]] for pneumonic plague.<ref name="pmid17255510">{{cite journal| author=Lathem WW, Price PA, Miller VL, Goldman WE| title=A plasminogen-activating protease specifically controls the development of primary pneumonic plague. | journal=Science | year= 2007 | volume= 315 | issue= 5811 | pages= 509-13 | pmid=17255510 | doi=10.1126/science.1137195 | pmc= | url=http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&tool=sumsearch.org/cite&retmode=ref&cmd=prlinks&id=17255510  }} </ref>  
* Among other things, these [[virulence]] factors are required for bacterial adhesion and injection of proteins into the host cell, invasion of bacteria into the host cell, and acquisition and binding of iron harvested from red blood cells.
* Y. pestis is thought to be descendant from ''Y. pseudotuberculosis'', differing only in the presence of specific virulence plasmids.
* A recent comprehensive and comparative [[proteomics]] analysis of ''Y. pestis'': strain KIM was recently performed <ref>{{cite journal | author = Hixson K ''et al.''.| title = Biomarker candidate identification in Yersinia pestis using organism-wide semiquantitative proteomics. | journal = Journal of Proteome Research | year = 2006 | volume = 5 | issue = 11 | pages = 3008-3017 | pmid = 16684765 }}</ref> , this analysis focused on the transition to a growth condition mimicking growth in host cells.


===Susceptibility===
Together, these [[plasmids]], and a [[pathogenicity island]] called HPI, encode several [[pathogenic]] [[proteins]], characteristic of [[Yersinia pestis]]. Among other things, these [[virulence factors]] are required of:
* [[Bacterial]] adhesion
* Injection of [[proteins]] into the host cell
* Invasion of the host cell (via a Type III secretion system)
* Acquisition and binding of [[iron]] from [[red blood cell]], via siderophores


* The traditional first line treatment for ''Y. pestis'' has been [[streptomycin]],<ref>{{cite journal | author=Wagle PM. | title=Recent advances in the treatment of bubonic plague | journal=Indian J Med Sci | year=1948 | volume=2 | pages=489&ndash;94 }}</ref><ref>{{cite journal | author=Meyer KF. | title=Modern therapy of plague | journal=JAMA | year=1950 | volume=144 | pages=982&ndash;5 | pmid = 14774219}}</ref> [[chloramphenicol]], [[tetracycline]],<ref>{{cite journal | author=Kilonzo BS, Makundi RH, Mbise TJ. | title= A decade of plague epidemiology and control in the Western Usambara mountains, north-east Tanzania | journal=Acta Tropica | year=1992 | volume=50 | pages=323&ndash;9 | pmid = 1356303}}</ref> and [[fluoroquinolones]].
A comprehensive and comparative [[proteomics]] analysis of [[Yersinia pestis]] strain KIM was performed in 2006.<ref>{{cite journal | author = Hixson K| title = Biomarker candidate identification in Yersinia pestis using organism-wide semiquantitative proteomics | journal = Journal of Proteome Research | year = 2006 | volume = 5 | issue = 11 | pages = 3008–3017 | pmid = 16684765  | doi = 10.1021/pr060179y | author-separator = , | display-authors = 1 | last2 = Adkins | first2 = Joshua N. | last3 = Baker | first3 = Scott E. | last4 = Moore | first4 = Ronald J. | last5 = Chromy | first5 = Brett A. | last6 = Smith | first6 = Richard D. | last7 = McCutchen-Maloney | first7 = Sandra L. | last8 = Lipton | first8 = Mary S. | last9 = Heffron | first9 = F}}</ref> The analysis focused on the transition to a growth condition mimicking growth in host cells.
* There is also good evidence to support the use of [[doxycycline]] or [[gentamicin]].<ref>{{cite journal | author=Mwengee W, Butler T, Mgema S, ''et al.'' | title=Treatment of plague with gentamicin or doxycycline in a randomized clinical trial in Tanzania | journal=Clin Infect Dis | year=2006 | volume=42 | pages=614&ndash;21 | pmid = 16447105 }}</ref>
* It should be noted that strains resistant to one or two agents specified above have been isolated: treatment should be guided by antibiotic sensitivities where available.  


==Treatment==
==Tropism==
===Medical Therapy===
[[Yersinia pestis]] shows [[tropism]] for [[lymphoid tissue]].


Antibiotic treatment alone is insufficient for some patients, who may also require circulatory, ventilatory, or renal support.
==Natural Reservoir==
Plague is primarily a disease of rodents. The [[infection]] is maintained in natural foci of the disease in wild rodent colonies, through [[transmission]] between rodents, by their flea ectoparasites.<ref name=CDC>{{cite web | title = Plague | url = http://www.cdc.gov/plague/healthcare/clinicians.html }}</ref>




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Image: Bubonic plague12.jpeg| Patient acquired a plague infection through abrasions on his upper right leg. <SMALL><SMALL>''[http://phil.cdc.gov/phil/home.asp From Public Health Image Library (PHIL).] ''<ref name=PHIL> {{Cite web | title = Public Health Image Library (PHIL) | url = http://phil.cdc.gov/phil/home.asp}}</ref></SMALL></SMALL>
Image: Bubonic plague12.jpeg| Patient acquired a plague infection through abrasions on his upper right leg. <SMALL><SMALL>''[http://phil.cdc.gov/phil/home.asp From Public Health Image Library (PHIL).] ''<ref name=PHIL> {{Cite web | title = Public Health Image Library (PHIL) | url = http://phil.cdc.gov/phil/home.asp}}</ref></SMALL></SMALL>
Image: Bubonic plague04.jpeg| Lung tissue sample revealed the histopathologic changes indicative of fatal human plague (125x mag). <SMALL><SMALL>''[http://phil.cdc.gov/phil/home.asp From Public Health Image Library (PHIL).] ''<ref name=PHIL> {{Cite web | title = Public Health Image Library (PHIL) | url = http://phil.cdc.gov/phil/home.asp}}</ref></SMALL></SMALL>
Image: Bubonic plague04.jpeg| Lung tissue sample revealed the histopathologic changes indicative of fatal human plague (125x mag). <SMALL><SMALL>''[http://phil.cdc.gov/phil/home.asp From Public Health Image Library (PHIL).] ''<ref name=PHIL> {{Cite web | title = Public Health Image Library (PHIL) | url = http://phil.cdc.gov/phil/home.asp}}</ref></SMALL></SMALL>
Image: Bubonic plague01.jpeg| Lung tissue sample revealed the histopathologic changes indicative of fatal human plague (125x mag). <SMALL><SMALL>''[http://phil.cdc.gov/phil/home.asp From Public Health Image Library (PHIL).] ''<ref name=PHIL> {{Cite web | title = Public Health Image Library (PHIL) | url = http://phil.cdc.gov/phil/home.asp}}</ref></SMALL></SMALL>
</SMALL>
 
Image: Bubonic plague01.jpeg| Purple-colored Yersinia pestis bacteria on the proventricular spines of a Xenopsylla cheopis flea. <SMALL><SMALL>''[http://phil.cdc.gov/phil/home.asp From Public Health Image Library (PHIL).] ''<ref name=PHIL> {{Cite web | title = Public Health Image Library (PHIL) | url = http://phil.cdc.gov/phil/home.asp}}</ref></SMALL>
</gallery>
</gallery>


== External Links ==
==References==
* A list of variant strains and information on synonyms (and much more) is available through the  [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=632 NCBI taxonomy browser].


==References==
{{Reflist|2}}
{{Reflist|2}}


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This page is about microbiologic aspects of the organism(s).  For clinical aspects of the disease, see Yersinia pestis infection.

Editor-In-Chief: C. Michael Gibson, M.S., M.D. [1]; Associate Editor(s)-in-Chief: Esther Lee, M.A.; Rim Halaby, M.D. [2]; João André Alves Silva, M.D. [3]

Overview

Yersinia pestis (Y. pestis), a rod-shaped facultative anaerobe with bipolar staining (giving it a safety pin appearance) causes the infection in mammals and humans.[1] The bacteria maintain their existence in a cycle involving rodents and their fleas. The genus Yersinia is gram-negative, bipolar staining coccobacilli, and, similarly to other Enterobacteriaceae, it has a fermentative metabolism. Y. pestis produces an antiphagocytic slime. The organism is motile when isolated, but becomes nonmotile in the mammalian host.

Taxonomy

Bacteria; Proteobacteria; Gammaproteobacteria; Enterobacteriales; Yersinia; Yersinia pestis

Biology

Yersinia pestis, Direct Fluorescent Antibody Stain (DFA), 200x Magnification Courtesy: Public Health Image Library (PHIL), Centers for Disease Control and Prevention (CDC)[2]
Photomicrograph of Yersinia (Pasteurella) pestis, sometimes referred to as Bacillus pestis. Courtesy: Public Health Image Library (PHIL), Centers for Disease Control and Prevention (CDC)[3]

Yersinia pestis is a nonmotile, non-spore-forming, Gram-negative, non-lactose fermenting, ovoid and "safety-pin-shaped" (bipolar appearance when stained) bacillus. It is an enterobacteriaceae commonly measuring 0.75x1.5 μm. On cell cultures, it grows on sheep-blood agar, as grayish translucent colonies. It also grows on MacConkey and nutrient-rich broths.[4]

Yersinia pestis only survives for a few hours on physical surfaces, being very sensitive to high temperatures, sunlight and disinfectants.[4][5][6]

Yersinia pestis is thought to have evolved from Yersinia pseudotuberculosis about 1500 - 2000 years ago. According to its behavior towards nitrate and glycerol, Yersinia pestis may be classified as:[4]

  • Biovar Antiqua
  • Nitrate reduction: positive
  • Glycerol reduction: positive
  • Biovar Medievalis
  • Nitrate reduction: negative
  • Glycerol use: positive
  • Biovar Orientalis
  • Nitrate reduction: positive
  • Glycerol use: negative

Genome

The complete genomic sequence is available for two of the three sub-species of yersinia pestis:

  • Strain KIM - Biovar Medievalis[7]
  • Strain CO92 - Biovar Orientalis[8]

As of 2006, the genomic sequence of a strain of biovar Antiqua has been completed.[9] Similar to the other pathogenic strains, there are signs of mutations causing loss of function. The chromosome of strain KIM is 4,600,755 base pairs long; the chromosome of strain CO92 is 4,653,728 base pairs long.

Similarly to other enterobacteriaceae (Yersinia pseudotuberculosis and Yersinia enterocolitica), Yersinia pestis is host to the plasmid pCD1. In addition, it also hosts two other plasmids, pPCP1 (also called pPla or pPst) and pMT1 (also called pFra) that are not carried by the other Yersinia species.

Together, these plasmids, and a pathogenicity island called HPI, encode several pathogenic proteins, characteristic of Yersinia pestis. Among other things, these virulence factors are required of:

  • Bacterial adhesion
  • Injection of proteins into the host cell
  • Invasion of the host cell (via a Type III secretion system)
  • Acquisition and binding of iron from red blood cell, via siderophores

A comprehensive and comparative proteomics analysis of Yersinia pestis strain KIM was performed in 2006.[11] The analysis focused on the transition to a growth condition mimicking growth in host cells.

Tropism

Yersinia pestis shows tropism for lymphoid tissue.

Natural Reservoir

Plague is primarily a disease of rodents. The infection is maintained in natural foci of the disease in wild rodent colonies, through transmission between rodents, by their flea ectoparasites.[12]


Gallery

References

  1. ↑ Collins FM (1996). Pasteurella, Yersinia, and Francisella. In: Baron's Medical Microbiology (Baron S et al, eds.) (4th ed.). Univ. of Texas Medical Branch. ISBN 0-9631172-1-1.
  2. ↑ "http://phil.cdc.gov/phil/details.asp". External link in |title= (help)
  3. ↑ "http://phil.cdc.gov/phil/details.asp". External link in |title= (help)
  4. ↑ 4.0 4.1 4.2 Koirala, Janak (2006). "Plague: Disease, Management, and Recognition of Act of Terrorism". Infectious Disease Clinics of North America. 20 (2): 273–287. doi:10.1016/j.idc.2006.02.004. ISSN 0891-5520.
  5. ↑ Perry RD, Fetherston JD (1997). "Yersinia pestis--etiologic agent of plague". Clin Microbiol Rev. 10 (1): 35–66. PMC 172914. PMID 8993858.
  6. ↑ Gage KL, Dennis DT, Orloski KA, Ettestad P, Brown TL, Reynolds PJ; et al. (2000). "Cases of cat-associated human plague in the Western US, 1977-1998". Clin Infect Dis. 30 (6): 893–900. doi:10.1086/313804. PMID 10852811.
  7. ↑ Deng W; et al. (2002). "Genome Sequence of Yersinia pestis KIM". Journal of Bacteriology. 184 (16): 4601&ndash, 4611. doi:10.1128/JB.184.16.4601-4611.2002. PMC 135232. PMID 12142430. Unknown parameter |author-separator= ignored (help)
  8. ↑ Parkhill J; et al. (2001). "Genome sequence of Yersinia pestis, the causative agent of plague". Nature. 413 (6855): 523&ndash, 527. doi:10.1038/35097083. PMID 11586360. Unknown parameter |author-separator= ignored (help)
  9. ↑ Chain PS; Hu P; Malfatti SA; et al. (2006). "Complete Genome Sequence of Yersinia pestis Strains Antiqua and Nepal516: Evidence of Gene Reduction in an Emerging Pathogen". J. Bacteriol. 188 (12): 4453–63. doi:10.1128/JB.00124-06. PMC 1482938. PMID 16740952. Unknown parameter |author-separator= ignored (help)
  10. ↑ Lathem WW, Price PA, Miller VL, Goldman WE (2007). "A plasminogen-activating protease specifically controls the development of primary pneumonic plague". Science. 315 (5811): 509–13. doi:10.1126/science.1137195. PMID 17255510.
  11. ↑ Hixson K; et al. (2006). "Biomarker candidate identification in Yersinia pestis using organism-wide semiquantitative proteomics". Journal of Proteome Research. 5 (11): 3008–3017. doi:10.1021/pr060179y. PMID 16684765. Unknown parameter |author-separator= ignored (help)
  12. ↑ "Plague".
  13. ↑ 13.00 13.01 13.02 13.03 13.04 13.05 13.06 13.07 13.08 13.09 13.10 13.11 13.12 13.13 "Public Health Image Library (PHIL)".
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