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{{About||the genus|Marburgvirus}}
{{Taxobox
<!-- Color parameter is not needed -- automatically assigned -->
| name = Marburg virus (MARV)
| image = Marburg virus.jpg
| image_caption = Transmission electron micrograph of Marburg virus
| virus_group = V
| ordo = ''[[Mononegavirales]]''
| familia = ''[[Filoviridae]]''
| genus = ''[[Marburgvirus]]''
| species = ''[[Marburg marburgvirus]]''
| type_species =
| subdivision_ranks = Species
| subdivision =
}}

'''Marburg virus''' ({{IPAc-en|ˈ|m|ɑr|b|ər|g|_|ˈ|v|aɪ|r|ə|s}} {{Respell|MAR|bərg}} {{Respell|VY|rəs}}<ref name=KuhnArch/>) is a [[hemorrhagic fever virus]] of the Filoviridae family of viruses and a member of the species ''[[Marburg marburgvirus]]'', genus ''[[Marburgvirus]]''. Marburg virus (MARV) causes [[Marburg virus disease]] in [[human]]s and nonhuman [[primates]], a form of [[viral hemorrhagic fever]].<ref name=Spickler>{{cite web|last1=Spickler|first1=Anna|title=Ebolavirus and Marburgvirus Infections|url=http://www.cfsph.iastate.edu/Factsheets/pdfs/viral_hemorrhagic_fever_filovirus.pdf}}</ref> The virus is considered to be extremely dangerous. The [[World Health Organization|WHO]] rates it as a Risk Group 4 Pathogen (requiring [[Biosafety_level#Biosafety_level_4|biosafety level 4-equivalent containment]]).<ref name=BMBL5>{{cite web|url=http://www.cdc.gov/biosafety/publications/bmbl5/|title=Biosafety in Microbiological and Biomedical Laboratories (BMBL) 5th Edition|accessdate=2011-10-16|last=US Department of Health and Human Services}}</ref> In the United States, the [[National Institutes of Health|NIH]]/[[National Institute of Allergy and Infectious Diseases]] ranks it as a Category A Priority Pathogen<ref name=PriorityPathogens>{{cite web|url=http://www.niaid.nih.gov/topics/biodefenserelated/biodefense/research/pages/cata.aspx |title=Biodefense and Emerging Infectious Diseases |accessdate=2011-10-16|publisher=US National Institute of Allergy and Infectious Diseases (NIAID), US National Institutes of Health (NIH)}}</ref> and the [[Centers for Disease Control and Prevention]] lists it as a [[Bioterrorism|Category A Bioterrorism Agent]].<ref name=CategoryBioterrorism>{{cite web|url=http://www.bt.cdc.gov/agent/agentlist-category.asp|title=Bioterrorism Agents/Diseases|accessdate=2011-10-16|last=US Centers for Disease Control and Prevention (CDC)}}</ref> It is also is listed as a biological agent for export control by the [[Australia Group]].<ref name=AustraliaGroup>{{cite web|url=http://www.australiagroup.net/en/biological_agents.html|title=List of Biological Agents for Export Control|accessdate=2011-10-16|last=The Australia Group}}</ref>

In 2009, expanded [[clinical trial]]s of an [[Ebola]] and Marburg [[vaccine]] began in [[Kampala]], [[Uganda]].<ref>Beth Skwarecki [http://www.medscape.com/viewarticle/831858 Ebola, Marburg DNA Vaccines Prove Safe in Phase 1 Trial] Medscape Medical News, September 17, 2014</ref><ref>[http://clinicaltrials.gov/show/NCT00997607 Evaluating an Ebola and a Marburg Vaccine in Uganda] [[U.S. Department of Health & Human Services]]</ref>
==Discovery==
Marburg virus was first described in 1967.<ref name=Siegert1967>{{cite pmid|4294540|noedit}}</ref> It was noticed during small outbreaks in the German cities [[Marburg]] and [[Frankfurt]] and the Yugoslav capital [[Belgrade]] in the 1960s. It was also misinterpreted as a pregnancy defect until is was disproved in 1992. German workers were accidentally exposed to tissues of infected [[grivet|grivet monkey]]s (''Chlorocebus aethiops'') at the city's former main industrial plant, the Behringwerke, then part of [[Hoechst AG|Hoechst]], and today of [[CSL Behring]]. During these outbreaks, 31 people became infected and seven of them died. MARV is a [[Select Agent]],<ref name=SelectAgents>{{cite web|url=http://www.selectagents.gov|title=National Select Agent Registry (NSAR)|accessdate=2011-10-16|last=US Animal and Plant Health Inspection Service (APHIS) and US Centers for Disease Control and Prevention (CDC)}}</ref>

===Nomenclature===
The virus is one of two members of the [[International Committee on Taxonomy of Viruses|species]] ''[[Marburg marburgvirus]]'', which is included in the [[International Committee on Taxonomy of Viruses|genus]] ''[[Marburgvirus]]'', [[International Committee on Taxonomy of Viruses|family]] ''[[Filoviridae]]'', [[International Committee on Taxonomy of Viruses|order]] ''[[Mononegavirales]]''. The name Marburg virus is derived from [[Marburg]] (the city in [[Hesse]], [[Germany]], where the virus was first discovered) and the [[Taxonomy (biology)|taxonomic]] [[suffix]] ''virus''.<ref name=KuhnArch>{{cite pmid|21046175|noedit}}</ref>

According to the rules for taxon naming established by the [[International Committee on Taxonomy of Viruses|International Committee on Taxonomy of Viruses (ICTV)]], the name Marburg virus is always to be [[Capitalization|capitalized]], but is never [[Italic type|italicized]], and may be [[Abbreviation|abbreviated]] (with MARV being the official abbreviation).

Marburg virus was first introduced under this name in 1967.<ref name=Siegert1967/> In 2005, the virus name was changed to Lake Victoria marburgvirus, which unfortunately was the same spelling as its species ''Lake Victoria marburgvirus''.<ref name=Feldmann2005>{{Cite book|last1=Feldmann|first1=H.|last2=Geisbert|first2=T. W.|last3=Jahrling|first3=P. B.|last4=Klenk|first4=H.-D.|last5=Netesov|first5=S. V.|last6=Peters|first6=C. J.|last7=Sanchez|first7=A.|last8=Swanepoel|first8=R.|last9=Volchkov|first9=V. E.| displayauthors = 8|chapter=Family Filoviridae|year=2005|editor-last=Fauquet|editor-first=C. M.|editor2-last=Mayo|editor2-first=M. A.|editor3-last=Maniloff|editor3-first=J.|editor4-last=Desselberger|editor4-first=U.|editor5-last=Ball|editor5-first=L. A.|title=Virus Taxonomy—Eighth Report of the International Committee on Taxonomy of Viruses|pages=645–653|publisher=Elsevier/Academic Press|location=San Diego, US|isbn=0-12-370200-3|postscript=<!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}}}}</ref><ref>{{cite journal|last1=Mayo|first1=M. A.|year = 2002|title = ICTV at the Paris ICV: results of the plenary session and the binomial ballot|journal = Archives of Virology|volume = 147|issue = 11|pages = 2254–60|doi=10.1007/s007050200052}}</ref> However, most scientific articles continued to refer to Marburg virus. Consequently, in 2010, the name Marburg virus was reinstated and the species name changed.<ref name=KuhnArch/> A previous abbreviation for the virus was MBGV.

==Human disease==
{{Main|Marburg virus disease}}
MARV is one of two marburgviruses that causes [[Marburg virus disease|Marburg virus disease (MVD)]] in humans (in the literature also often referred to as Marburg hemorrhagic fever, MHF). Both viruses fulfill the criteria for being a member of the species Marburg marburgvirus because their [[genome]]s diverge from the prototype Marburg marburgvirus or the Marburg virus variant Musoke (MARV/Mus) by <10% at the [[nucleotide]] level.<ref name=KuhnArch/>

===Recorded outbreaks===
{| class="sortable wikitable"
|+ [[Marburg virus disease|Marburg virus disease (MVD)]] outbreaks due to Marburg virus (MARV) infection
|-
| '''Year'''
| '''Geographic location'''
| '''Human Deaths/Cases (case-fatality rate)'''
|- valign="TOP"
| 1967
| [[Marburg]] and [[Frankfurt]], West [[Germany]], and [[Belgrade]], [[Yugoslavia]]
| 7/31 (23%)<ref name=Siegert1967/><ref name=Smith1967>{{cite pmid|4168558|noedit}}</ref><ref name=Kissling1968>{{cite pmid|4296724|noedit}}</ref><ref name=Martini1968>{{cite pmid|4966280|noedit}}</ref><ref name=Stille1968>{{cite pmid|4966281|noedit}}</ref><ref name=Bonin1969>{{cite pmid|5005859|noedit}}</ref><ref name=Jacob1971>{{cite pmid|5748997|noedit}}</ref><ref name= Stojkovic1971>{{Cite book|last1 = Stojkovic |first1 = L.|last2 = Bordjoski |first2 = M.|last3 = Gligic |first3 = A.| last4 = Stefanovic |first4 = Z.|year = 1971|chapter = Two Cases of Cercopithecus-Monkeys-Associated Haemorrhagic Fever |editor-last=Martini|editor-first=G. A.|editor2-last=Siegert|editor2-first=R.|title=Marburg Virus Disease|pages=24–33|publisher=Springer-Verlag|location=Berlin, Germany|isbn=978-0-387-05199-4|postscript = <!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}}}}</ref>
|- valign="TOP"
| 1975
| [[Rhodesia]] and [[Johannesburg]], [[South Africa]]
| 1/3 (33%)<ref name=Gear1975>{{cite pmid|811315|noedit}}</ref><ref name=Gear1977>{{cite pmid|406394|noedit}}</ref><ref name=Conrad1978>{{cite pmid|569445|noedit}}</ref>
|- valign="TOP"
| 1980
| [[Kenya]]
| 1/2 (50%)<ref name=Smith1982>{{cite pmid|6122054|noedit}}</ref>
|- valign="TOP"
| 1987
| [[Kenya]]
| 1/1 (100%)<ref>Marburg and Ebola viruses; Advances in Virus Research;
Volume 47, 1996, Pages 1–52</ref><ref>[http://www.cdc.gov/vhf/marburg/resources/outbreak-table.html Known Cases and Outbreaks of Marburg Hemorrhagic Fever, in Chronological Order]</ref>
|- valign="TOP"
| 1988
| [[Koltsovo, Novosibirsk Oblast|Koltsovo]], [[Soviet Union]]
| 1/1 (100%) [laboratory accident]<ref name=Beer1999>{{cite pmid|10024977|noedit}}</ref>
|- valign="TOP"
| 1990
| [[Koltsovo, Novosibirsk Oblast|Koltsovo]], [[Soviet Union]]
| 0/1 (0%) [laboratory accident]<ref name=Nikiforov1994>{{cite pmid|7941853|noedit}}</ref>
|- valign="TOP"
| 1998–2000
| [[Durba, Democratic Republic of the Congo|Durba]] and [[Watsa]], [[Democratic Republic of the Congo]]
| ? (A total of 154 cases and 128 deaths of marburgvirus infection were recorded during this outbreak. The case fatality was 83%. Two different marburgviruses, MARV and [[Ravn virus|Ravn virus (RAVV)]], cocirculated and caused disease. It has never been published how many cases and deaths were due to MARV or RAVV infection)<ref name=Bertherat1999>{{cite pmid|10546197|noedit}}</ref><ref name=Bausch2003>{{cite pmid|14720391|noedit}}</ref><ref name=Bausch2006>{{cite pmid|16943403|noedit}}</ref>
|- valign="TOP"
| 2004–2005
| [[Angola]]
| 227/252 (90%)<ref name=Hovette2005>{{cite pmid|16038348|noedit}}</ref><ref name=Ndayimirije2005>{{cite pmid|15917379|noedit}}</ref><ref name=Towner2006>{{cite pmid|16775337|noedit}}</ref><ref name=Jeffs2007>{{cite pmid|17940944|noedit}}</ref><ref name=Roddy2007>{{cite pmid|17940945|noedit}}</ref><ref name=Roddy2009>{{cite pmid|18838150|noedit}}</ref><ref name=Roddy2010>{{cite pmid|20441515|noedit}}</ref>
|- valign="TOP"
| 2007
| [[Uganda]]
| 1/3 (33%)<ref name=Towner2009>{{cite pmid|19649327|noedit}}</ref><ref name=Adjemian2011>{{cite pmid|21987753|noedit}}</ref>
|- valign="TOP"
| 2008
| [[Uganda]], [[Netherlands]], [[USA]]
| 1/2 (50%)<ref name=Timen2009>{{cite pmid|19751577|noedit}}</ref>
|- valign="TOP"
| 2012
| [[Uganda]]
| 9/18 (50%)<ref>{{cite web|title=Marburg hemorrhagic fever outbreak continues in Uganda|date=October 2012|url=http://www.healio.com/pediatrics/emerging-diseases/news/online/%7B52F1CE80-ACF7-4302-AB14-05428DDDA440%7D/Marburg-hemorrhagic-fever-outbreak-continues-in-Uganda-}}</ref>
|- valign="TOP"
| 2014
| [[Uganda]]
| 1/1 (100%)<ref>{{cite web|title=1st LD-Writethru: Deadly Marburg hemorrhagic fever breaks out in Uganda|date=October 5, 2014|url=http://www.china.org.cn/world/Off_the_Wire/2014-10/05/content_33686011.htm}}</ref><ref>{{cite news |last=Ntale |first=Samson |url=http://www.cnn.com/2014/10/07/health/uganda-marburg-death/index.html?hpt=wo_bn7 |title=99 in Uganda quarantined after Marburg virus death |work=CNN |date=October 8, 2014 |accessdate=2014-10-19 }}</ref>
|}

==Virology==

===Genome===
Like all [[Mononegavirales|mononegaviruses]], marburgvirions contain non-infectious, linear nonsegmented, single-stranded [[RNA]] [[genome]]s of negative polarity that possesses inverse-complementary 3' and 5' termini, do not possess a [[5' cap]], are not [[Polyadenylation|polyadenylated]], and are not [[Covalent bond|covalently]] linked to a [[protein]].<ref name=Fauquet2005>{{Cite book|last1=Pringle|first1=C. R.|chapter=Order Mononegavirales|year=2005|editor-last=Fauquet|editor-first=C. M.|editor2-last=Mayo|editor2-first=M. A.|editor3-last=Maniloff|editor3-first=J.|editor4-last=Desselberger|editor4-first=U.|editor5-last=Ball|editor5-first=L. A.|title=Virus Taxonomy—Eighth Report of the International Committee on Taxonomy of Viruses|pages=609–614|publisher=Elsevier/Academic Press|location=San Diego, US|isbn=0-12-370200-3|postscript=<!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}}}}</ref> Marburgvirus genomes are approximately 19 [[base pair|kb]] long and contain seven [[gene]]s in the order [[Three prime untranslated region|3'-UTR]]-''NP''-''VP35''-''VP40''-''GP''-''VP30''-''VP24''-''L''-[[Five prime untranslated region|5'-UTR]].<ref name = Kiley1982>{{cite pmid|7118520|noedit}}</ref> The genomes of the two different marburgviruses (MARV and RAVV) differ in [[nucleic acid sequence|sequence]].

===Structure===
[[File:Marburg em1986.png|thumb|CryoEM reconstruction of a section of the Marburg virus nucleocapsid. [[EMDB]] entry {{PDBe|EMD-1986}}<ref>{{cite PMID|22110401}}</ref>]]Like all [[Filoviridae|filoviruses]], marburgvirions are filamentous particles that may appear in the shape of a shepherd's crook or in the shape of a "U" or a "6", and they may be coiled, toroid, or branched.<ref name = Kiley1982/> Marburgvirions are generally 80&nbsp;nm in [[width]], but vary somewhat in length. In general, the median particle length of marburgviruses ranges from 795 to 828&nbsp;nm (in contrast to [[Ebolavirus|ebolavirions]], whose median particle length was measured to be 974–1,086&nbsp;nm ), but particles as long as 14,000&nbsp;nm have been detected in tissue culture.<ref name = Geisbert1995>{{cite pmid|8837880|noedit}}</ref> Marburgvirions consist of seven structural proteins. At the center is the [[helical]] [[ribonucleoprotein|ribonucleocapsid]], which consists of the genomic RNA wrapped around a [[polymer]] of [[nucleoprotein]]s (NP). Associated with the ribonucleoprotein is the [[RNA-dependent RNA polymerase]] (L) with the polymerase cofactor (VP35) and a transcription activator (VP30). The ribonucleoprotein is embedded in a matrix, formed by the major (VP40) and minor (VP24) matrix proteins. These particles are surrounded by a [[lipid bilayer|lipid membrane]] derived from the host cell membrane. The membrane anchors a glycoprotein (GP<sub>1,2</sub>) that projects 7 to 10&nbsp;nm spikes away from its surface. While nearly identical to ebolavirions in structure, marburgvirions are [[antigen]]ically distinct.

===Entry===

[[NPC1|Niemann–Pick C1]] (NPC1) cholesterol transporter protein appears to be essential for infection with both [[Ebola virus|Ebola]] and Marburg virus. Two independent studies reported in the same issue of ''[[Nature (journal)|Nature]]'' showed that [[Ebola virus]] cell entry and replication requires NPC1.<ref name="pmid21866103">{{cite journal | author = Carette JE, Raaben M, Wong AC, Herbert AS, Obernosterer G, Mulherkar N, Kuehne AI, Kranzusch PJ, Griffin AM, Ruthel G, Dal Cin P, Dye JM, Whelan SP, Chandran K, Brummelkamp TR | title = Ebola virus entry requires the cholesterol transporter Niemann-Pick C1 | journal = Nature | volume = 477 | issue = 7364 | pages = 340–3 |date=September 2011 | pmid = 21866103 | pmc = 3175325 | doi = 10.1038/nature10348 | laysummary = http://www.nytimes.com/2012/01/17/health/npc1-protein-may-give-ebola-its-opening.html | laysource = New York Times }}</ref><ref name="pmid21866101">{{cite journal | author = Côté M, Misasi J, Ren T, Bruchez A, Lee K, Filone CM, Hensley L, Li Q, Ory D, Chandran K, Cunningham J | title = Small molecule inhibitors reveal Niemann-Pick C1 is essential for Ebola virus infection | journal = Nature | volume = 477 | issue = 7364 | pages = 344–8 |date=September 2011 | pmid = 21866101 | pmc = 3230319 | doi = 10.1038/nature10380 | laysummary = http://www.nytimes.com/2012/01/17/health/npc1-protein-may-give-ebola-its-opening.html | laysource = New York Times }}</ref> When cells from patients lacking NPC1 were exposed to Ebola virus in the laboratory, the cells survived and appeared immune to the [[virus]], further indicating that Ebola relies on NPC1 to enter cells. This might imply that genetic mutations in the NPC1 gene in humans could make some people resistant to one of the deadliest known viruses affecting humans. The same studies described similar results with Marburg virus, showing that it also needs NPC1 to enter cells.<ref name="pmid21866103"/><ref name="pmid21866101"/> Furthermore, NPC1 was shown to be critical to [[filovirus]] entry because it mediates infection by binding directly to the [[viral envelope]] glycoprotein<ref name="pmid21866101"/> and that the second lysosomal domain of NPC1 mediates this binding.<ref name="pmid22395071">{{cite journal | author = Miller EH, Obernosterer G, Raaben M, Herbert AS, Deffieu MS, Krishnan A, Ndungo E, Sandesara RG, Carette JE, Kuehne AI, Ruthel G, Pfeffer SR, Dye JM, Whelan SP, Brummelkamp TR, Chandran K | title = Ebola virus entry requires the host-programmed recognition of an intracellular receptor | journal = EMBO Journal | volume = 31 | issue = 8 | pages = 1947–60 |date=March 2012 | pmid = 22395071 | pmc = 3343336 | doi = 10.1038/emboj.2012.53 }}</ref>

In one of the original studies, a [[small molecule]] was shown to inhibit Ebola virus infection by preventing the virus glycoprotein from binding to NPC1.<ref name="pmid21866101"/><ref name="pmid21959282">{{cite journal | author = Flemming A | title = Achilles heel of Ebola viral entry | journal = Nat Rev Drug Discov | volume = 10 | issue = 10 | pages = 731 |date=October 2011 | pmid = 21959282 | doi = 10.1038/nrd3568 }}</ref> In the other study, mice that were heterozygous for NPC1 were shown to be protected from lethal challenge with mouse-adapted Ebola virus.<ref name="pmid21866103"/> Together, these studies suggest NPC1 may be potential therapeutic target for an Ebola antiviral drug.

===Replication===
The marburg virus [[Biological life cycle|life cycle]] begins with virion attachment to specific cell-surface [[Receptor (biochemistry)|receptors]], followed by [[Lipid bilayer fusion|fusion]] of the virion envelope with cellular membranes and the concomitant release of the virus [[nucleocapsid]] into the [[cytosol]]. The virus RdRp partially uncoats the nucleocapsid and [[Transcription (genetics)|transcribes]] the [[genes]] into positive-stranded [[mRNA]]s, which are then [[translation (biology)|translated]] into structural and nonstructural [[proteins]]. Marburgvirus L binds to a single [[Promoter (biology)|promoter]] located at the 3' end of the genome. Transcription either terminates after a gene or continues to the next gene downstream. This means that genes close to the 3' end of the genome are transcribed in the greatest abundance, whereas those toward the 5' end are least likely to be transcribed. The gene order is therefore a simple but effective form of transcriptional regulation. The most abundant protein produced is the [[nucleoprotein]], whose [[concentration]] in the cell determines when L switches from gene transcription to genome replication. Replication results in full-length, positive-stranded antigenomes that are in turn transcribed into negative-stranded virus progeny genome copies. Newly synthesized structural proteins and genomes self-assemble and accumulate near the inside of the [[cell membrane]]. Virions [[Budding|bud]] off from the cell, gaining their envelopes from the cellular membrane they bud from. The mature progeny particles then infect other cells to repeat the cycle.<ref name=Feldmann2005/>

==Ecology==
In 2009, the successful isolation of infectious MARV was reported from caught healthy [[Rousettus aegyptiacus|Egyptian rousettes (''Rousettus aegyptiacus'')]].<ref name="Towner2009"/> This isolation, together with the isolation of infectious [[RAVV]],<ref name=Towner2009/> strongly suggests that [[Old World]] fruit [[bat]]s are involved in the natural maintenance of marburgviruses. Further studies are necessary to establish whether Egyptian rousettes are the actual hosts of MARV and RAVV or whether they get infected via contact with another animal and therefore serve only as intermediate hosts. Recently the first experimental infection study of ''Rousettus aegyptiacus'' with MARV provided further insight into the possible involvement of these bats in MARV ecology.<ref>{{Cite journal
| doi = 10.1371/journal.pone.0045479
| pmid = 23029039
| title = Virological and Serological Findings in Rousettus aegyptiacus Experimentally Inoculated with Vero Cells-Adapted Hogan Strain of Marburg Virus
| journal = PLoS ONE
| volume = 7
| issue = 9
| pages = e45479
| year = 2012
| last1 = Paweska | first1 = J. T.
| last2 = Jansen Van Vuren | first2 = P.
| last3 = Masumu | first3 = J.
| last4 = Leman | first4 = P. A.
| last5 = Grobbelaar | first5 = A. A.
| last6 = Birkhead | first6 = M.
| last7 = Clift | first7 = S.
| last8 = Swanepoel | first8 = R.
| last9 = Kemp | first9 = A.

| pmc = 3444458
}} {{open access}}</ref> Experimentally infected bats developed relatively low viremia lasting at least 5 days, but remained healthy and didn't develop any notable gross pathology. The virus also replicated to high titers in major organs (liver and spleen), and organs that might possibly be involved in virus transmission (lung, intestine, reproductive organ, salivary gland, kidney, bladder and mammary gland). The relatively long period of viremia noted in this experiment could possibly also facilitate mechanical transmission by blood sucking arthropods or infection of susceptible vertebrate hosts by direct contact with infected blood.

==Biological weapon==
The [[Soviet Union]] had an extensive offensive and defensive [[Biological warfare|biological weapon]]s program that included MARV.<ref name=Alibek1999>{{Cite book|last1=Alibek|first1=Steven|last2=Handelman|title=Biohazard: The Chilling True Story of the Largest Covert Biological Weapons Program in the World—Told from Inside by the Man Who Ran It|publisher=Random House|location=New York |isbn=0-385-33496-6|postscript=<!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}}}}</ref> At least three Soviet research institutes had MARV research programs during offensive times: the Virology Center of the Scientific-Research Institute for Microbiology in Zagorsk (today [[Sergiev Posad]]), the Scientific-Production Association "Vektor" (today the [[State Research Center of Virology and Biotechnology VECTOR|State Research Center of Virology and Biotechnology "Vektor"]]) in [[Koltsovo, Novosibirsk Oblast|Koltsovo]], and the Irkutsk Scientific-Research Anti-Plague Institute of Siberia and the Far East in [[Irkutsk]]. As most performed research was highly [[classified information|classified]], it remains unclear how successful the MARV program was. However, Soviet [[defection|defector]] [[Ken Alibek]] claimed that a weapon filled with MARV was tested at the [[Stepnagorsk Scientific and Technical Institute for Microbiology|Stepnogorsk Scientific Experimental and Production Base]] in [[Stepnogorsk]], [[Kazakh Soviet Socialist Republic]] (today [[Kazakhstan]]),<ref name=Alibek1999/> suggesting that the development of a MARV biological weapon had reached advanced stages. Independent confirmation for this claim is lacking. At least one laboratory accident with MARV, resulting in the death of Koltsovo researcher Nikolai Ustinov, occurred during offensive times in the Soviet Union and was first described in detail by Alibek.<ref name=Alibek1999/> After the [[dissolution of the Soviet Union]], MARV research continued in all three institutes.{{cn|date=July 2014}}

==In popular culture==
{{Unreferenced|section|date=June 2013}}
{{Cleanup-list|section=yes|date=October 2014}}
* In the non-fiction thriller, ''[[The Hot Zone]]'', [[Richard Preston]] describes several MARV infections.
* In the 2008 Indian science fiction movie ''[[Dasavathaaram]]'' by [[Kamal Haasan]], the plot features an intended bio weapon of "Ebola Marburg" virus.
* In the TV series [[Millennium (TV series)|''Millennium'']], at the end of Season 2, a "[[prion]] version" of MARV causes a disease outbreak in [[Seattle]], killing (amongst others) Frank Black's wife, Catherine. In the Season 3 episode "Collateral Damage", Peter Watt's daughter is infected with MARV by a Gulf War veteran who claims that the Millennium Group did the same to American soldiers during the first Gulf War.
* In the crossover event of the TV series ''[[Medical Investigation]]'', episode 17, and ''[[Third Watch]]'', season 6 episode 16, Marburg virus disease breaks out in [[New York City]], killing five of six infected people.
* In the ''[[Sarah Jane Smith]]'' series (Series Two), MARV is used as a weapon by a [[doomsday cult]].
* In the short story ''Hell Hath Enlarged Herself'' by [[Michael Marshall Smith]], one of the original scientists is infected with MARV in an attempt to test ImmunityWorks ver. 1.0.
* In the novel ''[[Microserfs]]'' by [[Douglas Coupland]], MARV is mentioned several times as a metaphor for the spread of information through the internet
* In the novel ''[[Resident Evil: Caliban Cove]]'', an insane scientist and former professor named Nicolas Griffith is referred to by Rebecca Chambers as having infected three men with MARV after they had been led to believe it was a harmless [[common cold]] virus.
* In the novel ''Pandora's Legion'' by [[Harold Coyle]] and [[Barrett Tillman]], an [[Al-Qaeda]] cell in [[Pakistan]] injects volunteers with MARV, who then board flights to major international airports in the western world where the large flow of people would facilitate the spreading of the virus into a [[pandemic]].
* In the TV series ''[[Body of Proof]]'', Season 2, episodes 18 and 19 include a MARV outbreak.
* In [[Mira Grant]]'s novel ''[[Feed (Grant novel)|Feed]]'', a modified Marburg virus that cures cancer combines with a virally transmitted cure for the common cold, resulting in a virulent viral plague that turns infected humans and animals into zombies.
* Motaba, the fictional deadly viral hemorrhagic fever, in the movie ''[[Outbreak (film)|Outbreak]]'', is based on MARV.
* In the video game ''[[Trauma Team]]'', the seventh chapter of the game, named "Patient Zero", has a storyline of a mass outbreak of the fictional Rosalia Virus, which has similar symptoms to the [[Ebola Virus]] and Marburg Virus.
* In the episode "The Order 23 Job" of the TV show ''[[Leverage (TV series)|Leverage]]'', the team's mark is led to believe that he is caught in an outbreak of weaponized Marburg virus made by the Soviets.
* In the episode "Death Is in the Air" of the TV show ''[[Psych]]'', the fictional Thornburg virus is based on the Marburg virus.
* In the episode "Small Sacrifices" of the TV show "[[House MD]]", the team explores Marburg as a diagnosis for a patient
*In the episode "The Promise" of the Canadian TV show ''[[ReGenesis]]'' Marburg was the subject of a war games exercise and a weaponized strain out of a lab in South Africa poses a potential threat.
* In the episode "Honor Among Thieves" of the TV show ''[[Person of Interest (TV series)|Person of Interest]]'', the Marburg virus is shown to be used as a potential bioterrorism agent to cause a pandemic starting in New York.
* In the episode "I Am the Apocalypse" of the TV show ''[[Chicago Fire (TV series)|Chicago Fire]]'', a man with Marburg virus attempts to start an outbreak in a Chicago hospital.
* In the TV series ''[[Bergerac (TV series)|Bergerac]]'', a potential Marburg outbreak is the subject of the episode "The Deadly Virus".

==References==
{{Reflist|20em}}

==Further reading==
{{Refbegin}}
* {{Cite book|last=Klenk|first=Hans-Dieter|title=Marburg and Ebola Viruses. Current Topics in Microbiology and Immunology, vol. 235|year=1999|publisher=Springer-Verlag|location=Berlin, Germany|isbn=978-3-540-64729-4|postscript=<!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}}}}
* {{Cite book|first1=Hans-Dieter|last1=Klenk|first2=Heinz|last2=Feldmann|title=Ebola and Marburg Viruses: Molecular and Cellular Biology|year=2004|publisher=Horizon Bioscience|location=Wymondham, Norfolk, UK|isbn=978-1-904933-49-6|postscript=<!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}}}}
* {{Cite book|last=Kuhn|first=Jens H.|title=Filoviruses: A Compendium of 40 Years of Epidemiological, Clinical, and Laboratory Studies. Archives of Virology Supplement, vol. 20|year=2008|publisher=SpringerWienNewYork|location=Vienna, Austria|isbn=978-3-211-20670-6|postscript=<!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}}}}
* {{Cite book|last1=Martini|first1=G. A.|last2=Siegert|first2=R.|title=Marburg Virus Disease|year=1971|publisher=Springer-Verlag|location=Berlin, Germany|isbn=978-0-387-05199-4}}
* {{Cite book|last1=Ryabchikova|first1=Elena I.|last2=Price|first2=Barbara B.|title=Ebola and Marburg Viruses: A View of Infection Using Electron Microscopy|year=2004|publisher=Battelle Press|location=Columbus, Ohio, US
|isbn=978-1-57477-131-2|postscript=<!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}}}}
{{Refend}}

==External links==
{{Commons category|Marburg virus}}
* [http://talk.ictvonline.org International Committee on Taxonomy of Viruses (ICTV)]
* [http://www.filovir.com FILOVIR - scientific resources for research on filoviruses]

{{Filoviridae}}

[[Category:Marburgviruses| ]]
[[Category:Animal virology]]
[[Category:Arthropod-borne viral fevers and viral haemorrhagic fevers]]
[[Category:Biological weapons]]
[[Category:Hemorrhagic fevers]]
[[Category:Tropical diseases]]
[[Category:Viral diseases]]
[[Category:Virus-related cutaneous conditions]]
[[Category:Zoonoses]]

Revision as of 22:26, 29 July 2015

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