View source for Zaire ebolavirus

{{short description|Species of virus affecting humans and animals}}
{{Use dmy dates|date=April 2017}}

{{About|the species Zaire ebolavirus|the genus|Ebolavirus|the disease|Ebola virus disease|other uses|Ebola (disambiguation)}}
{{Virusbox
| image = Ebola Virus - Electron Micrograph.tiff
| image_alt =
| image_caption = Colorized scanning electron micrograph of Ebola virus particles (green) found both as extracellular particles and budding particles from a chronically infected African Green Monkey Kidney cell (blue); 20,000x magnification|
| parent = Ebolavirus
| species = Zaire ebolavirus
| authority =
| synonyms =
| synonyms_ref =
| subdivision_ranks =
| subdivision =
}}

'''''Zaire ebolavirus''''', more commonly known as '''Ebola virus''' ({{IPAc-en|i|ˈ|b|oʊ|l|ə|,_|ɪ|-}}; '''EBOV'''), is one of six known species within the [[genus]] ''[[Ebolavirus]]''.<ref name=KuhnArch/> Four of the six known ebolaviruses, including EBOV, cause a severe and often fatal [[viral hemorrhagic fever|hemorrhagic fever]] in [[human]]s and other [[mammals]], known as [[Ebola virus disease]] (EVD). Ebola virus has caused the majority of human deaths from EVD, and was the cause of the [[Western African Ebola virus epidemic|2013–2016 epidemic in western Africa]],<ref>{{Cite journal |last1=Na |first1=Woonsung |last2=Park |first2=Nanuri |last3=Yeom |first3=Minju |last4=Song |first4=Daesub |date=4 December 2016 |title=Ebola outbreak in Western Africa 2014: what is going on with Ebola virus? |journal=Clinical and Experimental Vaccine Research |volume=4 |issue=1 |pages=17–22 |doi=10.7774/cevr.2015.4.1.17 |issn=2287-3651 |pmc=4313106 |pmid=25648530}}</ref> which resulted in at least {{#section:Ebola virus epidemic in West Africa|cases}} suspected cases and {{#section:Ebola virus epidemic in West Africa|deaths}} confirmed deaths.{{#section:Ebola virus epidemic in West Africa|caserefs}}<ref>{{Cite web |title=Ebola virus disease outbreak |url=https://www.who.int/csr/disease/ebola/en/ |url-status=live |archive-url=https://web.archive.org/web/20210323074126/https://www.who.int/health-topics/ebola |archive-date=23 March 2021 |access-date=4 December 2016 |website=World Health Organization}}</ref>

Ebola virus and its genus were both originally named for [[Zaire]] (now the [[Democratic Republic of the Congo]]), the country where it was [[List of Ebola outbreaks|first described]],<ref name=KuhnArch/> and was at first suspected to be a new "strain" of the closely related [[Marburg virus]].<ref name=Pattyn1977/><ref name=Bowen1977/> The virus was renamed "Ebola virus" in 2010 to avoid confusion. Ebola virus is the single member of the [[species]] ''Zaire ebolavirus'', which is assigned to the genus ''[[Ebolavirus]]'', [[family (biology)|family]] ''[[Filoviridae]]'', [[order (biology)|order]] ''[[Mononegavirales]]''. The members of the species are called Zaire ebolaviruses.<ref name="KuhnArch">{{Cite journal |display-authors=8 |vauthors=Kuhn JH, Becker S, Ebihara H, Geisbert TW, Johnson KM, Kawaoka Y, Lipkin WI, Negredo AI, Netesov SV |year=2010 |title=Proposal for a revised taxonomy of the family Filoviridae: Classification, names of taxa and viruses, and virus abbreviations |journal=Archives of Virology |volume=155 |issue=12 |pages=2083–2103 |doi=10.1007/s00705-010-0814-x |pmc=3074192 |pmid=21046175}}</ref><ref>{{Cite web |last=WHO |title=Ebola virus disease |url=https://www.who.int/mediacentre/factsheets/fs103/en/ |url-status=live |archive-url=https://web.archive.org/web/20141214011751/https://www.who.int/mediacentre/factsheets/fs103/en/ |archive-date=14 December 2014 |access-date=5 October 2020}}</ref> The natural reservoir of Ebola virus is believed to be [[bat]]s, particularly [[megabat|fruit bats]],<ref name="Quanmmen 2014-12-30">{{Cite web |last=Quammen |first=David |author-link=David Quammen |date=2014-12-30 |title=Insect-Eating Bat May Be Origin of Ebola Outbreak, New Study Suggests |url=http://news.nationalgeographic.com/news/2014/12/141230-ebola-virus-origin-insect-bats-meliandou-reservoir-host/ |url-status=dead |archive-url=https://web.archive.org/web/20141231124300/http://news.nationalgeographic.com/news/2014/12/141230-ebola-virus-origin-insect-bats-meliandou-reservoir-host/ |archive-date=31 December 2014 |access-date=2014-12-30 |website=news.nationalgeographic.com |publisher=[[National Geographic Society]] |location=Washington, DC}}</ref> and it is primarily transmitted between humans and from animals to humans through [[body fluid]]s.<ref name="NYT-20141027-NA">{{Cite news |last=Angier |first=Natalie |date=October 27, 2014 |title=Killers in a Cell but on the Loose – Ebola and the Vast Viral Universe |url=https://www.nytimes.com/2014/10/28/science/ebola-and-the-vast-viral-universe.html |url-status=live |archive-url=https://web.archive.org/web/20201112020413/https://www.nytimes.com/2014/10/28/science/ebola-and-the-vast-viral-universe.html |archive-date=12 November 2020 |access-date=October 27, 2014 |work=[[New York Times]]}}</ref>

The EBOV genome is a single-stranded RNA, approximately 19,000 [[nucleotide]]s long. It encodes seven structural [[protein]]s: [[nucleoprotein]] (NP), [[Cofactor (biochemistry)|polymerase cofactor]] (VP35), (VP40), GP, [[transcription activator]] (VP30), [[VP24]], and [[RNA-dependent RNA polymerase]] (L).<ref>{{Cite journal |last1=Nanbo |first1=Asuka |last2=Watanabe |first2=Shinji |last3=Halfmann |first3=Peter |last4=Kawaoka |first4=Yoshihiro |date=4 Feb 2013 |title=The spatio-temporal distribution dynamics of Ebola virus proteins and RNA in infected cells |journal=Scientific Reports |volume=3 |issue=1 |pages=1206 |bibcode=2013NatSR...3.1206N |doi=10.1038/srep01206 |pmc=3563031 |pmid=23383374}}</ref>

Because of its high [[fatality rate]] (up to 83 to 90 percent),<ref name="WHO2014">{{Cite web |date=March 2014 |title=Ebola virus disease Fact sheet N°103 |url=https://www.who.int/mediacentre/factsheets/fs103/en/ |url-status=live |archive-url=https://web.archive.org/web/20141214011751/https://www.who.int/mediacentre/factsheets/fs103/en/ |archive-date=14 December 2014 |access-date=12 April 2014 |website=World Health Organization}}</ref><ref name="Elsevier/Academic Press">{{Cite book |url=https://books.google.com/books?id=9Wy7Jgy5RWYC |title=Virus Taxonomy – Eighth Report of the International Committee on Taxonomy of Viruses |date=2005 |publisher=Elsevier/Academic Press |isbn=978-0080575483 |veditors=Fauquet CM, Mayo MA, Maniloff J, Desselberger U, Ball LA |location=Oxford |page=648 |access-date=7 February 2016 |archive-url=https://web.archive.org/web/20230114100058/https://books.google.com/books?id=9Wy7Jgy5RWYC |archive-date=14 January 2023 |url-status=live}}</ref> EBOV is also listed as a [[select agent]], [[World Health Organization]] Risk Group 4 Pathogen (requiring [[Biosafety level#Biosafety level 4|Biosafety Level 4-equivalent containment]]), a US [[National Institutes of Health]]/[[National Institute of Allergy and Infectious Diseases]] Category A Priority Pathogen, US CDC [[Centers for Disease Control and Prevention]] [[Bioterrorism|Category A Bioterrorism Agent]], and a Biological Agent for Export Control by the [[Australia Group]].{{citation needed|date=December 2016}}

==Structure==
[[File:Filovirus phylogenetic tree.svg|370px|thumb|[[Phylogenetic tree]] comparing ebolaviruses and marburgviruses. Numbers indicate percent confidence of branches.]]
[[File:Ebola virus Makona from West African Epidemic (43234028254).jpg|thumb|left|A colorized scanning electron micrograph of the virus]]
EBOV carries a [[negative-sense]] RNA genome in virions that are cylindrical/tubular, and contain [[viral envelope]], matrix, and nucleocapsid components. The overall cylinders are generally approximately 80&nbsp;[[nanometer|nm]] in diameter, and have a virally encoded [[glycoprotein]] (GP) projecting as 7–10&nbsp;nm long spikes from its lipid bilayer surface.<ref>{{Cite book |title=Ebola and Marburg Viruses – Molecular and Cellular Biology |publisher=Horizon Bioscience |year=2004 |isbn=978-0-9545232-3-7 |editor-last=Klenk |editor-first=H.-D. |location=Wymondham, Norfolk, UK |page=28 |editor-last2=Feldmann |editor-first2=H.}}</ref> The cylinders are of variable length, typically 800&nbsp;nm, but sometimes up to 1000&nbsp;nm long. The outer [[viral envelope]] of the virion is derived by budding from domains of host cell membrane into which the GP spikes have been inserted during their biosynthesis. Individual GP molecules appear with spacings of about 10&nbsp;nm. Viral proteins [[VP40]] and VP24 are located between the envelope and the nucleocapsid (see following), in the ''matrix space''.<ref name="pmid8219816">{{Cite book |last=Feldmann |first=H. K. |title=Unconventional Agents and Unclassified Viruses |work=Archives of Virology. Supplementum |year=1993 |isbn=978-3211824801 |series=Archives of Virology |volume=7 |pages=81–100 |chapter=Molecular biology and evolution of filoviruses |publisher=Springer |location=Vienna |doi=10.1007/978-3-7091-9300-6_8 |issn=0939-1983 |pmid=8219816}}</ref> At the center of the virion structure is the [[nucleocapsid]], which is composed of a series of viral proteins attached to an 18–19 kb linear, negative-sense RNA without 3′-[[polyadenylation]] or 5′-capping (see following); the RNA is helically wound and complexed with the NP, VP35, VP30, and L proteins; this helix has a diameter of 80&nbsp;nm.<ref>{{Cite journal |last1=Lee |first1=Jeffrey E |last2=Saphire |first2=Erica Ollmann |date=2009 |title=Ebolavirus glycoprotein structure and mechanism of entry |journal=Future Virology |volume=4 |issue=6 |pages=621–635 |doi=10.2217/fvl.09.56 |issn=1746-0794 |pmc=2829775 |pmid=20198110}}</ref><ref>{{Cite journal |vauthors=Falasca L, Agrati C, Petrosillo N, Di Caro A, Capobianchi MR, Ippolito G, Piacentini M |date=4 December 2016 |title=Molecular mechanisms of Ebola virus pathogenesis: focus on cell death |journal=Cell Death and Differentiation |volume=22 |issue=8 |pages=1250–1259 |doi=10.1038/cdd.2015.67 |issn=1350-9047 |pmc=4495366 |pmid=26024394}}</ref><ref>{{Cite journal |last1=Swetha |first1=Rayapadi G. |last2=Ramaiah |first2=Sudha |last3=Anbarasu |first3=Anand |last4=Sekar |first4=Kanagaraj |date=2016 |title=Ebolavirus Database: Gene and Protein Information Resource for Ebolaviruses |journal=Advances in Bioinformatics |volume=2016 |pages=1673284 |doi=10.1155/2016/1673284 |issn=1687-8027 |pmc=4848411 |pmid=27190508 |doi-access=free}}</ref>

The overall shape of the virions after purification and visualization (e.g., by [[ultracentrifugation]] and [[electron microscopy]], respectively) varies considerably; simple cylinders are far less prevalent than structures showing reversed direction, branches, and loops (e.g., U-, [[shepherd's crook]]-, 9-, or [[eye bolt]]-shapes, or other or circular/coiled appearances), the origin of which may be in the laboratory techniques applied.<ref name="Klenk2">{{Cite book |title=Ebola and Marburg Viruses: Molecular and Cellular Biology |publisher=Horizon Bioscience |year=2004 |isbn=978-1904933496 |editor-last=Klenk |editor-first=H.-D. |editor-last2=Feldmann |editor-first2=H.}}{{page needed|date=August 2020}}</ref><ref name="Hillman">{{Cite book |last=Hillman, H. |title=The Case for New Paradigms in Cell Biology and in Neurobiology |publisher=Edwin Mellen Press |year=1991}}</ref> The characteristic "threadlike" structure is, however, a more general morphologic characteristic of filoviruses (alongside their GP-decorated viral envelope, RNA nucleocapsid, etc.).<ref name="Klenk2" />

==Genome==
Each virion contains one molecule of linear, single-stranded, negative-sense RNA, 18,959 to 18,961 nucleotides in length.<ref>[https://www.ncbi.nlm.nih.gov/nuccore/674810549 Zaire ebolavirus isolate H.sapiens-wt/GIN/2014/Makona-Kissidougou-C15, complete genome] {{Webarchive|url=https://web.archive.org/web/20180124145827/https://www.ncbi.nlm.nih.gov/nuccore/674810549 |date=24 January 2018 }}, [[GenBank]]</ref> The 3′ terminus is not polyadenylated and the 5′ end is not capped. This viral genome codes for seven structural proteins and one non-structural protein. The gene order is 3′ – leader – NP – VP35 – VP40 – GP/sGP – VP30 – VP24 – L – trailer – 5′; with the leader and trailer being non-transcribed regions, which carry important signals to control transcription, replication, and packaging of the viral genomes into new virions. Sections of the NP, VP35 and the L genes from filoviruses have been identified as endogenous in the genomes of several groups of small mammals.<ref name="Cite pmid| 20569424">{{Cite journal |vauthors=Taylor D, Leach R, Bruenn J |year=2010 |title=Filoviruses are ancient and integrated into mammalian genomes |journal=BMC Evolutionary Biology |volume=10 |issue=1 |pages=193 |bibcode=2010BMCEE..10..193T |doi=10.1186/1471-2148-10-193 |pmc=2906475 |pmid=20569424 |doi-access=free}}</ref><ref name="Cite pmid| 20686665">{{Cite journal |last1=Belyi |first1=V. A. |last2=Levine |first2=A. J. |last3=Skalka |first3=A. M. |year=2010 |editor-last=Buchmeier |editor-first=Michael J. |title=Unexpected Inheritance: Multiple Integrations of Ancient Bornavirus and Ebolavirus/Marburgvirus Sequences in Vertebrate Genomes |journal=PLOS Pathogens |volume=6 |issue=7 |pages=e1001030 |doi=10.1371/journal.ppat.1001030 |pmc=2912400 |pmid=20686665 |doi-access=free}}</ref><ref name="cite journal|doi= 10.7717/peerj.556">{{Cite journal |vauthors=Taylor DJ, Ballinger MJ, Zhan JJ, Hanzly LE, Bruenn JA |year=2014 |title=Evidence that ebolaviruses and cuevaviruses have been diverging from marburgviruses since the Miocene |journal=PeerJ |volume=2 |pages=e556 |doi=10.7717/peerj.556 |pmc=4157239 |pmid=25237605 |doi-access=free}}</ref>

[[File:Ebola Genome structure.png|Genomic structure of Ebola virus, mostly after GenBank KJ660346.2]]

It was found that 472 nucleotides from the 3' end and 731 nucleotides from the 5' end are sufficient for replication of a viral "minigenome", though not sufficient for infection.<ref name="Klenk2" /> Virus sequencing from 78 patients with confirmed Ebola virus disease, representing more than 70% of cases diagnosed in Sierra Leone from late May to mid-June 2014,<ref name="New Yorker">{{Cite magazine |last=Richard Preston |date=27 October 2014 |title=The Ebola Wars |url=https://www.newyorker.com/magazine/2014/10/27/ebola-wars |url-status=live |archive-url=https://web.archive.org/web/20210125044952/https://www.newyorker.com/magazine/2014/10/27/ebola-wars |archive-date=25 January 2021 |access-date=20 October 2014 |magazine=[[The New Yorker]] |publisher=[[Condé Nast]] |location=New York}}</ref><ref name="Gire2014">{{Cite journal |last=Gire |first=Stephen K. |display-authors=etal |year=2014 |title=Genomic surveillance elucidates Ebola virus origin and transmission during the 2014 outbreak |journal=[[Science (journal)|Science]] |volume=345 |issue=6202 |pages=1369–1372 |bibcode=2014Sci...345.1369G |doi=10.1126/science.1259657 |pmc=4431643 |pmid=25214632}}</ref> provided evidence that the 2014 outbreak was no longer being fed by new contacts with its natural reservoir. Using [[third-generation sequencing]] technology, investigators were able to sequence samples as quickly as 48 hours.<ref>{{Cite journal |last=Check Hayden |first=Erika |date=2015-05-05 |title=Pint-sized DNA sequencer impresses first users |journal=Nature |language=en |volume=521 |issue=7550 |pages=15–16 |bibcode=2015Natur.521...15C |doi=10.1038/521015a |issn=0028-0836 |pmid=25951262 |doi-access=free}}</ref> Like other RNA viruses,<ref name="New Yorker" /> Ebola virus mutates rapidly, both within a person during the progression of disease and in the reservoir among the local human population.<ref name="Gire2014" /> The observed mutation rate of 2.0 x 10<sup>−3</sup> substitutions per site per year is as fast as that of seasonal [[influenza]].<ref>{{Cite journal |vauthors=Jenkins GM, Rambaut A, Pybus OG, Holmes EC |year=2002 |title=Rates of molecular evolution in RNA viruses: A quantitative phylogenetic analysis |journal=Journal of Molecular Evolution |volume=54 |issue=2 |pages=156–165 |bibcode=2002JMolE..54..156J |doi=10.1007/s00239-001-0064-3 |pmid=11821909 |s2cid=20759532}}</ref>

{|class=wikitable style="width:100%"
|+Proteins encoded by ''Zaire ebolavirus''
! Symbol !! Name !! UniProt !! Function
|-
| NP || Nucleoprotein || {{UniProt|P18272}} || Wraps genome for protection from nucleases and innate immunity.
|-
| VP35 || Polymerase cofactor VP35 || {{UniProt|Q05127}} || Polymerase cofactor; suppresses innate immunity by binding RNA.
|-
| VP40 || Matrix protein VP40 || {{UniProt|Q05128}} || Matrix.
|-
| GP || Envelope glycoprotein || {{UniProt|Q05320}} || Cleaved by host furin into GP1/2 to form envelope with spikes. Also makes shed GP as a decoy.
|-
| sGP || Pre-small/secreted glycoprotein || {{UniProt|P60170}} || Shares ORF with GP.  Cleaved by host furin into sGP (anti-inflammatory) and delta-peptide (viroporin).
|-
| ssGP || Super small secreted glycoprotein || {{UniProt|Q9YMG2}} || Shares ORF with GP; created by mRNA editing. Unknown function.
|-
| VP30 || Hexameric zinc-finger protein VP30 || {{UniProt|Q05323}} || Transcriptional activator.
|-
| VP24 || Membrane-associated protein VP24 || {{UniProt|Q05322}} || Blocks IFN-alpha/beta and IFN-gamma signaling.
|-
| L || RNA-directed RNA polymerase L || {{UniProt|Q05318}} || [[RNA replicase]].
|}

==Entry==
[[File:Protein NPC1 PDB 3GKH.png|thumb|NPC1]]

There are two candidates for host cell entry proteins. The first is a cholesterol transporter protein, the host-encoded Niemann–Pick C1 ([[NPC1]]), which appears to be essential for entry of Ebola virions into the host cell and for its ultimate replication.<ref name="pmid21866103">{{cite journal |vauthors=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 = 340–343 |date=September 2011 | pmid = 21866103 | pmc = 3175325 | doi = 10.1038/nature10348 | bibcode = 2011Natur.477..340C }}
* {{Cite news |last=Amanda Schaffer |date=January 16, 2012 |title=Key Protein May Give Ebola Virus Its Opening |url=https://www.nytimes.com/2012/01/17/health/npc1-protein-may-give-ebola-its-opening.html |url-access=subscription |url-status=live |archive-url=https://web.archive.org/web/20220220214545/https://www.nytimes.com/2012/01/17/health/npc1-protein-may-give-ebola-its-opening.html |archive-date=20 February 2022 |access-date=26 February 2017 |work=The New York Times}}</ref><ref name="pmid21866101">{{cite journal |vauthors=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 = 344–348 |date=September 2011 | pmid = 21866101 | pmc = 3230319 | doi = 10.1038/nature10380 | bibcode = 2011Natur.477..344C }}
* {{Cite news |last=Amanda Schaffer |date=January 16, 2012 |title=Key Protein May Give Ebola Virus Its Opening |url=https://www.nytimes.com/2012/01/17/health/npc1-protein-may-give-ebola-its-opening.html |url-access=subscription |url-status=live |archive-url=https://web.archive.org/web/20220220214545/https://www.nytimes.com/2012/01/17/health/npc1-protein-may-give-ebola-its-opening.html |archive-date=20 February 2022 |access-date=26 February 2017 |work=The New York Times}}</ref> In one study, mice with one copy of the NPC1 gene [[Gene knockout|removed]] showed an 80 percent survival rate fifteen days after exposure to mouse-adapted Ebola virus, while only 10 percent of unmodified mice survived this long.<ref name="pmid21866103" /> In another study, [[small molecule]]s were shown to inhibit Ebola virus infection by preventing [[viral envelope]] glycoprotein (GP) from binding to NPC1.<ref name="pmid21866101" /><ref name="pmid21959282">{{Cite journal |vauthors=Flemming A |date=October 2011 |title=Achilles heel of Ebola viral entry |journal=Nat Rev Drug Discov |volume=10 |issue=10 |pages=731 |doi=10.1038/nrd3568 |pmid=21959282 |s2cid=26888076 |doi-access=free}}</ref> Hence, NPC1 was shown to be critical to entry of this [[filovirus]], because it mediates infection by binding directly to viral GP.<ref name="pmid21866101" />

When cells from [[Niemann–Pick disease, type C|Niemann–Pick Type C]] individuals lacking this transporter were exposed to Ebola virus in the laboratory, the cells survived and appeared impervious to the virus, further indicating that Ebola relies on NPC1 to enter cells;<ref name="pmid21866103" /> mutations in the NPC1 gene in humans were conjectured as a possible mode to make some individuals resistant to this deadly viral disease. The same studies described similar results regarding NPC1's role in virus entry for [[Marburg virus]], a related [[filovirus]].<ref name="pmid21866103" /> A further study has also presented evidence that NPC1 is the critical receptor mediating Ebola infection via its direct binding to the viral GP, and that it is the second "lysosomal" domain of NPC1 that mediates this binding.<ref name="pmid22395071">{{Cite journal |vauthors=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 |date=March 2012 |title=Ebola virus entry requires the host-programmed recognition of an intracellular receptor |journal=EMBO Journal |volume=31 |issue=8 |pages=1947–1960 |doi=10.1038/emboj.2012.53 |pmc=3343336 |pmid=22395071}}</ref>

The second candidate is TIM-1 (a.k.a. [[HAVCR1]]).<ref>{{Cite journal |display-authors=etal |vauthors=Kondratowicz AS, Lennemann NJ, Sinn PL |date=May 2011 |title=T-cell immunoglobulin and mucin domain 1 (TIM-1) is a receptor for Zaire Ebolavirus and Lake Victoria Marburgvirus |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=108 |issue=20 |pages=8426–8431 |bibcode=2011PNAS..108.8426K |doi=10.1073/pnas.1019030108 |pmc=3100998 |pmid=21536871 |doi-access=free}}</ref> TIM-1 was shown to bind to the receptor binding domain of the EBOV glycoprotein, to increase the receptivity of [[Vero cells]]. Silencing its effect with siRNA prevented infection of [[Vero cells]]. TIM1 is expressed in tissues known to be seriously impacted by EBOV lysis (trachea, cornea, and conjunctiva). A monoclonal antibody against the IgV domain of TIM-1, ARD5, blocked EBOV binding and infection. Together, these studies suggest NPC1 and TIM-1 may be potential therapeutic targets for an Ebola anti-viral drug and as a basis for a rapid field diagnostic assay.{{citation needed|date=August 2014}}

==Replication==
[[File:178-EbolaVirusProteins EbolaProteins.png|thumb|300px|Cross-sectional drawing of the Ebola virus particle, with structures of the major proteins shown and labeled at the side. Pale circles represent domains too flexible to be observed in the experimental structure. Drawn by David Goodsell from PDB files 3csy, 4ldd, 4qb0, 3vne, 3fke, and 2i8b.]]

Being acellular, viruses such as Ebola do not replicate through any type of cell division; rather, they use a combination of host- and virally encoded enzymes, alongside host cell structures, to produce multiple copies of themselves. These then self-assemble into viral [[macromolecular assembly|macromolecular structures]] in the host cell.<ref name="BiomarkerDatabase">{{Cite book |last=Biomarker Database |url=http://biomarker.cdc.go.kr:8080/pathogen/pathogen_view_en.jsp?pclass=2&id=44 |title=Ebola virus |publisher=Korea National Institute of Health |access-date=2009-05-31 |archive-url=https://web.archive.org/web/20080422041525/http://biomarker.cdc.go.kr:8080/pathogen/pathogen_view_en.jsp?pclass=2&id=44 |archive-date=2008-04-22 |url-status=dead}}</ref> The virus completes a set of steps when infecting each individual cell. The virus begins its attack by attaching to host receptors through the glycoprotein (GP) surface [[peplomer]] and is [[Endocytosis|endocytosed]] into [[macropinosome]]s in the host cell.<ref>{{Cite journal |vauthors=Saeed MF, Kolokoltsov AA, Albrecht T, Davey RA |year=2010 |editor-last=Basler |editor-first=Christopher F. |title=Cellular Entry of Ebola Virus Involves Uptake by a Macropinocytosis-Like Mechanism and Subsequent Trafficking through Early and Late Endosomes |journal=PLOS Pathogens |volume=6 |issue=9 |pages=e1001110 |doi=10.1371/journal.ppat.1001110 |pmc=2940741 |pmid=20862315 |doi-access=free}}</ref> To penetrate the cell, the viral membrane fuses with [[Vesicle (biology)|vesicle]] membrane, and the [[nucleocapsid]] is released into the [[cytoplasm]]. Encapsidated, negative-sense genomic ssRNA is used as a template for the synthesis (3'–5') of polyadenylated, monocistronic mRNAs and, using the host cell's ribosomes, tRNA molecules, etc., the mRNA is translated into individual viral proteins.<ref>{{Cite journal |last=Mühlberger |first=Elke |date=4 December 2016 |title=Filovirus replication and transcription |journal=Future Virology |volume=2 |issue=2 |pages=205–215 |doi=10.2217/17460794.2.2.205 |issn=1746-0794 |pmc=3787895 |pmid=24093048}}</ref><ref>{{Cite book |last1=Feldmann |first1=H. |url=https://www.ncbi.nlm.nih.gov/books/NBK8129/ |title=Filoviruses |last2=Klenk |first2=H.-D. |date=1996 |work=Medical Microbiology |publisher=University of Texas Medical Branch at Galveston |isbn=978-0963117212 |access-date=4 December 2016 |archive-url=https://web.archive.org/web/20180909093501/https://www.ncbi.nlm.nih.gov/books/NBK8129/ |archive-date=9 September 2018 |url-status=live}}</ref><ref>{{Cite journal |last1=Lai |first1=Kang Yiu |last2=Ng |first2=Wing Yiu George |last3=Cheng |first3=Fan Fanny |date=28 November 2014 |title=Human Ebola virus infection in West Africa: a review of available therapeutic agents that target different steps of the life cycle of Ebola virus |journal=Infectious Diseases of Poverty |volume=3 |issue=1 |pages=43 |doi=10.1186/2049-9957-3-43 |issn=2049-9957 |pmc=4334593 |pmid=25699183 |doi-access=free}}</ref>

These viral proteins are processed: a glycoprotein precursor (GP0) is cleaved to GP1 and GP2, which are then heavily glycosylated using cellular enzymes and substrates. These two molecules assemble, first into heterodimers, and then into trimers to give the surface peplomers. Secreted glycoprotein (sGP) precursor is cleaved to sGP and delta peptide, both of which are released from the cell. As viral protein levels rise, a switch occurs from translation to replication. Using the negative-sense genomic RNA as a template, a complementary +ssRNA is synthesized; this is then used as a template for the synthesis of new genomic (-)ssRNA, which is rapidly encapsidated. The newly formed nucleocapsids and envelope proteins associate at the host cell's plasma membrane; [[Viral shedding|budding]] occurs, destroying the cell.{{citation needed|date=December 2016}}

==Ecology==
Ebola virus is a [[zoonosis|zoonotic]] pathogen. Intermediary hosts have been reported to be "various species of fruit bats&nbsp;... throughout central and sub-Saharan Africa". Evidence of infection in bats has been detected through molecular and serologic means. However, ebolaviruses have not been isolated in bats.<ref name="Quanmmen 2014-12-30" /><ref name="nejm52014">{{Cite journal |vauthors=Feldmann H |date=May 2014 |title=Ebola{{snd}}A Growing Threat? |journal=N. Engl. J. Med. |volume=371 |issue=15 |pages=1375–1378 |doi=10.1056/NEJMp1405314 |pmid=24805988 |s2cid=4657264 |doi-access=free}}</ref> End hosts are humans and great apes, infected through bat contact or through other end hosts. Pigs in the Philippines have been reported to be infected with [[Reston virus]], so other interim or amplifying hosts may exist.<ref name=nejm52014/> Ebola virus outbreaks tend to occur when temperatures are lower and humidity is higher than usual for Africa.<ref>{{Cite journal |last1=Ng |first1=S. |last2=Cowling |first2=B. |year=2014 |title=Association between temperature, humidity and ebolavirus disease outbreaks in Africa, 1976 to 2014 |journal=Eurosurveillance |volume=19 |issue=35 |pages=20892 |doi=10.2807/1560-7917.ES2014.19.35.20892 |pmid=25210981 |doi-access=free}}</ref> Even after a person recovers from the acute phase of the disease, Ebola virus survives for months in certain organs such as the eyes and testes.<ref>{{Cite web |date=2016 |title=Clinical care for survivors of Ebola virus disease |url=http://apps.who.int/iris/bitstream/10665/204235/1/WHO_EVD_OHE_PED_16.1_eng.pdf?ua=1 |url-status=live |archive-url=https://web.archive.org/web/20160831010912/http://apps.who.int/iris/bitstream/10665/204235/1/WHO_EVD_OHE_PED_16.1_eng.pdf?ua=1 |archive-date=31 August 2016 |access-date=4 December 2016 |publisher=World Health Organization}}</ref>

==Ebola virus disease==
{{main|Ebola virus disease}}
Zaire ebolavirus is one of the four ebolaviruses known to cause disease in humans. It has the highest [[case-fatality rate]] of these ebolaviruses, averaging 83 percent since the first outbreaks in 1976, although a fatality rate of up to 90 percent was recorded in one outbreak in the Republic of the Congo between December 2002 and April 2003. There have also been more outbreaks of Zaire ebolavirus than of any other ebolavirus. The first outbreak occurred on 26 August 1976 in [[Yambuku]].<ref>{{Cite web |title=Clinical Aspects of Ebola Virus Disease at the Ngaliema Hospital, Kinshasa, Zaire, 1976 |url=http://www.itg.be/internet/ebola/ebola-12.htm |url-status=dead |archive-url=https://web.archive.org/web/20140804113441/http://www.itg.be/internet/ebola/ebola-12.htm |archive-date=2014-08-04 |access-date=2014-06-24 |publisher=European Network for Diagnostics of "Imported" Viral Diseases (ENIVD) |vauthors=Isaacson M, Sureau P, Courteille G, Pattyn, SR}}</ref> The first recorded case was Mabalo Lokela, a 44‑year-old schoolteacher. The symptoms resembled [[malaria]], and subsequent patients received [[quinine]]. Transmission has been attributed to reuse of unsterilized needles and close personal contact, body fluids and places where the person has touched. During the 1976 Ebola outbreak in [[Zaire]], [[Ngoy Mushola]] travelled from [[Bumba, Democratic Republic of the Congo|Bumba]] to [[Yambuku]], where he recorded the first clinical description of the disease in his daily log:

''<blockquote>The illness is characterized with a high temperature of about 39°C, [[hematemesis]], diarrhea with blood, retrosternal abdominal pain, prostration with "heavy" articulations, and rapid evolution death after a mean of three days.<ref>{{Cite web |last=Bardi |first=Jason Socrates |title=Death Called a River |url=http://www.scripps.edu/newsandviews/e_20020114/ebola1.html |url-status=live |archive-url=https://web.archive.org/web/20140802170848/http://www.scripps.edu/newsandviews/e_20020114/ebola1.html |archive-date=2 August 2014 |access-date=9 October 2014 |website=The Scripps Research Institute}}</ref></blockquote>''

Since the first recorded clinical description of the disease during 1976 in Zaire, the recent Ebola outbreak that started in March 2014, in addition, reached epidemic proportions and has killed more than 8000 people as of January 2015. This outbreak was centered in West Africa, an area that had not previously been affected by the disease. The toll was particularly grave in three countries: Guinea, Liberia, and Sierra Leone. A few cases were also reported in countries outside of West Africa, all related to international travelers who were exposed in the most affected regions and later showed symptoms of Ebola fever after reaching their destinations.<ref>name: S. Reardan.; N Engl. J Med. (2014) " The first nine months of the epidemic and projection, Ebola virus disease in west Africa". archive of Ebola Response Team. 511(75.11):520</ref>

The severity of the disease in humans varies widely, from rapid fatality to mild illness or even asymptomatic response.<ref name="NYT Katze">{{Cite news |last=Gina Kolata |date=Oct 30, 2014 |title=Genes Influence How Mice React to Ebola, Study Says in 'Significant Advance' |url=https://www.nytimes.com/2014/10/31/health/genes-influence-ebola-infections-in-mice-study-suggests.html |url-status=live |archive-url=https://web.archive.org/web/20210309060020/https://www.nytimes.com/2014/10/31/health/genes-influence-ebola-infections-in-mice-study-suggests.html |archive-date=9 March 2021 |access-date=Oct 30, 2014 |work=[[New York Times]]}}</ref> Studies of outbreaks in the late twentieth century failed to find a correlation between the disease severity and the genetic nature of the virus. Hence the variability in the severity of illness was suspected to correlate with genetic differences in the victims. This has been difficult to study in animal models that respond to the virus with hemorrhagic fever in a similar manner as humans, because typical mouse models do not so respond, and the required large numbers of appropriate test subjects are not easily available. In late October 2014, a publication reported a study of the response to a mouse-adapted strain of Zaire ebolavirus presented by a genetically diverse population of mice that was bred to have a range of responses to the virus that includes fatality from hemorrhagic fever.<ref name="Rasmussen2014">{{Cite journal |last=Rasmussen |first=Angela L. |display-authors=etal |date=Oct 30, 2014 |title=Host genetic diversity enables Ebola hemorrhagic fever pathogenesis and resistance |journal=[[Science (journal)|Science]] |volume=346 |issue=6212 |pages=987–991 |bibcode=2014Sci...346..987R |doi=10.1126/science.1259595 |pmc=4241145 |pmid=25359852}}</ref>

=== Vaccine ===
{{Main|Ebola vaccine}}
In December 2016, a study found the [[VSV-EBOV]] vaccine to be 70–100% effective against the Zaire ebola virus (not the [[Sudan ebolavirus]]), making it the first vaccine against the disease.<ref name="TL-20161222">{{Cite journal |last=Henao-Restrepo |first=Ana Maria |display-authors=etal |date=22 December 2016 |title=Efficacy and effectiveness of an rVSV-vectored vaccine in preventing Ebola virus disease: final results from the Guinea ring vaccination, open-label, cluster-randomised trial (Ebola Ça Suffit!) |journal=[[The Lancet]] |volume=389 |issue=10068 |pages=505–518 |doi=10.1016/S0140-6736(16)32621-6 |pmc=5364328 |pmid=28017403}}</ref><ref name="CNN-2016122">{{Cite news |last=Berlinger |first=Joshua |date=22 December 2016 |title=Ebola vaccine gives 100% protection, study finds |url=http://www.cnn.com/2016/12/22/health/ebola-vaccine-study/index.html |url-status=live |archive-url=https://web.archive.org/web/20161227084931/http://www.cnn.com/2016/12/22/health/ebola-vaccine-study/index.html |archive-date=27 December 2016 |access-date=27 December 2016 |work=[[CNN]]}}</ref>  VSV-EBOV was approved by the U.S. [[Food and Drug Administration]] in December 2019.<ref name="FDA PR">{{Cite web |date=19 December 2019 |title=First FDA-approved vaccine for the prevention of Ebola virus disease, marking a critical milestone in public health preparedness and response |url=https://www.fda.gov/news-events/press-announcements/first-fda-approved-vaccine-prevention-ebola-virus-disease-marking-critical-milestone-public-health |url-status=live |archive-url=https://web.archive.org/web/20191220052152/https://www.fda.gov/news-events/press-announcements/first-fda-approved-vaccine-prevention-ebola-virus-disease-marking-critical-milestone-public-health |archive-date=20 December 2019 |access-date=19 December 2019 |website=U.S. [[Food and Drug Administration]] (FDA)}}</ref>

== History and nomenclature ==
[[File:Marburg virus.jpg|thumb|440 px|Marburg virus]]
Ebola virus was first identified as a possible new "strain" of [[Marburg virus]] in 1976.<ref name=Pattyn1977/><ref name=Bowen1977/><ref>{{Cite news |last=Brown |first=Rob |date=2014-07-18 |title=The virus detective who discovered Ebola |url=https://www.bbc.com/news/magazine-28262541 |url-status=live |archive-url=https://web.archive.org/web/20211026040315/https://www.bbc.com/news/magazine-28262541 |archive-date=26 October 2021 |access-date=21 June 2018 |work=BBC News}}</ref> The [[International Committee on Taxonomy of Viruses]] (ICTV) identifies Ebola virus as [[species]] ''Zaire ebolavirus'', which is part of the [[genus]] ''[[Ebolavirus]]'', [[Family (biology)|family]] ''[[Filoviridae]]'', [[Order (biology)|order]] ''[[Mononegavirales]]''. The name "Ebola virus" is derived from the [[Ebola River]]—a river that was at first thought to be in close proximity to the area in [[Democratic Republic of Congo]], previously called [[Zaire]], where the [[1976 Zaire Ebola virus outbreak]] occurred—and the [[Taxonomy (biology)|taxonomic]] [[suffix]] ''virus''.<ref name="KuhnArch" /><ref name="Pattyn1977">{{Cite journal |vauthors=Pattyn S, Jacob W, van der Groen G, Piot P, Courteille G |year=1977 |title=Isolation of Marburg-like virus from a case of haemorrhagic fever in Zaire |journal=Lancet |volume=309 |issue=8011 |pages=573–574 |doi=10.1016/s0140-6736(77)92002-5 |pmid=65663 |s2cid=33060636}}</ref><ref name="Bowen1977">{{Cite journal |vauthors=((Bowen ETW)), Lloyd G, Harris WJ, Platt GS, Baskerville A, Vella EE |year=1977 |title=Viral haemorrhagic fever in southern Sudan and northern Zaire. Preliminary studies on the aetiological agent |journal=Lancet |volume=309 |issue=8011 |pages=571–573 |doi=10.1016/s0140-6736(77)92001-3 |pmid=65662 |s2cid=3092094}}</ref><ref name="Johnson1977">{{Cite journal |vauthors=Johnson KM, Webb PA, Lange JV, Murphy FA |year=1977 |title=Isolation and partial characterisation of a new virus causing haemorrhagic fever in Zambia |journal=Lancet |volume=309 |issue=8011 |pages=569–571 |doi=10.1016/s0140-6736(77)92000-1 |pmid=65661 |s2cid=19368457}}</ref>

In 1998, the virus name was changed to "Zaire Ebola virus"<ref>{{Cite book |title=Virus Taxonomy{{snd}}Seventh Report of the International Committee on Taxonomy of Viruses |vauthors=Netesov SV, Feldmann H, Jahrling PB, Klenk HD, Sanchez A |publisher=Academic Press |year=2000 |isbn=978-0123702005 |veditors=((van Regenmortel MHV)), Fauquet CM, ((Bishop DHL)), Carstens EB, Estes MK, Lemon SM, Maniloff J, Mayo MA, McGeoch DJ, Pringle CR, Wickner RB |location=San Diego |pages=539–548 |chapter=Family Filoviridae}}</ref><ref>{{Cite journal |last=Pringle |first=C. R. |year=1998 |title=Virus taxonomy – San Diego 1998 |journal=Archives of Virology |volume=143 |issue=7 |pages=1449–1459 |doi=10.1007/s007050050389 |pmid=9742051 |s2cid=13229117}}</ref> and in 2002 to species ''Zaire ebolavirus''.<ref name="Feldmann2005">{{Cite book |title=Virus Taxonomy{{snd}}Eighth Report of the International Committee on Taxonomy of Viruses |vauthors=Feldmann H, Geisbert TW, Jahrling PB, Klenk HD, Netesov SV, Peters CJ, Sanchez A, Swanepoel R, Volchkov VE |publisher=Elsevier/Academic Press |year=2005 |isbn=978-0123702005 |veditors=Fauquet CM, Mayo MA, Maniloff J, Desselberger U, Ball LA |location=San Diego |pages=645–653 |chapter=Family Filoviridae}}</ref><ref name="Mayo02">{{Cite journal |last=Mayo |first=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=2254–2260 |doi=10.1007/s007050200052 |s2cid=43887711 |doi-access=free}}</ref> However, most scientific articles continued to refer to "Ebola virus" or used the terms "Ebola virus" and "''Zaire ebolavirus''" in parallel. Consequently, in 2010, a group of researchers recommended that the name "Ebola virus" be adopted for a subclassification within the species ''Zaire ebolavirus'', with the corresponding abbreviation EBOV.<ref name=KuhnArch/> Previous abbreviations for the virus were EBOV-Z (for "Ebola virus Zaire") and ZEBOV (for "Zaire Ebola virus" or "''Zaire ebolavirus''"). In 2011, the ICTV explicitly rejected a proposal (2010.010bV) to recognize this name, as ICTV does not designate names for subtypes, variants, strains, or other subspecies level groupings.<ref>{{Cite web |title=Replace the species name Lake Victoria marburgvirus with Marburg marburgvirus in the genus Marburgvirus |url=http://talk.ictvonline.org/files/ictv_official_taxonomy_updates_since_the_8th_report/m/vertebrate-official/4171.aspx |url-status=dead |archive-url=https://web.archive.org/web/20160305175803/http://talk.ictvonline.org/files/ictv_official_taxonomy_updates_since_the_8th_report/m/vertebrate-official/4171.aspx |archive-date=5 March 2016 |access-date=31 October 2014}}</ref> At present, ICTV does not officially recognize "Ebola virus" as a taxonomic rank, but rather continues to use and recommend only the species designation ''Zaire ebolavirus''.<ref>{{Cite web |last=International Committee on Taxonomy of Viruses |title=Virus Taxonomy: 2013 Release |url=http://ictvonline.org/virusTaxonomy.asp |url-status=live |archive-url=https://web.archive.org/web/20150710030353/http://ictvonline.org/virusTaxonomy.asp |archive-date=10 July 2015 |access-date=31 October 2014}}</ref> The [[Type (biology)|prototype]] Ebola virus, variant Mayinga (EBOV/May), was named for Mayinga N'Seka, a nurse who died during the 1976 Zaire outbreak.<ref name="KuhnArch" /><ref>{{Cite journal |vauthors=Wahl-Jensen V, Kurz SK, Hazelton PR, Schnittler HJ, Stroher U, Burton DR, Feldmann H |year=2005 |title=Role of Ebola Virus Secreted Glycoproteins and Virus-Like Particles in Activation of Human Macrophages |journal=Journal of Virology |volume=79 |issue=4 |pages=2413–2419 |doi=10.1128/JVI.79.4.2413-2419.2005 |pmc=546544 |pmid=15681442}}</ref><ref>{{Cite journal |vauthors=Kesel AJ, Huang Z, Murray MG, Prichard MN, Caboni L, Nevin DK, Fayne D, Lloyd DG, Detorio MA, Schinazi RF |year=2014 |title=Retinazone inhibits certain blood-borne human viruses including Ebola virus Zaire |journal=Antiviral Chemistry & Chemotherapy |volume=23 |issue=5 |pages=197–215 |doi=10.3851/IMP2568 |pmc=7714485 |pmid=23636868 |s2cid=34249020}}</ref>

The name ''Zaire ebolavirus'' is derived from ''[[Zaire]]'' and the [[Taxonomy (biology)|taxonomic]] [[suffix]] ''ebolavirus'' (which denotes an ebolavirus species and refers to the [[Ebola River]]).<ref name=KuhnArch/> According to the rules for taxon naming established by the [[International Committee on Taxonomy of Viruses]] (ICTV), the name ''Zaire ebolavirus'' is always to be [[Capitalization|capitalized]], [[Italic type|italicized]], and to be preceded by the word "species". The names of its members (Zaire ebolaviruses) are to be capitalized, are not italicized, and used without [[Article (grammar)|articles]].<ref name=KuhnArch/>

== Virus inclusion criteria ==
A virus of the genus ''[[Ebolavirus]]'' is a member of the species ''Zaire ebolavirus'' if:<ref name=KuhnArch/>
* it is endemic in the [[Democratic Republic of the Congo]], [[Gabon]], or the [[Republic of the Congo]]
* it has a genome with two or three [[gene overlap]]s (''VP35''/''VP40'', ''GP''/''VP30'', ''VP24''/''L'')
* it has a [[genomic sequence]] that differs from the [[type (biology)|type virus]] EBOV/May by less than 30%

== Evolution ==
''Zaire ebolavirus'' diverged from its ancestors between 1960 and 1976.<ref name=":0">{{Cite journal |last=Carroll |first=S.A. |date=2012 |title=Molecular Evolution of Viruses of the Family Filoviridae Based on 97 Whole-Genome Sequences |journal=Journal of Virology |volume=87 |issue=5 |pages=2608–2616 |doi=10.1128/JVI.03118-12 |pmc=3571414 |pmid=23255795}}</ref> The genetic diversity of ''Ebolavirus'' remained constant before 1900.<ref name=":0" /><ref name=":1">{{Cite journal |last=Li |first=Y.H. |date=2013 |title=Evolutionary history of Ebola virus |journal=Epidemiology and Infection |volume=142 |issue=6 |pages=1138–1145 |doi=10.1017/S0950268813002215 |pmc=9151191 |pmid=24040779 |s2cid=9873900}}</ref> Then, around the 1960s, most likely due to climate change or human activities, the genetic diversity of the virus dropped rapidly and most lineages became extinct.<ref name=":1" /> As the number of susceptible hosts declines, so does the effective population size and its genetic diversity. This genetic bottleneck effect has implications for the species' ability to cause [[Ebola virus disease]] in human hosts.{{citation needed|date=April 2019}}

A [[genetic recombination|recombination]] event between ''Zaire ebolavirus'' lineages likely took place between 1996 and 2001 in wild apes giving rise to recombinant progeny viruses.<ref name="Wittmann2007">Wittmann TJ, Biek R, Hassanin A, Rouquet P, Reed P, Yaba P, Pourrut X, Real LA, Gonzalez JP, Leroy EM.  "Isolates of Zaire ebolavirus from wild apes reveal genetic lineage and recombinants".  ''Proc Natl Acad Sci U S A''. 2007 Oct 23;104(43):17123–17127. Epub 2007 Oct 17. "Erratum" in: ''Proc Natl Acad Sci U S A''. 2007 Dec 4;104(49):19656.  {{PMID|17942693}}</ref>  These recombinant viruses appear to have been responsible for a series of outbreaks among humans in Central Africa in 2001–2003.<ref name = Wittmann2007/>

''Zaire ebolavirus'' – Makona variant caused the 2014 West Africa outbreak.<ref name=":2">{{Cite web |date=2017-08-02 |title=Outbreaks Chronology: Ebola Virus Disease |url=https://www.cdc.gov/vhf/ebola/outbreaks/history/chronology.html |url-status=live |archive-url=https://web.archive.org/web/20190517142302/https://www.cdc.gov/vhf/ebola/outbreaks/history/chronology.html |archive-date=17 May 2019 |access-date=2017-11-11 |website=Ebola Hemorrhagic Feve |publisher=CDC |language=en-us}}</ref> The outbreak was characterized by the longest instance of human-to-human transmission of the viral species.<ref name=":2" /> Pressures to adapt to the human host were seen at this time, however, no phenotypic changes in the virus (such as increased transmission, increased immune evasion by the virus) were seen.{{citation needed|date=April 2020}}

==In literature==
* [[Alex Kava]]'s 2008 crime novel, ''Exposed'', focuses on the virus as a serial killer's weapon of choice.<ref>{{Cite book |last=Kava |first=Alex |url=https://www.publishersweekly.com/978-0-7783-2557-4 |title=Fiction Book Review: 'Exposed' by Alex Kava, Author . Mira (332p) |date=October 2008 |publisher=PublishersWeekly.com |isbn=978-0778325574 |language=en |access-date=2021-11-07 |archive-url=https://web.archive.org/web/20211107222602/https://www.publishersweekly.com/978-0-7783-2557-4 |archive-date=7 November 2021 |url-status=live}}</ref>
* [[William Close]]'s 1995 ''Ebola: A Documentary Novel of Its First Explosion'' and 2002 ''Ebola: Through the Eyes of the People'' focused on individuals' reactions to the 1976 Ebola outbreak in Zaire.<ref>{{Cite book |last=Close |first=William T. |author-link=William Close |url=https://archive.org/details/eboladocumentary00clos |title=Ebola: A Documentary Novel of Its First Explosion |publisher=[[Ivy Books]] |year=1995 |isbn=978-0804114325 |location=New York |oclc=32753758 |url-access=registration}} At [[Google Books]].</ref><ref>{{Cite book |last=Grove |first=Ryan |url=https://www.amazon.com/Ebola-documentary-novel-first-explosion/dp/B007HEJSE6 |title=More about the people than the virus |date=2006-06-02 |work=Review of Close, William T., Ebola: A Documentary Novel of Its First Explosion |access-date=2014-09-17 |archive-url=https://web.archive.org/web/20141021075254/http://www.amazon.com/Ebola-documentary-novel-first-explosion/dp/B007HEJSE6 |archive-date=21 October 2014 |url-status=live}}</ref><ref>{{Cite book |last=Close |first=William T. |author-link=William Close |url=https://books.google.com/books?id=BX0gAQAAIAAJ |title=Ebola: Through the Eyes of the People |publisher=Meadowlark Springs Productions |year=2002 |isbn=978-0970337115 |location=Marbleton, Wyoming |oclc=49193962 |access-date=7 February 2016 |archive-url=https://web.archive.org/web/20230114100059/https://books.google.com/books?id=BX0gAQAAIAAJ |archive-date=14 January 2023 |url-status=live}} ''At'' [[Google Books]].</ref><ref>{{Cite web |last=Pink |first=Brenda |date=2008-06-24 |title=A fascinating perspective |url=https://www.amazon.com/Ebola-Through-William-T-Close/product-reviews/0970337116 |url-status=live |archive-url=https://web.archive.org/web/20141021075312/http://www.amazon.com/Ebola-Through-William-T-Close/product-reviews/0970337116 |archive-date=21 October 2014 |access-date=2014-09-17 |website=Review of Close, William T., Ebola: Through the Eyes of the People}}</ref>
* ''[[The Hot Zone|The Hot Zone: A Terrifying True Story]]'': A 1994 best-selling book by Richard Preston about Ebola virus and related viruses, including an account of the outbreak of an Ebolavirus in primates housed in a quarantine facility in Reston, Virginia, USA<ref>{{Cite book |last=Preston |first=Richard |title=The Hot Zone |date=1995 |publisher=Anchor |isbn=0385479565 |location=New York |oclc=32052009}}</ref>
* [[Tom Clancy]]'s 1996 novel, ''[[Executive Orders]]'', involves a [[Middle East]]ern terrorist attack on the United States using an airborne form of a deadly Ebola virus named "Ebola Mayinga".<ref>{{Cite book |last=Clancy |first=Tom |author-link=Tom Clancy |url=https://archive.org/details/executiveorders00clan |title=Executive Orders |publisher=Putnam |year=1996 |isbn=978-0399142185 |location=New York |oclc=34878804 |url-access=registration}}</ref><ref>{{Cite web |last=Stone |first=Oliver |author-link=Oliver Stone |date=1996-09-02 |title=Who's That in the Oval Office? |url=https://www.nytimes.com/books/97/08/17/bsp/20141.html |archive-url=https://web.archive.org/web/20090410091107/http://www.nytimes.com/books/97/08/17/bsp/20141.html |archive-date=2009-04-10 |access-date=2014-09-10 |website=The New York Times Company |department=Books News & Reviews}}</ref>

==References==
{{Reflist}}


==Further reading==
* {{Cite journal |last1=Pacheco |first1=Daniela Alexandra de Meneses Rocha |last2=Rodrigues |first2=Acácio Agostinho Gonçalves |last3=Silva |first3=Carmen Maria Lisboa da |date=October 2016 |title=Ebola virus – from neglected threat to global emergency state |journal=Revista da Associação Médica Brasileira |volume=62 |issue=5 |pages=458–467 |doi=10.1590/1806-9282.62.05.458 |pmid=27656857 |doi-access=free}}

==External links==
{{Wikispecies}}
{{Scholia}}
{{commons|Ebolavirus}}

* [http://viralzone.expasy.org/all_by_protein/207.html Ebolavirus molecular biology]
* [http://pdb101.rcsb.org/motm/178 Ebolavirus proteins (PDB-101)]
* [http://talk.ictvonline.org/default.aspx ICTV Files and Discussions{{snd}}Discussion forum and file distribution for the International Committee on Taxonomy of Viruses] {{Webarchive|url=https://web.archive.org/web/20111007192538/http://talk.ictvonline.org/default.aspx |date=7 October 2011 }}
* [https://web.archive.org/web/20190912070927/https://www.viprbrc.org/brc/home.spg?decorator=filo Genomic data on ''Ebola'' virus isolates and other members of the family ''Filoviridae'']
* [http://viralzone.expasy.org/all_by_species/207.html ViralZone: Ebola-like viruses] – Virological repository from the [[Swiss Institute of Bioinformatics]]
* [https://web.archive.org/web/20160119052327/http://www.viprbrc.org/brc/home.spg?decorator=filo_ebola Virus Pathogen Resource: Ebola Portal] – Genomic and other research data about Ebola and other human pathogenic viruses
* [https://web.archive.org/web/20141101141011/http://visualscience.ru/en/projects/ebola/poster The Ebola Virus] 3D model of the Ebola virus, prepared by Visual Science, Moscow.
* [http://www.filovir.com/ FILOVIR{{snd}}scientific resources for research on filoviruses] {{Webarchive|url=https://web.archive.org/web/20200730224033/http://www.filovir.com/ |date=30 July 2020 }}
* {{Cite web |title='Zaire ebolavirus' |url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&id=186538 |website=NCBI Taxonomy Browser |id=186538}}
* {{Cite web |title='Ebola virus sp.' |url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&id=205488 |website=NCBI Taxonomy Browser |id=205488}}

{{Medical resources 
| ICD11          = 
 | ICD10          =
 | ICD9           = 
 | DiseasesDB     = 
 | MedlinePlus    =
 | OMIM           =
 | eMedicineSubj  = 
 | eMedicineTopic = 
 | eMedicine_mult =
 | MeshName       = 
 | MeshNumber     = 
 | Orphanet       = 
 | Scholia        = Q8064876
 }}
{{Ebola|state=collapsed}}
{{Medicine}}
{{Taxonbar|from=Q8064876|from2=Q10538943}}
{{Authority control}}

[[Category:Ebolaviruses]]
[[Category:Ebola]]
[[Category:Primate diseases]]
[[Category:Biological weapons]]
[[Category:Biosafety level 4 pathogens]]
[[Category:Tropical diseases]]
[[Category:Zoonoses]]
[[Category:1976 in biology]]
[[Category:Animal viral diseases]]
[[Category:Arthropod-borne viral fevers and viral haemorrhagic fevers]]
[[Category:Hemorrhagic fevers]]
[[Category:Virus-related cutaneous conditions]]