# ICTV Virus Taxonomy Profile: Filoviridae

Jens H Kuhn, Gaya Amarasinghe, Christopher Basler, Sina Bavari, Alexander Bukreyev, Kartik Chandran, Ian Crozier, Olga Dolnik, John Dyer, Pierre B H Formenty, et al.

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Jens H Kuhn, Gaya Amarasinghe, Christopher Basler, Sina Bavari, Alexander Bukreyev, et al. ICTV Virus Taxonomy Profile: Filoviridae. Journal of General Virology, 2019, 100 (6), pp.911-912. 10.1099/jgv.0.001252. hal-04991614

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# ICTV Virus Taxonomy Profile: *Filoviridae*

Jens H. Kuhn,¹,* Gaya K. Amarasinghe,² Christopher F. Basler,³ Sina Bavari,⁴ Alexander Bukreyev,⁵ Kartik Chandran,⁶

Ian Crozier,⁷ Olga Dolnik,⁸ John M. Dye,⁴ Pierre B. H. Formenty,⁹ Anthony Griffiths,¹⁰ Roger Hewson,¹¹

Gary P. Kobinger,¹² Eric M. Leroy,¹³ Elke Mühlberger,¹⁰ Sergey V. Netesov (Нетёсов Сергей Викторович),¹⁴

Gustavo Palacios,⁴ Bernadett Pályi,¹⁵ Janusz T. Pawęska,¹⁶ Sophie J. Smither,¹⁷ Ayato Takada (高田礼人),¹⁸

Jonathan S. Towner,¹⁹ Victoria Wahl²⁰ and ICTV Report Consortium

## Abstract

Members of the family *Filoviridae* produce variously shaped, often filamentous, enveloped virions containing linear non-segmented, negative-sense RNA genomes of 15–19 kb. Several filoviruses (e.g., Ebola virus) are pathogenic for humans and are highly virulent. Several filoviruses infect bats (e.g., Marburg virus), whereas the hosts of most other filoviruses are unknown. This is a summary of the International Committee on Taxonomy of Viruses (ICTV) Report on *Filoviridae*, which is available at www.ictv.global/report/filoviridae.

<table><caption>**Table 1.** Characteristics of members of the family *Filoviridae*</caption><thead><tr><th>Typical member:</th><th>Marburg virus (DQ217792), species Marburg marburgvirus, genus Marburgvirus</th></tr></thead><tbody><tr><td>Virion</td><td>Enveloped, variously shaped but predominantly filamentous, typically with a single nucleocapsid</td></tr><tr><td>Genome</td><td>Approximately 15–19 kb of linear, negative-sense, non-segmented RNA</td></tr><tr><td>Replication</td><td>Antigenomic RNA is a replication intermediate. The genome and antigenome form ribonucleoprotein complexes, which serve as templates</td></tr><tr><td>Translation</td><td>From multiple 5'-capped and 3'-polyadenylated mRNAs</td></tr><tr><td>Host range</td><td>Primates (e.g., ebolaviruses, marburgviruses), bats (e.g., marburgviruses), domestic pigs (e.g., Reston virus) and probably fish (e.g., striaviruses, thamnoviruses) become naturally infected</td></tr><tr><td>Taxonomy</td><td>Realm Riboviria, phylum Negarnaviricota, subphylum Haploviricotina, class Mononegavirales; family includes multiple genera</td></tr></tbody></table>

## VIRION

Virions are enveloped and diverse in shape and can appear as branched, toroid, U- or 6-shaped, and long filamentous forms (Table 1, Fig. 1). Virions contain ribonucleoprotein (RNP) complexes composed of genomic RNA and, typically,

the structural proteins nucleoprotein (NP), polymerase co-factor (VP35), transcriptional activator (VP30), RNP-associated protein (VP24) and RNA-dependent RNA polymerase (L). The matrix protein (VP40) forms a regular layer beneath the viral envelope. Surface spikes formed by glycoproteins (GP₁,₂) are approximately 7 nm in diameter and

Received 7 March 2019; Accepted 8 March 2019; Published 25 April 2019

**Author affiliations:** ¹NIH/NIAID Integrated Research Facility at Fort Detrick, (IRF-Frederick), Frederick, MD 21702, USA; ²Washington University School of Medicine, St. Louis, MO 63110, USA; ³Center for Microbial Pathogenesis, Georgia State University, Atlanta GA 30303, USA; ⁴United States Army Medical Research Institute of Infectious Diseases, Frederick, MD 21702, USA; ⁵The University of Texas Medical Branch, Galveston, TX 77555, USA; ⁶Albert Einstein College of Medicine, Bronx, NY 10461, USA; ⁷Integrated Research Facility at Fort Detrick, Clinical Monitoring Research Program Directorate, Frederick National Laboratory for Cancer Research supported by the National Cancer Institute, Frederick, MD 21702, USA; ⁸Institute of Virology, Philipps University Marburg, 35043 Marburg, Germany; ⁹World Health Organization, CH-1211 Geneva, Switzerland; ¹⁰Department of Microbiology and National Emerging Infectious Diseases Laboratories, Boston University School of Medicine, Boston, MA 02118, USA; ¹¹Public Health England, Porton Down, Wiltshire, SP4 0JG Salisbury, UK; ¹²Université Laval, Québec City, QC G1V 0A6, Canada; ¹³Infectious Diseases and Vectors Unit, Institut de Recherche pour le Développement, 911 Av Agropolis, 34394 Montpellier, France; ¹⁴Novosibirsk State University, Novosibirsk, Novosibirsk Oblast, 630090, Russia; ¹⁵National Biosafety Laboratory, National Public Health Center, Budapest, Hungary; ¹⁶National Institute for Communicable Diseases of the National Health Laboratory Service, 2131 Sandringham-Johannesburg, Gauteng, South Africa; ¹⁷Defence Science and Technology Laboratory, Porton Down, Salisbury, Wiltshire SP4 0JQ, UK; ¹⁸Division of Global Epidemiology, Hokkaido University Research Center for Zoonosis Control, 001-0020 Sapporo, Japan; ¹⁹Viral Special Pathogens Branch, Centers for Disease Control and Prevention, Atlanta, GA 30333, USA; ²⁰National Biodefense Analysis and Countermeasures Center, Frederick, MD 21702, USA.

*Correspondence: Jens H. Kuhn, kuhnjens@mail.nih.gov

Keywords: *Filoviridae*; filovirus; ICTV Report; ebolavirus; marburgvirus; taxonomy.

Abbreviations: L, RNA-dependent RNA polymerase; NP, nucleoprotein; RNP, ribonucleoprotein.
<figure><img src="image_4.png" /></figure>

<figure><img src="image_5.png" /><figcaption>**Fig. 1.** Electron microscopic images of Marburg virus particles budding from infected Vero E6 cells, (a) scanning EM, (b) transmission EM. Images are colourized for clarity. Courtesy of John G. Bernbaum and Jiro Wada, IRF-Frederick.</figcaption></figure>

<figure><img src="image_6.png" /><figcaption>**Fig. 2.** Schematic representation of the genome organization of Marburg virus. Courtesy of Jiro Wada, IRF-Frederick.</figcaption></figure>

cover the virion surface at approximately 10-nm intervals [1–3]. Some filoviruses do not have discernable glycoproteins and may have different RNP complexes [4].

## GENOME

Filovirus genomes are approximately 15–19 kb (Fig. 2) without a 5'-cap or 3'-poly(A). Terminal leader and trailer sequences contain the replication and transcription promoters. Marburg-virus genomes contain seven separate, continuous open reading frames (ORFs) flanked by 3'- and 5'-terminal non-coding regions that contain transcription initiation and termination sites. These ORFs encode the virion structural proteins. Cuevavirus and ebolavirus genomes encode homologues of the marburgvirus structural proteins. However, the marburgvirus *GP* gene encodes only $GP_{1,2}$, whereas the primary expression product of cuevavirus and ebolavirus *GP* gene transcription is a soluble glycoprotein. Co-transcriptional editing is used to express $GP_{1,2}$ and additional non-structural proteins. Striaviruses and thamnoviruses encode some, but not all, marburgvirus protein homologues and several proteins of unknown function [4, 5].

## REPLICATION

Virus proteins are translated from mRNAs that are synthesized by successive, polar transcription from RNP complexes containing genomic RNA. Replication occurs in the cytoplasm through the synthesis of RNP complexes containing antigDNAs that are templates for genomic RNA production. Replication and transcription enzymes include L and VP35. VP30 serves as a transcription enhancer for ebolaviruses and

probably cuevaviruses, but its function in marburgvirus infection is less defined. Virion assembly, including acquisition of the $GP_{1,2}$-containing lipid envelope, occurs by VP40-mediated budding at the plasma membrane [4, 5].

## TAXONOMY

Filoviruses form a family in the haploviricotine order *Mononegavirales*. Within this order, filoviruses are most closely related to members of the families *Paramyxoviridae*, *Pneumoviridae* and *Sunviridae*. The family *Filoviridae* includes multiple genera for viruses that differ in geographic and host range and genomic organization.

## RESOURCES

Full ICTV Report on the family *Filoviridae*: www.ictv.global/report/filoviridae.

### Funding information

Production of this summary, the online chapter and associated resources was funded by a grant from the Wellcome Trust (WT108418A1A). This work was supported in part through Battelle Memorial Institute's prime contract with the US National Institute of Allergy and Infectious Diseases (NIAID) under Contract No. HHSN272200700016I (J.H.K.). This project was funded in part with federal funds from the National Cancer Institute, National Institutes of Health under Contract No. HHSN261200800001E. This work was also funded in part under Contract No. HSHQDC-15-C-00064 awarded by Department of Homeland Security (DHS S&T) for the management and operation of the National Biodefense Analysis and Countermeasures Center (NBACC), a federally funded research and development center (V.W.). The views and conclusions contained in this document are those of the authors and should not be interpreted as necessarily representing the official policies, either expressed or implied, of the US Department of the Army, the US Department of Defense, the US Department of Health and Human Services, or of the institutions and companies affiliated with the authors. In no event shall any of these entities have any responsibility or liability for any use, misuse, inability to use or reliance upon the information contained herein. The US departments do not endorse any products or commercial services mentioned in this publication.

### Acknowledgements

Members of the ICTV (10th) Report Consortium are Elliot J. Lefkowitz, Andrew J. Davison, Stuart G. Siddell, Peter Simmonds, Sead Sabanadzovic, Donald B. Smith, Richard J. Orton and Jens H. Kuhn. We thank Laura Bollinger (IFR-Frederick) for critically editing the manuscript and the associated report chapter.

### Conflicts of interest

The authors declare that there are no conflicts of interest.

### References

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