Rhabdoviruses Daisy L Jennings, Wildlife Zoonoses and Vector Borne Diseases Research Group, Animal and Plant Health Agency, Surrey, UK Denise A Marston, Wildlife Zoonoses and Vector Borne Diseases Research Group, Animal and Plant Health Agency, Surrey, UK Ashley C Banyard, Wildlife Zoonoses and Vector Borne Diseases Research Group, Animal and Plant Health Agency, Surrey, UK Anthony R Fooks, Wildlife Zoonoses and Vector Borne Diseases Research Group, Animal and Plant Health Agency, Surrey, UK Based in part on the previous versions of this eLS article ‘Rhab- doviruses’ (2003, 2006, 2013). Advanced article Article Contents • Classification • Plant-adapted Rhabdoviruses • Assembly, Envelopment and Budding • Pathobiology and Epidemiology of Some Clinically Important Rhabdoviruses • Pathobiology and Epidemiology of Lyssaviruses Online posting date: 15 th December 2017 The family Rhabdoviridae is classified within the order Mononegavirales and is made up of 18 different genera and one other virus awaiting clas- sification into this family. Generally, Rhabdovirus virions contain a nonsegmented negative-sense single-stranded ribonucleic acid (RNA) genome that, in its most basic form, encodes five proteins: the nucleocapsid protein (N), the phosphopro- tein (P), the matrix protein (M), the glycoprotein (G) and the RNA-dependent RNA polymerase protein (L). All Rhabdoviruses have a distinctive bullet-shaped morphology; however, two recently recognised genera have been shown to contain viruses with bipartite genomes. This diverse group of over 165 viruses can infect a range of species including mammals, reptiles, birds, fish, insects and plants. Several of these viruses cause notable economic losses to both the agriculture and aqua- culture sector. However, only the lyssaviruses and the vesiculoviruses are able to infect both animals and humans to cause clinical disease, with rabies virus being the most important human pathogen. Classification The order Mononegavirales The International Committee on Taxonomy of Viruses (ICTV) 2016 Release (http://www.ictvonline.org/virusTaxonomy.asp) eLS subject area: Virology How to cite: Jennings, Daisy L; Marston, Denise A; Banyard, Ashley C; and Fooks, Anthony R (December 2017) Rhabdoviruses. In: eLS. John Wiley & Sons, Ltd: Chichester. DOI: 10.1002/9780470015902.a0001085.pub4 included the order Mononegavirales expanding to encompass 69 novel species and application of non-Latinised binomial species names throughout the entire order (Amarasinghe et al., 2017). The order Mononegavirales contains eight virus families, including the Bornaviridae, the Filoviridae, the Paramyxoviridae and the Rhabdoviridae families. Generally, all viruses within these families contain linear, nonsegmented and negative-sense single-stranded ribonucleic acid (RNA) genomes. These genomes act as templates for two distinct processes: (1) transcription of viral messenger ribonucleic acids (mRNAs) and (2) transcrip- tion of a full-length positive-sense copy of the virus genome, termed the antigenome, which acts as the template for replication to produce negative-sense progeny genomes (Whelan et al., 2004). Neither the negative-sense genome nor the positive-sense antigenome is considered infectious as they require viral proteins to drive transcription of viral messenger RNA and initiate the replicative cycle. The minimal replicative unit for all rhab- doviruses appears to be the genomic ribonucleic acid (gRNA) tightly wound as a helical structure of 30–70 nm, encapsidated with N protein and in contact with P and L. This complex of proteins with nucleic acid is termed the ribonucleoprotein (RNP) complex. The generation of this complex in vitro is the basis for reverse genetics techniques whereby the virus is recovered following transfection of the deoxyribonucleic acid (DNA). Virus release occurs through budding from the plasma membrane, whereas virus replication and transcription are gen- erally cytoplasmic for members of the order Mononegavirales. Exceptions to this are the bornaviruses, the nucleorhabdoviruses and the dichorhaviruses, which perform both replication and transcription of their genetic material in the nucleus. Other features shared by the members of the order Mononegavirales include a helical nucleocapsid, the apparent inability to undergo genetic reassortment and a highly conserved gene order – with the N, P and M genes, always at or near the 3 ′ terminus, and the L gene always the furthest from the genome promoter. This is most likely an evolutionary adaptation that controls protein expression levels. See also: Bornaviruses; Filoviruses; RNA Virus Genomes Generally within this family of viruses, genomes encode 5–10 genes, the position of which is conserved to control gene This article is © 2017 Crown Copyright in the UK and © 2017 John Wiley & Sons, Ltd in the rest of the world. eLS © 2017, John Wiley & Sons, Ltd. www.els.net 1