Orbivirus

Orbivirus
Negatively stained bluetongue virus–like particle that caused a cytopathic effect in BHK-21 cells. Scale bar = 50 nm
Virus classification
Group: Group III (dsRNA)
Order: Unassigned
Family: Reoviridae
Subfamily: Sedoreovirinae
Genus: Orbivirus
Type Species

The genus Orbivirus is a member of the Reoviridae family, in the subfamily Sedoreovirinae. Unlike the other reoviruses, orbiviruses are arboviruses. This genus currently contains 22 species (including the type species Bluetongue virus) and at least 130 different serotypes.[1][2] Orbiviruses can infect and replicate within a wide range of arthropod and vertebrate hosts. Orbiviruses are named after their characteristic doughnut shaped capsomers (Orbi in Latin means ring).

Many orbiviruses are transmitted by ticks or haematophagus insect vectors (Culicoides, mosquitoes and sand flies) and have a wide host range that includes cattle, goats and sheep, wild ruminants, equids, camelids, marsupials, sloths, bats, birds, large canine and feline carnivores and humans.

The three economically most important orbiviruses are Bluetongue virus, African horse sickness virus and epizootic hemorrhagic disease virus all of which are transmitted by Culicoides species.

Virion

The virons are non-enveloped particles that are between 70-80 nm in diameter. The virus particles are spherical in appearance and have icosahedral symmetry.[3] An outer and an inner capsid layer surround the genome, and have T=13 and T=2 symmetry, respectively.[2] The viron is constructed of two concentric protein shells, the sub-core layer which contain 120 copies/particle of the VP3 and the core-surface layer composed of 780 copies/particle of the VP7. VP1, VP4 and VP6 are minor enzymatic proteins that are packaged along with the ten genome segments within the central space of the virus core. The orbivirus outer-capsid layer is composed of two additional structural proteins (VP2 and VP5) which mediate cell-attachment and penetration during initiation of infection. The outer-capsid proteins are more variable than the core proteins and most of the non-structural proteins and the specificity of their reactions with neutralising antibodies determines the virus serotype.

Genome

These viruses have double stranded RNA genomes and therefore are classified as Class III viruses. Their genome is linear and is segmented into 10 segments of various lengths. One copy of each gene segment is packaged per virion. In most cases each gene segment encodes a single open reading frame (ORF). The genome encodes 7 major structural proteins (VP1-VP7) and 3 major non structural proteins (NS1-NS3). Exceptions to the one gene one protein rule are segment 9 (Seg-9) and segment 10 (Seg-10) both of which encode two nearly identical proteins initiated from in-phase AUG codons close together near the upstream termini (VP6 and VP6a encoded by Seg-9: NS3 and NS3a encoded by Seg-10)

An open reading frame spans almost the entire length of genome segment-9 and encodes VP6 (the viral helicase). A second open reading frame (OrfX) is also present on this segment and encodes a fourth non structural protein (NS4), which was predicted from sequence analysis of various orbiviruses including Segment 9 of Great Island virus which contained a long NS4 ORF (approximately 21kDa). The existence of NS4 was experimentally confirmed in both insect-borne and tick-borne orbiviruses in 2011.[4]

NS1 is the most abundant protein in BTV infected cells. It forms tubules that may be involved in translocation of progeny virus particles to the cell membrane. NS2 is phosphorylated by cellular kinases and is an important matrix protein of the granular viral inclusion bodies that form within the cytoplasm of infected cells. These viral inclusion bodies act as the centres of viral replication. The membrane glycoproteins NS3 and NS3a are expressed in large amounts in insect cells but not in mammalian cells. They are involved in the release of progeny virus particles from infected cells and may be involved in determination of both vector competence and virulence.

Genus Structure Symmetry Capsid Genomic Arrangement Genomic Segmentation
OrbivirusIcosahedralT=13, T=2Non-EnvelopedLinearSegmented

Replication

Many orbiviruses preferentially infect vascular endothelial cells. Orbiviruses enter the host cell by endocytosis and the outer capsid is subsequently removed. The whole cycle of viral replication takes place within the cytoplasm of the host cell. Transcription of the viral genome into mRNA occurs within the core particle and mRNA is translated into proteins using the host cell ribosomes. Viral proteins are synthesized 2–14 days after initial infection. New virons self-assemble within the cytoplasm and are then released from the host cell by budding. During the budding process they transiently acquire a lipid envelope which can be detected for a short period of time following their release but this is subsequently lost.

Genus Host Details Tissue Tropism Entry Details Release Details Replication Site Assembly Site Transmission
OrbivirusVertebrates; mosquitoes; midges; gnats; sandflies; ticksvascular endothelial cellsCell receptor endocytosisCell deathCytoplasmCytoplasmArthropod bite

Pathogenesis

Orbiviruses primarily cause diseases in animals. The different Orbivirus species have different host specificities. Orbiviruses are vector-borne pathogens transmitted between vertebrate hosts by vectors such as mosquitoes, midges, gnats, sandflies and ticks. Bluetongue virus (BTV) is an Orbivirus that causes bluetongue disease in sheep, cattle, goats and wild ungulates. BTV has been in the forefront of molecular studies for last three decades and now represents one of the best understood viruses at the molecular and structural levels.[5][6] Other species of Orbiviruses are responsible for other diseases of animals such as African Horse sickness and Equine encephalosis virus.[7]

Classification

ICTV Taxonomy

Group: dsRNA

[1]

Serogroups

This genus has been divided into several (at least 14) serogroups.[8][9] The serogroups have been divided in some cases into subgroups. A number of these viruses have yet to be assigned to a serogroup. The serogroups are differentiated on the basis of a fourfold or greater difference in antibody based tests. These tests include ELISAs and complement fixation tests.

Species

There are currently 22 recognized virus species as well as 13 unassigned viruses in this genus

Species (with constituent viruses):

Vector groups

These viruses are known to be transmitted by midges (Culicoides), mosquitoes and ticks. Those viruses transmitted by a particular type of vector seem to be related both genetically and serologically. These groups are

Midge vector group:

Mosquito vector group:

Tick vector group:

The vector(s) of the St Croix river virus are not known and based on its genome sequence this virus does not appear to group with any other vector group.

Notes

The tick group may be ancestral to the other groups.[10]

History

In 1719, African horse sickness virus caused the first major recorded orbivirus epidemic, killing 1,500 animals. The most historically significant outbreak of orbivirus occurred in 1854-1855, when AHSV infected 70,000 horses. AHSV was discovered to be a virus in 1900 and bluetongue disease followed shortly thereafter in 1905. Outbreaks have occurred sporadically in the 20th and 21st centuries.[3]

See also

See also

References

  1. 1 2 ICTV. "Virus Taxonomy: 2014 Release". Retrieved 15 June 2015.
  2. 1 2 "Viral Zone". ExPASy. Retrieved 15 June 2015.
  3. 1 2 Knipe, David M. (2013-06-17). Fields Virology. Wolters Kluwer Health. ISBN 9781451105636.
  4. ↑ Belhouchet M, Mohd Jaafar F, Firth AE, Grimes JM, Mertens PP, Attoui H (2011) Detection of a fourth orbivirus non-structural protein. PLoS One 6(10):e25697.
  5. ↑ Roy P (2008). "Molecular Dissection of Bluetongue Virus". Animal Viruses: Molecular Biology. Caister Academic Press. ISBN 978-1-904455-22-6.
  6. ↑ Roy P (2008). "Structure and Function of Bluetongue Virus and its Proteins". Segmented Double-stranded RNA Viruses: Structure and Molecular Biology. Caister Academic Press. ISBN 978-1-904455-21-9.
  7. ↑ Viljoen and Huismans (1989) The Characterization of Equine Encephalosis Virus and the Development of Genomic Probes. J. gen. Virol. 70, 2007-2015.
  8. ↑ Dilcher M, Hasib L, Lechner M, Wieseke N, Middendorf M, Marz M, Koch A, Spiegel M, Dobler G, Hufert FT, Weidmann M (2000) Genetic characterization of Tribeč virus and Kemerovo virus, two tick-transmitted human-pathogenic Orbiviruses. Virology
  9. ↑ Warrell DA, Cox TM, Firth JD (2003) Oxford textbook of Medicine. Oxford
  10. ↑ Belaganahalli MN, Maan S, Maan NS, Tesh R, Attoui H, Mertens PP (2011) Umatilla virus genome sequencing and phylogenetic analysis: identification of stretch lagoon orbivirus as a new member of the Umatilla virus species. PLoS One 6(8):e23605.
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