Orbivirus | |
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Cryo-EM of the protein structure of a bluetongue virus capsid | |
Negatively stained Bluetongue virus –like particle that caused a cytopathic effect in BHK-21 cells. Scale bar = 50 nm | |
Virus classification | |
(unranked): | Virus |
Realm: | Riboviria |
Kingdom: | Orthornavirae |
Phylum: | Duplornaviricota |
Class: | Resentoviricetes |
Order: | Reovirales |
Family: | Reoviridae |
Subfamily: | Sedoreovirinae |
Genus: | Orbivirus |
Orbivirus is a genus of double-stranded RNA viruses in the family Reoviridae and subfamily Sedoreovirinae . Unlike other reoviruses, orbiviruses are arboviruses. They can infect and replicate within a wide range of arthropod and vertebrate hosts. Orbiviruses are named after their characteristic doughnut-shaped capsomers (orbis 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. The genus contains 22 species and at least 130 different serotypes. [1] [2]
In 1719, African horse sickness virus (AHSV) 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]
The virons are nonenveloped particles that are 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 subcore 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 10 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.
These viruses have double-stranded RNA genomes, so 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 seven major structural proteins (VP1–VP7) and three major nonstructural 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 ORF spans almost the entire length of genome segment 9 and encodes VP6 (the viral helicase). A second ORF (OrfX) is also present on this segment and encodes a fourth nonstructural protein (NS4), which was predicted from sequence analysis of various orbiviruses including segment 9 of Great Island virus which contained a long NS4 ORF (around 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 bluetongue virus 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 numbers 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.
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.
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]
The genus Orbivirus contains the following species: [1]
The genus is divided into several (at least 14) serogroups. [9] [10] The serogroups are divided in some cases into subgroups. A number of member 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.
Member viruses are transmitted by midges ( Culicoides ), mosquitoes, and ticks. The viruses transmitted by a particular type of vector are generally related both genetically and serologically.
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.
The tick group may be ancestral to the other groups. [12]
Bluetongue disease is a noncontagious, insect-borne, viral disease of ruminants, mainly sheep and less frequently cattle, yaks, goats, buffalo, deer, dromedaries, and antelope. It is caused by Bluetongue virus (BTV). The virus is transmitted by the midges Culicoides imicola, Culicoides variipennis, and other culicoids.
Flaviviridae is a family of enveloped positive-strand RNA viruses which mainly infect mammals and birds. They are primarily spread through arthropod vectors. The family gets its name from the yellow fever virus; flavus is Latin for "yellow", and yellow fever in turn was named because of its propensity to cause jaundice in victims. There are 89 species in the family divided among four genera. Diseases associated with the group include: hepatitis (hepaciviruses), hemorrhagic syndromes, fatal mucosal disease (pestiviruses), hemorrhagic fever, encephalitis, and the birth defect microcephaly (flaviviruses).
Alphavirus is a genus of RNA viruses, the sole genus in the Togaviridae family. Alphaviruses belong to group IV of the Baltimore classification of viruses, with a positive-sense, single-stranded RNA genome. There are 32 alphaviruses, which infect various vertebrates such as humans, rodents, fish, birds, and larger mammals such as horses, as well as invertebrates. Alphaviruses that could infect both vertebrates and arthropods are referred dual-host alphaviruses, while insect-specific alphaviruses such as Eilat virus and Yada yada virus are restricted to their competent arthropod vector. Transmission between species and individuals occurs mainly via mosquitoes, making the alphaviruses a member of the collection of arboviruses – or arthropod-borne viruses. Alphavirus particles are enveloped, have a 70 nm diameter, tend to be spherical, and have a 40 nm isometric nucleocapsid.
Aphthovirus is a viral genus of the family Picornaviridae. Aphthoviruses infect split-hooved animals, and include the causative agent of foot-and-mouth disease, Foot-and-mouth disease virus (FMDV). There are seven FMDV serotypes: A, O, C, SAT 1, SAT 2, SAT 3 and Asia 1, and four non-FMDV serotypes belonging to three additional species Bovine rhinitis A virus (BRAV), Bovine rhinitis B virus (BRBV) and Equine rhinitis A virus (ERAV).
African horse sickness (AHS) is a highly infectious and deadly disease caused by African horse sickness virus. It commonly affects horses, mules, and donkeys. It is caused by a virus of the genus Orbivirus belonging to the family Reoviridae. This disease can be caused by any of the nine serotypes of this virus. AHS is not directly contagious, but is known to be spread by insect vectors.
Erbovirus is a genus of viruses in the order Picornavirales, in the family Picornaviridae. Horses serve as natural hosts. There is only one species in this genus: Erbovirus A. Diseases associated with this genus include: upper respiratory tract disease with viremia and fecal shedding. Viruses belonging to the genus Erbovirus have been isolated in horses with acute upper febrile respiratory disease. The structure of the Erbovirus virion is icosahedral, having a diameter of 27–30 nm.
Coltivirus is a genus of viruses that infects vertebrates and invertebrates. It includes the causative agent of Colorado tick fever. Colorado tick fever virus can cause a fever, chills, headache, photophobia, myalgia, arthralgia, and lethargy. Children, in particular, may develop a hemorrhagic disease. Leukopenia with both lymphocytes and neutrophils is very common for Colorado tick fever virus. In either case, the infection can lead to encephalitis or meningitis.
Orthonairovirus is a genus of viruses in the family Nairoviridae of the order Bunyavirales that include viruses with circular, negative-sense single stranded RNA. It got its name from the Nairobi sheep disease that affects the gastrointestinal tracts of sheep and goats. The vast majority, and perhaps all viruses in this genus are tick-borne viruses that can have human or other vertebrate hosts.
Orthobunyavirus is a genus of the Peribunyaviridae family in the order Bunyavirales. There are currently ~170 viruses recognised in this genus. These have been assembled into 103 species and 20 serogroups.
Double-stranded RNA viruses are a polyphyletic group of viruses that have double-stranded genomes made of ribonucleic acid. The double-stranded genome is used to transcribe a positive-strand RNA by the viral RNA-dependent RNA polymerase (RdRp). The positive-strand RNA may be used as messenger RNA (mRNA) which can be translated into viral proteins by the host cell's ribosomes. The positive-strand RNA can also be replicated by the RdRp to create a new double-stranded viral genome.
Vesivirus is a genus of viruses, in the family Caliciviridae. Swine, sea mammals, and felines serve as natural hosts. There are two species in this genus. Diseases associated with this genus include: respiratory disease, Feline calicivirus (FCV); conjunctivitis, and respiratory disease.
Equine encephalosis virus (EEV) is a species of virus the Orbivirus genus, and a member of the Reoviridae family, related to African horse sickness virus (AHSV) and Bluetongue virus (BTV).
Sedoreovirinae is a subfamily of the Reoviridae family of viruses. Viruses in this subfamily are distinguished by the absence of a turreted protein on the inner capsid to produce a smooth surface.
Epizootic hemorrhagic disease virus, often abbreviated to EHDV, is a species of the genus Orbivirus, a member of the family Reoviridae. It is the causative agent of epizootic hemorrhagic disease, an acute, infectious, and often fatal disease of wild ruminants. In North America, the most severely affected ruminant is the white-tailed deer, although it may also infect mule deer, black-tailed deer, elk, bighorn sheep, and pronghorn antelope. It is often mistakenly referred to as “bluetongue virus” (BTV), another Orbivirus that like EHDV causes the host to develop a characteristic blue tongue due to systemic hemorrhaging and lack of oxygen in the blood. Despite showing clinical similarities, these two viruses are genetically distinct.
Avibirnavirus is a genus of viruses in family Birnaviridae. There is a single species in this genus: Infectious bursal disease virus, which infects chickens and other fowl. It causes severe inflammation of the bursa of Fabricius, and causes considerable morbidity and mortality.
Umatilla virus(UMAV) is a dsRNA virus in the family Reoviridae, subfamily Sedoreovirinae, and the genus Orbivirus. This arbovirus was first isolated in 1969 in Umatilla County, Oregon in a group of Culex pipiens mosquitoes. The viral host is the Passer domesticus bird with the vectors being Culex mosquitoes.
Mammalian orthoreovirus (MRV) is a double-stranded RNA virus. It is a part of the family Reoviridae, as well as the subfamily Spinareovirinae. As seen in the name, the Mammalian Ortheoreovirus infects numerous mammalian species and vertebrates which serve as natural hosts. Some diseases that occur as a result of this virus or are associated with this virus include mild upper respiratory illness, and gastrointestinal illness. Examples of these are: upper respiratory tract syndromes, gastroenteritis, biliary atresia, obstructive hydrocephalus, jaundice, alopecia, conjunctivitis, and ‘oily hair’ associated with steatorrhea.
Modoc virus (MODV) is a rodent-associated flavivirus. Small and enveloped, MODV contains positive single-stranded RNA. Taxonomically, MODV is part of the Flavivirus genus and Flaviviridae family. The Flavivirus genus includes nearly 80 viruses, both vector-borne and no known vector (NKV) species. Known flavivirus vector-borne viruses include Dengue virus, Yellow Fever virus, tick-borne encephalitis virus, and West Nile virus.
Orthornavirae is a kingdom of viruses that have genomes made of ribonucleic acid (RNA), those genomes encoding an RNA-dependent RNA polymerase (RdRp). The RdRp is used to transcribe the viral RNA genome into messenger RNA (mRNA) and to replicate the genome. Viruses in this kingdom also share a number of characteristics involving evolution, including high rates of genetic mutations, recombinations, and reassortments.
Rio Negro virus is an alphavirus that was first isolated in Argentina in 1980. The virus was first called Ag80-663 but was renamed to Rio Negro virus in 2005. It is a former member of the Venezuelan equine encephalitis complex (VEEC), which are a group of alphaviruses in the Americas that have the potential to emerge and cause disease. Río Negro virus was recently reclassified as a distinct species. Closely related viruses include Mucambo virus and Everglades virus.