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''Bunyavirales'' is an order of segmented negative-strand RNA viruses with mainly tripartite genomes. Member viruses infect
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s,
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
s,
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
ns, and
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s. It is the only order in the class ''Ellioviricetes''. The name ''Bunyavirales'' derives from
Bunyamwera Bunyamwera is a town in Bundibugyo District, Uganda. It lies just outside of Rwenzori Mountains National Park, which is to the south. The town is north of Kagugu, southeast of Bunyana, and southwest of Butama. Other nearby settlements incl ...
, where the original
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specim ...
''
Bunyamwera orthobunyavirus ''Bunyamwera orthobunyavirus'' (BUNV) is a negative-sense, single-stranded enveloped RNA virus. It is assigned to the '' Orthobunyavirus'' genus, in the ''Bunyavirales'' order. ''Bunyamwera orthobunyavirus'' can infect both humans and ''Aedes ...
'' was first discovered. ''Ellioviricetes'' is named in honor of late virologist Richard M. Elliott for his early work on bunyaviruses. Bunyaviruses belong to the fifth group of the
Baltimore classification system Baltimore classification is a system used to classify viruses based on their manner of messenger RNA (mRNA) synthesis. By organizing viruses based on their manner of mRNA production, it is possible to study viruses that behave similarly as a d ...
, which includes viruses with a
negative-sense In molecular biology and genetics, the sense of a nucleic acid molecule, particularly of a strand of DNA or RNA, refers to the nature of the roles of the strand and its complement in specifying a sequence of amino acids. Depending on the context, ...
, single-stranded RNA genome. They have an enveloped, spherical virion. Though generally found in
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s or rodents, certain viruses in this order occasionally infect humans. Some of them also infect plants. In addition, there is a group of bunyaviruses whose replication is restricted to arthropods and is known as insect-specific bunyaviruses. A majority of bunyaviruses are vector-borne. With the exception of
Hantaviruses ''Orthohantavirus'' is a genus of single-stranded, enveloped, negative-sense RNA viruses in the family '' Hantaviridae'' within the order ''Bunyavirales''. Members of this genus may be called orthohantaviruses or simply hantaviruses. Orthohantav ...
and
Arenavirus An arenavirus is a bisegmented ambisense RNA virus that is a member of the family ''Arenaviridae''. These viruses infect rodents and occasionally humans. A class of novel, highly divergent arenaviruses, properly known as reptarenaviruses, have ...
es, all viruses in the ''Bunyavirales'' order are transmitted by arthropods (mosquitos, tick, or sandfly). Hantaviruses are transmitted through contact with
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
feces. Incidence of infection is closely linked to vector activity, for example, mosquito-borne viruses are more common in the summer. Human infections with certain members of ''Bunyavirales'', such as '' Crimean-Congo hemorrhagic fever orthonairovirus'', are associated with high levels of morbidity and mortality, consequently handling of these viruses is done in biosafety level 4 laboratories. They are also the cause of severe fever with thrombocytopenia syndrome.
Hantavirus ''Orthohantavirus'' is a genus of single-stranded, enveloped, negative-sense RNA viruses in the family '' Hantaviridae'' within the order ''Bunyavirales''. Members of this genus may be called orthohantaviruses or simply hantaviruses. Orthohantav ...
es are another medically important member of the order ''Bunyvirales''. They are found worldwide, and are relatively common in
Korea Korea ( ko, 한국, or , ) is a peninsular region in East Asia. Since 1945, it has been divided at or near the 38th parallel, with North Korea (Democratic People's Republic of Korea) comprising its northern half and South Korea (Republic ...
,
Scandinavia Scandinavia; Sámi languages: /. ( ) is a subregion in Northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. In English usage, ''Scandinavia'' most commonly refers to Denmark, Norway, and S ...
(including
Finland Finland ( fi, Suomi ; sv, Finland ), officially the Republic of Finland (; ), is a Nordic country in Northern Europe. It shares land borders with Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of Bot ...
),
Russia Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It is the largest country in the world, with its internationally recognised territory covering , and encompassing one-ei ...
, western North America and parts of South America. Hantavirus infections are associated with high fever, lung edema, and pulmonary failure. The mortality rate varies significantly depending on the form, being up to 50% in New World hantaviruses (the Americas), up to 15% in Old World hantaviruses (Asia and Europe), and as little as 0.1% in Puumala virus (mostly Scandinavia). The antibody reaction plays an important role in decreasing levels of viremia.


Virology


Structure

Bunyavirus morphology is somewhat similar to that of the '' Paramyxoviridae'' family; ''Bunyavirales'' form enveloped, spherical virions with diameters of 80–120 nm. These viruses contain no matrix proteins.


Genome

Bunyaviruses have bi- or tripartite
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
s consisting of a large (L) and small(s), or large (L), medium (M), and small (S) RNA segment. These RNA segments are single-stranded, and exist in a helical formation within the virion. Besides, they exhibit a pseudo-circular structure due to each segment's complementary ends. The L segment encodes the
RNA-dependent RNA polymerase RNA-dependent RNA polymerase (RdRp) or RNA replicase is an enzyme that catalyzes the replication of RNA from an RNA template. Specifically, it catalyzes synthesis of the RNA strand complementary to a given RNA template. This is in contrast to ...
, necessary for viral RNA replication and mRNA synthesis. The M segment encodes the viral
glycoprotein Glycoproteins are proteins which contain oligosaccharide chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glyco ...
s, which project from the viral surface and aid the virus in attaching to and entering the host cell. The S segment encodes the
nucleocapsid A capsid is the protein shell of a virus, enclosing its genetic material. It consists of several oligomeric (repeating) structural subunits made of protein called protomers. The observable 3-dimensional morphological subunits, which may or may ...
protein (N). Most bunyaviruses have a negative-sense L and M segment. The S segment of the genus '' Phlebovirus'', and both M and S segment of the genus ''
Tospovirus ''Orthotospovirus'' is a genus of negative-strand RNA viruses, in the family ''Tospoviridae'' of the order ''Bunyavirales,'' which infects plants. Tospoviruses take their name from the species '' Tomato spotted wilt orthotospovirus'' (TSWV) which ...
'' are ambisense. Ambisense means that some of the genes on the RNA strand are negative sense and others are positive sense. The ambisense S segment codes for the viral
nucleoprotein Nucleoproteins are proteins conjugated with nucleic acids (either DNA or RNA). Typical nucleoproteins include ribosomes, nucleosomes and viral nucleocapsid proteins. Structures Nucleoproteins tend to be positively charged, facilitating int ...
(N) in the negative sense and a nonstructural protein (NSs) in the positive sense. The ambisense M segment codes for
glycoprotein Glycoproteins are proteins which contain oligosaccharide chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glyco ...
(GP) in the negative sense and a nonstructural protein (NSm) in the positive sense. The total genome size ranges from 10.5 to 22.7 kbp.


Life cycle

The ambisense genome requires two rounds of transcription to be carried out. First, the negative-sense RNA is transcribed to produce mRNA and a full-length replicative intermediate. From this intermediate, a subgenomic mRNA encoding the small segment nonstructural protein is produced while the polymerase produced following the first round of transcription can now replicate the full-length RNA to produce viral genomes. Bunyaviruses replicate in the
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
, while the viral proteins transit through the ER and
Golgi apparatus The Golgi apparatus (), also known as the Golgi complex, Golgi body, or simply the Golgi, is an organelle found in most eukaryotic cells. Part of the endomembrane system in the cytoplasm, it packages proteins into membrane-bound vesicles i ...
. Mature virions bud from the Golgi apparatus into vesicles which are transported to the cell surface.


Transmission

Bunyaviruses infect
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s,
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
s,
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
ns, and
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s. Plants can host bunyaviruses from the families ''Tospoviridae'' and ''Fimoviridae'' (e.g. tomato, pigeonpea, melon, wheat, raspberry, redbud, and rose). Members of some families are insect-specific, for example the phasmavirids, first isolated from phantom midges, and since identified in diverse insects including
moths Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
, wasps and bees, and other
true flies Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
.


Taxonomy

There are 477 virus species recognised in this order. The phylogenetic tree diagram provides a full list of member species and the hosts which they infect. The order is organized into the following 12 families: * '' Arenaviridae'' * ''
Cruliviridae Cruliviridae is a family of virus in the order of Bunyavirales. Unlike other families in the order, this family is unique in the sense that it is only used to contain crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon wh ...
'' * '' Fimoviridae'' * ''
Hantaviridae ''Hantaviridae'' is a family of viruses in the order ''Bunyavirales''. It is named for the Hantan River area in South Korea where an early outbreak of one of its species was observed. Taxonomy * '' Actantavirinae'' ** '' Actinovirus'' * '' Aga ...
'' * ''
Leishbunyaviridae ''Leishbunyaviridae'' is a family of negative-strand RNA viruses belonging to the ''Bunyavirales'' order, which infect protozoans Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukar ...
'' * '' Mypoviridae'' * ''
Nairoviridae ''Nairoviridae'' is a family of viruses in the order ''Bunyavirales ''Bunyavirales'' is an order of segmented negative-strand RNA viruses with mainly tripartite genomes. Member viruses infect arthropods, plants, protozoans, and vertebrates. I ...
'' * '' Peribunyaviridae'' * ''
Phasmaviridae Phasmaviridae is a family of viruses with negative stranded RNA genomes associated with insect hosts. They are a member of the order ''Bunyavirales''. Phasmaviruses were first discovered in phantom midges of the genus '' Chaoborus'' in 2014. T ...
'' * ''
Phenuiviridae ''Phenuiviridae'' is a family of negative-strand RNA viruses in the order '' Bunyavirales''. Ruminants, camels, humans, and mosquitoes serve as natural hosts. Member genus ''Phlebovirus'' is the only genus of the family that has viruses that caus ...
'' * ''
Tospoviridae ''Orthotospovirus'' is a genus of negative-strand RNA viruses, in the family '' Tospoviridae'' of the order ''Bunyavirales,'' which infects plants. Tospoviruses take their name from the species '' Tomato spotted wilt orthotospovirus'' (TSWV) whic ...
'' * '' Wupedeviridae''


Diseases in humans

Bunyaviruses that cause disease in humans include: *
California encephalitis virus ''California encephalitis orthobunyavirus'' type strain California encephalitis virus was discovered in Kern County, California, and causes encephalitis in humans. Encephalitis is an acute inflammation of the brain that can cause minor symptoms, ...
,
Jamestown Canyon virus Jamestown Canyon encephalitis is an infectious disease caused by the Jamestown Canyon virus, an orthobunyavirus of the California serogroup. It is mainly spread during the summer by different mosquito species in the United States and Canada. ...
, La Crosse encephalitis virus, Oropouche orthobunyavirus, and Snowshoe hare virus (vector: mosquitoes; family: Peribunyaviridae); *
Hantavirus ''Orthohantavirus'' is a genus of single-stranded, enveloped, negative-sense RNA viruses in the family '' Hantaviridae'' within the order ''Bunyavirales''. Members of this genus may be called orthohantaviruses or simply hantaviruses. Orthohantav ...
reservoir: small mammals or rodents (vector: aerosolized excreta from these mammals; family: Hantaviridae); *
Crimean–Congo hemorrhagic fever Crimean–Congo hemorrhagic fever (CCHF) is a viral disease. Symptoms of CCHF may include fever, muscle pains, headache, vomiting, diarrhea, and bleeding into the skin. Onset of symptoms is less than two weeks following exposure. Complicatio ...
reservoir and (vector: ticks; amplifying hosts and vector: small mammals, domestic mammals; family:
Nairoviridae ''Nairoviridae'' is a family of viruses in the order ''Bunyavirales ''Bunyavirales'' is an order of segmented negative-strand RNA viruses with mainly tripartite genomes. Member viruses infect arthropods, plants, protozoans, and vertebrates. I ...
); * Rift Valley fever (reservoir: bats; vector: mosquitoes; amplifying hosts: small and domestic mammals; family:
Phenuiviridae ''Phenuiviridae'' is a family of negative-strand RNA viruses in the order '' Bunyavirales''. Ruminants, camels, humans, and mosquitoes serve as natural hosts. Member genus ''Phlebovirus'' is the only genus of the family that has viruses that caus ...
); * Bwamba Fever (reservoir: monkeys; vector: mosquitoes; amplifying hosts: donkeys; family: Peribunyaviridae); * Severe fever with thrombocytopenia syndrome (vector: ticks); *
Lassa fever Lassa fever, also known as Lassa hemorrhagic fever (LHF), is a type of viral hemorrhagic fever caused by the Lassa virus. Many of those infected by the virus do not develop symptoms. When symptoms occur they typically include fever, weakness, ...
and Argentine hemorrhagic fever (reservoir: rodents; vector: aerosolized excreta from these mammals; family: Arenaviridae). Bunyaviruses have segmented genomes, making them capable of rapid recombination and increasing the risk of outbreak. The bunyavirus that causes severe fever with thrombocytopenia syndrome can undergo recombination both by reassortment of
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
segments and by intragenic
homologous recombination Homologous recombination is a type of genetic recombination in which genetic information is exchanged between two similar or identical molecules of double-stranded or single-stranded nucleic acids (usually DNA as in cellular organisms but may ...
. Bunyaviridae are transmitted by hematophagous arthropods including mosquitoes, midges, flies, and ticks. The viral incubation period is about 48 hours. Symptomatic infection typically causes non-specific
flu-like Influenza-like illness (ILI), also known as flu-like syndrome or flu-like symptoms, is a medical diagnosis of possible influenza or other illness causing a set of common symptoms. These include fever, shivering, chills, malaise, dry cough, loss ...
symptoms with fever lasting for about three days. Because of their
non-specific symptoms Signs and symptoms are the observed or detectable signs, and experienced symptoms of an illness, injury, or condition. A sign for example may be a higher or lower temperature than normal, raised or lowered blood pressure or an abnormality showin ...
, Bunyavirus infections are frequently mistaken for other illnesses. For example, Bwamba fever is often mistaken for malaria.


Prevention

Prevention depends on the reservoir, amplifying hosts and how the viruses are transmitted, i.e. the vector, whether ticks or mosquitoes and which animals are involved. Preventive measures include general hygiene, limiting contact with vector saliva, urine, feces, or bedding. There is no licensed vaccine for bunyaviruses. As precautions Cache Valley virus and Hantavirus research are conducted in BSL-2 (or higher), Rift Valley Fever virus research is conducted in BSL-3 (or higher), Congo-Crimean Hemorrhagic Fever virus research is conducted in BSL-4 laboratories.


Timeline

1940s:
Crimean–Congo hemorrhagic fever Crimean–Congo hemorrhagic fever (CCHF) is a viral disease. Symptoms of CCHF may include fever, muscle pains, headache, vomiting, diarrhea, and bleeding into the skin. Onset of symptoms is less than two weeks following exposure. Complicatio ...
is discovered in
Russia Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It is the largest country in the world, with its internationally recognised territory covering , and encompassing one-ei ...
1951: 3,000 cases of
Hantavirus ''Orthohantavirus'' is a genus of single-stranded, enveloped, negative-sense RNA viruses in the family '' Hantaviridae'' within the order ''Bunyavirales''. Members of this genus may be called orthohantaviruses or simply hantaviruses. Orthohantav ...
were reported in South Korea in 1951, a time when UN forces were fighting on the 38th parallel during the Korean War 1956: Cache Valley virus isolated in '' Culiseta inornata'' mosquitoes in Utah 1960:
La Crosse virus La Crosse encephalitis is an encephalitis caused by an arbovirus (the La Crosse virus) which has a mosquito vector ('' Ochlerotatus triseriatus'' synonym ''Aedes'' ''triseriatus''). La Crosse encephalitis virus (LACV) is one of a group of mo ...
was first recognized in a fatal case of encephalitis in
La Crosse, Wisconsin La Crosse is a city in the U.S. state of Wisconsin and the county seat of La Crosse County. Positioned alongside the Mississippi River, La Crosse is the largest city on Wisconsin's western border. La Crosse's population as of the 2020 census wa ...
1977: Rift Valley Fever virus caused approximately 200,000 cases and 598 deaths in
Egypt Egypt ( ar, مصر , ), officially the Arab Republic of Egypt, is a List of transcontinental countries, transcontinental country spanning the North Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via a land bridg ...
2017: Bunyavirales order is created


References


External links


Viralzone: Bunyaviridae





Bunyaviridae Genomes
��database search results from th
Viral Bioinformatics Resource Center

Virus Pathogen Database and Analysis Resource (ViPR): Bunyaviridae
* {{Authority control Hemorrhagic fevers Virus orders