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''Bunyaviricetes'' is a class of segmented
negative-strand RNA virus Negative-strand RNA viruses (−ssRNA viruses) are a group of related viruses that have Sense (molecular biology), negative-sense, single-stranded genomes made of ribonucleic acid (RNA). They have genomes that act as complementary strands from w ...
es with mainly tripartite genomes. Member viruses infect
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s,
plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s,
protozoa Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
ns, and
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s. The name ''Bunyaviricetes'' derives from Bunyamwera, where the virus Bunyamwera virus was first discovered. Bunyaviruses belong to the fifth group of the Baltimore classification system, which includes viruses with a negative-sense, single-stranded RNA genome. They have an enveloped, spherical virion. Though generally found in
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s or rodents, certain viruses in this class 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 and
Arenavirus An arenavirus is a bi- or trisegmented 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 reptarenavirus ...
es, all viruses in the ''Bunyaviricetes'' class 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 (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
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 ''Bunyaviricetes'', such as Crimean-Congo hemorrhagic fever virus, 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 viruses that includes all hantaviruses (family ''Hantaviridae'') that cause disease in humans. Orthohantaviruses, hereafter referred to as hantaviruses, are naturally found primarily in rodents. In general, each ...
es are another medically important member of the class ''Bunyaviricetes''. They are found worldwide, and are relatively common in
Korea Korea is a peninsular region in East Asia consisting of the Korean Peninsula, Jeju Island, and smaller islands. Since the end of World War II in 1945, it has been politically Division of Korea, divided at or near the 38th parallel north, 3 ...
,
Scandinavia Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...
(including
Finland Finland, officially the Republic of Finland, is a Nordic country in Northern Europe. It borders Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of Bothnia to the west and the Gulf of Finland to the south, ...
),
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
, 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 ''Paramyxoviridae'' (from Ancient Greek, Greek ''para-'' “by the side of” and ''myxa'' “mucus”) is a family of negative-strand RNA viruses in the order ''Mononegavirales''. Vertebrates serve as natural hosts. Diseases associated with this ...
'' family; bunyaviruses form enveloped, spherical virions with diameters of 80–120 nm. These viruses contain no matrix proteins. Instead, the viral surface glycoproteins which form a continuous layer on the virion surface are thought to play a role in the formation of new virions by budding from a cell membrane.


Genome

Bunyaviruses have bi- or tripartite
genome 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 genes, other functional regions of the genome such as ...
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 self-replication, replication of RNA from an RNA template. Specifically, it catalyzes synthesis of the RNA strand Complementarity (molecular biology), compleme ...
, necessary for viral RNA replication and mRNA synthesis. The M segment encodes the viral
glycoprotein Glycoproteins are proteins which contain oligosaccharide (sugar) 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 a ...
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 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'' 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 (N) in the negative sense and a
nonstructural protein In virology, a nonstructural protein is a protein encoded by a virus but that is not part of the viral particle. They typically include the various enzymes and transcription factors the virus uses to replicate itself, such as a viral protease ( 3CL ...
(NSs) in the positive sense. The ambisense M segment codes for
glycoprotein Glycoproteins are proteins which contain oligosaccharide (sugar) 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 a ...
(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 The cytoplasm describes all the material within a eukaryotic or prokaryotic cell, enclosed by the cell membrane, including the organelles and excluding the nucleus in eukaryotic cells. The material inside the nucleus of a eukaryotic cell a ...
, 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 Cell (biology), cells. Part of the endomembrane system in the cytoplasm, it protein targeting, packages proteins ...
. Mature virions bud from the Golgi apparatus into vesicles which are transported to the cell surface.


Transmission

Bunyaviruses infect
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s,
plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s,
protozoa Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
ns, and
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
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 group of insects that includes all members of the order Lepidoptera that are not butterflies. They were previously classified as suborder Heterocera, but the group is paraphyletic with respect to butterflies (suborder Rhopalocera) a ...
, wasps and bees, and other true flies.


Taxonomy

The class ''Bunyaviricetes'' contains two orders which contain 16 families. These taxa are listed hereafter (-''virales'' denotes order and -''viridae'' denotes family): * '' Elliovirales'' ** '' Cruliviridae'' ** '' Fimoviridae'' ** '' Hantaviridae'' ** '' Peribunyaviridae'' ** '' Phasmaviridae'' ** '' Tospoviridae'' ** '' Tulasviridae'' * '' Hareavirales'' ** '' Arenaviridae'' ** '' Discoviridae'' ** '' Konkoviridae'' ** '' Leishbuviridae'' ** '' Mypoviridae'' ** '' Nairoviridae'' ** '' Phenuiviridae'' ** '' Tosoviridae'' ** '' Wupedeviridae''


Diseases in humans

Bunyaviruses that cause disease in humans include: * California encephalitis virus, Jamestown Canyon virus, La Crosse encephalitis virus, Oropouche virus, and Snowshoe hare virus (vector: mosquitoes; family: ''Peribunyaviridae''); *
Hantavirus ''Orthohantavirus'' is a genus of viruses that includes all hantaviruses (family ''Hantaviridae'') that cause disease in humans. Orthohantaviruses, hereafter referred to as hantaviruses, are naturally found primarily in rodents. In general, each ...
reservoir: small mammals or rodents (vector: aerosolized excreta from these mammals; family: ''Hantaviridae''); * Crimean–Congo hemorrhagic fever reservoir and (vector: ticks; amplifying hosts and vector: small mammals, domestic mammals; family: '' Nairoviridae''); * Rift Valley fever (reservoir: bats; vector: mosquitoes; amplifying hosts: small and domestic mammals; family: '' Phenuiviridae''); * 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, is a type of viral hemorrhagic fever caused by the Lassa virus. Many of those infected by the virus asymptomatic, do not develop symptoms. When symptoms occur they typically include fever, wea ...
and
Argentine hemorrhagic fever Argentine hemorrhagic fever (AHF) or O'Higgins disease, also known in Argentina as mal de los rastrojos (stubble disease) is a hemorrhagic fever and zoonotic infectious disease occurring in Argentina. It is caused by the Junín virus (an arenavi ...
(reservoir: rodents; vector: aerosolized excreta from these mammals; family: ''Arenaviridae''). Bunyaviruses have segmented genomes, making them capable of rapid reassortment and increasing the risk of outbreak. The bunyavirus that causes severe fever with thrombocytopenia syndrome can undergo recombination both by reassortment of
genome 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 genes, other functional regions of the genome such as ...
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 Cell (biology), cellular organi ...
. Bunyaviruses 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 symptoms with fever lasting for about three days. Because of their non-specific symptoms, 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 is discovered in
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
1951: 3,000 cases of
Hantavirus ''Orthohantavirus'' is a genus of viruses that includes all hantaviruses (family ''Hantaviridae'') that cause disease in humans. Orthohantaviruses, hereafter referred to as hantaviruses, are naturally found primarily in rodents. In general, each ...
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 was first recognized in a fatal case of encephalitis in
La Crosse, Wisconsin La Crosse ( ) is a city in La Crosse County, Wisconsin, United States, and its county seat. Positioned alongside the Mississippi River, La Crosse is the largest city on Wisconsin's western border. La Crosse's population was 52,680 as of the 202 ...
1977: Rift Valley Fever virus caused approximately 200,000 cases and 598 deaths in
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
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 classes