Double-stranded RNA viruses (dsRNA viruses) are a
polyphyletic
A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
group of
virus
A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea.
Since Dmitri Ivanovsky's 1 ...
es that have
double-stranded
A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DN ...
genomes made of
ribonucleic acid
Ribonucleic acid (RNA) is a polymeric molecule essential in various biological roles in coding, decoding, regulation and expression of genes. RNA and deoxyribonucleic acid ( DNA) are nucleic acids. Along with lipids, proteins, and carbohydra ...
. The double-stranded genome is used to transcribe a
positive-strand RNA by the viral
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 t ...
(RdRp). The positive-strand RNA may be used as
messenger RNA
In molecular biology, messenger ribonucleic acid (mRNA) is a single-stranded molecule of RNA that corresponds to the genetic sequence of a gene, and is read by a ribosome in the process of synthesizing a protein.
mRNA is created during the p ...
(mRNA) which can be
translated
Translation is the communication of the meaning of a source-language text by means of an equivalent target-language text. The English language draws a terminological distinction (which does not exist in every language) between ''transla ...
into viral proteins by the
host cell's ribosome
Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to ...
s. The positive-strand RNA can also be replicated by the RdRp to create a new double-stranded viral genome.
Double-stranded RNA viruses are classified in two separate phyla ''
Duplornaviricota
''Duplornaviricota'' is a phylum of RNA viruses, which contains all double-stranded RNA viruses, except for those in phylum ''Pisuviricota.'' Characteristic of the group is a viral capsid composed of 60 homo- or heterodimers of capsid protein on ...
'' and ''
Pisuviricota
''Pisuviricota'' is a phylum of RNA viruses that includes all positive-strand and double-stranded RNA viruses that infect eukaryotes and are not members of the phylum ''Kitrinoviricota,'' ''Lenarviricota'' or ''Duplornaviricota.'' The name of t ...
'' (specifically class ''
Duplopiviricetes
''Durnavirales'' is an order of double-stranded RNA viruses which infect eukaryotes. The name of the group derives from Italian ''duplo'' which means double (a reference to double-stranded), ''rna'' for the type of virus, and -''virales'' which i ...
''), which are in the kingdom ''
Orthornavirae
''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 repli ...
'' and
realm
A realm is a community or territory over which a sovereign rules. The term is commonly used to describe a monarchical or dynastic state. A realm may also be a subdivision within an empire, if it has its own monarch, e.g. the German Empire.
Etym ...
''
Riboviria
''Riboviria'' is a realm of viruses that includes all viruses that use a homologous RNA-dependent polymerase for replication. It includes RNA viruses that encode an RNA-dependent RNA polymerase, as well as reverse-transcribing viruses (with eithe ...
''. The two groups do not share a common dsRNA virus ancestor. Double-stranded RNA viruses evolved two separate times from
positive-strand RNA virus
Positive-strand RNA viruses (+ssRNA viruses) are a group of related viruses that have positive-sense, single-stranded genomes made of ribonucleic acid. The positive-sense genome can act as messenger RNA (mRNA) and can be directly translated int ...
es. In the
Baltimore classification
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 di ...
system, dsRNA viruses belong to Group III.
Virus group members vary widely in host range (
animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
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 exclud ...
s,
fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
, and
bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
), genome segment number (one to twelve), and
virion
A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea.
Since Dmitri Ivanovsky's 1 ...
organization (
T-number,
capsid
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 ...
layers, or turrets). Double-stranded RNA viruses include the
rotavirus
''Rotavirus'' is a genus of double-stranded RNA viruses in the family ''Reoviridae''. Rotaviruses are the most common cause of diarrhoeal disease among infants and young children. Nearly every child in the world is infected with a rotavirus a ...
es, known globally as a common cause of
gastroenteritis
Gastroenteritis, also known as infectious diarrhea and gastro, is an inflammation of the gastrointestinal tract including the stomach and intestine. Symptoms may include diarrhea, vomiting, and abdominal pain. Fever, lack of energy, and dehydra ...
in young children, and
bluetongue virus
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 ...
, an economically significant pathogen of cattle and sheep. The family ''
Reoviridae
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
'' is the largest and most diverse dsRNA virus family in terms of host range.
Classification
Two clades of dsRNA viruses exist: the phylum ''Duplornaviricota'' and the class ''Duplopiviricetes'', which is in the phylum ''Pisuviricota''. Both are included in the kingdom ''Orthornavirae'' in the realm ''Riboviria''. Based on phylogenetic analysis of RdRp, the two clades do not share a common dsRNA ancestor but are instead separately descended from different positive-sense, single-stranded RNA viruses. In the
Baltimore classification
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 di ...
system, which groups viruses together based on their manner of mRNA synthesis, dsRNA viruses are group III.
''Duplornaviricota''
''
Duplornaviricota
''Duplornaviricota'' is a phylum of RNA viruses, which contains all double-stranded RNA viruses, except for those in phylum ''Pisuviricota.'' Characteristic of the group is a viral capsid composed of 60 homo- or heterodimers of capsid protein on ...
'' contains most dsRNA viruses, including
reoviruses
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
, which infect a diverse range of eukaryotes, and
cystoviruses, which are the only dsRNA viruses known to infect prokaryotes. Apart from RdRp, viruses in ''Duplornaviricota'' also share icosahedral capsids that contain 60 homo- or heterodimers of the capsid protein organized on a pseudo T=2 lattice. The phylum is divided into three classes: ''
Chrymotiviricetes
''Ghabrivirales'' is an order of double-stranded RNA viruses. It is the only order in the class ''Chrysmotiviricetes''. The name of the class is a portmanteau of member families: ''chrys''oviridae, ''m''egabirnaviridae, and ''toti''viridae; and - ...
'', which primarily contains fungal and protozoan viruses, ''Resentoviricetes'', which contains reoviruses, and ''Vidaverviricetes'', which contains cystoviruses.
''Duplopiviricetes''
The class ''
Duplopiviricetes
''Durnavirales'' is an order of double-stranded RNA viruses which infect eukaryotes. The name of the group derives from Italian ''duplo'' which means double (a reference to double-stranded), ''rna'' for the type of virus, and -''virales'' which i ...
'' is the second clade of dsRNA viruses and is in the phylum ''
Pisuviricota
''Pisuviricota'' is a phylum of RNA viruses that includes all positive-strand and double-stranded RNA viruses that infect eukaryotes and are not members of the phylum ''Kitrinoviricota,'' ''Lenarviricota'' or ''Duplornaviricota.'' The name of t ...
'', which also contains positive-sense single-stranded RNA viruses. ''Duplopiviricetes'' mostly contains plant and fungal viruses and includes the following four families: ''
Amalgaviridae
''Amalgaviridae'' is a family of double-stranded RNA viruses. Member viruses infect plants and are transmitted vertically via seeds. The name derives from ''amalgam'' (blend, mix) which refers to amalgaviruses possessing characteristics of both ...
'', ''
Hypoviridae
''Hypovirus'' is a genus of viruses, in the family ''Hypoviridae''. Fungi serve as natural hosts. There are four species in this genus. Infection reduces the virulence of its parasitic host, making it a hyperparasite useful for blight control.
...
'', ''
Partitiviridae
''Partitiviridae'' is a family of double-stranded RNA viruses. Plants, fungi, and protozoa serve as natural hosts. It has been suggested that they can also infect bacteria. The name comes from the Latin ''partitius,'' which means divided, and ...
'', and ''
Picobirnaviridae
''Picobirnavirus'' is a genus of double-stranded RNA viruses. It is the only genus in the family ''Picobirnaviridae''. Although amniotes, especially mammals, were thought to serve as hosts, it has been recently suggested that these viruses migh ...
''.
Notes on selected species
Reoviridae
''
Reoviridae
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
'' are currently classified into nine
genera
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
. The genomes of these viruses consist of 10 to 12 segments of
dsRNA
Ribonucleic acid (RNA) is a polymeric molecule essential in various biological roles in coding, decoding, regulation and expression of genes. RNA and deoxyribonucleic acid ( DNA) are nucleic acids. Along with lipids, proteins, and carbohydra ...
, each generally encoding one
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
. The mature virions are non-enveloped. Their capsids, formed by multiple proteins, have
icosahedral
In geometry, an icosahedron ( or ) is a polyhedron with 20 faces. The name comes and . The plural can be either "icosahedra" () or "icosahedrons".
There are infinitely many non- similar shapes of icosahedra, some of them being more symmetrica ...
symmetry and are arranged generally in concentric layers. A distinguishing feature of the dsRNA viruses, irrespective of their family association, is their ability to carry out
transcription
Transcription refers to the process of converting sounds (voice, music etc.) into letters or musical notes, or producing a copy of something in another medium, including:
Genetics
* Transcription (biology), the copying of DNA into RNA, the fir ...
of the dsRNA segments, under appropriate conditions, within the capsid. In all these viruses, the
enzyme
Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. A ...
s required for endogenous transcription are thus part of the virion structure.
Orthoreoviruses
The
orthoreovirus
''Orthoreovirus'' is a genus of viruses, in the family ''Reoviridae'', in the subfamily ''Spinareovirinae''. Vertebrates serve as natural hosts. There are ten species in this genus. Diseases associated with this genus include mild upper respirato ...
es (
reovirus
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
es) are the prototypic members of the virus ''
Reoviridae
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
'' family and representative of the turreted members, which comprise about half the genera. Like other members of the family, the reoviruses are non-enveloped and characterized by concentric capsid shells that encapsidate a segmented dsRNA
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 ge ...
. In particular, reovirus has eight structural proteins and ten segments of dsRNA. A series of uncoating steps and conformational changes accompany cell entry and replication. High-resolution structures are known for almost all of the proteins of mammalian reovirus (MRV), which is the best-studied genotype. Electron cryo-
microscopy
Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of micr ...
(cryoEM) and X-ray
crystallography
Crystallography is the experimental science of determining the arrangement of atoms in crystalline solids. Crystallography is a fundamental subject in the fields of materials science and solid-state physics (condensed matter physics). The wor ...
have provided a wealth of structural information about two specific MRV strains, type 1 Lang (T1L) and type 3 Dearing (T3D).
Cypovirus
The cytoplasmic polyhedrosis viruses (CPVs) form the genus
Cypovirus
''Cypovirus'', short for cytoplasmic polyhedrosis virus, is a genus of double-stranded RNA viruses in the family ''Reoviridae'' and subfamily ''Spinareovirinae''. Cypoviruses have only been isolated from insects. Diseases associated with this ge ...
of the family ''
Reoviridae
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
''. CPVs are classified into 14 species based on the
electrophoretic
Electrophoresis, from Ancient Greek ἤλεκτρον (ḗlektron, "amber") and φόρησις (phórēsis, "the act of bearing"), is the motion of dispersed particles relative to a fluid under the influence of a spatially uniform electric fie ...
migration profiles of their genome segments. Cypovirus has only a single capsid shell, which is similar to the orthoreovirus inner core. CPV exhibits striking capsid stability and is fully capable of endogenous RNA transcription and processing. The overall folds of CPV proteins are similar to those of other reoviruses. However, CPV proteins have insertional domains and unique structures that contribute to their extensive intermolecular interactions. The CPV turret protein contains two
methylase
Methyltransferases are a large group of enzymes that all methylate their substrates but can be split into several subclasses based on their structural features. The most common class of methyltransferases is class I, all of which contain a Rossm ...
domains with a highly conserved
helix
A helix () is a shape like a corkscrew or spiral staircase. It is a type of smooth space curve with tangent lines at a constant angle to a fixed axis. Helices are important in biology, as the DNA molecule is formed as two intertwined helices, ...
-pair/β-sheet/helix-pair sandwich fold but lacks the β-barrel flap present in orthoreovirus
λ2. The stacking of turret protein functional domains and the presence of constrictions and A spikes along the mRNA release pathway indicate a mechanism that uses pores and channels to regulate the highly coordinated steps of RNA transcription, processing, and release.
Rotavirus
Rotavirus
''Rotavirus'' is a genus of double-stranded RNA viruses in the family ''Reoviridae''. Rotaviruses are the most common cause of diarrhoeal disease among infants and young children. Nearly every child in the world is infected with a rotavirus a ...
is the most common cause of acute
gastroenteritis
Gastroenteritis, also known as infectious diarrhea and gastro, is an inflammation of the gastrointestinal tract including the stomach and intestine. Symptoms may include diarrhea, vomiting, and abdominal pain. Fever, lack of energy, and dehydra ...
in infants and young children worldwide. This virus contains a dsRNA genome and is a member of the ''
Reoviridae
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
'' family. The genome of rotavirus consists of eleven segments of dsRNA. Each genome segment codes for one protein with the exception of segment 11, which codes for two proteins. Among the twelve proteins, six are structural and six are non-structural proteins.
It is a double-stranded RNA non-enveloped virus
Bluetongue virus
The members of 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 verte ...
'' within the ''
Reoviridae
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
'' family are
arthropod borne viruses and are responsible for high morbidity and mortality in
ruminant
Ruminants (suborder Ruminantia) are ungulate, hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principally ...
s.
Bluetongue virus
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 ...
(BTV) which causes disease in livestock (
sheep
Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus ''Ovis'', in everyday usage it almost always refers to domesticated s ...
,
goat
The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
,
cattle
Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult mal ...
) has been in the forefront of molecular studies for the last three decades and now represents the best understood
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 verte ...
at the molecular and structural levels. BTV, like other members of the family, is a complex non-enveloped virus with seven structural proteins and a RNA genome consisting of 10 variously sized dsRNA segments.
Phytoreoviruses
Phytoreovirus
''Phytoreovirus'' is a genus of viruses, in the family ''Reoviridae'', in the subfamily ''Sedoreovirinae''. They are non-turreted reoviruses that are major agricultural pathogens, particularly in Asia. Oryza sativa for RDV and RGDV, dicotyledono ...
es are non-turreted
reoviruses
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. They lack lipid envelopes and package their segmented genome within multi-layered ...
that are major agricultural pathogens, particularly in Asia. One member of this family,
Rice Dwarf Virus
Rice dwarf virus (RDV) is a plant pathogenic virus of the family Reoviridae
''Reoviridae'' is a family of double-stranded RNA viruses. Member viruses have a wide host range, including vertebrates, invertebrates, plants, protists and fungi. The ...
(RDV), has been extensively studied by
electron cryomicroscopy and
x-ray crystallography
X-ray crystallography is the experimental science determining the atomic and molecular structure of a crystal, in which the crystalline structure causes a beam of incident X-rays to diffract into many specific directions. By measuring the angles ...
. From these analyses, atomic models of the capsid proteins and a plausible model for capsid assembly have been derived. While the structural proteins of RDV share no sequence similarity to other proteins, their folds and the overall capsid structure are similar to those of other ''Reoviridae''.
''Saccharomyces cerevisiae'' virus L-A
The
L-A dsRNA virus of the
yeast
Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constitut ...
''
Saccharomyces cerevisiae
''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have been o ...
'' has a single 4.6 kb genomic segment that encodes its major coat protein, Gag (76 kDa) and a Gag-Pol fusion protein (180 kDa) formed by a -1 ribosomal frameshift. L-A can support the replication and