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''Reovirales'' is an order of double-stranded RNA viruses. Member viruses, called reoviruses, have a wide host range, including
vertebrates 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 ...
,
invertebrates Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum ...
, plants,
protists A protist ( ) or protoctist is any Eukaryote, eukaryotic organism that is not an animal, Embryophyte, land plant, or fungus. Protists do not form a Clade, natural group, or clade, but are a Paraphyly, paraphyletic grouping of all descendants o ...
and fungi. They lack lipid envelopes and package their segmented genome within multi-layered
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 m ...
s. Lack of a lipid envelope has allowed three-dimensional structures of these large complex viruses (diameter ~60–100 nm) to be obtained, revealing a structural and likely evolutionary relationship to the cystovirus family of
bacteriophage A bacteriophage (), also known informally as a phage (), is a virus that infects and replicates within bacteria. The term is derived . Bacteriophages are composed of proteins that Capsid, encapsulate a DNA or RNA genome, and may have structu ...
. Reoviruses can affect the gastrointestinal system (such as
rotavirus Rotaviruses are the most common cause of diarrhea, diarrhoeal disease among infants and young children. Nearly every child in the world is infected with a rotavirus at least once by the age of five. Immunity (medical), Immunity develops with ...
es) and
respiratory tract The respiratory tract is the subdivision of the respiratory system involved with the process of conducting air to the alveoli for the purposes of gas exchange in mammals. The respiratory tract is lined with respiratory epithelium as respirato ...
. The name "reo-" is an
acronym An acronym is a type of abbreviation consisting of a phrase whose only pronounced elements are the initial letters or initial sounds of words inside that phrase. Acronyms are often spelled with the initial Letter (alphabet), letter of each wor ...
for "''r''espiratory ''e''nteric ''o''rphan" viruses''.'' The term " orphan virus" refers to the fact that some of these viruses have been observed not associated with any known disease. Even though viruses in the order ''Reovirales'' have more recently been identified with various diseases, the original name is still used. Reovirus infections occur often in humans, but most cases are mild or subclinical.
Rotavirus Rotaviruses are the most common cause of diarrhea, diarrhoeal disease among infants and young children. Nearly every child in the world is infected with a rotavirus at least once by the age of five. Immunity (medical), Immunity develops with ...
es, however, can cause severe
diarrhea Diarrhea (American English), also spelled diarrhoea or diarrhœa (British English), is the condition of having at least three loose, liquid, or watery bowel movements in a day. It often lasts for a few days and can result in dehydration d ...
and intestinal distress in children, and lab studies in mice have implicated orthoreoviruses in the expression of coeliac disease in pre-disposed individuals. The virus can be readily detected in
feces Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the ...
, and may also be recovered from pharyngeal or nasal secretions, urine, cerebrospinal fluid, and blood. Despite the ease of finding reoviruses in clinical specimens, their role in human disease or treatment is still uncertain. Some viruses of this order, such as
phytoreovirus ''Phytoreovirus'' is a genus of viruses, in the order ''Reovirales'', in the family ''Sedoreoviridae''. They are non-turreted reoviruses that are major agricultural pathogens, particularly in Asia. Oryza sativa for RDV and RGDV, dicotyledonous f ...
es and oryzaviruses, infect plants. Most of the plant-infecting reoviruses are transmitted between plants by insect vectors. The viruses replicate in both the plant and the insect, generally causing disease in the plant, but little or no harm to the infected insect.


Structure

Reoviruses are non-enveloped and have an icosahedral
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 m ...
composed of an outer ( T=13) and inner (T=2) protein shell. Ultrastructure studies show that virion capsids are composed of two or three separate layers which depends on species type. The innermost layer (core) has T=1 icosahedral symmetry and is composed of 60 different types of structural proteins. The core contains the genome segments, each of them encode a variety enzyme structure which is required for transcription. The core is covered by capsid layer T=13 icosahedral symmetry. Reoviruses have a unique structure which contains a glycosylated spike protein on the surface.


Genome

The genomes of viruses in the order ''Reovirales'' contain 9–12 segments which are grouped into three categories corresponding to their size: L (large), M (medium) and S (small). Segments range from about 0.2 to 3 kbp and each segment encodes 1–3 proteins (10–14 proteins in total). Proteins of viruses in the order ''Reovirales'' are denoted by the Greek character corresponding to the segment it was translated from (the L segment encodes for λ proteins, the M segment encodes for μ proteins and the S segment encodes for σ proteins).


Life cycle

Viruses in the order ''Reovirales'' have genomes consisting of segmented, double-stranded RNA (dsRNA). Because of this, replication occurs exclusively in the cytoplasm, and the virus encodes several proteins which are needed for replication and conversion of the dsRNA genome into positive-sense RNAs. The virus can enter the host cell via a receptor on the cell surface. The receptor is not known but is thought to include sialic acid and junctional adhesion molecules (JAMs). The virus is partially uncoated by proteases in the endolysosome, where the capsid is partially digested to allow further cell entry. The core particle then enters the cytoplasm by a yet unknown process where the genome is transcribed conservatively causing an excess of positive-sense strands, which are 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 ...
templates to synthesize negative-sense strands. The genome of the rotavirus is divided into 11 segments. These segments are associated with the VP1 molecule which is responsible for RNA synthesis. In early events, the selection process occurs so that the entry of the 11 different RNA segments go in the cell. This procedure is performed by newly synthesized RNAs. This event ensures that one each of the 11 different RNA segments is received. In late events, the transcription process occurs again but this time is not capped unlike the early events. For virus different amounts of RNAs are required therefore during the translation step there is a control machinery. There are the same quantities of RNA segments but different quantities of proteins. The reason for this is that the RNA segments are not translated at the same rate. Viral particles begin to assemble in the cytoplasm 6–7 hours after infection. Translation takes place by leaky scanning, suppression of termination, and ribosomal skipping. The virus exits the host cell by monopartite non-tubule guided viral movement, cell to cell movement, and existing in occlusion bodies after cell death and remaining infectious until finding another host.


Multiplicity reactivation

Multiplicity reactivation (MR) is the process by which two or more virus genomes, each containing inactivating genome damage, can interact within an infected cell to form a viable virus genome. McClain and Spendlove demonstrated MR for three types of reovirus after exposure to ultraviolet irradiation. In their experiments, reovirus particles were exposed to doses of UV-light that would be lethal in single infections. However, when two or more inactivated viruses were allowed to infect individual host cells MR occurred and viable progeny were produced. As they stated, multiplicity reactivation by definition involves some type of repair. Michod et al. reviewed numerous examples of MR in different viruses, and suggested that MR is a common form of sexual interaction in viruses that provides the benefit of recombinational repair of genome damages.


Taxonomy

The order ''Reovirales'' is the sole order in the class ''Resentoviricetes'', which belongs to the phylum '' Duplornaviricota''. ''Reovirales'' contains two families: '' Sedoreoviridae'' and '' Spinareoviridae''.


Therapeutic applications

Although reoviruses are mostly nonpathogenic in humans, these viruses have served as very productive experimental models for studies of viral pathogenesis. Newborn mice are extremely sensitive to reovirus infections and have been used as the preferred experimental system for studies of reovirus pathogenesis. Reoviruses have been demonstrated to have oncolytic (cancer-killing) properties, encouraging the development of reovirus-based therapies for cancer treatment. Reolysin is a formulation of reovirus ( Mammalian orthoreovirus serotype 3-dearing strain) that is currently in clinical trials for the treatment of various cancers, including studies currently developed to investigate the role of Reolysin combined with other immunotherapies.


See also

* Double-stranded RNA viruses * Oncolytic virus * Orphan virus


References


External links

{{Authority control Virus orders Riboviria