Nidovirales | |
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Life cycle of nidoviruses | |
Virus classification | |
(unranked): | Virus |
Realm: | Riboviria |
Kingdom: | Orthornavirae |
Phylum: | Pisuviricota |
Class: | Pisoniviricetes |
Order: | Nidovirales |
Nidovirales is an order of enveloped, positive-strand RNA viruses which infect vertebrates and invertebrates. Host organisms include mammals, birds, reptiles, amphibians, fish, arthropods, molluscs, and helminths. [1] The order includes the families Coronaviridae , Arteriviridae, Roniviridae, Tobaniviridae , and Mesoniviridae . [2]
Member viruses have a viral envelope and a positive-sense, single-stranded RNA genome which is capped and polyadenylated. [3] Nidoviruses are named for the Latin nidus, meaning nest, as all viruses in this order produce a 3' co-terminal nested set of subgenomic mRNAs during infection. [4]
Nidoviruses have a viral envelope and a positive-sense, single-stranded RNA genome which is capped and polyadenylated. [3] The group expresses structural proteins separately from the nonstructural ones. The structural proteins are encoded at the 3' region of the genome and are expressed from a set of subgenomic mRNAs.[ citation needed ]
Member viruses encode one main proteinase and between one and three accessory proteinases which are mainly involved in expressing the replicase gene. These proteinases are also responsible for activating or inactivating specific proteins at the correct time in the virus life cycle, ensuring replication occurs at the right time.[ citation needed ]
Nidoviruses can be distinguished from other RNA viruses by a constellation of seven conserved domains—5'-TM2-3CLpro-TM3-RdRp-Zm-HEL1-NendoU-3'—with the first three being encoded in ORF1a and the remaining four in ORF1b. TM2 and TM3 and transmembrane domains; RdRp is the RNA-dependent RNA polymerase; Zm is a Zn-cluster binding domain fused with a helicase (HEL1); 3CLpro is a 3C-like protease; and NendoU is an uridylate-specific endonuclease. The 3CLpro has a catalytic His-Cys dyad, and is related to the SARS coronavirus main proteinase (Mpro).[ citation needed ]
Most, but not all, nidovirus subgenomic RNAs contain a 5′ leader sequence derived from the 5′ end of the genomic RNA. The frameshift that generates ORF1b frameshift occurs at a UUUAAAC heptanucleotide 'slippery' sequence located upstream of the ORF1a stop codon and a putative RNA pseudoknot structure.[ citation needed ]
Many proteins have been identified on the genomes of Nidovirales, but their function has not yet been determined. Other enzymes that may be present in the genome include papain-like proteases, ADP-ribose/poly(ADP-ribose)-binding or ADP-ribose 1''-phosphate phosphatase activities and cyclic nucleotide phosphodiesterase.[ citation needed ]
The order Nidovirales can be divided into two clades depending on the size of the genome: those with large genomes (26.3–31.7 kilobases) which included the Coronaviridae and Roniviridae (the large nidoviruses) and those with small genomes (the small nidoviruses)—a clade that includes the distantly related Arteriviridae (12.7–15.7 kb).[ citation needed ]
The large nidoviruses encode both a 2'-O-methyltransferase and a 3'–5' exoribonuclease (ExoN)—the latter being very unusual for an RNA virus. They also encode a superfamily 1 helicase, uridylate-specific endonuclease (an enzyme unique to nidoviruses) and several proteases.[ citation needed ]
Nidoviruses as a group have the largest RNA genomes of viruses. Group member planarian secretory cell nidovirus (PSCNV) has the largest known nonsegmented RNA genome of 41.1kb. [6] Its host is the planarian flatworm. [7]
The following suborders and families are recognized (-virineae denotes suborders and -viridae denotes families): [8]
An RNA virus is a virus—other than a retrovirus—that has ribonucleic acid (RNA) as its genetic material. The nucleic acid is usually single-stranded RNA (ssRNA) but it may be double-stranded (dsRNA). Notable human diseases caused by RNA viruses include the common cold, influenza, SARS, MERS, COVID-19, Dengue virus, hepatitis C, hepatitis E, West Nile fever, Ebola virus disease, rabies, polio, mumps, and measles.
Virus classification is the process of naming viruses and placing them into a taxonomic system similar to the classification systems used for cellular organisms.
Coronaviruses are a group of related RNA viruses that cause diseases in mammals and birds. In humans and birds, they cause respiratory tract infections that can range from mild to lethal. Mild illnesses in humans include some cases of the common cold, while more lethal varieties can cause SARS, MERS and COVID-19. In cows and pigs they cause diarrhea, while in mice they cause hepatitis and encephalomyelitis.
Severe-acute-respiratory-syndrome–related coronavirus is a species of virus consisting of many known strains. Two strains of the virus have caused outbreaks of severe respiratory diseases in humans: severe acute respiratory syndrome coronavirus 1, which caused the 2002–2004 outbreak of severe acute respiratory syndrome (SARS), and severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), which is causing the ongoing pandemic of COVID-19. There are hundreds of other strains of SARSr-CoV, which are only known to infect non-human mammal species: bats are a major reservoir of many strains of SARSr-CoV; several strains have been identified in Himalayan palm civets, which were likely ancestors of SARS-CoV-1.
Rhabdoviridae is a family of negative-strand RNA viruses in the order Mononegavirales. Vertebrates, invertebrates, plants, fungi and protozoans serve as natural hosts. Diseases associated with member viruses include rabies encephalitis caused by the rabies virus, and flu-like symptoms in humans caused by vesiculoviruses. The name is derived from Ancient Greek rhabdos, meaning rod, referring to the shape of the viral particles. The family has 40 genera, most assigned to three subfamilies.
Coronaviridae is a family of enveloped, positive-strand RNA viruses which infect amphibians, birds, and mammals. The group includes the subfamilies Letovirinae and Orthocoronavirinae; the members of the latter are known as coronaviruses.
Arteriviridae is a family of enveloped, positive-strand RNA viruses in the order Nidovirales which infect vertebrates. Host organisms include equids, pigs, Possums, nonhuman primates, and rodents. The family includes, for example, equine arteritis virus in horses which causes mild-to-severe respiratory disease and reproductive failure, porcine reproductive and respiratory syndrome virus type 1 and type 2 in pigs which causes a similar disease, simian hemorrhagic fever virus which causes a highly lethal fever, lactate dehydrogenase–elevating virus which affects mice, and wobbly possum disease virus.
Torovirus is a genus of enveloped, positive-strand RNA viruses in the order Nidovirales and family Tobaniviridae. They primarily infect vertebrates, especially cattle, pigs, and horses. Diseases associated with this genus include gastroenteritis, which commonly presents in mammals. Torovirus is the only genus in the monotypic subfamily Torovirinae. Torovirus is also a monotypic taxon, containing only one subgenus, Renitovirus.
Picornavirales is an order of viruses with vertebrate, invertebrate, protist and plant hosts. The name has a dual etymology. First, picorna- is an acronym for poliovirus, insensitivity to ether, coxsackievirus, orphan virus, rhinovirus, and ribonucleic acid. Secondly, pico-, meaning extremely small, combines with RNA to describe these very small RNA viruses. The order comprises viruses that historically are referred to as picorna-like viruses.
The 3C-like protease (3CLpro) or main protease (Mpro), formally known as C30 endopeptidase or 3-chymotrypsin-like protease, is the main protease found in coronaviruses. It cleaves the coronavirus polyprotein at eleven conserved sites. It is a cysteine protease and a member of the PA clan of proteases. It has a cysteine-histidine catalytic dyad at its active site and cleaves a Gln–(Ser/Ala/Gly) peptide bond.
Mesoniviridae is a family of enveloped, positive-strand RNA viruses in the order Nidovirales which infect mosquitoes. The family is named after the size of the genomes relative to other nidoviruses, with meso- coming from the Greek word mesos, which means medium, and -ni being an abbreviation of nido.
Positive-strand RNA 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 into viral proteins by the host cell's ribosomes. Positive-strand RNA viruses encode an RNA-dependent RNA polymerase (RdRp) which is used during replication of the genome to synthesize a negative-sense antigenome that is then used as a template to create a new positive-sense viral genome.
Alphaabyssovirus is a genus of positive-strand RNA viruses in the order Nidovirales which infect sea hares. The genus is monotypic. It contains only the subgenus Aplyccavirus, which contains only one species, Aplysia abyssovirus 1. Alphaabyssovirus is also the only member of the subfamily Tiamatvirinae, which in turn is the only member of family Abyssoviridae, which likewise is the only member of the Abnidovirineae suborder. Aplysia abyssovirus 1 was first isolated from a sample from a California sea hare.
Alphamononivirus is a genus of enveloped, positive-strand RNA viruses in the order Nidovirales which infect planarian flatworms. Member virus planarian secretory cell nidovirus (PSCNV) has the largest known nonsegmented RNA genome of 41.1kb of any RNA virus. The genus is monotypic. It contains the subgenus Dumedivirus, which contains only one species, Planidovirus 1. Alphamononivirus is also the only member of the subfamily Mononivirinae, which in turn is the only member of family Mononiviridae, which likewise is the only member of the Monidovirineae suborder.
Orthornavirae is a kingdom of viruses that have genomes made of ribonucleic acid (RNA), including genes which encode 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 share a number of characteristics which promote rapid evolution, including high rates of genetic mutation, recombination, and reassortment.
Piscanivirinae is a virus subfamily of the family Tobaniviridae within the order Nidovirales which comprises different fish viruses. The virions have a viral envelope and a positive-sense single-stranded RNA genome which is linear and unsegmented.
ORF1ab refers collectively to two open reading frames (ORFs), ORF1a and ORF1b, that are conserved in the genomes of nidoviruses, a group of viruses that includes coronaviruses. The genes express large polyproteins that undergo proteolysis to form several nonstructural proteins with various functions in the viral life cycle, including proteases and the components of the replicase-transcriptase complex (RTC). Together the two ORFs are sometimes referred to as the replicase gene. They are related by a programmed ribosomal frameshift that allows the ribosome to continue translating past the stop codon at the end of ORF1a, in a -1 reading frame. The resulting polyproteins are known as pp1a and pp1ab.
Planarian secretory cell nidovirus (PSCNV) is a virus of the species Planidovirus 1, a nidovirus notable for its extremely large genome. At 41.1 kilobases, it is the largest known genome of an RNA virus. It was discovered by inspecting the transcriptomes of the planarian flatworm Schmidtea mediterranea and is the first known RNA virus infecting planarians. It was first described in 2018.
The nidoviral papain-like protease is a papain-like protease protein domain encoded in the genomes of nidoviruses. It is expressed as part of a large polyprotein from the ORF1a gene and has cysteine protease enzymatic activity responsible for proteolytic cleavage of some of the N-terminal viral nonstructural proteins within the polyprotein. A second protease also encoded by ORF1a, called the 3C-like protease or main protease, is responsible for the majority of further cleavages. Coronaviruses have one or two papain-like protease domains; in SARS-CoV and SARS-CoV-2, one PLPro domain is located in coronavirus nonstructural protein 3 (nsp3). Arteriviruses have two to three PLP domains. In addition to their protease activity, PLP domains function as deubiquitinating enzymes (DUBs) that can cleave the isopeptide bond found in ubiquitin chains. They are also "deISGylating" enzymes that remove the ubiquitin-like domain interferon-stimulated gene 15 (ISG15) from cellular proteins. These activities are likely responsible for antagonizing the activity of the host innate immune system. Because they are essential for viral replication, papain-like protease domains are considered drug targets for the development of antiviral drugs against human pathogens such as MERS-CoV, SARS-CoV, and SARS-CoV-2.
Nsp12 is a non-structural protein in the Coronavirus genome. Its gene is part of the ORF1ab reading frame and it is part of the pp1ab polyprotein; it is cleaved by 3CLpro.