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a group of viruses including those causing influenza
any of a group of RNA viruses including those that cause influenza and mumps

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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2014/07/28 16:04:34」(JST)

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The Orthomyxoviruses (orthos, Greek for "straight"; myxa, Greek for "mucus")^[1] are a family of RNA viruses that includes six genera: Influenzavirus A, Influenzavirus B, Influenzavirus C, Isavirus, Thogotovirus and Quaranjavirus. The first three genera contain viruses that cause influenza in vertebrates, including birds (see also avian influenza), humans, and other mammals. Isaviruses infect salmon; thogotoviruses infect vertebrates and invertebrates, such as mosquitoes and sea lice.^[2]^[3]^[4]

The three genera of Influenzavirus, which are identified by antigenic differences in their nucleoprotein and matrix protein, infect vertebrates as follows:

Influenzavirus A infects humans, other mammals, and birds, and causes all flu pandemics
Influenzavirus B infects humans and seals
Influenzavirus C infects humans, pigs and dogs.

Classification

In a phylogenetic-based taxonomy, the category "RNA virus" includes the category "negative-sense ssRNA virus", which includes the Order "Mononegavirales", and the Family "Orthomyxovirus" (among others). The genera-associated species and serotypes of Orthomyxovirus are shown in the following table.

Orthomyxovirus Genera, Species, and Serotypes
Genus	Species (* indicates type species)	Serotypes or Subtypes	Hosts
Influenzavirus A	Influenza A virus*	H1N1, H1N2, H2N2, H3N1, H3N2, H3N8, H5N1, H5N2, H5N3, H5N8, H5N9, H7N1, H7N2, H7N3, H7N4, H7N7, H7N9, H9N2, H10N7	Human, pig, bird, horse
Influenzavirus B	Influenza B virus*	Victoria, Yamagata^[5]	Human, seal
Influenzavirus C	Influenza C virus*		Human, pig, dog
Isavirus	Infectious Salmon Anemia Virus*		Atlantic salmon
Thogotovirus	Thogotovirus*		Tick, mosquito, mammal (including human)
Thogotovirus	Dhori Virus	Batken Virus, Dhori Virus, Jos Virus	Tick, mosquito, mammal (including human)
Quarja Viruses ^[6]
Quarja Viruses ^[6]	Quaranfil Virus, Johnston Atoll Virus

Types

There are three genera of influenza virus: Influenzavirus A, Influenzavirus B and Influenzavirus C. Each genus includes only one species, or type: Influenza A virus, Influenza B virus, and Influenza C virus, respectively. Influenza A and C infect multiple species, while influenza B almost exclusively infects humans.^[7]^[8]

Influenza A

Influenza A viruses are further classified, based on the viral surface proteins hemagglutinin (HA or H) and neuraminidase (NA or N). Sixteen H subtypes (or serotypes) and nine N subtypes of influenza A virus have been identified.

Diagram of influenza nomenclature

Further variation exists; thus, specific influenza strain isolates are identified by a standard nomenclature specifying virus type, geographical location where first isolated, sequential number of isolation, year of isolation, and HA and NA subtype.^[9]^[10]

Examples of the nomenclature are:

A/Brisbane/59/2007 (H1N1)
A/Moscow/10/99 (H3N2).

The type A viruses are the most virulent human pathogens among the three influenza types and cause the most severe disease. The serotypes that have been confirmed in humans, ordered by the number of known human pandemic deaths, are:

H1N1 caused "Spanish Flu" in 1918, "Swine flu" in 2009.^[11]
H2N2 caused "Asian Flu".
H3N2 caused "Hong Kong Flu".
H5N1 is a pandemic threat.
H7N7 has unusual zoonotic potential.^[12]
H1N2 is endemic in humans and pigs.
H9N2, H7N2, H7N3, H10N7.

Known flu pandemics^[13]^[14]^[15]
Name of pandemic	Date	Deaths	Case fatality rate	Subtype involved	Pandemic Severity Index
1889–1890 flu pandemic (Asiatic or Russian Flu)^[16]	1889–1890	1 million	0.15%	possibly H3N8 or H2N2	NA
1918 flu pandemic (Spanish flu)^[17]	1918–1920	20 to 100 million	2%	H1N1	5
Asian Flu	1957–1958	1 to 1.5 million	0.13%	H2N2	2
Hong Kong Flu	1968–1969	0.75 to 1 million	<0.1%	H3N2	2
Russian flu	1977–1978	no accurate count	N/A	H1N1	N/A
2009 flu pandemic^[18]^[19]	2009–2010	18,000	0.03%	H1N1	NA

Influenza B

Influenza B virus is almost exclusively a human pathogen, and is less common than influenza A. The only other animal known to be susceptible to influenza B infection is the seal.^[20] This type of influenza mutates at a rate 2–3 times lower than type A^[21] and consequently is less genetically diverse, with only one influenza B serotype.^[7] As a result of this lack of antigenic diversity, a degree of immunity to influenza B is usually acquired at an early age. However, influenza B mutates enough that lasting immunity is not possible.^[22] This reduced rate of antigenic change, combined with its limited host range (inhibiting cross species antigenic shift), ensures that pandemics of influenza B do not occur.^[23]

Influenza C

The influenza C virus infects humans and pigs, and can cause severe illness and local epidemics.^[24] However, influenza C is less common than the other types and usually seems to cause mild disease in children.^[25]^[26]

Virology

Morphology

Structure of the influenza virion. The hemagglutinin (HA) and neuraminidase (NA) proteins are shown on the surface of the particle. The viral RNAs that make up the genome are shown as red coils inside the particle and bound to Ribonuclear Proteins (RNPs).

The virion is pleomorphic; the envelope can occur in spherical and filamentous forms. In general, the virus's morphology is spherical with particles 50 to 120 nm in diameter, or filamentous virions 20 nm in diameter and 200 to 300 (–3000) nm long. There are some 500 distinct spike-like surface projections of the envelope each projecting 10 to 14 nm from the surface with some types (i.e. hemagglutinin esterase (HEF)) densely dispersed over the surface, and with others (i.e. hemagglutinin (HA)) spaced widely apart.

The major glycoprotein (HA) is interposed irregularly by clusters of neuraminidase (NA), with a ratio of HA to NA of about 4–5 to 1.

Lipoprotein membranes enclose the nucleocapsids; nucleoproteins of different size classes with a loop at each end; the arrangement within the virion is uncertain. The nucleocapsids are filamentous and fall in the range of 50 to 130 nm long and 9 to 15 nm in diameter. They have a helical symmetry.

Genome

Viruses of this family contain 6 to 8 segments of linear negative-sense single stranded RNA.^[27]

The total genome length is 12000–15000 nucleotides (nt). The largest segment 2300–2500 nt; of second largest 2300–2500 nt; of third 2200–2300 nt; of fourth 1700–1800 nt; of fifth 1500–1600 nt; of sixth 1400–1500 nt; of seventh 1000–1100 nt; of eighth 800–900 nt. Genome sequence has terminal repeated sequences; repeated at both ends. Terminal repeats at the 5'-end 12–13 nucleotides long. Nucleotide sequences of 3'-terminus identical; the same in genera of same family; most on RNA (segments), or on all RNA species. Terminal repeats at the 3'-end 9–11 nucleotides long. Encapsidated nucleic acid is solely genomic. Each virion may contain defective interfering copies.

Structure

The following applies for Influenza A viruses, although other influenza strains are very similar in structure:^[28]

The influenza A virus particle or virion is 80–120 nm in diameter and usually roughly spherical, although filamentous forms can occur.^[29] Unusually for a virus, the influenza A genome is not a single piece of nucleic acid; instead, it contains eight pieces of segmented negative-sense RNA (13.5 kilobases total), which encode 11 proteins (HA, NA, NP, M1, M2, NS1, NEP, PA, PB1, PB1-F2, PB2).^[30] The best-characterised of these viral proteins are hemagglutinin and neuraminidase, two large glycoproteins found on the outside of the viral particles. Neuraminidase is an enzyme involved in the release of progeny virus from infected cells, by cleaving sugars that bind the mature viral particles. By contrast, hemagglutinin is a lectin that mediates binding of the virus to target cells and entry of the viral genome into the target cell.^[31] The hemagglutinin (H) and neuraminidase (N) proteins are targets for antiviral drugs.^[32] These proteins are also recognised by antibodies, i.e. they are antigens.^[13] The responses of antibodies to these proteins are used to classify the different serotypes of influenza A viruses, hence the H and N in H5N1.

Replication cycle

Invasion and replication of the influenza virus. The steps in this process are discussed in the text.

Typically, influenza is transmitted from infected mammals through the air by coughs or sneezes, creating aerosols containing the virus, and from infected birds through their droppings. Influenza can also be transmitted by saliva, nasal secretions, feces and blood. Infections occur through contact with these bodily fluids or with contaminated surfaces. Flu viruses can remain infectious for about one week at human body temperature, over 30 days at 0 °C (32 °F), and indefinitely at very low temperatures (such as lakes in northeast Siberia). They can be inactivated easily by disinfectants and detergents.^[33]^[34]^[35]

The viruses bind to a cell through interactions between its hemagglutinin glycoprotein and sialic acid sugars on the surfaces of epithelial cells in the lung and throat (Stage 1 in infection figure).^[36] The cell imports the virus by endocytosis. In the acidic endosome, part of the haemagglutinin protein fuses the viral envelope with the vacuole's membrane, releasing the viral RNA (vRNA) molecules, accessory proteins and RNA-dependent RNA polymerase into the cytoplasm (Stage 2).^[37] These proteins and vRNA form a complex that is transported into the cell nucleus, where the RNA-dependent RNA polymerase begins transcribing complementary positive-sense cRNA (Steps 3a and b).^[38] The cRNA is either exported into the cytoplasm and translated (step 4), or remains in the nucleus. Newly synthesised viral proteins are either secreted through the Golgi apparatus onto the cell surface (in the case of neuraminidase and hemagglutinin, step 5b) or transported back into the nucleus to bind vRNA and form new viral genome particles (step 5a). Other viral proteins have multiple actions in the host cell, including degrading cellular mRNA and using the released nucleotides for vRNA synthesis and also inhibiting translation of host-cell mRNAs.^[39]

Negative-sense vRNAs that form the genomes of future viruses, RNA-dependent RNA transcriptase, and other viral proteins are assembled into a virion. Hemagglutinin and neuraminidase molecules cluster into a bulge in the cell membrane. The vRNA and viral core proteins leave the nucleus and enter this membrane protrusion (step 6). The mature virus buds off from the cell in a sphere of host phospholipid membrane, acquiring hemagglutinin and neuraminidase with this membrane coat (step 7).^[40] As before, the viruses adhere to the cell through hemagglutinin; the mature viruses detach once their neuraminidase has cleaved sialic acid residues from the host cell.^[36] After the release of new influenza virus, the host cell dies.

Since RNA proofreading enzymes are absent, the RNA-dependent RNA transcriptase makes a single nucleotide insertion error roughly every 10 thousand nucleotides, which is the approximate length of the influenza vRNA. Hence, nearly every newly manufactured influenza virus will contain a mutation in its genome.^[41] The separation of the genome into eight separate segments of vRNA allows mixing (reassortment) of the genes if more than one variety of influenza virus has infected the same cell (superinfection). The resulting alteration in the genome segments packaged into viral progeny confers new behavior, sometimes the ability to infect new host species or to overcome protective immunity of host populations to its old genome (in which case it is called an antigenic shift).^[13]

Viability and disinfection

Mammalian influenza viruses tend to be labile, but can survive several hours in mucus.^[42] Avian influenza virus can survive for 100 days in distilled water at room temperature, and 200 days at 17 °C (63 °F). The avian virus is inactivated more quickly in manure, but can survive for up to 2 weeks in feces on cages. Avian influenza viruses can survive indefinitely when frozen.^[42] Influenza viruses are susceptible to bleach, 70% ethanol, aldehydes, oxidizing agents, and quaternary ammonium compounds. They are inactivated by heat of 133 °F (56 °C) for minimum of 60 minutes, as well as by low pH <2.^[42]

Vaccination and prophylaxis

Vaccines and drugs are available for the prophylaxis and treatment of influenza virus infections. Vaccines are composed of either inactivated or live attenuated virions of the H1N1 and H3N2 human influenza A viruses, as well as those of influenza B viruses. Because the antigenicities of the wild viruses evolve, vaccines are reformulated annually by updating the seed strains. However, when the antigenicities of the seed strains and wild viruses do not match, vaccines fail to protect the vaccinees. In addition, even when they do match, escape mutants are often generated. Drugs available for the treatment of influenza include amantadine, which inhibits the uncoating of virions by interfering with M2, and oseltamivir (marketed under the brand name Tamiflu), which inhibits the release of virions from infected cells by interfering with NA. However, escape mutants are often generated for the former drug and less frequently for the latter drug.^[43]

References

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^ Jones LD, Nuttall PA (1989). "Non-viraemic transmission of Thogoto virus: influence of time and distance". Trans. R. Soc. Trop. Med. Hyg. 83 (5): 712–4. doi:10.1016/0035-9203(89)90405-7. PMID 2617637.
^ Barry Ely (1999). "Infectious Salmon Anaemia". Mill Hill Essays. National Institute for Medical Research. Archived from the original on 2007-08-24. Retrieved 2007-09-14.
^ Raynard RS, Murray AG, Gregory A (2001). "Infectious salmon anaemia virus in wild fish from Scotland". Dis. Aquat. Org. 46 (2): 93–100. doi:10.3354/dao046093. PMID 11678233.
^ Differentiation of Influenza B Virus Lineages Yamagata and Victoria by Real-Time PCR, in: Journal of Clinical Microbiology, Jan. 2013, Vol. 51, Issue 1, by B. Biere, B. Bauer, B. Schweiger
^ ICTV Taxonomy History, ICTV, 2014, retrieved 6 June 2006
^ ^a ^b Hay A, Gregory V, Douglas A, Lin Y (Dec 29, 2001). "The evolution of human influenza viruses" (PDF). Philos Trans R Soc Lond B Biol Sci 356 (1416): 1861–70. doi:10.1098/rstb.2001.0999. PMC 1088562. PMID 11779385.
^ "Avian Influenza (Bird Flu)". Centers for Disease Control and Prevention. Retrieved 2007-09-15.
^ Atkinson W, Hamborsky J, McIntyre L, Wolfe S, ed. (2007). Epidemiology and Prevention of Vaccine-Preventable Diseases (10th ed.). Washington DC: Centers for Disease Control and Prevention.
^ "Avian Influenza (Bird Flu): Implications for Human Disease". Center for Infectious Disease Research & Policy, University of Minnesota. 2007-06-27. Retrieved 2007-09-14.
^ Wang TT, Palese P (June 2009). "Unraveling the Mystery of Swine Influenza Virus". Cell 137 (6): 983–5. doi:10.1016/j.cell.2009.05.032. PMID 19524497.
^ Fouchier R, Schneeberger P, Rozendaal F, Broekman J, Kemink S, Munster V, Kuiken T, Rimmelzwaan G, Schutten M, Van Doornum G, Koch G, Bosman A, Koopmans M, Osterhaus A (2004). "Avian influenza A virus (H7N7) associated with human conjunctivitis and a fatal case of acute respiratory distress syndrome". Proc Natl Acad Sci USA 101 (5): 1356–61. Bibcode:2004PNAS..101.1356F. doi:10.1073/pnas.0308352100. PMC 337057. PMID 14745020.
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^ "ECDC Daily Update – Pandemic (H1N1) 2009 – January 18, 2010". European Centre for Disease Prevention and Control. 2010-01-18. Retrieved 2010-01-18.
^ Osterhaus A, Rimmelzwaan G, Martina B, Bestebroer T, Fouchier R (2000). "Influenza B virus in seals". Science 288 (5468): 1051–3. Bibcode:2000Sci...288.1051O. doi:10.1126/science.288.5468.1051. PMID 10807575.
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^ http://www.ictvdb.org/Ictv/index.htm
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^ Cros J, Palese P (September 2003). "Trafficking of viral genomic RNA into and out of the nucleus: influenza, Thogoto and Borna disease viruses". Virus Res 95 (1–2): 3–12. doi:10.1016/S0168-1702(03)00159-X. PMID 12921991.
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^ Drake J (May 1, 1993). "Rates of spontaneous mutation among RNA viruses". Proc Natl Acad Sci USA 90 (9): 4171–5. Bibcode:1993PNAS...90.4171D. doi:10.1073/pnas.90.9.4171. PMC 46468. PMID 8387212.
^ ^a ^b ^c http://www.cfsph.iastate.edu/Factsheets/pdfs/influenza.pdf, p. 7
^ Yoshiyuki Suzuki (July 3, 2006). "Natural selection on the influenza virus genome". Molecular Biology and Evolution 23 (10): 1902–1911. doi:10.1093/molbev/msl050. PMID 16818477.

Hoyle, L. (1969). "The Influenza Viruses". Virology Monographs (Springer-Verlag) 4. ISBN 3-211-80892-2. ISSN 0083-6591. OCLC 4053391.

External links

Wikispecies has information related to: Orthomyxoviridae

Health-EU Portal EU work to prepare a global response to influenza.
Influenza Research Database Database of influenza genomic sequences and related information.
European Commission—Public Health EU coordination on Pandemic (H1N1) 2009
3D Influenza-virus-related structures from the EM Data Bank(EMDB)

English Journal

Long noncoding RNAs (lncRNAs) dynamics evidence immunomodulation during ISAV-Infected Atlantic salmon (Salmo salar).

Boltaña S1, Valenzuela-Miranda D1, Aguilar A1, Mackenzie S2, Gallardo-Escárate C1.
Scientific reports.Sci Rep.2016 Mar 4;6:22698. doi: 10.1038/srep22698.
Despite evidence for participation in the host response to infection, the roles of many long non-coding RNAs (lncRNAs) remain unknown. Therefore, the aims of this study were to identify lncRNAs in Atlantic salmon (Salmo salar) and evaluate their transcriptomic regulation during ISA virus (ISAV) infe
PMID 26939752

Influenza A virus infection in dogs: Epizootiology, evolution and prevention - A review.

Xie X1, Ma K1, Liu Y1.
Acta veterinaria Hungarica.Acta Vet Hung.2016 Mar;64(1):125-39. doi: 10.1556/004.2016.014.
Canine influenza virus (CIV) is an enveloped virus belonging to the genus Influenza virus A within the family Orthomyxoviridae. Prior to 2004, only sporadic outbreaks of canine influenza had been observed in dog populations around the world. However, in 2004 an H3N8 influenza virus of equine origin
PMID 26919150

Infectious Salmon Anaemia Virus (ISAV) RNA Binding Protein Encoded by Segment 8 ORF2 and Its Interaction with ISAV and Intracellular Proteins.

Olsen CM1, Markussen T2, Thiede B3, Rimstad E4.
Viruses.Viruses.2016 Feb 18;8(2). pii: E52. doi: 10.3390/v8020052.
Infectious salmon anaemia virus (ISAV) is an orthomyxovirus infecting salmonid fish. The virus is adapted to low temperature and has a replication optimum between 10-15 °C. In this study the subcellular localization and protein interactions for the protein encoded by the largest open reading frame
PMID 26901217

Japanese Journal

昨今のインフルエンザの流行動向とこれからの治療展望 (特集これからのインフルエンザ診療 : 的確な診断・適正な治療・先手の予防)

感染と抗菌薬 16(3), 201-206, 2013-09
NAID 40019810643

Generation of congenic mouse strains by introducing the virus-resistant genes, Mx1 and Oas1b, of feral mouse-derived inbred strain MSM/Ms into the common strain C57BL/6J

Japanese Journal of Veterinary Research 57(2), 89-99, 2009-08
NAID 120001596861

Detection of type B influenza virus genes from biopsied gastric mucosa

Journal of gastroenterology 37(2), 101-105, 2002-02-01
NAID 10010843639

Related Pictures

★リンクテーブル★

リンク元	「オルソミクソウイルス」「myxovirus」

「オルソミクソウイルス」

Orthomyxovirus
Virus classification
Group:	Group V ((-)ssRNA)
Order:	Unassigned
Family:	Orthomyxoviridae
Genera
	Influenzavirus A; Influenzavirus B; Influenzavirus C; Isavirus; Quaranjavirus; Thogotovirus;

　　[★]

英: orthomyxovirus
関: ミクソウイルス、インフルエンザウイルス、オルトミクソウイルス科

「myxovirus」

　　[★]

ミクソウイルス

関: influenza virus、Orthomyxoviridae、orthomyxovirus

[1] International Committee on Taxonomy of Viruses Index of Viruses — Orthomyxovirus (2006). In: ICTVdB—The Universal Virus Database, version 4. Büchen-Osmond, C (Ed), Columbia University, New York, USA.

[pmid2617637-2] Jones LD, Nuttall PA (1989). "Non-viraemic transmission of Thogoto virus: influence of time and distance". Trans. R. Soc. Trop. Med. Hyg. 83 (5): 712–4. doi:10.1016/0035-9203(89)90405-7. PMID 2617637.

[3] Barry Ely (1999). "Infectious Salmon Anaemia". Mill Hill Essays. National Institute for Medical Research. Archived from the original on 2007-08-24. Retrieved 2007-09-14.

[pmid11678233-4] Raynard RS, Murray AG, Gregory A (2001). "Infectious salmon anaemia virus in wild fish from Scotland". Dis. Aquat. Org. 46 (2): 93–100. doi:10.3354/dao046093. PMID 11678233.

[5] Differentiation of Influenza B Virus Lineages Yamagata and Victoria by Real-Time PCR, in: Journal of Clinical Microbiology, Jan. 2013, Vol. 51, Issue 1, by B. Biere, B. Bauer, B. Schweiger

[ICTV_Taxonomy_History-6] ICTV Taxonomy History, ICTV, 2014, retrieved 6 June 2006

[hay-7] Hay A, Gregory V, Douglas A, Lin Y (Dec 29, 2001). "The evolution of human influenza viruses" (PDF). Philos Trans R Soc Lond B Biol Sci 356 (1416): 1861–70. doi:10.1098/rstb.2001.0999. PMC 1088562. PMID 11779385.

[8] "Avian Influenza (Bird Flu)". Centers for Disease Control and Prevention. Retrieved 2007-09-15.

[9] Atkinson W, Hamborsky J, McIntyre L, Wolfe S, ed. (2007). Epidemiology and Prevention of Vaccine-Preventable Diseases (10th ed.). Washington DC: Centers for Disease Control and Prevention.

[10] "Avian Influenza (Bird Flu): Implications for Human Disease". Center for Infectious Disease Research & Policy, University of Minnesota. 2007-06-27. Retrieved 2007-09-14.

[11] Wang TT, Palese P (June 2009). "Unraveling the Mystery of Swine Influenza Virus". Cell 137 (6): 983–5. doi:10.1016/j.cell.2009.05.032. PMID 19524497.

[12] Fouchier R, Schneeberger P, Rozendaal F, Broekman J, Kemink S, Munster V, Kuiken T, Rimmelzwaan G, Schutten M, Van Doornum G, Koch G, Bosman A, Koopmans M, Osterhaus A (2004). "Avian influenza A virus (H7N7) associated with human conjunctivitis and a fatal case of acute respiratory distress syndrome". Proc Natl Acad Sci USA 101 (5): 1356–61. Bibcode:2004PNAS..101.1356F. doi:10.1073/pnas.0308352100. PMC 337057. PMID 14745020.

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