Mononegavirales

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Order Mononegavirales
Virus classification
Group: Group V ((-)ssRNA)
Order: Mononegavirales
Families

The order Mononegavirales is the taxonomic home of numerous related viruses. Members of the order that are commonly known are, for instance, Ebola virus, human respiratory syncytial virus, measles virus, mumps virus, Nipah virus, and rabies virus. All of these viruses cause significant disease in humans. Many very important pathogens of nonhuman animals and plants are also members of this order.

Use of term[edit]

The order Mononegavirales is a virological taxon that was created in 1991[1][2] and emended in 1995,[3] 1997,[4] 2000,[5] 2005,[6] 2011,[7] and 2016.[8] The name Mononegavirales is derived from the Greek adjective μóνος [monos] (alluding to the monopartite and single-stranded genomes of most mononegaviruses), the Latin verb negare (alluding to the negative polarity of these genomes), and the taxonomic suffix -virales (denoting a viral order).[9] The order currently includes the seven virus families Bornaviridae, Mymonaviridae, Filoviridae, Nyamiviridae, Paramyxoviridae, Pneumoviridae, Rhabdoviridae, and Sunviridae.[7][9][10][11][12]

Note[edit]

Mononegavirales is pronounced ˌmɒnəˌnɛgəviː’rɑ:lɨz (IPA) or mo-nuh-ne-guh-vee-rah-liz in English phonetic notation.[9] According to the rules for taxon naming established by the International Committee on Taxonomy of Viruses (ICTV), the name Mononegavirales is always to be capitalized, italicized, and never abbreviated.[7] The names of the order's physical members (mononegaviruses/mononegavirads) are to be written in lower case, are not italicized, and used without articles.[7][9]

Order inclusion criteria[edit]

A virus is a member of the order Mononegavirales if[7][9]

  • its genome is a linear, typically (but not always) nonsegmented, single-stranded, non-infectious RNA of negative polarity; possesses inverse-complementary 3' and 5' termini; is not covalently linked to a protein;
  • its genome has the characteristic gene order 3'-UTR–core protein genes–envelope protein genes–RNA-dependent RNA polymerase gene–5'-UTR;
  • it produces 5–10 distinct mRNAs from its genome via polar sequential transcription from a single promoter located at the 3' end of the genome; mRNAs are 5' capped and polyadenylated;
  • it replicates by synthesizing complete antigenomes;
  • it forms infectious helical ribonucleocapsids as the templates for the synthesis of mRNAs, antigenomes, and genomes;
  • it encodes an RNA-dependent RNA polymerase (RdRp) that is highly homologous to those of other mononegaviruses; and/or
  • it forms enveloped virions with a molecular mass of 300–1,000×106; an S20W of 550–>1,045; and a buoyant density in CsCl of 1.18–1.22 g/cm3.

Order organization[edit]

The order includes seven accepted families that include numerous genera, consisting of many different species. The order has expanded considerably during recent years due to the discovery of many novel agents that were found to be phylogenetically diverse from already known mononegaviruses. Novel taxa (genera and/or species) had to be proposed, some of which have by now been accepted by the ICTV, and others that are in various stages of the suggestion/proposal/consideration process. The table below provides an overview of the current ICTV-approved composition of the order.

Order Mononegavirales: families, genera, species, and their viruses
Family Genus Species Virus (Abbreviation)
Bornaviridae Bornavirus Elapid 1 bornavirus Loveridge’s garter snake virus 1 (LGSV-1)
Mammalian 1 bornavirus* Borna disease virus 1 (BoDV-1)
Borna disease virus 2 (BoDV-2)
Passeriform 1 bornavirus canary bornavirus 1 (CnBV-1)
canary bornavirus 2 (CnBV-2)
canary bornavirus 3 (CnBV-3))
Passeriform 2 bornavirus estrildid finch bornavirus 1 (EsBV-1)
Psittaciform 1 bornavirus parrot bornavirus 1 (PaBV-1)
parrot bornavirus 2 (PaBV-2)
parrot bornavirus 3 (PaBV-3)
parrot bornavirus 4 (PaBV-4)
parrot bornavirus 7 (PaBV-7)
Psittaciform 2 bornavirus parrot bornavirus 5 (PaBV-5)
Waterbird 1 bornavirus aquatic bird bornavirus 1 (ABBV-1)
aquatic bird bornavirus 2 (ABBV-2)
Filoviridae Cuevavirus Lloviu cuevavirus* Lloviu virus (LLOV)
Ebolavirus Bundibugyo ebolavirus Bundibugyo virus (BDBV)
Reston ebolavirus Reston virus (RESTV)
Sudan ebolavirus Sudan virus (SUDV)
Taï Forest ebolavirus Taï Forest virus (TAFV)
Zaire ebolavirus* Ebola virus (EBOV)
Marburgvirus Marburg marburgvirus* Marburg virus (MARV)
Ravn virus (RAVV)
Mymonaviridae Sclerotimonavirus Sclerotinia sclerotimonavirus* Sclerotinia sclerotiorum negative-stranded RNA virus 1 (SsNSRV-1)
Nyamiviridae Nyavirus Midway nyavirus Midway virus (MIDWV)
Nyamanini nyavirus* Nyamanini virus (NYMV)
Sierra Nevada nyavirus Sierra Nevada virus (SNVV)
Socyvirus Soybean cyst nematode socyvirus* soybean cyst nematode virus 1 (SbCNV-1)
Paramyxoviridae Aquaparamyxovirus Atlantic salmon paramyxovirus* Atlantic salmon paramyxovirus (AsaPV)
Avulavirus Avian paramyxovirus 2 avian paramyxovirus 2 (APMV-2)
Avian paramyxovirus 3 avian paramyxovirus 3 (APMV-3)
Avian paramyxovirus 4 avian paramyxovirus 4 (APMV-4)
Avian paramyxovirus 5 avian paramyxovirus 5 (APMV-5)
Avian paramyxovirus 6 avian paramyxovirus 6 (APMV-6)
Avian paramyxovirus 7 avian paramyxovirus 7 (APMV-7)
Avian paramyxovirus 8 avian paramyxovirus 8 (APMV-8)
Avian paramyxovirus 9 avian paramyxovirus 9 (APMV-9)
Avian paramyxovirus 10 avian paramyxovirus 10 (APMV-10)
Avian paramyxovirus 11 avian paramyxovirus 11 (APMV-11)
Avian paramyxovirus 12 avian paramyxovirus 12 (APMV-12)
Newcastle disease virus* avian paramyxovirus 1 (APMV-1)
Ferlavirus Fer-de-Lance paramyxovirus* Fer-de-Lance virus (FDLV)
Henipavirus Cedar henipavirus Cedar virus (CedV)
Ghanaian bat henipavirus Kumasi virus (KV)
Hendra virus* Hendra virus (HeV)
Mojiang henipavirus Mòjiāng virus (MojV)
Nipah virus Nipah virus (NiV)
Morbillivirus Canine distemper virus canine distemper virus (CDV)
Cetacean morbillivirus cetacean morbillivirus (CeMV)
Feline morbillivirus feline morbillivirus (FeMV)
Measles virus* measles virus (MeV)
Peste-des-petits-ruminants virus peste-des-petits-ruminants virus (PPRV)
Phocine distemper virus phocine distemper virus (PDV)
Rinderpest virus rinderpest virus (RPV)
Respirovirus Bovine parainfluenza virus 3 bovine parainfluenza virus 3 (BPIV-3)
Human parainfluenza virus 1 human parainfluenza virus 1 (HPIV-1)
Human parainfluenza virus 3 human parainfluenza virus 3 (HPIV-3)
Porcine parainfluenzavirus 1 porcine parainfluenza virus 1 (PPIV-1)
Sendai virus* Sendai virus (SeV)
Rubulavirus Human parainfluenza virus 2 human parainfluenza virus 2 (HPIV-2)
Human parainfluenza virus 4 human parainfluenza virus 4a (HPIV-4a)
human parainfluenza virus 4b (HPIV-4b)
Mapuera virus Mapuera virus (MapV)
Mumps virus* mumps virus (MuV)
bat mumps virus (BMV)
Parainfluenza virus 5 parainfluenza virus 5 (PIV-5)
Porcine rubulavirus La Piedad Michoacán Mexico virus (LPMV)
Simian virus 41 simian virus 41 (SV-41)
Pneumoviridae Metapneumovirus Avian metapneumovirus* avian metapneumovirus (AMPV)
Human metapneumovirus human metapneumovirus (HMPV)
Orthopneumovirus Bovine respiratory syncytial virus bovine respiratory syncytial virus (BRSV)
Human respiratory syncytial virus* human respiratory syncytial virus A2 (HRSV-A2)
human respiratory syncytial virus B1 (HRSV-B1)
human respiratory syncytial virus S2 (HRSV-S2)
Murine pneumonia virus murine pneumonia virus (MPV)
Rhabdoviridae Cytorhabdovirus Alfalfa dwarf cytorhabdovirus alfalfa dwarf virus (ADV)
Barley yellow striate mosaic cytorhabdovirus barley yellow striate mosaic virus (BYSMV)
Broccoli necrotic yellows cytorhabdovirus broccoli necrotic yellows virus (BNYV)
Festuca leaf streak cytorhabdovirus festuca leaf streak virus (FLSV)
Lettuce necrotic yellows cytorhabdovirus* lettuce necrotic yellows virus (LNYV)
Lettuce yellow mottle cytorhabdovirus lettuce yellow mottle virus (LYMoV)
Northern cereal mosaic cytorhabdovirus northern cereal mosaic virus (NCMV)
Sonchus cytorhabdovirus sonchus virus (SonV)
Strawberry crinkle cytorhabdovirus strawberry crinkle virus (SCV)
Wheat American striate mosaic cytorhabdovirus wheat American striate mosaic virus (WASMV)
Dichorhavirus Coffee ringspot dichorhavirus coffee ringspot virus (CoRSV)
Orchid fleck dichorhavirus* orchid fleck virus (OFV)
Ephemerovirus Adelaide River ephemerovirus Adelaide River virus (ARV)
Berrimah ephemerovirus Berrimah virus (BRMV)
Bovine fever ephemerovirus* bovine ephemeral fever virus (BEFV)
Kotonkan ephemerovirus kotonkan virus (KOTV)
Obodhiang ephemerovirus Obodhiang virus (OBOV)
Lyssavirus Aravan lyssavirus Aravan virus (ARAV)
Australian bat lyssavirus Australian bat lyssavirus (ABLV)
Bokeloh bat lyssavirus Bokeloh bat lyssavirus (BBLV)
Duvenhage lyssavirus Duvenhage virus (DUVV)
European bat 1 lyssavirus European bat lyssavirus 1 (EBLV-1)
European bat 2 lyssavirus European bat lyssavirus 2 (EBLV-2)
Ikoma lyssavirus Ikoma lyssavirus (IKOV)
Irkut lyssavirus Irkut virus (IRKV)
Khujand lyssavirus Khujand virus (KHUV)
Lagos bat lyssavirus Lagos bat virus (LBV)
Mokola lyssavirus Mokola virus (MOKV)
Rabies lyssavirus* rabies virus (RABV)
Shimoni bat lyssavirus Shimoni bat virus (SHIBV)
West Caucasian bat lyssavirus West Caucasian bat virus (WCBV)
Novirhabdovirus Hirame novirhabdovirus Hirame rhabdovirus (HIRV)
Oncorhynchus 1 novirhabdovirus* infectious hematopoietic necrosis virus (IHNV)
Oncorhynchus 2 novirhabdovirus viral hemorrhagic septicemia virus (VHSV)
Snakehead novirhabdovirus snakehead rhabdovirus (SHRV)
Nucleorhabdovirus Datura yellow vein nucleorhabdovirus datura yellow vein virus (DYVV)
Eggplant mottled dwarf nucleorhabdovirus eggplant mottled dwarf virus (EMDV)
Maize fine streak nucleorhabdovirus maize fine streak virus (MSFV)
Maize Iranian mosaic nucleorhabdovirus maize Iranian mosaic virus (MIMV)
Maize mosaic nucleorhabdovirus maize mosaic virus (MMV)
Potato yellow dwarf nucleorhabdovirus* potato yellow dwarf virus (PYDV)
Rice yellow stunt nucleorhabdovirus rice yellow stunt virus (RYSV)
rice transitory yellowing virus (RTYV)
Sonchus yellow net nucleorhabdovirus sonchus yellow net virus (SYNV)
Sowthistle yellow vein nucleorhabdovirus sowthistle yellow vein virus (SYVV)
Taro vein chlorosis nucleorhabdovirus taro vein chlorosis virus (TaVCV)
Perhabdovirus Anguillid perhabdovirus eel virus European X (EVEX)
Perch perhabdovirus* perch rhabdovirus (PRV)
Sea trout perhabdovirus lake trout rhabdovirus (LTRV)
Sigmavirus Drosophila affinis sigmavirus Drosophila affinis sigmavirus (DAffSV)
Drosophila ananassae sigmavirus Drosophila ananassae sigmavirus (DAnaSV)
Drosophila immigrans sigmavirus Drosophila immigrans sigmavirus (DImmSV)
Drosophila melanogaster sigmavirus* Drosophila melanogaster sigmavirus (DMelSV)
Drosophila obscura sigmavirus Drosophila obscura sigmavirus (DObsSV)
Drosophila tristis sigmavirus Drosophila tristis sigmavirus (DTriSV)
Muscina stabulans sigmavirus Muscina stabulans sigmavirus (MStaSV)
Sprivivirus Carp sprivivirus* spring viremia of carp virus (SVCV)
Pike fry sprivivirus grass carp rhabdovirus (GrCRV)
pike fry rhabdovirus (PFRV)
Tench rhabdovirus (TenRV)
Tibrovirus Coastal Plains tibrovirus Coastal Plains virus (CPV)
Tibrogargan tibrovirus* Bivens Arm virus (BAV)
Tibrogargan virus (TIBV)
Tupavirus Durham tupavirus* Durham virus (DURV)
Tupaia tupavirus tupaia virus (TUPV)
Varicosavirus Lettuce big-vein associated varicosavirus* lettuce big-vein associated virus (LBVaV)
Vesiculovirus Alagoas vesiculovirus vesicular stomatitis Alagoas virus (VSAV)
Carajas vesiculovirus Carajás virus (CJSV)
Chandipura vesiculovirus Chandipura virus (CHPV)
Cocal vesiculovirus Cocal virus (COCV)
Indiana vesiculovirus* vesicular stomatitis Indiana virus (VSIV)
Isfahan vesiculovirus Isfahan virus (ISFV)
Maraba vesiculovirus Maraba virus (MARAV)
New Jersey vesiculovirus vesicular stomatitis New Jersey virus (VSNJV)
Piry vesiculovirus Piry virus (PIRYV)
Unassigned Flanders virus Flanders virus (FLAV)
Ngaingan virus Ngaingan virus (NGAV)
Wongabel virus Wongabel virus (WONV)
Sunviridae Sunshinevirus Reptile sunshinevirus 1* Sunshine Coast virus (SunCV)
Unassigned Anphevirus Xincheng anphevirus* Xīnchéng mosquito virus (XcMV)
Unassigned Arlivirus Lishi arlivirus* Líshí spider virus 2 (LsSV-2)
Unassigned Chengtivirus Tacheng chengtivirus* Tǎchéng tick virus 6 (TcTV-6)
Unassigned Crustavirus Wenzhou crustavirus* Wēnzhōu crab virus 1 (WzCV-1)
Unassigned Wastrivirus Sanxia wastrivirus* Sānxiá water strider virus 4 (SxWSV-4)

Table legend: "*" denotes type species.

Life cycle[edit]

The mononegavirus life cycle begins with virion attachment to specific cell-surface receptors, followed by fusion of the virion envelope with cellular membranes and the concomitant release of the virus nucleocapsid into the cytosol. The virus RdRp partially uncoats the nucleocapsid and transcribes the genes into positive-stranded mRNAs, which are then translated into structural and nonstructural proteins. Mononegavirus RdRps bind to a single promoter located at the 3' end of the genome. Transcription either terminates after a gene or continues to the next gene downstream. This means that genes close to the 3' end of the genome are transcribed in the greatest abundance, whereas those toward the 5' end are least likely to be transcribed. The gene order is therefore a simple but effective form of transcriptional regulation. The most abundant protein produced is the nucleoprotein, whose concentration in the cell determines when the RdRp switches from gene transcription to genome replication. Replication results in full-length, positive-stranded antigenomes that are in turn transcribed into negative-stranded virus progeny genome copies. Newly synthesized structural proteins and genomes self-assemble and accumulate near the inside of the cell membrane. Virions bud off from the cell, gaining their envelopes from the cellular membrane they bud from. The mature progeny particles then infect other cells to repeat the cycle.[6]

Paleovirology[edit]

Mononegaviruses have a history that dates back several tens of million of years. Mononegavirus "fossils" have been discovered in the form of mononegavirus genes or gene fragments integrated into mammalian genomes. For instance, bornavirus gene "fossils" have been detected in the genomes of bats, fish, hyraxes, marsupials, primates, rodents, ruminants, and elephants.[13][14][15] Filovirus gene "fossils" have been detected in the genomes of bats, rodents, shrews, tenrecs, and marsupials.[14][15][16] A Midway virus "fossil" was found in the genome of zebrafish.[14] Finally, rhabdovirus "fossils" were found in the genomes of mosquitoes, ticks, and plants.[15][17][18]

References[edit]

  1. ^ "The order Mononegavirales". Archives of Virology. 117 (1–2): 137–140. 1991. PMID 2006902. 
  2. ^ Pringle, C. R. (1991), "Order Mononegavirales", in Francki, R. I. B.; Fauquet, C. M.; Knudson, D. L.; et al., Classification and Nomenclature of Viruses-Fifth Report of the International Committee on Taxonomy of Viruses. Archives of Virology Supplement, vol. 2, Vienna, Austria: Springer, pp. 239–41, ISBN 0-387-82286-0 
  3. ^ Bishop, D. H. L.; Pringle, C. R. (1995), "Order Mononegavirales", in Murphy, F. A.; Fauquet, C. M.; Bishop, D. H. L.; Ghabrial, S. A.; Jarvis, A. W.; Martelli, G. P.; Mayo, M. A.; Summers, M. D., Virus Taxonomy—Sixth Report of the International Committee on Taxonomy of Viruses. Archives of Virology Supplement, vol. 10, Vienna, Austria: Springer, pp. 265–267, ISBN 3-211-82594-0 
  4. ^ Pringle, C. R. (1997). "The order Mononegavirales--current status". Archives of Virology. 142 (11): 2321–2326. PMID 9672597. 
  5. ^ Pringle, C. R. (2000), "Order Mononegavirales", in van Regenmortel, M. H. V.; Fauquet, C. M.; Bishop, D. H. L.; Carstens, E. B.; Estes, M. K.; Lemon, S. M.; Maniloff, J.; Mayo, M. A.; McGeoch, D. J.; Pringle, C. R.; Wickner, R. B., Virus Taxonomy—Seventh Report of the International Committee on Taxonomy of Viruses, San Diego, USA: Academic Press, pp. 525–530, ISBN 0-12-370200-3 
  6. ^ a b Pringle, C. R. (2005), "Order Mononegavirales", in Fauquet, C. M.; Mayo, M. A.; Maniloff, J.; Desselberger, U.; Ball, L. A., Virus Taxonomy—Eighth Report of the International Committee on Taxonomy of Viruses, San Diego, USA: Elsevier/Academic Press, pp. 609–614, ISBN 0-12-370200-3 
  7. ^ a b c d e Easton, A.; Pringle, C. R. (2011), "Order Mononegavirales", in King, Andrew M. Q.; Adams, Michael J.; Carstens, Eric B.; et al., Virus Taxonomy—Ninth Report of the International Committee on Taxonomy of Viruses, London, UK: Elsevier/Academic Press, pp. 653–657, ISBN 978-0-12-384684-6 
  8. ^ Afonso, Claudio L.; Amarasinghe, Gaya K.; Bányai, Krisztián; Bào, Yīmíng; Basler, Christopher F.; Bavari, Sina; Bejerman, Nicolás; Blasdell, Kim R.; Briand, François-Xavier (2016-08-01). "Taxonomy of the order Mononegavirales: update 2016". Archives of Virology. 161 (8): 2351–2360. doi:10.1007/s00705-016-2880-1. ISSN 1432-8798. PMC 4947412free to read. PMID 27216929. 
  9. ^ a b c d e Kuhn, J. H.; Becker, S.; Ebihara, H.; Geisbert, T. W.; Johnson, K. M.; Kawaoka, Y.; Lipkin, W. I.; Negredo, A. I.; Netesov, S. V.; Nichol, S. T.; Palacios, G.; Peters, C. J.; Tenorio, A.; Volchkov, V. E.; Jahrling, P. B. (2010). "Proposal for a revised taxonomy of the family Filoviridae: Classification, names of taxa and viruses, and virus abbreviations". Archives of Virology. 155 (12): 2083–2103. doi:10.1007/s00705-010-0814-x. PMC 3074192free to read. PMID 21046175. 
  10. ^ Vetten, H. J.; Haenni, A. -L. (2006). "Taxon-specific suffixes for vernacular names". Archives of Virology. 151 (6): 1249–1250. doi:10.1007/s00705-006-0743-x. PMID 16721512. 
  11. ^ Kuhn, J. H.; Bekal, S.; Caì, Y. N. N.; Clawson, A. N.; Domier, L. L.; Herrel, M.; Jahrling, P. B.; Kondo, H.; Lambert, K. N.; Mihindukulasuriya, K. A.; Nowotny, N.; Radoshitzky, S. R.; Schneider, U.; Staeheli, P.; Suzuki, N.; Tesh, R. B.; Wang, D.; Wang, L. F.; Dietzgen, R. G. (2013). "Nyamiviridae: Proposal for a new family in the order Mononegavirales". Archives of Virology. 158 (10): 2209–2226. doi:10.1007/s00705-013-1674-y. PMID 23636404. 
  12. ^ Afonso, Claudio L.; Amarasinghe, Gaya K.; Bányai, Krisztián; Bào, Yīmíng; Basler, Christopher F.; Bavari, Sina; Bejerman, Nicolás; Blasdell, Kim R.; Briand, François-Xavier (2016-08-01). "Taxonomy of the order Mononegavirales: update 2016". Archives of Virology. 161 (8): 2351–2360. doi:10.1007/s00705-016-2880-1. ISSN 1432-8798. PMC 4947412free to read. PMID 27216929. 
  13. ^ Horie, M.; Honda, T.; Suzuki, Y.; Kobayashi, Y.; Daito, T.; Oshida, T.; Ikuta, K.; Jern, P.; Gojobori, T.; Coffin, J. M.; Tomonaga, K. (2010). "Endogenous non-retroviral RNA virus elements in mammalian genomes". Nature. 463 (7277): 84–87. doi:10.1038/nature08695. PMC 2818285free to read. PMID 20054395. 
  14. ^ a b c Belyi, V. A.; Levine, A. J.; Skalka, A. M. (2010). Buchmeier, Michael J., ed. "Unexpected Inheritance: Multiple Integrations of Ancient Bornavirus and Ebolavirus/Marburgvirus Sequences in Vertebrate Genomes". PLoS Pathogens. 6 (7): e1001030. doi:10.1371/journal.ppat.1001030. PMC 2912400free to read. PMID 20686665. 
  15. ^ a b c Katzourakis, A.; Gifford, R. J. (2010). Malik, Harmit S., ed. "Endogenous Viral Elements in Animal Genomes". PLoS Genetics. 6 (11): e1001191. doi:10.1371/journal.pgen.1001191. PMC 2987831free to read. PMID 21124940. 
  16. ^ Taylor, D.; Leach, R.; Bruenn, J. (2010). "Filoviruses are ancient and integrated into mammalian genomes". BMC Evolutionary Biology. 10: 193. doi:10.1186/1471-2148-10-193. PMC 2906475free to read. PMID 20569424. 
  17. ^ Chiba, S.; Kondo, H.; Tani, A.; Saisho, D.; Sakamoto, W.; Kanematsu, S.; Suzuki, N. (2011). Nagy, Peter D, ed. "Widespread Endogenization of Genome Sequences of Non-Retroviral RNA Viruses into Plant Genomes". PLoS Pathogens. 7 (7): e1002146. doi:10.1371/journal.ppat.1002146. PMC 3136472free to read. PMID 21779172. 
  18. ^ Fort, P.; Albertini, A.; Van-Hua, A.; Berthomieu, A.; Roche, S.; Delsuc, F.; Pasteur, N.; Capy, P.; Gaudin, Y.; Weill, M. (2011). "Fossil Rhabdoviral Sequences Integrated into Arthropod Genomes: Ontogeny, Evolution, and Potential Functionality". Molecular Biology and Evolution. 29 (1): 381–390. doi:10.1093/molbev/msr226. PMID 21917725. 

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