"CoronavirusTo this itemtransferIt is done. For the higher class Coronaviridae, seeCoronavirus family" For coronaviruses that infect humansHuman coronavirus" For the new coronavirus (SARS-CoV-2019) discovered in 2, seeSARS coronavirus 2" Regarding the impact of the new coronavirus (SARS-CoV-2) epidemic,Pandemic of New Coronavirus Infection (2019-)"Please refer to the.

Orthocoronavirus subfamily(Orthocoronavirinae / Coronavirus〈Coronavirus〉) isgenomeAsRibonucleic acid (RNA) WithSingle-stranded plus-strand RNA virusso,mammalian,birdsTosickcauseウ イ ル スOne of a group of^[1].Nidovirus eyesCoronavirus familyBelongs to^[2][3].

The viruses included are from the Coronavirus familyAmphibianIt is a so-called alpha to delta coronavirus, except that it infects. Prior to 2018, it was called "Coronavirus subfamily", and before 2009 it was called "Coronavirus genus".SortAs a name, in 2018 ``Orthocoronavirus subfamilyWas renamed to ", but as a general nameCoronavirusIs still used.

Outline

If you just say coronavirus,Coronavirus family, Or this orthocoronavirus subfamily. Since the Coronavirus family includes the Retovirus subfamily in addition to the coronaviruses from alpha to delta, the actual classification range is Coronavirus=Orthocoronavirus subfamily.

Orthocoronavirus is a virus particle surfaceEnvelope (membrane structure)Has long spike-shaped spike protein protrusions (S protein, about 20 nm)コ ロ ナ((I.e.Similar to^[1].. Spiral symmetricNucleocapsidWithEnvelope virusIs. Polymorphic, size 80-220 in diameternmIs about^[1]..Genome size is about 26-32Kilobase (kb) in knownRNA virusIs the largest^[4].

Symptoms depend on the type of organism, and in the case of chickensUpper respiratory tractCausing disease and in the case of cattle and pigsdiarrheacause.

Human capitalThen,coldincludingRespiratory infectionscause.SARS coronavirus (SARS-CoV),MERS coronavirus (MERS-CoV) andSARS coronavirus 2 It is lethal in viruses of the type (SARS-CoV-2).To prevent human coronavirus infectionvaccineHas not been offered yet^[5].

Name

The name of "coronavirus" isLatin Ofcrown(コ ロ ナ)andGreek language OfcrownorLight crown(Round light circle),corollaMeans κορώνη(korṓnē Derived from^[6][7].

This name iselectronic microscopebyVirionIt is derived from the characteristic appearance of (virus particles having infectivity). The virion has the edge of a large spherical surface protrusion,Sun coronaCreate an image reminiscent of. The subfamily name "ortho" coronavirus comes from the Greek word ορθός (orthos), which means "regular".

History

The first to be discovered in this group was the chicken reported in 1931.Infectious bronchitis virusIs^[8].. Mouse hepatitis virus and swine infectious gastroenteritis virus were also reported in the 1940s.^{[Note 1]}.

As a human pathogen, from a human who caught a cold in 1960,Human coronavirusB814 was discovered. This strain was later lost because it was difficult to culture. In the 1960scoldOf a human patientNasal cavityからHuman coronavirus 229EandHuman coronavirus OC43Two viruses were discovered^[9].. Initially, the relationship between these viruses was unclear and was not called coronavirus. Human coronavirus was originally called human respiratory virus.

In the late 1960s, electron micrographs pointed out similarities in structure.^[10]Started to be called coronavirus in 1968^[11].. In 1971CoronavirusWas summarized as.

In 2009, the classification was advanced due to the progress of molecular phylogenetic analysis. The genus Coronavirus was dismantled and a new Delta Coronavirus was installed from Alpha. In addition, a subfamily of torovirus (later independent as the family of torovirus) was discovered as a virus closely related to coronavirus, and as a group of coronaviruses from alpha to delta.Coronavirus subfamilyWas set.

In 2018, there was another move, with the Torovirus subfamily becoming independent of the Torovirus family, while the Retovirus subfamily was set up as a new group of the Coronavirus family. At this time the coronavirus subfamilyOrthocoronavirus subfamilyIt has been renamed (meaning true coronavirus) and is still in use today.

Changes in classification of coronaviruses and related strains

1971

2009

2018

Sort

Virus classificationIn the Orthocoronavirus subfamily together with the Retovirus subfamilyCoronavirus familyinclude^[12].. Orthocoronavirus contains 4 genera of 45 genera.

Orthocoronavirus subfamily

Alpha coronavirus

• Alpha coronavirus (Alphacoronavirus) Type:
  • seed:Bat coronavirus CDPHE15, Bat coronavirus HKU10, Rhinolophus ferrumequinum alphacoronavirus HuB-2013,Human coronavirus 229E,Lucheng Rn rat coronavirus, Mink coronavirus 1, Miniopterus bat coronavirus 1, Miniopterus bat coronavirus HKU8, Myotis ricketti alphacoronavirus Sax-2011, Nyctalus velutinus alphacoronavirus SC-2013, Pipistrellus kuhlii coronavirus 3398, (Porcine epidemic diarrhea virus),Scotophilus bat coronavirus 512, Rhinolophus bat coronavirus HKU2, Human coronavirus NL63, NL63-related bat coronavirus strain BtKYNL63-9b, Sorex araneus coronavirus T14, Suncus murinus coronavirus X74, Alpha Coronavirus 1

Beta coronavirus

• Beta coronavirus (betacoronavirus) Type: Mouse Coronavirus
  • seed:Beta Coronavirus 1,China Rattus coronavirus HKU24,Human coronavirus HKU1,Myodes coronavirus 2JL14, Bat Hp-betacoronavirus Zhejiang2013, Hedgehog coronavirus 1,MERS coronavirus,Pipistrellus bat coronavirus HKU5, Tylonycteris bat coronavirus HKU4, Eidolon bat coronavirus C704, Rousettus bat coronavirus GCCDC1, Rousettus bat coronavirus HKU9,SARS-related coronavirus

Gamma coronavirus

• Gamma coronavirus (Gammacoronavirus) Type:Chicken infectious bronchitis virus
  • seed:Goose coronavirus CB17, Beluga whale coronavirus SW1, Chicken infectious bronchitis virus,Avian coronavirus 9203, Duck coronavirus 2714

Delta coronavirus

• Delta coronavirus (Deltacoronavirus) Type: Bulbul coronavirus HKU11
  • seed:Wigeon coronavirus HKU20, Bulbul coronavirus HKU11,Common moorhen coronavirus HKU21, Coronavirus HKU15, Munia coronavirus HKU13, White-eye coronavirus HKU16, Night heron coronavirus HKU19

Construction

Viruses belonging to the subfamily Orthocoronavirus are particulate, and their surface is a lipid bilayer membrane similar to that of cells in general. When photographed with an electron microscope, the appearance of many spike proteins growing on the surface looks like a crown. This lipid bilayer membrane, called the envelope, has a function of recognizing the host cell to be infected with spikes.HemagglutininProtein embedded (especially beta coronavirus subgroup A member)^[13].. Inside the envelope is the viral genome. The genome is a piece of RNA wrapped in protein, and the state in which this genomic RNA is wrapped in protein is called nucleocapsid.

genome

Orthocoronavirus subfamily (and most otherNidovirus eyes), one of its features is that its genome is RNA rather than DNA. The genomic RNA remains intact in the host cellMessenger RNA(Messenger RNA, mRNA,proteinTo翻 訳Can be doneNucleotide sequenceWith information and structureRNAIt has an array structure that functions as^[6].. mRNA consists of several mRNAs that extend from the '3 side to the '5 side of genomic RNA with different lengths, and the '5 end of each mRNA is at the 5'end of genomic RNA.Leader arrayhave^[6].

Viral protein翻 訳Is generally present at the '5 end of each mRNAOpen reading frame Translated only from (ORF)^[6].. Genomic RNA (mRNA-1) consists of two ORFs (encoding 5kDa in 20a and 2b) at about 1 kb at the 1'end.Pseudo knot called (pseudoknot, Pn)Nucleic acid tertiary structurehave^[6].. There are cases where translation is terminated only by the 1a protein, and cases where Pn synthesizes the 1a + 1b fusion protein. The 1a + 1b protein is cleaved into 16 regulatory proteins,Protease(Peptide bondHydrolase),RNA polymeraseWork as^[6].

protein

Virus particle structureProteinThe highest amount of nucleoprotein (N) binds to genomic RNA. It binds to genomic RNA,NucleocapsidBecome^[6].. Wrap the nucleocapsidenvelopeHas a crown-like process characteristic of coronavirusSpike protein (S),Integral membrane protein (M),Envelope protein There is (E)^[6].

Proliferation process

Orthocoronavirus propagates by infecting animal cells. The process generally consists of three stages: infection (1, 2), replication (3-5), and release (6).

1. Spike protein S exposed on the surface of the viral envelope and, depending on the species,HemagglutininThe protein HE recognizes and binds to molecules on the surface of target cells.
2. The spike protein undergoes cleavage and activation by a host protease such as TMPRSS2.
3. Direct fusion of the viral envelope with the cell membrane of the target cell, orEndocytosisThe virus is taken up by the cells. In the case of direct fusion, the viral genome was directly introduced into the cell, but if incorporated by endocytosis, the virus was once included.EndosomeAre produced in the cell, where the viral genome is introduced by fusing the endosomal membrane with the virus. In the endosome,Proton pumpThe internal pH is lowered withLysosomeIs blocked by the virus as it is transported to.
4. Since coronavirus has a positive-stranded single-stranded RNA as its genome, it remains intact in the cytoplasm of the target cell.mFunctions as.Of the target cellRibosomeTo produce a viral protein containing RNA synthase.The viral RNA synthase does not replicate other than the viral genomic sequence, but replicates as a minus-strand RNA using only the viral genomic RNA as a template.This prepares the virus to replicate only its own RNA sequence.
5. Positive-strand RNA is synthesized for each gene from the negative-strand viral genomic RNA, and they bind to the ribosome of the target cell to produce viral proteins from each.In addition, the positive-strand genome that constitutes the virus is replicated from the negative-strand genome.Its genome is the same as the RNA sequence that has invaded the cell.
6. The produced viral protein N binds to the plus-strand genomic RNANucleocapsidOf the target cellsEndoplasmic reticulum Imported into (ER).Viral membrane protein M, spike protein S, and hemagglutinin HE are integrated into the endoplasmic reticulum membrane of target cells.The membrane of the endoplasmic reticulum becomes the membrane of the virus as it is.The virus is made from the nucleocapsid and the membrane of the endoplasmic reticulum (which becomes the envelope).
7. From the endoplasmic reticulumGolgi bodyViaExocytosisReleases the virus extracellularly from the target cell.The viral spike protein is then released in a manner that clings to the exocytosis membrane.This replicates the virus.

A specific example of SARS coronavirus, defined on SReceptorThe binding domain is the viral cellular receptorAngiotensin converting enzyme 2 (ACE2) Mediates binding to^[14].

Evolutionary process

The latest of all coronaviruses (=orthocoronavirus subfamily)Nearest common ancestor (MRCA) is believed to have existed in 8000 BC, but the MRCA for some models dates back more than 5500 million years.BatWithLong-term co-evolutionSuggest^[15]. (English edition）MRCA was around 2400 BC,(English edition）Around 3300 BC,(English edition）Is around 2800 BC,(English edition）Is believed to be around 3000 BC. Flying warm-blooded vertebrate bats and birds are the hosts that promote the evolution and spread of coronaviruses for coronavirus gene sources (alpha coronavirus and beta coronavirus bats, gamma coronavirus and delta coronavirus birds). Ideal as^[16].

Because of this, many human coronaviruses originate from bats^[17].. Human coronavirus NL63 had a common ancestor with the bat coronavirus (ARCoV 1190) between 1449 and 2 AD^[18].. Human coronavirus 229E also had a common ancestor with the bat coronavirus (GhanaGrp1686 Bt CoV) between 1800 and 1^[19].. In a more recent example, the alpaca coronavirus and human coronavirus 1960E diverged before 229.^[20].. MERS coronavirus emerged in humans from bats as an intermediate host via camels^[21].. The MERS coronavirus is associated with several bat coronaviruses and appears to have diverged from these species several centuries ago^[22].

In particular, the SARS coronavirus has a deeper relationship with the bat coronavirus than other human coronaviruses and diverged only around 1986.^[23].. The evolutionary pathways of Keen bat coronaviruses and SARS coronaviruses suggest that SARS-associated coronaviruses may have co-evolved in bats for long periods of time. SARS coronavirus ancestors were first infected with leaf-nose bats of the bat family Batidae. After that, it was infected with horseshoe bats of the family Bombyx bat family, further civet cat, and finally humans.^[24][25].

Unlike other beta coronaviruses, beta coronavirus 1 and the subgenus of the enbecovirus, bovine coronavirus, are thought to be of murine origin rather than bat origin^[17][26].. In the 1790s, equine coronavirus diverged from bovine coronavirus across species^[27]

Another interspecific transmission occurred in the late 1890s, when human coronavirus OC43 diverged from bovine coronavirus.^[27][28].. It is speculated that the 1890 influenza outbreak may have been caused by the spillover of this virus, not the influenza virus, due to the fact that the pathogen has not been actually identified, and due to its timing and its neurological symptoms.^[29].. Human coronavirus OC43 causes respiratory disease and is suspected to be involved in neurological diseases^[30].. The most common genotypes now emerge in the 1950s^[31].

Mouse hepatitis virus infecting mouse liver and central nervous system is systematically associated with human coronavirus OC43 and bovine coronavirus^[32].. Human coronavirus HKU1 also originates in rodents^[17].

Human coronavirus

Coronaviruses that infect humans are known to be 4 types of cold syndrome and 2 types of severe pneumonia virus (SARS-CoV, MERS-CoV) that are transmitted from animals.^[33], And 2 types including SARS-CoV-7 (as of March 2020)^[34]Is. The following alpha coronaviruses and beta coronaviruses are known^{[34][Note 2]}.

• (English edition）
  • Human coronavirus 229E : coldPathogen of
  • Human coronavirus NL63 : coldPathogen of
• (English edition）
  • Human coronavirus HKU1 : coldPathogen of
  • Human coronavirus OC43 : coldPathogen of
  • SARS coronavirus (SARS-CoV):Severe acute respiratory syndrome (SARS_Thurs) Pathogen
  • MERS coronavirus (MERS-CoV):Middle East respiratory syndrome (MERS_Mars) Pathogen
  • SARS coronavirus 2 (SARS-CoV-2):New coronavirus infection (COVID-19_{Covit Nineteen}) Pathogen

Animal coronavirus

Coronavirus家畜,Laboratory animal,Pet,Wild animalsAnd so onAnimalAnd cause various diseases^[33].Dog,cat,cow,pig,Chicken,A horse,camelLivestock, such asBeluga,Giraffe,ferret,Sunkus,Bat,SparrowA unique coronavirus has also been detected from^[33].

家畜

• Coronavirus family - Coronaviridae
  • (English edition） - Alphacoronavirus
    • Pedako virus subgenus (Pedacovirus)
      • (English edition） : Swine epidemic diarrheaVirus (PEDV): lethal^[33].. Almost 100% of pigs die due to dehydration associated with diarrhea, but the symptoms are milder as the age increases.^[1].
    • Tegacovirus subgenus (Tegacovirus)
      • (English edition） : Pig infectious gastroenteritisVirus (TGEV): lethal^[33].. Death rate is almost 7% under 100 days of age^[1].
  • (English edition） - betacoronavirus
    • (English edition） - Embecovirus
      • Porcine hemagglutinating encephalomyelitis virus (Porcine hemagglutinating encephalomyelitis virusHEV): Adult pigs have mild symptoms^[1].
      • Cow coronavirus^[1] - Bovine coronavirus
  • (English edition） - Gammacoronavirus
    • Igacovirus subgenus
      • (English edition） - Avian coronavirus
        • Chicken infectious bronchitis virus - Avian infectious bronchitis virus (IBV): Lethal^[33].. The mortality rate is higher for younger children^[1].
• Pig respiratory coronavirus (Porcine respiratory coronavirus; PRCoV): Asymptomatic and shows no symptoms^[1].
• Horse coronavirus: mortality rate of about 25%^[1]
• Turkey Coronavirus Enteritis: High Mortality in Young Birds^[1].

Laboratory animal

• Mouse hepatitis virus (MHV): Lethal^[33]

Pet

• Cat infectious peritonitisVirus (FIPV): Lethal^[33]

footnote

注 釈

1. ^ However, these viruses were merely isolated as pathogens, and were not characterized by electron microscopy.
2. ^ (- Human enteric coronavirus 4408 (HECV-4408): is excluded)

Source

1. ^ ^{a b c d e f g h i j k} Coronavirus disease of livestock and poultry National Research and Development Agency National Agriculture and Food Research Institute Animal Health Research Division April 2003, 4
2. ^ AMQ King, ed (2011). “Family Coronaviridae". Ninth Report of the International Committee on Taxonomy of Viruses. Elsevier, Oxford. Pp. 806-828. ISBN 978-0-12-384684-6 
3. ^ ICTV Master Species List 2009-v10 (xls)-International Committee on Taxonomy of Viruses (24 August 2010)
4. ^ “Homology-Based Identification of a Mutation in the Coronavirus RNA-Dependent RNA Polymerase That Confers Resistance to Multiple Mutagens”. Journal of Virology 90 (16): 7415-28. (August 2016). two:10.1128 / JVI.00080-16. PMC: 4984655. PMID 27279608. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4984655/"CoVs also have the largest known RNA virus genomes, ranging from 27 to 34 kb (31, 32), and increased fidelity in CoVs is likely required for the maintenance of these large genomes (14)." 
5. ^ Eiji Kusumi Coronaviruses can "defend" if you know them correctly Toyo Keizai 2020/02/01 5:30
6. ^ ^{a b c d e f g h} Fumihiro TaguchiCoronavirus Virus Vol. 61, No. 2, pp. 205-210, 2011
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10. ^ https://www.ncbi.nlm.nih.gov/pmc/articles/PMC224637/
11. ^ https://www.nature.com/articles/220650b0
12. ^ Virus Taxonomy: 2019 ReleaseICTV
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16. ^ “Discovery of seven novel Mammalian and avian coronaviruses in the genus deltacoronavirus supports bat coronaviruses as the gene source of alphacoronavirus and betacoronavirus and avian coronaviruses as the gene source of gammacoronavirus and deltacoronavirus”. Journal of Virology 86 (7): 3995-4008. (April 2012). two:10.1128 / JVI.06540-11. PMC: 3302495. PMID 22278237. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3302495/. 
17. ^ ^{a b c} “Molecular Evolution of Human Coronavirus Genomes”. Trends in Microbiology 25 (1): 35-48. (January 2017). two:10.1016 / j.tim.2016.09.001. PMC: 7111218. PMID 27743750. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7111218/"Specifically, all HCoVs are thought to have a bat origin, with the exception of lineage A beta-CoVs, which may have reservoirs in rodents [2]." 
18. ^ “Evidence supporting a zoonotic origin of human coronavirus strain NL63”. Journal of Virology 86 (23): 12816-25. (December 2012). two:10.1128 / JVI.00906-12. PMC: 3497669. PMID 22993147. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3497669/"If these predictions are correct, this observation suggests that HCoV-NL63 may have originated from bats between 1190 and 1449 CE." 
19. ^ “Distant relatives of severe acute respiratory syndrome coronavirus and close relatives of human coronavirus 229E in bats, Ghana”. Emerging Infectious Diseases 15 (9): 1377-84. (September 2009). two:10.3201 / eid1509.090224. PMC: 2819850. PMID 19788804. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2819850/"The most recent common ancestor of hCoV-229E and GhanaBt-CoVGrp1 existed in ≈1686-1800 AD." 
20. ^ “Identification and characterization of a novel alpaca respiratory coronavirus most closely related to the human coronavirus 229E”. V 4 (12): 3689-700. (December 2012). two:10.3390 / v4123689. PMC: 3528286. PMID 23235471. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3528286/. 
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22. ^ “Genetic characterization of Betacoronavirus lineage C viruses in bats reveals marked sequence divergence in the spike protein of pipistrellus bat coronavirus HKU5 in Japanese pipistrelle: implications for the origin of the novel Middle East respiratory syndrome coronavirus”. Journal of Virology 87 (15): 8638-50. (August 2013). two:10.1128 / JVI.01055-13. PMC: 3719811. PMID 23720729. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3719811/. 
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References

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• Ryo Harasawa "New classification of animal viruses (2005)", "Veterinary Animal Husbandry Bulletin" 58, 921-931 2005 ISSN 0447-0192
• Supervised by Akira Mijo "Veterinary Infection Color Atlas" published by Bunagaido 2006 ISBN 4830032030
• "Standard Microbiology" Osamu Nakagome, Shigeru Kamiya (edit),Medical school, February 2015, 2, 15th edition.National bibliographic number:22540957.ISBN 978-4-260-02046-6.NCID BB18056640.OCLC 904535631.
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• Cryo-EM structure of the 2019-nCoV spike in the prefusion conformation, Daniel Wrapp, Nianshuang Wang, Kizzmekia S. Corbett, Jory A. Goldsmith, Ching-Lin Hsieh, Olubukola Abiona, Barney S. Graham, Jason S. McLellan1, Science, 19 Feb 2020: eabb2507, DOI: 10.1126 / science.abb2507

外部 リンク

• What is coronavirus - National Institute of Infectious Diseases