Gammacoronavirus‡
‡This chapter was reprinted from the first edition of the Springer Index of Viruses. Taxonomy and classification of the virus species described in this chapter may have changed.
Virion
Morphology: | Spherical |
Envelope: | Yes |
Diameter (nm): | 120–160 |
Length (nm): | |
Structural components: | Nucleocapsid, core, envelope |
Buoyant density (g/mL): | 1.23–1.24 |
Buoyant density method: | CsCl |
Lipid composition: | Envelope lipids are derived from cytoplasma membrane of host cell |
Additional information: | Surface projections made by the spike (S) protein |
Genome
Nucleic acid: | RNA | |
Strandedness: | ||
Polarity: | ||
Configuration: | ||
Segment organization: | Segment no. 1 (kb): | 26.4–31.7 |
One segment(s): | 26.4–31.7 (kb) total (calculated) | |
G + C content (%): | 37.6–41.8 | |
mRNA transcripts: | 7–10 | |
Open reading frames: | 7–10 | |
Additional information: | The genome contains a leader at the 5′ end and a poly(A) tail; genes are arranged in the order 5′-replicase-S-E-M-N-3′, with a variable number of other genes that are believed to be non-structural |
Replication
Entry mechanism: | Receptor-mediated endocytosis |
Site of transcription: | Cytoplasm |
Transcriptase: | Virus-encoded RNA-dependent RNA polymerase |
Site of genome replication: | Cytoplasm |
Replicase: | Virus-encoded RNA-dependent RNA polymerase |
Replication intermediate: | Negative-strand RNA intermediate |
Site of virion assembly: | Cytoplasm, the intermediate compartment |
Egress mechanism: | Budding through the pre-Golgi and Golgi |
Additional information: | Only the membrane (M) and envelope (E) proteins are required for the production of virus-like particles (VLPs) |
History
Year of event | Event | References |
---|---|---|
1930 | First report of infectious bronchitis in poultry in North Dakota | Schalk AE, Hawn MC (1931) J Am Vet Med Assoc 78:413–422 |
1936 | First demonstration of a coronavirus disease, infectious bronchitis (IB), in domestic fowl | Beach JR, Schalm OW (1936) Poult Sci 15:199–206 |
1937 | First cultivation of infectious bronchitis virus in chick embryos | Beaudette ER, Hudson CB (1937) J Am Vet Med Assoc 90:51–58 |
1949 | “Stunting and curling” in chick embryos is an IB pathognomonic lesion | Fabricant J (1951) Cornell Vet 39:414–431 |
1973 | Coronavirus-like (turkey coronavirus, TuCoV) particles identified by electron microscopy in turkeys with bluecomb disease | Ritchie JM et al (1973) |
1975 | ICTV approves Coronaviridae family with one genus, Coronavirus | Tyrrell DAJ et al (1975) |
1977 | Coronavirus (IBV) RNA shown to be infectious | Lomniczi B (1977) |
1980 | Coronavirus (IBV) subgenomic mRNAs form a 3′coterminal nested set | Stern DI, Kennedy SI (1980) |
1987 | First report on a coronavirus genome (IBV) complete sequence. | Boursnell MEG et al (1987) |
1987 | Frameshift in translation of replicase gene experimentally demonstrated in vitro | Brierley AS et al (2000) |
1996 | An IBV-related coronavirus is isolated from pheasants | Gough AE et al (1996) |
1996 | ICTV recognises Coronaviridae as containing two genera: Coronavirus and Torovirus | Cavanagh JG et al (1997) |
1996 | ICTV recognises the order Nidovirales containing families Coronaviridae and Arteriviridae | Cavanagh JG et al (1997) |
2005 | Design of wide-spectrum inhibitors of coronavirus main protease | Yang Y et al (2005) |
2008 | Identification of a novel coronavirus (beluga whale coronavirus, BWCoV) from a beluga whale | Mihindukulasuriya KA et al (2008) |
2009 | ICTV recognises the family Coronaviridae as containing two subfamilies, Coronavirinae and Torovirinae, with the former including three genera | Carstens K et al (2010) |
2009 | According to the new taxonomy, genus Coronavirus is replaced by genera Alpha-, Beta- and Gammacoronavirus, corresponding to the old antigenic groups | Carstens K (2010) |
2009 | IBV, TuCoV, PhCoV, BWCoV and related viruses are recognised as host variants of a unique species, Avian coronavirus, of the genus Betacoronavirus | Carstens K (2010) |
2009 | Identification in wild birds of new putative members of the genus Gammacoronavirus | Woo J et al (2009) |
Genus Members
Species name | Synonyms | Wild-type strains/isolates | Natural host range | Experimental host range | Membership status |
---|---|---|---|---|---|
Avian coronavirus | Avian infectious bronchitis virus (IBV); Turkey coronavirus (TuCoV); Pheasant coronavirus (PhCoV); Duck coronavirus (DuCoV); Goose coronavirus (GoCoV); Pigeon coronavirus (PgCoV); Quail coronavirus (QCoV) | IBV:Massachusetts M41;4/91;H; TuCoV:Minnesota; PhCoV:ph/UK/6/99; PgCoV:03/653; QCoV:Italy/Elvia/2005 | Domestic fowl, turkeys, other domestic birds | Quail, gulls, suckling mice (IBV); chicken (TuCoV) | Type species |
Beluga Whale coronavirus SW1 (BWCoV-SW1) | SW1 | Beluga whales (Delphinapterus leucas) | Approved member | ||
Asian leopard cat coronavirus (ALCCoV) | Guangxi/F230/2006 | Asian leopard cats (Prionailurus bengalensis) | Tentative member | ||
Black-headed gull coronavirus | CIR-66152; CIR-66146; CIR-66187; CIR-66185 | Black-headed gull (Chroicocephalus ridibundus) | Tentative member | ||
Brent goose coronavirus | KR-70 | Russian Brent goose (Branta bernicla) | Tentative member | ||
Bulbul coronavirus HKU11 (BuCoV-HKU11) | 796; 934 | Bulbuls (Pycnonotus jocosus) | Tentative member | ||
Chinese ferret badger coronavirus Guanxi/2006 (CFBCoV-Guanxi/2006) | Guangxi/F247/2006; Guangxi/F250/2006 | Chinese ferret badgers (Melogale moschata) | Tentative member | ||
Glaucous gull coronavirus | PBA-173 | Glaucous gull (Larus hyperboreus) | Tentative member | ||
Glaucous-winged gull coronavirus | CIR-66002 | Glaucous-winged gull (Larus glaucescens) | Tentative member | ||
Munia coronavirus HKU13 (Munia coronavirus HKU13) | 3514 | 3,514 | Tentative member | ||
Pintail coronavirus PBA-124 | 10; 15; 25; PBA-124 | Pintail (Anas acuta) | Tentative member | ||
Rock sandpiper coronavirus | CIR-65885; CIR-65828; CIR-65824; CIR-665821 | Rock sandpiper (Calidris ptilocnemis) | Tentative member | ||
Snow goose coronavirus | WIR-159 | Snow goose coronavirus (Anser caerulescens) | Tentative member | ||
Thrush coronavirus HKU12 (ThCoV-HKU12) | 600 | Thrushes (Turdus hortulorum, Turdus merula) | Tentative member | ||
Western sandpiper coronavirus | KR-28 | Western sandpiper (Calidris mauri) | Tentative member |
Nucleotide Sequences
Genomic region | Species | Strain | Nucleotides | Access number | References |
---|---|---|---|---|---|
Complete genome | Avian coronavirus (IBV) | Beaudette | 27,608 | M95169 | Boursnell ME et al (1987) |
Complete genome | Avian coronavirus (TuCoV) | TuCoV-540 | 27,771 | EU022525 | Cao R et al (2008) |
Complete genome | BuCoV-HKU11 | 796 | 26,476 | FJ376620 | Woo J et al (2009) |
Complete genome | ThCoV-HKU12 | 600 | 26,396 | NC_011549 | Woo J et al (2009) |
Complete genome | MuCoV-HKU13 | 3514 | 26,552 | NC_011550 | Woo J et al (2009) |
Complete genome | BWCoV-SW1 | SW1 | 31,686 | NC_010646 | Mihindukulasuriya KA et al (2008) |
Genomic very 3′ end | Avian coronavirus (PhCoV) | ph/UK/602/95 | 443 | AJ619604 | Cavanagh PR et al (2002) |
Replicase (partial) | Avian coronavirus (QCoV) | Italy/Elvia/2005 | 429 | EF446156 | Circella M et al (2007) |
Replicase (partial) | Avian coronavirus (PgCoV) | 03/428-1 | 208 | AJ854131 | Jonassen CM et al (2005) |
Replicase (partial) | Avian coronavirus (GCoV) | 03/586-1 | 208 | AJ854114 | Jonassen CM et al (2005) |
replicase (partial) | Avian coronavirus (DCoV) | 03/1094 | 208 | AJ8541 | Jonassen CM et al (2005) |
Genomic 3′ end | ALCCoV | Guangxi/F230/2006 | 12,801 | EF584908 | Dong PD et al (2007) |
Replicase (partial) | CBFCoV-Guanxi/2006 | Guangxi/F247/2006 | 1,179 | EF584909 | Dong PD et al (2007) |
Replicase (partial) | Snow goose coronavirus | WIR-159 | 561 | GU396690 | Muradrasoli S et al (2010) |
Replicase (partial) | Rock sandpiper coronavirus | CIR-65885 | 562 | GU396689 | Muradrasoli, et al (2010) |
Replicase (partial) | Black-headed gull coronavirus | CIR-66152 | 562 | GU396686 | Muradrasoli S et al (2010) |
Replicase (partial) | Glaucous-winged gull coronavirus | CIR-66002 | 562 | GU396682 | Muradrasoli S et al (2010) |
Replicase (partial) | Glaucous gull coronavirus | PBA-173 | 541 | GU396674 | Muradrasoli, et al (2010) |
Replicase (partial) | Brent goose coronavirus | KR-70 | 531 | GU396676 | Muradrasoli S et al (2010) |
Replicase (partial) | Western sandpiper coronavirus | KR-28 | 535 | GU396675 | Muradrasoli S et al (2010) |
Replicase (partial) | Pintail coronavirus | PBA-124 | 542 | GU396673 | Muradrasoli S et al (2010) |
Proteins
Protein name | Protein name abbreviation | Number of amino acids | Molecular weight (kDa) | Time of expression | Accession numbers | Additional information |
---|---|---|---|---|---|---|
Polyprotein 1ab (replicase complex) | pp1ab | 6262–6663 | 740–800 | Throughout | NP_066134; ACH72792; YP_001941164; ACJ12052; ACJ12043; ACJ12061; YP_001876435 | Encoded by two ORFs, 1a and 1b; pseudoknot involved in frameshifting; cleaved to several products, including an RNA-dependent RNA polymerase |
Spike glycoprotein | S | 1156–1472 | 150–220 | Throughout | ADC79651; ABF61518; YP_001941166; YP_001876437; ACJ12044; YP_002308506; YP_002308497 | Highly glycosylated; forms homotrimers; cleaved to S1 and S2 subunits in some members |
Membrane protein | M | 217–260 | 217–264 | Throughout | ACD93207; ACZ17397; AAF35863; YP_001941170; YP_002308508; YP_002308499; ACJ12046; YP_001876439 | N-linked or O-linked glycans; triple-spanning |
Envelope protein | E | 82–110 | 9–12 | Throughout | CAZ86711; CAZ86702; ADQ57408; ABW81430; YP_002308498; ACJ12045; YP_002308507; ABQ39959; YP_001876438 | Essential for virion assembly; E plus M forms virus-like particles |
Nucleocapsid protein | N | 343–417 | 50–60 | Throughout | AAB24054; AAB48157; AAV88459; AAF23873; CAI40312; ACJ12057; ACJ12048; ACJ12066; ABW87831 | Highly basic phosphoprotein; forms a helical nucleocapsid |
IBV non-structural protein 3a | IBVns3a | 57–61 | 6.7 | Throughout | AAT70773; CAA42115; ABW81428; ACC60267 | Unique to Avian coronavirus; absent in some strains; not essential for replication |
IBV non-structural protein 3b | IBVns3b | 62–72 | 7.4 | Throughout | AAT70774; CAA42488; CAA43123; ABQ84824; ACV87274; ACV87252 | Unique to Avian coronavirus; not essential for replication |
IBV non-structural protein 5a | IBVns5a | 65 | 7.4 | Throughout | AAA70240; NP_040836; ABW75145; ACV87248; CAF22050 | Unique to Avian coronavirus; not essential for replication |
IBV non-structural protein 5b | IBVns5b | 80–90 | 9.5 | Throughout | ABG36793; ABQ84764; ABQ84859; ACC60264; ACV87269 | Unique to Avian coronavirus; not essential for replication |
MuCoV/BuCoV/ThCoV non-structural protein 6 | MuCoV/BuCoV/ThCoV ns6 | 91–108 | 10.3–12.2 | YP_002308500; ACJ12047; YP_002308509 | Putative accessory protein unique to MuCoV/BuCoV/ThCoV; gene located between M and N genes | |
MuCoV/BuCoV/ThCoV non-structural protein 7a | MuCoV/BuCoV/ThCoV ns7a | 123 | 13.8 | ACJ12040; YP_002308502; YP_002308511 | putative accessory protein unique to MuCoV/BuCoV/ThCoV; gene located downstream of N gene | |
MuCoV/BuCoV/ThCoV non-structural protein 7b | MuCoV/BuCoV/ThCoV ns7b | 83–85 | 9.1–9.3 | ACJ12041; YP_002308503; YP_002308512 | Putative accessory protein unique to MuCoV/BuCoV/ThCoV; gene located downstream of N gene | |
MuCoV/BuCoV/ThCoV non-structural protein 7c | MuCoV/BuCoV/ThCoV ns7C | 77–94 | 9.1–11.2 | ACJ12042; YP_002308513; YP_002308504 | Putative accessory protein unique to MuCoV/BuCoV/ThCoV; gene located downstream of N gene | |
BWCoV-SW1 non-structural protein 5a | BWCoV-SW1 ns5a | 138 | 15.6 | YP_001876440 | Putative accessory protein unique to BWCoV-SW1; gene located between M and N genes | |
BWCoV-SW1 non-structural protein 5b | BWCoV-SW1 ns5b | 172 | 19.4 | YP_001876441 | Putative accessory protein unique to BWCoV-SW1; gene located between M and N genes | |
BWCoV-SW1 non-structural protein 5c | BWCoV-SW1 ns5c | 175 | 20.1 | YP_001876442 | Putative accessory protein unique to BWCoV-SW1; gene located between M and N genes | |
BWCoV-SW1 non-structural protein 6 | BWCoV-SW1 ns6 | 228 | 25.3 | YP_001876443 | Putative accessory protein unique to BWCoV-SW1; gene located between M and N genes | |
BWCoV-SW1 non-structural protein 7 | BWCoV-SW1 ns7 | 161 | 18.4 | YP_001876444 | putative accessory protein unique to BWCoV-SW1; gene located between M and N genes | |
BWCoV-SW1 non-structural protein 8 | BWCoV-SW1 ns8 | 59 | 6.8 | YP_001876445 | Putative accessory protein unique to BWCoV-SW1; gene located between M and N genes | |
BWCoV-SW1 non-structural protein 9 | BWCoV-SW1 ns9 | 152 | 17.3 | YP_001876446 | Putative accessory protein unique to BWCoV-SW1; gene located between M and N genes | |
BWCoV-SW1 non-structural protein 10 | BWCoV-SW1 ns10 | 210 | 23.7 | YP_001876447 | Putative accessory protein unique to BWCoV-SW1; gene located between M and N genes |
Biology
Species | Permissive cell lines | Tissue tropism | Cytopathic effects | Additional information |
---|---|---|---|---|
Avian coronavirus (IBV) | Chicken eggs and kidney cells; Vero (Beaudette strain) | Respiratory epithelium; also kidney, oviduct, gut | Embryonic stunting/curling (egg); syncytia (cells) | Can cause viral persistence; some strains highly nephropathogenic; some strains haemagglutinating |
Avian coronavirus (TuCoV) | Embryonated turkey and chicken eggs | Intestinal epithelium, bursa of Fabricius | Embryonic stunting/curling | Amniotic inoculation is required |
Avian coronavirus (PhCoV) | Embryonated chicken eggs | Respiratory tissues and kidneys | Embryonic stunting/curling | Allantoic route inoculation |
Diseases
Disease | Causative agent | Affected organisms | Disease CHARACTERISTICS | Transmission route/vector | Treatment | Geographic distribution |
---|---|---|---|---|---|---|
Infectious bronchitis of poultry | Avian coronavirus (IBV) | Chickens | Sneezing, coughing; nephritis; declined egg production | Aerosol | No specific treatment | World-wide |
Bluecomb disease, mud fever | Avian coronavirus (TuCoV) | Turkeys | Enteritis, wet droppings, depression | Faecal-oral | No specific treatment | World-wide |
Pheasant respiratory and kidney disease | Avian coronavirus (PhCoV) | Pheasants | Rapid death, sneezing, poor hatchability, small size, variable color-pale buff to greenish brown | Aerosol | No specific treatment | World-wide |
Diagnosis
Method | Species | Sample material | Detection target | References |
---|---|---|---|---|
Electron microscopy | Avian coronavirus (IBV, TuCoV, PhCoV) | Respiratory and/or cloacal specimens, kidneys | Particle morphology | Reagan RL et al (1948) |
Cultivation in embryonated chicken eggs | Avian coronavirus (IBV, TuCoV, PhCoV) | Respiratory and/or cloacal specimens, kidneys | Embryonic alterations (stunting and curling) | Cunningham CH, Stuart HO (1947) |
Agar gel precipitin test | Avian coronavirus (IBV, TuCoV, PhCoV) | Respiratory and/or cloacal specimens, kidneys | Viral antigens | Lohr M (1981) |
Immunofluorescence assay | Avian coronavirus (IBV, TuCoV, PhCoV) | Tracheal, intestinal, renal sections | Viral antigens | Yagyu K, Ohta S (1990) |
Immunoperoxidase assay | Avian coronavirus (IBV, TuCoV, PhCoV) | Tracheal, intestinal, renal sections | Viral antigens | Naqi S (1990) |
Enzyme-linked immunosorbent assay with monoclonal antibodies | Avian coronavirus (IBV, TuCoV, PhCoV) | Respiratory and/or cloacal specimens, kidneys | Viral antigens | Naqi S et al (1993) |
RT-PCR amplification of the spike-protein gene | Avian coronavirus (IBV, TuCoV, PhCoV) | Respiratory and/or cloacal specimens, kidneys | Viral rna | Lin J et al (1991) |
Nested RT-PCR amplification of the nucleocapsid-protein gene | Avian coronavirus (IBV, TuCoV, PhCoV) | Respiratory and/or cloacal specimens, kidneys | Viral rna | Falcone DJ et al (1997) |
Real-time RT-PCR amplification of the 5′-UTR | Avian coronavirus (IBV, TuCoV, PhCoV) | Respiratory and/or cloacal specimens, kidneys | Viral rna | Callison B et al (2006) |
Real-time polymerase chain reaction/high-resolution melt curve analysis for IBV strain differentiation | Avian coronavirus (IBV) | Respiratory and/or cloacal specimens, kidneys | Viral rna | Hewson I et al (2010) |
Vaccine Strains
Strain | Attenuation process | Additional information | References |
---|---|---|---|
Avian coronavirus IBV many strains | Inactivated vacciness | Poor effective | Cavanagh D (2007) |
Avian coronavirus IBV many strains | Passage in embryonated domestic fowl eggs | Highly effective; may induce sterilising immunisation | Cavanagh D (2007) |
Vector Constructs
Vector name | Backbone strain | Application | Insertion capacity (kb) | Additional information | References |
---|---|---|---|---|---|
IBV CD-61 | Avian coronavirus (IBV Beaudette) | Expression | 1 | Transcription regulated by ibv control sequences | Penzes et al (1996) |
vNotI/IBV(FL) | Avian coronavirus (IBV Beaudette) | Reverse genetics | 27.6 | Recombinant vaccinia virus containing the full-length genome of mhv | Casais R et al (2001) |
IBV c-DNA | Avian coronavirus (IBV Beaudette) | Reverse genetics | 27.6 | Full-length genome infectious clone constructed through ligation of five inserts | Youn JG et al (2005) |
Transmission electron micrograph, negative staining of purified virus. Length of bar (nm): 100 (Courtesy of Dr. A. Lavazza, Istituto Zooprofilattico di Lombardia ed Emilia Romagna, Italy)
HE protein is present only in some Betacoronaviruses (Courtesy of Dr Viviana Tarallo, Department of Veterinary Public Health, Valenzano, Italy)
Numbers above bars = ORFs; L leader; UTR untranslated region; Rep replicase; (A)n poly A (Modified from Springer Index of Viruses, 1st edition, with permission)
References
Carstens (2010)
Cavanagh (1997)
Enjuanes L, Siddell SG, Spaan WJ (1998) Coronaviruses and arteriviruses. Plenum, New York
Enjuanes L, Brian D, Cavanagh D, Holmes K, Lai MMC, Laude H, Masters P et al (1999a) Coronaviridae. In: Murphy FA et al (eds) Virus taxonomy. Academic, New York
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Holmes KV, Lai MMC (1996) Coronaviridae: the viruses and their replication. In: Fields BN, Knipe DM, Howley PM (eds) Fundamental virology. Academic, New York
Lai and Cavanagh (1997)
Perlman S, Gallagher T, Snijder EJ (2008) Nidovirales. ASM Press, Washington, DC
Siddell SG (1995) In: Fraenkel-Conrat H, Wagner RR (eds) The coronaviridae. Plenum, New York
Sturman and Holmes (1983)