Coronavirus

Coronaviruses are a group of related viruses that cause diseases in mammals and birds. In humans, coronaviruses cause respiratory tract infections that can range from mild to lethal. Mild illnesses include some cases of the common cold (which is caused also by certain other viruses, predominantly rhinoviruses), while more lethal varieties can cause SARS, MERS, and COVID-19. Symptoms in other species vary: in chickens, they cause an upper respiratory tract disease, while in cows and pigs they cause diarrhea. There are as yet no vaccines or antiviral drugs to prevent or treat human coronavirus infections.

This article is about the group of viruses. For the disease involved in the 2019–20 coronavirus pandemic, see Coronavirus disease 2019. For the virus that causes this disease, see Severe acute respiratory syndrome coronavirus 2.

Orthocoronavirinae
Transmission electron micrograph (TEM) of avian infectious bronchitis virus
Illustration of the morphology of coronaviruses; the club-shaped viral spike peplomers, colored red, create the look of a corona surrounding the virion when observed with an electron microscope.
Virus classification
(unranked): Virus
Realm: Riboviria
Phylum: incertae sedis
Order: Nidovirales
Family: Coronaviridae
Subfamily: Orthocoronavirinae
Genera[1]
  • Alphacoronavirus
  • Betacoronavirus
  • Gammacoronavirus
  • Deltacoronavirus
Synonyms[2][3][4]
  • Coronavirinae

Coronaviruses constitute the subfamily Orthocoronavirinae, in the family Coronaviridae, order Nidovirales, and realm Riboviria.[5][6] They are enveloped viruses with a positive-sense single-stranded RNA genome and a nucleocapsid of helical symmetry. The genome size of coronaviruses ranges from approximately 26 to 32 kilobases, one of the largest among RNA viruses.[7] They have characteristic club-shaped spikes that project from their surface, which in electron micrographs create an image reminiscent of the solar corona, from which their name derives.[8]

Etymology

The name "coronavirus" is derived from Latin corona, meaning "crown" or "wreath", itself a borrowing from Greek κορώνη korṓnē, "garland, wreath".[9][10] The name was coined by June Almeida and David Tyrrell who first observed and studied human coronaviruses.[11] The word was first used in print in 1968 by an informal group of virologists in the journal Nature to designate the new family of viruses.[8] The name refers to the characteristic appearance of virions (the infective form of the virus) by electron microscopy, which have a fringe of large, bulbous surface projections creating an image reminiscent of the solar corona or halo.[8][11] This morphology is created by the viral spike peplomers, which are proteins on the surface of the virus.[12]

History
Transmission electron micrograph of infectious bronchitis viruses

Coronaviruses were first discovered in the 1930s when an acute respiratory infection of domesticated chickens was shown to be caused by infectious bronchitis virus (IBV).[13] Arthur Schalk and M.C. Hawn described in 1931 a new respiratory infection of chickens in North Dakota. The infection of new-born chicks was characterized by gasping and listlessness. The mortality rate of the chicks was 40–90%.[14] Fred Beaudette and Charles Hudson six years later successfully isolated and cultivated the infectious bronchitis virus which caused the disease.[15] In the 1940s, two more animal coronaviruses, mouse hepatitis virus (MHV) and transmissible gastroenteritis virus (TGEV), were isolated.[16] It was not realized at the time that these three different viruses were related.[17]

Human coronaviruses were discovered in the 1960s.[18][19] They were isolated using two different methods in the United Kingdom and the United States.[20] E.C. Kendall, Malcom Byone, and David Tyrrell working at the Common Cold Unit of the British Medical Research Council in 1960 isolated from a boy a novel common cold virus B814.[21][22][23] The virus was not able to be cultivated using standard techniques which had successfully cultivated rhinoviruses, adenoviruses and other known common cold viruses. In 1965, Tyrrell and Byone successfully cultivated the novel virus by serially passing it through organ culture of human embryonic trachea.[24] The new cultivating method was introduced to the lab by Bertil Hoorn.[25] The isolated virus when intranasally inoculated into volunteers caused a cold and was inactivated by ether which indicated it had a lipid envelope.[21][26] Around the same time, Dorothy Hamre and John Procknow at the University of Chicago isolated a novel cold virus 229E from medical students, which they grew in kidney tissue culture. The novel virus 229E, like the virus strain B814, when inoculated into volunteers caused a cold and was inactivated by ether.[27]

Transmission electron micrograph of organ cultured coronavirus OC43

The two novel strains B814 and 229E were subsequently imaged by electron microscopy in 1967 by Scottish virologist June Almeida at St. Thomas Hospital in London.[28][29] Almeida through electron microscopy was able to show that B814 and 229E were morphologically related by their distinctive club-like spikes. Not only were they related with each other, but they were morphologically related to infectious bronchitis virus (IBV).[30] A research group at the National Institute of Health the same year was able to isolate another member of this new group of viruses using organ culture and named the virus strain OC43 (OC for organ culture).[31] Like B814, 229E, and IBV, the novel cold virus OC43 had distinctive club-like spikes when observed with the electron microscope.[32][33]

The IBV-like novel cold viruses were soon shown to be also morphologically related to the mouse hepatitis virus.[16] This new group of IBV-like viruses came to be known as coronaviruses after their distinctive morphological appearance.[8] Human coronavirus 229E and human coronavirus OC43 continued to be studied in subsequent decades.[34][35] The coronavirus strain B814 was lost. It is not known which present human coronavirus it was.[36] Other human coronaviruses have since been identified, including SARS-CoV in 2003, HCoV NL63 in 2004, HCoV HKU1 in 2005, MERS-CoV in 2012, and SARS-CoV-2 in 2019.[37][38] There have also been a large number of animal coronaviruses identified since the 1960s.[5]

Microbiology

Structure
Cross-sectional model of a coronavirus

Coronaviruses are large pleomorphic spherical particles with bulbous surface projections.[39] The average diameter of the virus particles is around 120 nm (.12 μm). The diameter of the envelope is ~80 nm (.08 μm) and the spikes are ~20 nm (.02 μm) long. The envelope of the virus in electron micrographs appears as a distinct pair of electron dense shells.[40][41]

The viral envelope consists of a lipid bilayer where the membrane (M), envelope (E) and spike (S) structural proteins are anchored.[42] A subset of coronaviruses (specifically the members of betacoronavirus subgroup A) also have a shorter spike-like surface protein called hemagglutinin esterase (HE).[5]

Inside the envelope, there is the nucleocapsid, which is formed from multiple copies of the nucleocapsid (N) protein, which are bound to the positive-sense single-stranded RNA genome in a continuous beads-on-a-string type conformation.[41][43] The lipid bilayer envelope, membrane proteins, and nucleocapsid protect the virus when it is outside the host cell.[44]

Genome
Schematic representation of the genome organization and functional domains of S protein for SARS-CoV and MERS-CoV

Coronaviruses contain a positive-sense, single-stranded RNA genome. The genome size for coronaviruses ranges from 26.4 to 31.7 kilobases.[7] The genome size is one of the largest among RNA viruses. The genome has a 5′ methylated cap and a 3′ polyadenylated tail.[41]

The genome organization for a coronavirus is 5′-leader-UTR-replicase/transcriptase-spike (S)-envelope (E)-membrane (M)-nucleocapsid (N)-3′UTR-poly (A) tail. The open reading frames 1a and 1b, which occupy the first two-thirds of the genome, encode the replicase/transcriptase polyprotein. The replicase/transcriptase polyprotein self cleaves to form nonstructural proteins.[41]

The later reading frames encode the four major structural proteins: spike, envelope, membrane, and nucleocapsid.[45] Interspersed between these reading frames are the reading frames for the accessory proteins. The number of accessory proteins and their function is unique depending on the specific coronavirus.[41]

Replication cycle

Entry
The life cycle of a coronavirus

Infection begins when the viral spike (S) glycoprotein attaches to its complementary host cell receptor. After attachment, a protease of the host cell cleaves and activates the receptor-attached spike protein. Depending on the host cell protease available, cleavage and activation allows the virus to enter the host cell by endocytosis or direct fusion of the viral envelop with the host membrane.[46]

On entry into the host cell, the virus particle is uncoated, and its genome enters the cell cytoplasm.[41] The coronavirus RNA genome has a 5′ methylated cap and a 3′ polyadenylated tail, which allows the RNA to attach to the host cell's ribosome for translation.[41] The host ribosome translates the initial overlapping open reading frame of the virus genome and forms a long polyprotein. The polyprotein has its own proteases which cleave the polyprotein into multiple nonstructural proteins.[41]

Replication

A number of the nonstructural proteins coalesce to form a multi-protein replicase-transcriptase complex (RTC). The main replicase-transcriptase protein is the RNA-dependent RNA polymerase (RdRp). It is directly involved in the replication and transcription of RNA from an RNA strand. The other nonstructural proteins in the complex assist in the replication and transcription process. The exoribonuclease nonstructural protein, for instance, provides extra fidelity to replication by providing a proofreading function which the RNA-dependent RNA polymerase lacks.[47]

One of the main functions of the complex is to replicate the viral genome. RdRp directly mediates the synthesis of negative-sense genomic RNA from the positive-sense genomic RNA. This is followed by the replication of positive-sense genomic RNA from the negative-sense genomic RNA.[41] The other important function of the complex is to transcribe the viral genome. RdRp directly mediates the synthesis of negative-sense subgenomic RNA molecules from the positive-sense genomic RNA. This is followed by the transcription of these negative-sense subgenomic RNA molecules to their corresponding positive-sense mRNAs.[41]

Release

The replicated positive-sense genomic RNA becomes the genome of the progeny viruses. The mRNAs are gene transcripts of the last third of the virus genome after the initial overlapping reading frame. These mRNAs are translated by the host's ribosomes into the structural proteins and a number of accessory proteins.[41] RNA translation occurs inside the endoplasmic reticulum. The viral structural proteins S, E, and M move along the secretory pathway into the Golgi intermediate compartment. There, the M proteins direct most protein-protein interactions required for assembly of viruses following its binding to the nucleocapsid.[48] Progeny viruses are then released from the host cell by exocytosis through secretory vesicles.[48]

Transmission

The interaction of the coronavirus spike protein with its complement host cell receptor is central in determining the tissue tropism, infectivity, and species range of the virus.[49][50] The SARS coronavirus, for example, infects human cells by attaching to the angiotensin-converting enzyme 2 (ACE2) receptor.[51]

Classification
Phylogenetic tree of coronaviruses

The scientific name for coronavirus is Orthocoronavirinae or Coronavirinae.[2][3][4] Coronaviruses belong to the family of Coronaviridae, order Nidovirales, and realm Riboviria.[5][6] They are divided into alphacoronaviruses and betacoronaviruses which infect mammals – and gammacoronaviruses and deltacoronaviruses which primarily infect birds.[52][53]

  • Genus: Alphacoronavirus;[54] type species: Alphacoronavirus 1 (TGEV)
    • Species: Alphacoronavirus 1, Human coronavirus 229E, Human coronavirus NL63, Miniopterus bat coronavirus 1, Miniopterus bat coronavirus HKU8, Porcine epidemic diarrhea virus, Rhinolophus bat coronavirus HKU2, Scotophilus bat coronavirus 512
  • Genus Betacoronavirus;[55] type species: Murine coronavirus (MHV)
    • Species: Betacoronavirus 1 (Bovine Coronavirus, Human coronavirus OC43), Hedgehog coronavirus 1, Human coronavirus HKU1, Middle East respiratory syndrome-related coronavirus, Murine coronavirus, Pipistrellus bat coronavirus HKU5, Rousettus bat coronavirus HKU9, Severe acute respiratory syndrome-related coronavirus (SARS-CoV, SARS-CoV-2), Tylonycteris bat coronavirus HKU4
  • Genus Gammacoronavirus;[15] type species: Avian coronavirus (IBV)
    • Species: Avian coronavirus, Beluga whale coronavirus SW1
  • Genus Deltacoronavirus; type species: Bulbul coronavirus HKU11
    • Species: Bulbul coronavirus HKU11, Porcine coronavirus HKU15

Origin
Origins of human coronaviruses with possible intermediate hosts

The most recent common ancestor (MRCA) of all coronaviruses is estimated to have existed as recently as 8000 BCE, although some models place the common ancestor as far back as 55 million years or more, implying long term coevolution with bat and avian species.[56] The most recent common ancestor of the alphacoronavirus line has been placed at about 2400 BCE, the betacoronavirus line at 3300 BCE, the gammacoronavirus line at 2800 BCE, and the deltacoronavirus line at about 3000 BCE. Bats and birds, as warm-blooded flying vertebrates, are an ideal natural reservoir for the coronavirus gene pool (bats the reservoir for alphacoronavirus and betacoronavirus – and birds the reservoir for gammacoronavirus and deltacoronavirus). The large number of host bat and avian species, and their global range, has enabled extensive evolution and dissemination of coronaviruses.[57]

Many human coronavirus have their origin in bats.[58] The human coronavirus NL63 shared a common ancestor with a bat coronavirus (ARCoV.2) between 1190–1449 CE.[59] The human coronavirus 229E shared a common ancestor with bat coronavirus (GhanaGrp1 Bt CoV) between 1686–1800 CE.[60] More recently, alpaca coronavirus and human coronavirus 229E diverged sometime before 1960.[61] MERS-CoV emerged in humans from bats through the intermediate host of camels.[62] MERS-CoV, although related to several bat coronavirus species, appears to have diverged from these several centuries ago.[63] The most closely related bat coronavirus and SARS-CoV diverged in 1986.[64] A possible path of evolution, of SARS coronavirus and keen bat coronaviruses, suggests that SARS related coronaviruses coevolved in bats for a long time. The ancestors of SARS-CoV first infected leaf-nose bats of the genus Hipposideridae; subsequently, they spread to horseshoe bats in the species Rhinolophidae, and then to civets, and finally to humans.[65][66]

Unlike other betacoronaviruses, bovine coronavirus of the species Betacoronavirus 1 and subgenus Embecovirus is thought to have originated in rodents and not in bats.[58][67] In the 1790s, equine coronavirus diverged from the bovine coronavirus after a cross-species jump.[68] Later in the 1890s, human coronavirus OC43 diverged from bovine coronavirus after another cross-species spillover event.[69][68] It is speculated that the flu pandemic of 1890 may have been caused by this spillover event, and not by the influenza virus, because of the related timing, neurological symptoms, and unknown causative agent of the pandemic.[70] Human coronavirus OC43 besides causing respiratory infections is also suspected of playing a role in neurological diseases.[71] In the 1950s, the human coronavirus OC43 began to diverge into its present genotypes.[72] Phylogentically, mouse hepatitis virus (Murine coronavirus), which infects the mouse's liver and the central nervous system,[73] is related to human coronavirus OC43 and bovine coronavirus. Human coronavirus HKU1, like the aforementioned viruses, also has its origins in rodents.[58]

Infection in humans
Illustration of SARSr-CoV virion

Coronaviruses vary significantly in risk factor. Some can kill more than 30% of those infected, such as MERS-CoV, and some are relatively harmless, such as the common cold.[41] Coronaviruses can cause colds with major symptoms, such as fever, and a sore throat from swollen adenoids.[74] Coronaviruses can cause pneumonia (either direct viral pneumonia or secondary bacterial pneumonia) and bronchitis (either direct viral bronchitis or secondary bacterial bronchitis).[75] The human coronavirus discovered in 2003, SARS-CoV, which causes severe acute respiratory syndrome (SARS), has a unique pathogenesis because it causes both upper and lower respiratory tract infections.[75]

Six species of human coronaviruses are known, with one species subdivided into two different strains, making seven strains of human coronaviruses altogether. Four of these coronaviruses continually circulate in the human population and produce the generally mild symptoms of the common cold in adults and children worldwide: -OC43, -HKU1, HCoV-229E, -NL63.[76] Coronaviruses cause about 15% of commons colds.[77] The majority of colds are caused by rhinoviruses.[78] The four mild coronaviruses have a seasonal incidence occurring in the winter months in temperate climates.[79][80] There is no preference towards a particular season in tropical climates.[81]

Seasonal distribution of HCoV-NL63 in Germany shows a preferential detection from November to March

Four human coronaviruses produce symptoms that are generally mild:

  1. Human coronavirus OC43 (HCoV-OC43), β-CoV
  2. Human coronavirus HKU1 (HCoV-HKU1), β-CoV
  3. Human coronavirus 229E (HCoV-229E), α-CoV
  4. Human coronavirus NL63 (HCoV-NL63), α-CoV

Three human coronaviruses produce symptoms that are potentially severe:

  1. Middle East respiratory syndrome-related coronavirus (MERS-CoV), β-CoV
  2. Severe acute respiratory syndrome coronavirus (SARS-CoV), β-CoV
  3. Severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), β-CoV

Outbreaks of human coronavirus diseases

Severe acute respiratory syndrome (SARS)
Main article: Severe acute respiratory syndrome
Characteristics of human coronavirus strains
MERS-CoV, SARS-CoV, SARS-CoV-2,
and related diseases
MERS-CoVSARS-CoVSARS-CoV-2
DiseaseMERSSARSCOVID-19
Outbreaks2012, 2015,
2018		2002–20042019–2020
pandemic
Epidemiology
Date of first
identified case		June
2012		November
2002		December
2019[82]
Location of first
identified case		Jeddah,
Saudi Arabia		Shunde,
China		Wuhan,
China
Age average5644[83][lower-alpha 1]56[84]
Sex ratio (M:F)3.3:10.8:1[85]1.6:1[84]
Confirmed cases24948096[86]2,897,645[87][lower-alpha 2]
Deaths858774[86]202,880[87][lower-alpha 2]
Case fatality rate37%9.2%7.0%[87]
Symptoms
Fever98%99–100%87.9%[88]
Dry cough47%29–75%67.7%[88]
Dyspnea72%40–42%18.6%[88]
Diarrhea26%20–25%3.7%[88]
Sore throat21%13–25%13.9%[88]
Ventilatory use24.5%[89]14–20%4.1%[90]
Notes
  1. Based on data from Hong Kong.
  2. Data as of 26 April 2020.

In 2003, following the outbreak of severe acute respiratory syndrome (SARS) which had begun the prior year in Asia, and secondary cases elsewhere in the world, the World Health Organization (WHO) issued a press release stating that a novel coronavirus identified by a number of laboratories was the causative agent for SARS. The virus was officially named the SARS coronavirus (SARS-CoV). More than 8,000 people were infected, about ten percent of whom died.[51]

Middle East respiratory syndrome (MERS)

In September 2012, a new type of coronavirus was identified, initially called Novel Coronavirus 2012, and now officially named Middle East respiratory syndrome coronavirus (MERS-CoV).[91][92] The World Health Organization issued a global alert soon after.[93] The WHO update on 28 September 2012 said the virus did not seem to pass easily from person to person.[94] However, on 12 May 2013, a case of human-to-human transmission in France was confirmed by the French Ministry of Social Affairs and Health.[95] In addition, cases of human-to-human transmission were reported by the Ministry of Health in Tunisia. Two confirmed cases involved people who seemed to have caught the disease from their late father, who became ill after a visit to Qatar and Saudi Arabia. Despite this, it appears the virus had trouble spreading from human to human, as most individuals who are infected do not transmit the virus.[96] By 30 October 2013, there were 124 cases and 52 deaths in Saudi Arabia.[97]

After the Dutch Erasmus Medical Centre sequenced the virus, the virus was given a new name, Human Coronavirus—Erasmus Medical Centre (HCoV-EMC). The final name for the virus is Middle East respiratory syndrome coronavirus (MERS-CoV). The only U.S. cases (both survived) were recorded in May 2014.[98]

In May 2015, an outbreak of MERS-CoV occurred in the Republic of Korea, when a man who had traveled to the Middle East, visited four hospitals in the Seoul area to treat his illness. This caused one of the largest outbreaks of MERS-CoV outside the Middle East.[99] As of December 2019, 2,468 cases of MERS-CoV infection had been confirmed by laboratory tests, 851 of which were fatal, a mortality rate of approximately 34.5%.[100]

Coronavirus disease 2019 (COVID-19)
Main article: Coronavirus disease 2019

In December 2019, a pneumonia outbreak was reported in Wuhan, China.[101] On 31 December 2019, the outbreak was traced to a novel strain of coronavirus,[102] which was given the interim name 2019-nCoV by the World Health Organization (WHO),[103][104][105] later renamed SARS-CoV-2 by the International Committee on Taxonomy of Viruses.

As of 26 April 2020, there have been at least 202,880[87] confirmed deaths and more than 2,897,645[87] confirmed cases in the coronavirus pneumonia pandemic. The Wuhan strain has been identified as a new strain of Betacoronavirus from group 2B with approximately 70% genetic similarity to the SARS-CoV.[106] The virus has a 96% similarity to a bat coronavirus, so it is widely suspected to originate from bats as well.[107][108] The pandemic has resulted in travel restrictions and nationwide lockdowns in several countries.

Infection in other animals

Coronaviruses have been recognized as causing pathological conditions in veterinary medicine since the 1930s.[16] Except for avian infectious bronchitis, the major related diseases have mainly an intestinal location.[109]

Diseases caused

Coronaviruses primarily infect the upper respiratory and gastrointestinal tract of mammals and birds. They also cause a range of diseases in farm animals and domesticated pets, some of which can be serious and are a threat to the farming industry. In chickens, the infectious bronchitis virus (IBV), a coronavirus, targets not only the respiratory tract but also the urogenital tract. The virus can spread to different organs throughout the chicken.[110] Economically significant coronaviruses of farm animals include porcine coronavirus (transmissible gastroenteritis coronavirus, TGE) and bovine coronavirus, which both result in diarrhea in young animals. Feline coronavirus: two forms, feline enteric coronavirus is a pathogen of minor clinical significance, but spontaneous mutation of this virus can result in feline infectious peritonitis (FIP), a disease associated with high mortality. Similarly, there are two types of coronavirus that infect ferrets: Ferret enteric coronavirus causes a gastrointestinal syndrome known as epizootic catarrhal enteritis (ECE), and a more lethal systemic version of the virus (like FIP in cats) known as ferret systemic coronavirus (FSC).[111] There are two types of canine coronavirus (CCoV), one that causes mild gastrointestinal disease and one that has been found to cause respiratory disease. Mouse hepatitis virus (MHV) is a coronavirus that causes an epidemic murine illness with high mortality, especially among colonies of laboratory mice.[112] Sialodacryoadenitis virus (SDAV) is highly infectious coronavirus of laboratory rats, which can be transmitted between individuals by direct contact and indirectly by aerosol. Acute infections have high morbidity and tropism for the salivary, lachrymal and harderian glands.[113]

A HKU2-related bat coronavirus called swine acute diarrhea syndrome coronavirus (SADS-CoV) causes diarrhea in pigs.[114]

Prior to the discovery of SARS-CoV, MHV had been the best-studied coronavirus both in vivo and in vitro as well as at the molecular level. Some strains of MHV cause a progressive demyelinating encephalitis in mice which has been used as a murine model for multiple sclerosis. Significant research efforts have been focused on elucidating the viral pathogenesis of these animal coronaviruses, especially by virologists interested in veterinary and zoonotic diseases.[115]

Domestic animals
  • Infectious bronchitis virus (IBV) causes avian infectious bronchitis.
  • Porcine coronavirus (transmissible gastroenteritis coronavirus of pigs, TGEV).[116][117]
  • Bovine coronavirus (BCV), responsible for severe profuse enteritis in of young calves.
  • Feline coronavirus (FCoV) causes mild enteritis in cats as well as severe Feline infectious peritonitis (other variants of the same virus).
  • the two types of canine coronavirus (CCoV) (one causing enteritis, the other found in respiratory diseases).
  • Turkey coronavirus (TCV) causes enteritis in turkeys.
  • Ferret enteric coronavirus causes epizootic catarrhal enteritis in ferrets.
  • Ferret systemic coronavirus causes FIP-like systemic syndrome in ferrets.[118]
  • Pantropic canine coronavirus.
  • Rabbit enteric coronavirus causes acute gastrointestinal disease and diarrhea in young European rabbits. Mortality rates are high.[119]
  • Porcine epidemic diarrhea virus (PED or PEDV), has emerged around the world.[120]

See also

References
  1. "Virus Taxonomy: 2018b Release". International Committee on Taxonomy of Viruses (ICTV). March 2019. Archived from the original on 2018-03-04. Retrieved 2020-01-24.
  2. "2017.012-015S" (xlsx). International Committee on Taxonomy of Viruses (ICTV). October 2018. Archived from the original on 2019-05-14. Retrieved 2020-01-24.
  3. "ICTV Taxonomy history: Orthocoronavirinae". International Committee on Taxonomy of Viruses (ICTV). Retrieved 2020-01-24.
  4. Fan Y, Zhao K, Shi ZL, Zhou P (March 2019). "Bat Coronaviruses in China". Viruses. 11 (3): 210. doi:10.3390/v11030210. PMC 6466186. PMID 30832341.
  5. de Groot RJ, Baker SC, Baric R, Enjuanes L, Gorbalenya AE, Holmes KV, Perlman S, Poon L, Rottier PJ, Talbot PJ, Woo PC, Ziebuhr J (2011). "Family Coronaviridae". In King AM, Lefkowitz E, Adams MJ, Carstens EB, International Committee on Taxonomy of Viruses, International Union of Microbiological Societies. Virology Division (eds.). Ninth Report of the International Committee on Taxonomy of Viruses. Oxford: Elsevier. pp. 806–28. ISBN 978-0-12-384684-6.
  6. International Committee on Taxonomy of Viruses (2010-08-24). "ICTV Master Species List 2009—v10" (xls).
  7. Woo PC, Huang Y, Lau SK, Yuen KY (August 2010). "Coronavirus genomics and bioinformatics analysis". Viruses. 2 (8): 1804–20. doi:10.3390/v2081803. PMC 3185738. PMID 21994708. Coronaviruses possess the largest genomes [26.4 kb (ThCoV HKU12) to 31.7 kb (SW1)] among all known RNA viruses (Figure 1) [2,13,16].
  8. Almeida JD, Berry DM, Cunningham CH, Hamre D, Hofstad MS, Mallucci L, McIntosh K, Tyrrell DA (November 1968). "Virology: Coronaviruses". Nature. 220 (5168): 650. Bibcode:1968Natur.220..650.. doi:10.1038/220650b0. [T]here is also a characteristic "fringe" of projections 200 A long, which are rounded or petal shaped ... This appearance, recalling the solar corona, is shared by mouse hepatitis virus and several viruses recently recovered from man, namely strain B814, 229E and several others.
  9. Definition of Coronavirus by Merriam-Webster, Merriam-Webster, archived from the original on 2020-03-23, retrieved 2020-03-24
  10. Definition of Corona by Merriam-Webster, Merriam-Webster, archived from the original on 2020-03-24, retrieved 2020-03-24
  11. Tyrrell, David Arthur John; Fielder, Michael (2002). Cold Wars: The Fight Against the Common Cold. Oxford University Press. p. 96. ISBN 978-0-19-263285-2. We looked more closely at the appearance of the new viruses and noticed that they had a kind of halo surrounding them. Recourse to a dictionary produced the Latin equivalent, corona, and so the name coronavirus was born.
  12. Sturman LS, Holmes KV (1983-01-01). Lauffer MA, Maramorosch K (eds.). "The molecular biology of coronaviruses". Advances in Virus Research. 28: 35–112. doi:10.1016/s0065-3527(08)60721-6. ISBN 9780120398287. PMC 7131312. PMID 6362367. [T]hese viruses displayed a characteristic fringe of large, distinctive, petal-shaped peplomers or spikes which resembled a crown, like the corona spinarum in religious art; hence the name coronaviruses.
  13. Estola, T. (1970). "Coronaviruses, a New Group of Animal RNA Viruses". Avian Diseases. 14 (2): 330–336. doi:10.2307/1588476. ISSN 0005-2086. JSTOR 1588476.
  14. Fabricant, Julius (1998). "The Early History of Infectious Bronchitis". Avian Diseases. 42 (4): 648–650. doi:10.2307/1592697. ISSN 0005-2086. JSTOR 1592697.
  15. Decaro, Nicola (2011), Tidona, Christian; Darai, Gholamreza (eds.), "Gammacoronavirus", The Springer Index of Viruses, Springer, pp. 403–413, doi:10.1007/978-0-387-95919-1_58, ISBN 978-0-387-95919-1, PMC 7176155, retrieved 2020-04-25
  16. McIntosh K (1974). Arber W, Haas R, Henle W, Hofschneider PH, Jerne NK, Koldovský P, Koprowski H, Maaløe O, Rott R (eds.). "Coronaviruses: A Comparative Review". Current Topics in Microbiology and Immunology / Ergebnisse der Mikrobiologie und Immunitätsforschung. Berlin, Heidelberg: Springer: 87. doi:10.1007/978-3-642-65775-7_3. ISBN 978-3-642-65775-7. Cite journal requires |journal= (help)
  17. "Il était une fois les coronavirus". Réalités Biomédicales (in French). 2020-03-27. Retrieved 2020-04-18.
  18. Kahn JS, McIntosh K (November 2005). "History and recent advances in coronavirus discovery". The Pediatric Infectious Disease Journal. 24 (11 Suppl): S223–7, discussion S226. doi:10.1097/01.inf.0000188166.17324.60. PMID 16378050.
  19. Mahase, Elisabeth (2020-04-16). "Covid-19: First coronavirus was described in The BMJ in 1965". BMJ. 369: m1547. doi:10.1136/bmj.m1547. ISSN 1756-1833. PMID 32299810.
  20. Monto, Arnold S. (1984). "Coronaviruses". In Evans, Alfred S. (ed.). Viral Infections of Humans. Viral Infections of Humans: Epidemiology and Control. Springer US. pp. 151–165. doi:10.1007/978-1-4684-4727-9_7. ISBN 978-1-4684-4727-9.
  21. Kendall, E. J.; Bynoe, M. L.; Tyrrell, D. A. (1962-07-14). "Virus isolations from common colds occurring in a residential school". British Medical Journal. 2 (5297): 82–86. doi:10.1136/bmj.2.5297.82. ISSN 0007-1447. PMC 1925312. PMID 14455113.
  22. Richmond, Caroline (2005-06-18). "David Tyrrell". BMJ : British Medical Journal. 330 (7505): 1451. doi:10.1136/bmj.330.7505.1451. ISSN 0959-8138. PMC 558394.
  23. Group, British Medical Journal Publishing (1969-06-28). "Obituary Notices". Br Med J. 2 (5660): 827–829. doi:10.1136/bmj.2.5660.827. ISSN 0007-1447.
  24. Tyrrell, D. a. J.; Bynoe, M. L. (1965-06-05). "Cultivation of a Novel Type of Common-cold Virus in Organ Cultures". Br Med J. 1 (5448): 1467–1470. doi:10.1136/bmj.1.5448.1467. ISSN 0007-1447. PMC 2166670. PMID 14288084.
  25. Tyrrell, David Arthur John; Fielder, Michael (2002). Cold Wars: The Fight Against the Common Cold. Oxford University Press. pp. 93–95. ISBN 978-0-19-263285-2.
  26. Hagan, William Arthur; Bruner, Dorsey William; Gillespie, James Howard; Timoney, John Francis; Scott, Fredric W.; Barlough, Jeffrey E. (1988). Hagan and Bruner's Microbiology and Infectious Diseases of Domestic Animals: With Reference to Etiology, Epizootiology, Pathogenesis, Immunity, Diagnosis, and Antimicrobial Susceptibility. Cornell University Press. p. 440. ISBN 978-0-8014-1896-9.
  27. Hamre, Dorothy; Procknow, John J. (1966-01-01). "A New Virus Isolated from the Human Respiratory Tract". Proceedings of the Society for Experimental Biology and Medicine. 121 (1): 190–193. doi:10.3181/00379727-121-30734. ISSN 0037-9727. PMID 4285768.
  28. "The woman who discovered the first coronavirus".
  29. Almeida, Joyce (2008-06-26). "June Almeida (née Hart)". BMJ. 336 (7659): 1511.1–1511. doi:10.1136/bmj.a434. ISSN 0959-8138. PMC 2440895.
  30. Almeida, June D.; Tyrrell, D. A. J. (1967). "The Morphology of Three Previously Uncharacterized Human Respiratory Viruses that Grow in Organ Culture". Journal of General Virology. 1 (2): 175–178. doi:10.1099/0022-1317-1-2-175. ISSN 0022-1317. PMID 4293939.
  31. McIntosh, K; Becker, W B; Chanock, R M (December 1967). "Growth in suckling-mouse brain of "IBV-like" viruses from patients with upper respiratory tract disease". Proceedings of the National Academy of Sciences of the United States of America. 58 (6): 2268–2273. Bibcode:1967PNAS...58.2268M. doi:10.1073/pnas.58.6.2268. ISSN 0027-8424. PMC 223830. PMID 4298953.
  32. McIntosh, K; Dees, J H; Becker, W B; Kapikian, A Z; Chanock, R M (April 1967). "Recovery in tracheal organ cultures of novel viruses from patients with respiratory disease". Proceedings of the National Academy of Sciences of the United States of America. 57 (4): 933–940. Bibcode:1967PNAS...57..933M. doi:10.1073/pnas.57.4.933. ISSN 0027-8424. PMC 224637. PMID 5231356.
  33. Times, Harold M. Schmeck Jr Special To the New York (1967-05-05). "Six Newly Discovered Viruses May Explain Cold; Strains Are Similar to Germ That Causes a Bronchial Infection in Chickens Believed to Be New Group". The New York Times. ISSN 0362-4331. Retrieved 2020-04-25.
  34. Myint, Steven H. (1995), Siddell, Stuart G. (ed.), The Coronaviridae, The Viruses, Springer US, pp. 389–401, doi:10.1007/978-1-4899-1531-3_18, ISBN 978-1-4899-1531-3 Missing or empty |title= (help); |chapter= ignored (help)
  35. Geller C, Varbanov M, Duval RE (November 2012). "Human coronaviruses: insights into environmental resistance and its influence on the development of new antiseptic strategies". Viruses. 4 (11): 3044–68. doi:10.3390/v4113044. PMC 3509683. PMID 23202515.
  36. Monto, Arnold S.; Cowling, Benjamin J.; Peiris, J. S. Malik (2014-02-27). "Coronaviruses". Viral Infections of Humans: 199–223. doi:10.1007/978-1-4899-7448-8_10. ISBN 978-1-4899-7447-1. PMC 7122465. The other OC strains and B814 that could not be adapted to mouse brain resisted adaptation to cell culture as well; these distinct viruses have since been lost and may actually have been rediscovered recently.
  37. Su S, Wong G, Shi W, Liu J, Lai AC, Zhou J, et al. (June 2016). "Epidemiology, Genetic Recombination, and Pathogenesis of Coronaviruses". Trends in Microbiology. 24 (6): 490–502. doi:10.1016/j.tim.2016.03.003. PMC 7125511. PMID 27012512.
  38. Zhu N, Zhang D, Wang W, Li X, Yang B, Song J, et al. (February 2020). "A Novel Coronavirus from Patients with Pneumonia in China, 2019". The New England Journal of Medicine. 382 (8): 727–733. doi:10.1056/NEJMoa2001017. PMC 7092803. PMID 31978945.
  39. Goldsmith CS, Tatti KM, Ksiazek TG, Rollin PE, Comer JA, Lee WW, et al. (February 2004). "Ultrastructural characterization of SARS coronavirus". Emerging Infectious Diseases. 10 (2): 320–26. doi:10.3201/eid1002.030913. PMC 3322934. PMID 15030705. Virions acquired an envelope by budding into the cisternae and formed mostly spherical, sometimes pleomorphic, particles that averaged 78 nm in diameter (Figure 1A).
  40. Neuman BW, Adair BD, Yoshioka C, Quispe JD, Orca G, Kuhn P, et al. (August 2006). "Supramolecular architecture of severe acute respiratory syndrome coronavirus revealed by electron cryomicroscopy". Journal of Virology. 80 (16): 7918–28. doi:10.1128/JVI.00645-06. PMC 1563832. PMID 16873249. Particle diameters ranged from 50 to 150 nm, excluding the spikes, with mean particle diameters of 82 to 94 nm; Also See Figure 1 for double shell.
  41. Fehr AR, Perlman S (2015). "Coronaviruses: an overview of their replication and pathogenesis". In Maier HJ, Bickerton E, Britton P (eds.). Coronaviruses. Methods in Molecular Biology. 1282. Springer. pp. 1–23. doi:10.1007/978-1-4939-2438-7_1. ISBN 978-1-4939-2438-7. PMC 4369385. PMID 25720466. See section: Virion Structure.
  42. Lai MM, Cavanagh D (1997). "The molecular biology of coronaviruses". Advances in Virus Research. 48: 1–100. doi:10.1016/S0065-3527(08)60286-9. ISBN 9780120398485. PMC 7130985. PMID 9233431.
  43. Chang CK, Hou MH, Chang CF, Hsiao CD, Huang TH (March 2014). "The SARS coronavirus nucleocapsid protein—forms and functions". Antiviral Research. 103: 39–50. doi:10.1016/j.antiviral.2013.12.009. PMC 7113676. PMID 24418573. See Figure 4c.
  44. Neuman BW, Kiss G, Kunding AH, Bhella D, Baksh MF, Connelly S, et al. (April 2011). "A structural analysis of M protein in coronavirus assembly and morphology". Journal of Structural Biology. 174 (1): 11–22. doi:10.1016/j.jsb.2010.11.021. PMC 4486061. PMID 21130884. See Figure 10.
  45. Snijder EJ, Bredenbeek PJ, Dobbe JC, Thiel V, Ziebuhr J, Poon LL, et al. (August 2003). "Unique and conserved features of genome and proteome of SARS-coronavirus, an early split-off from the coronavirus group 2 lineage". Journal of Molecular Biology. 331 (5): 991–1004. doi:10.1016/S0022-2836(03)00865-9. PMC 7159028. PMID 12927536. See Figure 1.
  46. Simmons G, Zmora P, Gierer S, Heurich A, Pöhlmann S (December 2013). "Proteolytic activation of the SARS-coronavirus spike protein: cutting enzymes at the cutting edge of antiviral research". Antiviral Research. 100 (3): 605–14. doi:10.1016/j.antiviral.2013.09.028. PMC 3889862. PMID 24121034. See Figure 2.
  47. Sexton NR, Smith EC, Blanc H, Vignuzzi M, Peersen OB, Denison MR (August 2016). "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. doi:10.1128/JVI.00080-16. PMC 4984655. PMID 27279608. Finally, these results, combined with those from previous work (33, 44), suggest that CoVs encode at least three proteins involved in fidelity (nsp12-RdRp, nsp14-ExoN, and nsp10), supporting the assembly of a multiprotein replicase-fidelity complex, as described previously (38).
  48. Fehr AR, Perlman S (2015). "Coronaviruses: an overview of their replication and pathogenesis". In Maier HJ, Bickerton E, Britton P (eds.). Coronaviruses. Methods in Molecular Biology. 1282. Springer. pp. 1–23. doi:10.1007/978-1-4939-2438-7_1. ISBN 978-1-4939-2438-7. PMC 4369385. PMID 25720466. See section: Coronavirus Life Cycle—Assembly and Release
  49. Masters PS (2006-01-01). "The molecular biology of coronaviruses". Advances in Virus Research. Academic Press. 66: 193–292. doi:10.1016/S0065-3527(06)66005-3. ISBN 978-0120398690. PMC 7112330. PMID 16877062. Nevertheless, the interaction between S protein and receptor remains the principal, if not sole, determinant of coronavirus host species range and tissue tropism.
  50. Cui J, Li F, Shi ZL (March 2019). "Origin and evolution of pathogenic coronaviruses". Nature Reviews. Microbiology. 17 (3): 181–92. doi:10.1038/s41579-018-0118-9. PMC 7097006. PMID 30531947. Different SARS-CoV strains isolated from several hosts vary in their binding affinities for human ACE2 and consequently in their infectivity of human cells 76, 78 (Fig. 6b)
  51. Li F, Li W, Farzan M, Harrison SC (September 2005). "Structure of SARS coronavirus spike receptor-binding domain complexed with receptor". Science. 309 (5742): 1864–68. Bibcode:2005Sci...309.1864L. doi:10.1126/science.1116480. PMID 16166518.
  52. Wertheim JO, Chu DK, Peiris JS, Kosakovsky Pond SL, Poon LL (June 2013). "A case for the ancient origin of coronaviruses". Journal of Virology. 87 (12): 7039–45. doi:10.1128/JVI.03273-12. PMC 3676139. PMID 23596293. Alphacoronaviruses and betacoronaviruses are found exclusively in mammals, whereas gammacoronaviruses and deltacoronaviruses primarily infect birds.
  53. Nextstrain, phylogenetic tree of Beta-CoV
  54. Decaro, Nicola (2011), Tidona, Christian; Darai, Gholamreza (eds.), "Alphacoronavirus", The Springer Index of Viruses, Springer, pp. 371–383, doi:10.1007/978-0-387-95919-1_56, ISBN 978-0-387-95919-1, PMC 7176201, retrieved 2020-04-25
  55. Decaro, Nicola (2011), Tidona, Christian; Darai, Gholamreza (eds.), "Betacoronavirus", The Springer Index of Viruses, Springer, pp. 385–401, doi:10.1007/978-0-387-95919-1_57, ISBN 978-0-387-95919-1, PMC 7176184, retrieved 2020-04-25
  56. Wertheim JO, Chu DK, Peiris JS, Kosakovsky Pond SL, Poon LL (June 2013). "A case for the ancient origin of coronaviruses". Journal of Virology. 87 (12): 7039–45. doi:10.1128/JVI.03273-12. PMC 3676139. PMID 23596293.
  57. Woo PC, Lau SK, Lam CS, Lau CC, Tsang AK, Lau JH, et al. (April 2012). "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. doi:10.1128/JVI.06540-11. PMC 3302495. PMID 22278237.
  58. Forni D, Cagliani R, Clerici M, Sironi M (January 2017). "Molecular Evolution of Human Coronavirus Genomes". Trends in Microbiology. 25 (1): 35–48. doi:10.1016/j.tim.2016.09.001. PMC 7111218. PMID 27743750. 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].
  59. Huynh J, Li S, Yount B, Smith A, Sturges L, Olsen JC, et al. (December 2012). "Evidence supporting a zoonotic origin of human coronavirus strain NL63". Journal of Virology. 86 (23): 12816–25. doi:10.1128/JVI.00906-12. PMC 3497669. PMID 22993147. If these predictions are correct, this observation suggests that HCoV-NL63 may have originated from bats between 1190 and 1449 CE.
  60. Pfefferle S, Oppong S, Drexler JF, Gloza-Rausch F, Ipsen A, Seebens A, et al. (September 2009). "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. doi:10.3201/eid1509.090224. PMC 2819850. PMID 19788804. The most recent common ancestor of hCoV-229E and GhanaBt-CoVGrp1 existed in ≈1686–1800 AD.
  61. Crossley BM, Mock RE, Callison SA, Hietala SK (December 2012). "Identification and characterization of a novel alpaca respiratory coronavirus most closely related to the human coronavirus 229E". Viruses. 4 (12): 3689–700. doi:10.3390/v4123689. PMC 3528286. PMID 23235471.
  62. Forni D, Cagliani R, Clerici M, Sironi M (January 2017). "Molecular Evolution of Human Coronavirus Genomes". Trends in Microbiology. 25 (1): 35–48. doi:10.1016/j.tim.2016.09.001. PMC 7111218. PMID 27743750.
  63. Lau SK, Li KS, Tsang AK, Lam CS, Ahmed S, Chen H, et al. (August 2013). "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. doi:10.1128/JVI.01055-13. PMC 3719811. PMID 23720729.
  64. Vijaykrishna D, Smith GJ, Zhang JX, Peiris JS, Chen H, Guan Y (April 2007). "Evolutionary insights into the ecology of coronaviruses". Journal of Virology. 81 (8): 4012–20. doi:10.1128/jvi.02605-06. PMC 1866124. PMID 17267506.
  65. Gouilh MA, Puechmaille SJ, Gonzalez JP, Teeling E, Kittayapong P, Manuguerra JC (October 2011). "SARS-Coronavirus ancestor's foot-prints in South-East Asian bat colonies and the refuge theory". Infection, Genetics and Evolution. 11 (7): 1690–702. doi:10.1016/j.meegid.2011.06.021. PMC 7106191. PMID 21763784.
  66. Cui J, Han N, Streicker D, Li G, Tang X, Shi Z, et al. (October 2007). "Evolutionary relationships between bat coronaviruses and their hosts". Emerging Infectious Diseases. 13 (10): 1526–32. doi:10.3201/eid1310.070448. PMC 2851503. PMID 18258002.
  67. Lau SK, Woo PC, Li KS, Tsang AK, Fan RY, Luk HK, et al. (March 2015). "Discovery of a novel coronavirus, China Rattus coronavirus HKU24, from Norway rats supports the murine origin of Betacoronavirus 1 and has implications for the ancestor of Betacoronavirus lineage A". Journal of Virology. 89 (6): 3076–92. doi:10.1128/JVI.02420-14. PMC 4337523. PMID 25552712.
  68. Bidokhti MR, Tråvén M, Krishna NK, Munir M, Belák S, Alenius S, Cortey M (September 2013). "Evolutionary dynamics of bovine coronaviruses: natural selection pattern of the spike gene implies adaptive evolution of the strains". The Journal of General Virology. 94 (Pt 9): 2036–2049. doi:10.1099/vir.0.054940-0. PMID 23804565. See Table 1
  69. Vijgen L, Keyaerts E, Moës E, Thoelen I, Wollants E, Lemey P, et al. (February 2005). "Complete genomic sequence of human coronavirus OC43: molecular clock analysis suggests a relatively recent zoonotic coronavirus transmission event". Journal of Virology. 79 (3): 1595–604. doi:10.1128/jvi.79.3.1595-1604.2005. PMC 544107. PMID 15650185.
  70. Vijgen L, Keyaerts E, Moës E, Thoelen I, Wollants E, Lemey P, et al. (February 2005). "Complete genomic sequence of human coronavirus OC43: molecular clock analysis suggests a relatively recent zoonotic coronavirus transmission event". Journal of Virology. 79 (3): 1595–604. doi:10.1128/JVI.79.3.1595-1604.2005. PMC 544107. PMID 15650185. However, it is tempting to speculate about an alternative hypothesis, that the 1889-1890 pandemic may have been the result of interspecies transmission of bovine coronaviruses to humans, resulting in the subsequent emergence of HCoV-OC43.
  71. Corman VM, Muth D, Niemeyer D, Drosten C (2018). "Hosts and Sources of Endemic Human Coronaviruses". Advances in Virus Research. 100: 163–188. doi:10.1016/bs.aivir.2018.01.001. ISBN 9780128152010. PMC 7112090. PMID 29551135.
  72. Lau SK, Lee P, Tsang AK, Yip CC, Tse H, Lee RA, et al. (November 2011). "Molecular epidemiology of human coronavirus OC43 reveals evolution of different genotypes over time and recent emergence of a novel genotype due to natural recombination". Journal of Virology. 85 (21): 11325–37. doi:10.1128/JVI.05512-11. PMC 3194943. PMID 21849456.
  73. Schaumburg CS, Held KS, Lane TE (May 2008). "Mouse hepatitis virus infection of the CNS: a model for defense, disease, and repair". Frontiers in Bioscience. 13 (13): 4393–406. doi:10.2741/3012. PMC 5025298. PMID 18508518.
  74. Liu P, Shi L, Zhang W, He J, Liu C, Zhao C, et al. (November 2017). "Prevalence and genetic diversity analysis of human coronaviruses among cross-border children". Virology Journal. 14 (1): 230. doi:10.1186/s12985-017-0896-0. PMC 5700739. PMID 29166910.
  75. Forgie S, Marrie TJ (February 2009). "Healthcare-associated atypical pneumonia". Seminars in Respiratory and Critical Care Medicine. 30 (1): 67–85. doi:10.1055/s-0028-1119811. PMID 19199189.
  76. Corman VM, Muth D, Niemeyer D, Drosten C (2018). "Hosts and Sources of Endemic Human Coronaviruses". Advances in Virus Research. 100: 163–188. doi:10.1016/bs.aivir.2018.01.001. ISBN 978-0-12-815201-0. PMID 29551135.
  77. Pelczar (2010). Microbiology: Application Based Approach. p. 656. ISBN 978-0-07-015147-5. Archived from the original on 2016-05-16.
  78. Russell La Fayette Cecil; Lee Goldman; Andrew I. Schafer (2012), Goldman's Cecil Medicine, Expert Consult Premium Edition (24 ed.), Elsevier Health Sciences, pp. 2103–, ISBN 978-1-4377-1604-7, archived from the original on 2016-05-04
  79. Charlton CL, Babady E, Ginocchio CC, Hatchette TF, Jerris RC, Li Y, et al. (January 2019). "Practical Guidance for Clinical Microbiology Laboratories: Viruses Causing Acute Respiratory Tract Infections". Clinical Microbiology Reviews. 32 (1). doi:10.1128/CMR.00042-18. PMC 6302358. PMID 30541871. See Figure 1.
  80. Monto AS, DeJonge P, Callear AP, Bazzi LA, Capriola S, Malosh RE, et al. (April 2020). "Coronavirus occurrence and transmission over 8 years in the HIVE cohort of households in Michigan". The Journal of Infectious Diseases: jiaa161. doi:10.1093/infdis/jiaa161. PMID 32246136.
  81. Abdul-Rasool S, Fielding BC (May 2010). "Understanding Human Coronavirus HCoV-NL63". The Open Virology Journal. 4: 76–84. doi:10.2174/1874357901004010076. PMC 2918871. PMID 20700397.
  82. Wang C, Horby PW, Hayden FG, Gao GF (February 2020). "A novel coronavirus outbreak of global health concern". Lancet. 395 (10223): 470–473. doi:10.1016/S0140-6736(20)30185-9. PMID 31986257.
  83. Lau EH, Hsiung CA, Cowling BJ, Chen CH, Ho LM, Tsang T, et al. (March 2010). "A comparative epidemiologic analysis of SARS in Hong Kong, Beijing and Taiwan". BMC Infectious Diseases. 10: 50. doi:10.1186/1471-2334-10-50. PMC 2846944. PMID 20205928.
  84. "Old age, sepsis tied to poor COVID-19 outcomes, death". CIDRAP, University of Minnesota. Retrieved 2020-03-29.
  85. Karlberg J, Chong DS, Lai WY (February 2004). "Do men have a higher case fatality rate of severe acute respiratory syndrome than women do?". American Journal of Epidemiology. 159 (3): 229–31. doi:10.1093/aje/kwh056. PMID 14742282.
  86. "Summary of probable SARS cases with onset of illness from 1 November 2002 to 31 July 2003". World Health Organization. April 2004.
  87. "COVID-19 Dashboard by the Center for Systems Science and Engineering (CSSE) at Johns Hopkins University (JHU)". ArcGIS. Johns Hopkins University. Retrieved 2020-04-25.
  88. "Report of the WHO-China Joint Mission on Coronavirus Disease 2019 (COVID-19)" (PDF). World Health Organization. February 2020.
  89. Oh MD, Park WB, Park SW, Choe PG, Bang JH, Song KH, et al. (March 2018). "Middle East respiratory syndrome: what we learned from the 2015 outbreak in the Republic of Korea". The Korean Journal of Internal Medicine. 33 (2): 233–246. doi:10.3904/kjim.2018.031. PMC 5840604. PMID 29506344.
  90. Ñamendys-Silva SA (March 2020). "Respiratory support for patients with COVID-19 infection". The Lancet. Respiratory Medicine. doi:10.1016/S2213-2600(20)30110-7. PMID 32145829.
  91. Doucleef M (2012-09-26). "Scientists Go Deep On Genes Of SARS-Like Virus". Associated Press. Archived from the original on 2012-09-27. Retrieved 2012-09-27.
  92. Falco M (2012-09-24). "New SARS-like virus poses medical mystery". CNN Health. Archived from the original on 2013-11-01. Retrieved 2013-03-16.
  93. "New SARS-like virus found in Middle East". Al-Jazeera. 2012-09-24. Archived from the original on 2013-03-09. Retrieved 2013-03-16.
  94. Kelland K (2012-09-28). "New virus not spreading easily between people: WHO". Reuters. Archived from the original on 2012-11-24. Retrieved 2013-03-16.
  95. Nouveau coronavirus—Point de situation : Un nouveau cas d'infection confirmé Archived 8 June 2013 at the Wayback Machine (Novel coronavirus—Status report: A new case of confirmed infection) 12 May 2013, social-sante.gouv.fr
  96. "MERS Transmission". Centers for Disease Control and Prevention (CDC). 2019-08-02. Archived from the original on 2019-12-07. Retrieved 2019-12-10.
  97. "Novel coronavirus infection". World Health Association. 2013-05-22. Archived from the original on 2013-06-07. Retrieved 2013-05-23.
  98. "MERS in the U.S." Center for Disease Control. 2019-08-02. Archived from the original on 2019-12-15. Retrieved 2019-12-10.
  99. Sang-Hun C (2015-06-08). "MERS Virus's Path: One Man, Many South Korean Hospitals". The New York Times. Archived from the original on 2017-07-15. Retrieved 2017-03-01.
  100. "Middle East respiratory syndrome coronavirus (MERS-CoV)". WHO. Archived from the original on 2019-10-18. Retrieved 2019-12-10.
  101. The Editorial Board (2020-01-29). "Is the World Ready for the Coronavirus?—Distrust in science and institutions could be a major problem if the outbreak worsens". The New York Times. Retrieved 2020-01-30.
  102. "WHO Statement Regarding Cluster of Pneumonia Cases in Wuhan, China". www.who.int. 2020-01-09. Archived from the original on 2020-01-14. Retrieved 2020-01-10.
  103. "Laboratory testing of human suspected cases of novel coronavirus (nCoV) infection. Interim guidance, 10 January 2020" (PDF). Archived (PDF) from the original on 2020-01-20. Retrieved 2020-01-14.
  104. "Novel Coronavirus 2019, Wuhan, China". www.cdc.gov (CDC). 2020-01-23. Archived from the original on 2020-01-20. Retrieved 2020-01-23.
  105. "2019 Novel Coronavirus infection (Wuhan, China): Outbreak update". Canada.ca. 2020-01-21.
  106. Hui DS, I Azhar E, Madani TA, Ntoumi F, Kock R, Dar O, et al. (February 2020). "The continuing 2019-nCoV epidemic threat of novel coronaviruses to global health—The latest 2019 novel coronavirus outbreak in Wuhan, China". International Journal of Infectious Diseases. 91: 264–66. doi:10.1016/j.ijid.2020.01.009. PMC 7128332. PMID 31953166.
  107. Cohen J (2020-01-26). "Wuhan seafood market may not be source of novel virus spreading globally". ScienceMag American Association for the Advancement of Science. (AAAS). Archived from the original on 2020-01-27. Retrieved 2020-01-29.
  108. Eschner K (2020-01-28). "We're still not sure where the COVID-19 really came from". Popular Science. Archived from the original on 2020-01-30. Retrieved 2020-01-30.
  109. Murphy FA, Gibbs EP, Horzinek MC, Studdart MJ (1999). Veterinary Virology. Boston: Academic Press. pp. 495–508. ISBN 978-0-12-511340-3.
  110. Bande F, Arshad SS, Bejo MH, Moeini H, Omar AR (2015). "Progress and challenges toward the development of vaccines against avian infectious bronchitis". Journal of Immunology Research. 2015: 424860. doi:10.1155/2015/424860. PMC 4411447. PMID 25954763.
  111. Murray J (2014-04-16). "What's New With Ferret FIP-like Disease?" (xls). Archived from the original on 2014-04-24. Retrieved 2014-04-24.
  112. Weiss SR, Navas-Martin S (December 2005). "Coronavirus pathogenesis and the emerging pathogen severe acute respiratory syndrome coronavirus". Microbiology and Molecular Biology Reviews. 69 (4): 635–64. doi:10.1128/MMBR.69.4.635-664.2005. PMC 1306801. PMID 16339739.
  113. "Rat Coronavirus—an overview". www.ScienceDirect.com Topics.
  114. Zhou P, Fan H, Lan T, Yang XL, Shi WF, Zhang W, et al. (April 2018). "Fatal swine acute diarrhoea syndrome caused by an HKU2-related coronavirus of bat origin". Nature. 556 (7700): 255–58. Bibcode:2018Natur.556..255Z. doi:10.1038/s41586-018-0010-9. PMC 7094983. PMID 29618817.
  115. Tirotta E, Carbajal KS, Schaumburg CS, Whitman L, Lane TE (July 2010). "Cell replacement therapies to promote remyelination in a viral model of demyelination". Journal of Neuroimmunology. 224 (1–2): 101–07. doi:10.1016/j.jneuroim.2010.05.013. PMC 2919340. PMID 20627412.
  116. Cruz JL, Sola I, Becares M, Alberca B, Plana J, Enjuanes L, Zuñiga S (June 2011). "Coronavirus gene 7 counteracts host defenses and modulates virus virulence". PLOS Pathogens. 7 (6): e1002090. doi:10.1371/journal.ppat.1002090. PMC 3111541. PMID 21695242.
  117. Cruz JL, Becares M, Sola I, Oliveros JC, Enjuanes L, Zúñiga S (September 2013). "Alphacoronavirus protein 7 modulates host innate immune response". Journal of Virology. 87 (17): 9754–67. doi:10.1128/JVI.01032-13. PMC 3754097. PMID 23824792.
  118. "Merck Veterinary Manual". Merck Veterinary Manual. Archived from the original on 2019-12-13. Retrieved 2020-01-24.
  119. "Enteric Coronavirus". Diseases of Research Animals. Archived from the original on 2019-07-01. Retrieved 2020-01-24.
  120. Wei X, She G, Wu T, Xue C, Cao Y (February 2020). "PEDV enters cells through clathrin-, caveolae-, and lipid raft-mediated endocytosis and traffics via the endo-/lysosome pathway". Veterinary Research. 51 (1): 10. doi:10.1186/s13567-020-0739-7. PMC 7011528. PMID 32041637.

Further reading
Classification
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