Coronavirus - Wikipedia https://en.wikipedia.

org/wiki/Coronavirus

Coronavirus
Coronaviruses are a group of related viruses that
cause diseases in mammals and birds. In humans, Orthocoronavirinae
coronaviruses cause respiratory tract infections that
can range from mild to lethal. Mild illnesses include
some cases of the common cold (which has other
possible causes, 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 yet to be vaccines or antiviral drugs to
Transmission electron micrograph
prevent or treat human coronavirus infections.
(TEM) of avian infectious bronchitis
Coronaviruses constitute the subfamily virus
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]

Illustration of the morphology of

Contents coronaviruses; the club-shaped viral
spike peplomers, colored red, create
Discovery
the look of a corona surrounding the
Etymology
virion when observed with an electron
Structure microscope.
Genome
Virus classification
Life cycle
(unranked): Virus
Entry
Replication Realm: Riboviria
Release Phylum: incertae sedis
Transmission Order: Nidovirales
Taxonomy Family: Coronaviridae

1 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

Evolution Subfamily: Orthocoronavirinae

Human coronaviruses Genera[1]
Outbreaks of human coronavirus
diseases Alphacoronavirus
Severe acute respiratory Betacoronavirus
syndrome (SARS)
Gammacoronavirus
Middle East respiratory syndrome
(MERS) Deltacoronavirus
Coronavirus disease 2019 Synonyms[2][3][4]
(COVID-19)
Other animals Coronavirinae
Diseases caused
Domestic animals
See also
References
Further reading

Discovery
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).[9] 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%.[10] Fred Beaudette and Charles Hudson six years later successfully isolated and cultivated
the infectious bronchitis virus which caused the disease.[11] In the 1940s, two more animal
coronaviruses, mouse hepatitis virus (MHV) and transmissible gastroenteritis virus (TGEV), were
isolated.[12] It was not realized at the time that these three different viruses were related.[13]

Human coronaviruses were discovered in the 1960s.[14][15] They were isolated using two different
methods in the United Kingdom and the United States.[16] E.C. Kendall, David Tyrrell and Malcom
Byone working at the Common Cold Unit of the British Medical Research Council in 1960 isolated
from a boy a novel common cold virus B814.[17][18][19] The virus was not able to be cultivated using
standards techniques which had successfully cultivated rhinoviruses, adenoviruses and other known
common cold viruses. In 1965, Tyrrell and Byone successfully cultivated the novel virus by serial
passaging it through organ culture of human embryonic trachea.[20] The isolated virus when
intranasally inoculated into volunteers caused a cold and was inactivated by ether which indicated it
had a lipid envelope.[17][21] 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.[22]

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.[23][24] Almeida through electron

2 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

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).[25] 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).[26] Like B814, 229E, and IBV, the novel cold virus
OC43 had distinctive club-like spikes when observed with the electron microscope.[27][28]

The IBV-like novel cold viruses were soon shown to be also morphologically related to the mouse
hepatitis virus.[12] 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.[29][30] The coronavirus strain B814 was lost. It is not
known which present human coronavirus it was.[31] Other human coronaviruses have since been
identified, including SARS-CoV in 2003, HCoV NL63 in 2004, HKU1 in 2005, MERS-CoV in 2012,
and SARS-CoV-2 in 2019.[32][33] There have also been a large number of animal coronaviruses
identified since the 1960s.[5]

Etymology
The name "coronavirus" is derived from Latin corona, meaning "crown" or "wreath", itself a
borrowing from Greek κορώνη korṓnē, "garland, wreath".[34][35] The name was first used in 1968 by
an informal group of virologists in the journal Nature to designate the new family of viruses. 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 a crown or of a solar corona. This morphology is created by the viral spike peplomers, which are
proteins on the surface of the virus.[8][36]

Structure
Coronaviruses are large pleomorphic spherical particles with
bulbous surface projections.[37] 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.
[38][39]
Cross-sectional model of a
The viral envelope consists of a lipid bilayer where the coronavirus
membrane (M), envelope (E) and spike (S) structural proteins
are anchored.[40] 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.[39][41] The lipid bilayer envelope, membrane
proteins, and nucleocapsid protect the virus when it is outside the host cell.[42]

3 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

Genome
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.[39]

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 Schematic representation of the
frames 1a and 1b, which occupy the first two-thirds of the genome organization and
genome, encode the replicase/transcriptase polyprotein. The functional domains of S protein
replicase/transcriptase polyprotein self cleaves to form for SARS-CoV and MERS-CoV
nonstructural proteins.[39]

The later reading frames encode the four major structural proteins: spike, envelope, membrane, and
nucleocapsid.[43] 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.[39]

Life cycle

Entry

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.[44]

On entry into the host cell, the virus particle is uncoated, and
its genome enters the cell cytoplasm.[39] The coronavirus The life cycle of a 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.[39] 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.[39]

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

4 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

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.[45]

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.[39] 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.[39]

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.[39] 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.[46] Progeny viruses are then released from the host
cell by exocytosis through secretory vesicles.[46]

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.[47][48] The SARS
coronavirus, for example, infects human cells by attaching to the angiotensin-converting enzyme 2
(ACE2) receptor.[49]

Taxonomy
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.[50]

Phylogenetic tree of
coronaviruses

5 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

Genus: Alphacoronavirus
Species: 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;[51] type species: Murine coronavirus
Species: Betacoronavirus 1 (Bovine Coronavirus, Human coronavirus
OC43), Human coronavirus HKU1, 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, Middle East respiratory syndrome-
related coronavirus, Hedgehog coronavirus 1 (EriCoV)
Genus Gammacoronavirus; type species: Infectious bronchitis virus
Species: Beluga whale coronavirus SW1, Infectious bronchitis virus
Genus Deltacoronavirus; type species: Bulbul coronavirus HKU11
Species: Bulbul coronavirus HKU11, Porcine coronavirus HKU15

Evolution
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.[52] 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 Origins of human coronaviruses
deltacoronavirus line at about 3000 BCE. Bats and birds, as with possible intermediate
warm-blooded flying vertebrates, are an ideal natural reservoir hosts
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.[53]

Many human coronavirus have their origin in bats.[54] The human coronavirus NL63 shared a
common ancestor with a bat coronavirus (ARCoV.2) between 1190–1449 CE.[55] The human
coronavirus 229E shared a common ancestor with bat coronavirus (GhanaGrp1 Bt CoV) between
1686–1800 CE.[56] More recently, alpaca coronavirus and human coronavirus 229E diverged
sometime before 1960.[57] MERS-CoV emerged in humans from bats through the intermediate host of
camels.[58] MERS-CoV, although related to several bat coronavirus species, appears to have diverged
from these several centuries ago.[59] The most closely related bat coronavirus and SARS-CoV
diverged in 1986.[60] A possible path of evolution, of SARS coronavirus and keen bat coronaviruses,

6 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

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.[61][62]

Unlike other betacoronaviruses, bovine coronavirus of the species Betacoronavirus 1 and subgenus
Embecovirus is thought to have originated in rodents and not in bats.[54][63] In the 1790s, equine
coronavirus diverged from the bovine coronavirus after a cross-species jump.[64] Later in the 1890s,
human coronavirus OC43 diverged from bovine coronavirus after another cross-species spillover
event.[65][64] 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.[66] Human coronavirus OC43 besides causing respiratory
infections is also suspected of playing a role in neurological diseases.[67] In the 1950s, the human
coronavirus OC43 began to diverge into its present genotypes.[68] Phylogentically, mouse hepatitis
virus (Murine coronavirus), which infects the mouse's liver and the central nervous system,[69] is
related to human coronavirus OC43 and bovine coronavirus. Human coronavirus HKU1, like the
aforementioned viruses, also has its origins in rodents.[54]

Human coronaviruses
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.[39]
Coronaviruses can cause colds with major symptoms, such as
fever, and a sore throat from swollen adenoids.[70]
Coronaviruses can cause pneumonia (either direct viral
pneumonia or secondary bacterial pneumonia) and bronchitis
(either direct viral bronchitis or secondary bacterial
bronchitis).[71] The human coronavirus discovered in 2003,
SARS-CoV, which causes severe acute respiratory syndrome Illustration of SARSr-CoV virion
(SARS), has a unique pathogenesis because it causes both
upper and lower respiratory tract infections.[71]

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.[72] Coronaviruses cause about
15% of commons colds.[73] The majority of colds are caused by rhinoviruses.[74] The four mild
coronaviruses have a seasonal incidence occurring in the winter months in temperate climates.[75][76]
There is no preference towards a particular season in tropical climates.[77]

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

7 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

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 Seasonal distribution of HCoV-
coronavirus 2 (SARS-CoV-2), β-CoV NL63 in Germany shows a
preferential detection from
November to March
Outbreaks of human coronavirus
diseases

Severe acute respiratory syndrome (SARS)

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.[49]

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).[87][88] The
World Health Organization issued a global alert soon after.[89] The WHO update on 28 September
2012 said the virus did not seem to pass easily from person to person.[90] 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.[91] 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.[92] By 30 October 2013, there were 124 cases and 52 deaths in Saudi
Arabia.[93]

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.[94]

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.[95] 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%.[96]

8 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

Coronavirus disease Characteristics of human coronavirus strains

2019 (COVID-19) MERS-CoV, SARS-CoV, SARS-CoV-2,
and related diseases
In December 2019, a MERS-CoV SARS-CoV SARS-CoV-2
pneumonia outbreak was
reported in Wuhan, Disease MERS SARS COVID-19
China. [97] On 31 December 2012, 2015, 2019–2020
2019, the outbreak was Outbreaks
2018
2002–2004
pandemic
traced to a novel strain of
coronavirus,[98] which was Epidemiology
given the interim name 2019- Date of first June November December
nCoV by the World Health identified case 2012 2002 2019[78]
Organization (WHO),
[99][100][101] later renamed Location of first Jeddah, Shunde, Wuhan,
identified case Saudi Arabia China China
SARS-CoV-2 by the
International Committee on Age average 56 44[79][a] 56[80]
Taxonomy of Viruses. Some
Sex ratio (M:F) 3.3:1 0.8:1[81] 1.6:1[80]
researchers have suggested
the Huanan Seafood Confirmed cases 2494 8096[82] 2,614,040[83][b]
Wholesale Market may not
be the original source of viral Deaths 858 774[82] 182,004[83][b]
transmission to humans. Case fatality rate 37% 9.2% 7.0%[83]
[102][103]
Symptoms
As of 22 April 2020, there
Fever 98% 99–100% 87.9%[84]
have been at least 182,004[83]
confirmed deaths and more Dry cough 47% 29–75% 67.7%[84]
than 2,614,040[83] confirmed Dyspnea 72% 40–42% 18.6%[84]
cases in the coronavirus
pneumonia pandemic. The Diarrhea 26% 20–25% 3.7%[84]
Wuhan strain has been
Sore throat 21% 13–25% 13.9%[84]
identified as a new strain of
Betacoronavirus from group Ventilatory use 24.5%[85] 14–20% 4.1%[86]
2B with approximately 70%
genetic similarity to the Notes
SARS-CoV.[104] The virus a. Based on data from Hong Kong.
has a 96% similarity to a bat b. Data as of 22 April 2020.
coronavirus, so it is widely
suspected to originate from
bats as well.[102][105] The pandemic has resulted in travel restrictions and nationwide lockdowns in
several countries.

Other animals
Coronaviruses have been recognized as causing pathological conditions in veterinary medicine since
the 1930s.[12] Except for avian infectious bronchitis, the major related diseases have mainly an
intestinal location.[106]

9 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

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.[107] 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).[108] 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.[109]
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.[110]

A HKU2-related bat coronavirus called swine acute diarrhea syndrome coronavirus (SADS-CoV)
causes diarrhea in pigs.[111]

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.[112]

Domestic animals
Infectious bronchitis virus (IBV) causes avian infectious bronchitis.
Porcine coronavirus (transmissible gastroenteritis coronavirus of pigs, TGEV).
[113][114]

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. [115]
Pantropic canine coronavirus.
Rabbit enteric coronavirus causes acute gastrointestinal disease and diarrhea

10 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

in young European rabbits. Mortality rates are high.[116]

Porcine epidemic diarrhea virus (PED or PEDV), has emerged around the
world.[117]

See also
Bat-borne virus
Zoonosis

References
1. "Virus Taxonomy: 2018b Release" (https://talk.ictvonline.org/taxonomy/).
International Committee on Taxonomy of Viruses (ICTV). March 2019.
Archived (https://web.archive.org/web/20180304035352/https://talk.ictvonlin
e.org/taxonomy/) from the original on 2018-03-04. Retrieved 2020-01-24.
2. "2017.012-015S" (https://talk.ictvonline.org/ictv/proposals/2017.012_015S.A.v
1.Nidovirales.zip) (xlsx). International Committee on Taxonomy of Viruses
(ICTV). October 2018. Archived (https://web.archive.org/web/2019051416283
6/https://talk.ictvonline.org/ictv/proposals/2017.012_015S.A.v1.Nidovirales.zi
p) from the original on 2019-05-14. Retrieved 2020-01-24.
3. "ICTV Taxonomy history: Orthocoronavirinae" (https://talk.ictvonline.org//taxo
nomy/p/taxonomy-history?taxnode_id=201851847). 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" (htt
ps://www.ncbi.nlm.nih.gov/pmc/articles/PMC6466186). Viruses. 11 (3): 210.
doi:10.3390/v11030210 (https://doi.org/10.3390%2Fv11030210).
PMC 6466186 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6466186).
PMID 30832341 (https://pubmed.ncbi.nlm.nih.gov/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" (http://talk.ictvonline.org/files/ictv_documents/m/msl/
1231/download.aspx) (xls).
7. Woo PC, Huang Y, Lau SK, Yuen KY (August 2010). "Coronavirus genomics and
bioinformatics analysis" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3185
738). Viruses. 2 (8): 1804–20. doi:10.3390/v2081803 (https://doi.org/10.339
0%2Fv2081803). PMC 3185738 (https://www.ncbi.nlm.nih.gov/pmc/articles/P
MC3185738). PMID 21994708 (https://pubmed.ncbi.nlm.nih.gov/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]."

11 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

8. Almeida JD, Berry DM, Cunningham CH, Hamre D, Hofstad MS, Mallucci L,
McIntosh K, Tyrrell DA (November 1968). "Virology: Coronaviruses" (https://w
ww.nature.com/articles/220650b0). Nature. 220 (5168): 650.
doi:10.1038/220650b0 (https://doi.org/10.1038%2F220650b0). "[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. Estola, T. (1970). "Coronaviruses, a New Group of Animal RNA Viruses" (http://
www.jstor.org/stable/1588476). Avian Diseases. 14 (2): 330–336.
doi:10.2307/1588476 (https://doi.org/10.2307%2F1588476). ISSN 0005-2086
(https://www.worldcat.org/issn/0005-2086).
10. Fabricant, Julius (1998). "The Early History of Infectious Bronchitis" (http://ww
w.jstor.org/stable/1592697). Avian Diseases. 42 (4): 648–650.
doi:10.2307/1592697 (https://doi.org/10.2307%2F1592697). ISSN 0005-2086
(https://www.worldcat.org/issn/0005-2086).
11. Decaro N (2011). Tidona C, Darai G (eds.). Gammacoronavirus‡. The Springer
Index of Viruses. Springer. pp. 403–413. doi:10.1007/978-0-387-95919-1_58
(https://doi.org/10.1007%2F978-0-387-95919-1_58).
ISBN 978-0-387-95919-1.
12. 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" (https://link.springer.com/chapter/10.1007/978-3-642-6
5775-7_3). Current Topics in Microbiology and Immunology / Ergebnisse der
Mikrobiologie und Immunitätsforschung. Current Topics in Microbiology and
Immunology / Ergebnisse der Mikrobiologie und Immunitätsforschung. Berlin,
Heidelberg: Springer: 87. doi:10.1007/978-3-642-65775-7_3 (https://doi.org/1
0.1007%2F978-3-642-65775-7_3). ISBN 978-3-642-65775-7.
13. "Il était une fois les coronavirus" (https://www.lemonde.fr/blog/realitesbiomedi
cales/2020/03/27/il-etait-une-fois-les-coronavirus). Réalités Biomédicales (in
French). 2020-03-27. Retrieved 2020-04-18.
14. 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 (h
ttps://doi.org/10.1097%2F01.inf.0000188166.17324.60). PMID 16378050 (htt
ps://pubmed.ncbi.nlm.nih.gov/16378050).
15. Mahase, Elisabeth (2020-04-16). "Covid-19: First coronavirus was described in
The BMJ in 1965" (https://www.bmj.com/content/369/bmj.m1547). BMJ. 369.
doi:10.1136/bmj.m1547 (https://doi.org/10.1136%2Fbmj.m1547).
ISSN 1756-1833 (https://www.worldcat.org/issn/1756-1833). PMID 32299810
(https://pubmed.ncbi.nlm.nih.gov/32299810).
16. Monto, Arnold S. (1984), Evans, Alfred S. (ed.), "Coronaviruses" (https://doi.or
g/10.1007/978-1-4684-4727-9_7), Viral Infections of Humans: Epidemiology
and Control, Springer US, pp. 151–165, doi:10.1007/978-1-4684-4727-9_7 (htt
ps://doi.org/10.1007%2F978-1-4684-4727-9_7), ISBN 978-1-4684-4727-9,
retrieved 2020-04-22

12 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

17. Kendall, E. J.; Bynoe, M. L.; Tyrrell, D. A. (1962-07-14). "Virus isolations from
common colds occurring in a residential school" (https://www.ncbi.nlm.nih.go
v/pubmed/14455113). British Medical Journal. 2 (5297): 82–86.
doi:10.1136/bmj.2.5297.82 (https://doi.org/10.1136%2Fbmj.2.5297.82).
ISSN 0007-1447 (https://www.worldcat.org/issn/0007-1447). PMC 1925312 (ht
tps://www.ncbi.nlm.nih.gov/pmc/articles/PMC1925312). PMID 14455113 (http
s://pubmed.ncbi.nlm.nih.gov/14455113).
18. Richmond, Caroline (2005-06-18). "David Tyrrell" (https://www.ncbi.nlm.nih.go
v/pmc/articles/PMC558394/). BMJ : British Medical Journal. 330 (7505): 1451.
ISSN 0959-8138 (https://www.worldcat.org/issn/0959-8138).
19. Group, British Medical Journal Publishing (1969-06-28). "Obituary Notices" (htt
ps://www.bmj.com/content/2/5660/827). Br Med J. 2 (5660): 827–829.
doi:10.1136/bmj.2.5660.827 (https://doi.org/10.1136%2Fbmj.2.5660.827).
ISSN 0007-1447 (https://www.worldcat.org/issn/0007-1447).
20. Tyrrell, D. a. J.; Bynoe, M. L. (1965-06-05). "Cultivation of a Novel Type of
Common-cold Virus in Organ Cultures" (https://www.bmj.com/content/1/5448/
1467). Br Med J. 1 (5448): 1467–1470. doi:10.1136/bmj.1.5448.1467 (https://
doi.org/10.1136%2Fbmj.1.5448.1467). ISSN 0007-1447 (https://www.worldca
t.org/issn/0007-1447). PMID 14288084 (https://pubmed.ncbi.nlm.nih.gov/142
88084).
21. 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 (https://books.google.com/books?id=UtxUbXOfAF
UC&pg=PA440&lpg=PA440&dq=Ether:+Enveloped+viruses+are+susceptibl
e+to+ether.&source=bl&ots=K49miLyxmO&sig=ACfU3U0sscygOn67LgTmSk
FPFGJKi9l8eQ&hl=en&sa=X&ved=2ahUKEwighYDpxPvoAhUXvp4KHTviCGcQ6
AEwEHoECC8QAQ#v=onepage&q=Ether:%20Enveloped%20viruses%20are%
20susceptible%20to%20ether.&f=false). Cornell University Press. p. 440.
ISBN 978-0-8014-1896-9.
22. Hamre, Dorothy; Procknow, John J. (1966-01-01). "A New Virus Isolated from
the Human Respiratory Tract" (https://journals.sagepub.com/doi/abs/10.3181/
00379727-121-30734). Proceedings of the Society for Experimental Biology
and Medicine. 121 (1): 190–193. doi:10.3181/00379727-121-30734 (https://d
oi.org/10.3181%2F00379727-121-30734). ISSN 0037-9727 (https://www.world
cat.org/issn/0037-9727).
23. "The woman who discovered the first coronavirus" (https://www.bbc.com/new
s/uk-scotland-52278716).
24. Almeida, Joyce (2008-06-26). "June Almeida (née Hart)" (https://www.bmj.com
/content/336/7659/1511.1). BMJ. 336 (7659): 1511–1511.
doi:10.1136/bmj.a434 (https://doi.org/10.1136%2Fbmj.a434).
ISSN 0959-8138 (https://www.worldcat.org/issn/0959-8138).

13 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

25. Almeida, June D.; Tyrrell, D. A. J. (1967). "The Morphology of Three Previously
Uncharacterized Human Respiratory Viruses that Grow in Organ Culture" (http
s://www.microbiologyresearch.org/content/journal/jgv/10.1099/0022-1317-1-2
-175). Journal of General Virology,. 1 (2): 175–178.
doi:10.1099/0022-1317-1-2-175 (https://doi.org/10.1099%2F0022-1317-1-2-1
75). ISSN 0022-1317 (https://www.worldcat.org/issn/0022-1317).
26. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC223
830/). Proceedings of the National Academy of Sciences of the United States
of America. 58 (6): 2268–2273. ISSN 0027-8424 (https://www.worldcat.org/iss
n/0027-8424). PMID 4298953 (https://pubmed.ncbi.nlm.nih.gov/4298953).
27. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC224637/).
Proceedings of the National Academy of Sciences of the United States of
America. 57 (4): 933–940. ISSN 0027-8424 (https://www.worldcat.org/issn/00
27-8424). PMID 5231356 (https://pubmed.ncbi.nlm.nih.gov/5231356).
28. "Six Newly Discovered Viruses May Explain Cold; Strains Are Similar to Germ
That Causes a Bronchial Infection in Chickens Believed to Be New Group" (htt
p://timesmachine.nytimes.com/timesmachine/1967/05/05/107189792.html?p
ageNumber=25). timesmachine.nytimes.com. Retrieved 2020-04-22.
29. Myint, Steven H. (1995), Siddell, Stuart G. (ed.), "Human Coronavirus
Infections" (https://doi.org/10.1007/978-1-4899-1531-3_18), The
Coronaviridae, The Viruses, Springer US, pp. 389–401,
doi:10.1007/978-1-4899-1531-3_18#enumeration (https://doi.org/10.1007%2
F978-1-4899-1531-3_18%23enumeration), ISBN 978-1-4899-1531-3,
retrieved 2020-04-22
30. Geller C, Varbanov M, Duval RE (November 2012). "Human coronaviruses:
insights into environmental resistance and its influence on the development
of new antiseptic strategies" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3
509683). Viruses. 4 (11): 3044–68. doi:10.3390/v4113044 (https://doi.org/10.
3390%2Fv4113044). PMC 3509683 (https://www.ncbi.nlm.nih.gov/pmc/article
s/PMC3509683). PMID 23202515 (https://pubmed.ncbi.nlm.nih.gov/2320251
5).
31. Monto, Arnold S.; Cowling, Benjamin J.; Peiris, J. S. Malik (2014-02-27).
"Coronaviruses" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7122465/).
Viral Infections of Humans: 199–223. doi:10.1007/978-1-4899-7448-8_10 (htt
ps://doi.org/10.1007%2F978-1-4899-7448-8_10). PMC 7122465 (https://www.
ncbi.nlm.nih.gov/pmc/articles/PMC7122465). "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."

14 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

32. Su S, Wong G, Shi W, Liu J, Lai AC, Zhou J, et al. (June 2016). "Epidemiology,
Genetic Recombination, and Pathogenesis of Coronaviruses" (https://www.ncb
i.nlm.nih.gov/pmc/articles/PMC7125511). Trends in Microbiology. 24 (6):
490–502. doi:10.1016/j.tim.2016.03.003 (https://doi.org/10.1016%2Fj.tim.201
6.03.003). PMC 7125511 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7125
511). PMID 27012512 (https://pubmed.ncbi.nlm.nih.gov/27012512).
33. 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" (https://ww
w.ncbi.nlm.nih.gov/pmc/articles/PMC7092803). The New England Journal of
Medicine. 382 (8): 727–733. doi:10.1056/NEJMoa2001017 (https://doi.org/10.
1056%2FNEJMoa2001017). PMC 7092803 (https://www.ncbi.nlm.nih.gov/pmc/
articles/PMC7092803). PMID 31978945 (https://pubmed.ncbi.nlm.nih.gov/319
78945).
34. Definition of Coronavirus by Merriam-Webster (https://www.merriam-webster.c
om/dictionary/coronavirus), Merriam-Webster, archived (https://web.archive.o
rg/web/20200323161218/https://www.merriam-webster.com/dictionary/coron
avirus) from the original on 2020-03-23, retrieved 2020-03-24
35. Definition of Corona by Merriam-Webster (https://www.merriam-webster.com/
dictionary/corona), Merriam-Webster, archived (https://web.archive.org/web/2
0200324161709/https://www.merriam-webster.com/dictionary/corona) from
the original on 2020-03-24, retrieved 2020-03-24
36. 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 (https://doi.org/10.1016%2Fs00
65-3527%2808%2960721-6). PMID 6362367 (https://pubmed.ncbi.nlm.nih.go
v/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."
37. Goldsmith CS, Tatti KM, Ksiazek TG, Rollin PE, Comer JA, Lee WW, et al.
(February 2004). "Ultrastructural characterization of SARS coronavirus" (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC3322934). Emerging Infectious
Diseases. 10 (2): 320–26. doi:10.3201/eid1002.030913 (https://doi.org/10.32
01%2Feid1002.030913). PMC 3322934 (https://www.ncbi.nlm.nih.gov/pmc/art
icles/PMC3322934). PMID 15030705 (https://pubmed.ncbi.nlm.nih.gov/15030
705). "Virions acquired an envelope by budding into the cisternae and formed
mostly spherical, sometimes pleomorphic, particles that averaged 78 nm in
diameter (Figure 1A)."
38. 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" (https://www.ncbi.nlm.nih.g
ov/pmc/articles/PMC1563832). Journal of Virology. 80 (16): 7918–28.
doi:10.1128/JVI.00645-06 (https://doi.org/10.1128%2FJVI.00645-06).
PMC 1563832 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1563832).
PMID 16873249 (https://pubmed.ncbi.nlm.nih.gov/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."

15 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

39. Fehr AR, Perlman S (2015). Maier HJ, Bickerton E, Britton P (eds.).
"Coronaviruses: an overview of their replication and pathogenesis" (https://w
ww.ncbi.nlm.nih.gov/pmc/articles/PMC4369385). Methods in Molecular
Biology. Springer. 1282: 1–23. doi:10.1007/978-1-4939-2438-7_1 (https://doi.
org/10.1007%2F978-1-4939-2438-7_1). ISBN 978-1-4939-2438-7.
PMC 4369385 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4369385).
PMID 25720466 (https://pubmed.ncbi.nlm.nih.gov/25720466). "See section:
Virion Structure."
40. 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
(https://doi.org/10.1016%2FS0065-3527%2808%2960286-9).
ISBN 9780120398485. PMID 9233431 (https://pubmed.ncbi.nlm.nih.gov/9233
431).
41. 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 (https://doi.org/10.1016%2Fj.
antiviral.2013.12.009). PMID 24418573 (https://pubmed.ncbi.nlm.nih.gov/244
18573). "See Figure 4c."
42. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4486061).
Journal of Structural Biology. 174 (1): 11–22. doi:10.1016/j.jsb.2010.11.021 (h
ttps://doi.org/10.1016%2Fj.jsb.2010.11.021). PMC 4486061 (https://www.ncbi.
nlm.nih.gov/pmc/articles/PMC4486061). PMID 21130884 (https://pubmed.ncb
i.nlm.nih.gov/21130884). "See Figure 10."
43. 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 (https://doi.org/10.1016%2FS0022-283
6%2803%2900865-9). PMID 12927536 (https://pubmed.ncbi.nlm.nih.gov/129
27536). "See Figure 1."
44. 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" (https://www.ncbi.nlm.nih.gov/pmc/a
rticles/PMC3889862). Antiviral Research. 100 (3): 605–14.
doi:10.1016/j.antiviral.2013.09.028 (https://doi.org/10.1016%2Fj.antiviral.201
3.09.028). PMC 3889862 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3889
862). PMID 24121034 (https://pubmed.ncbi.nlm.nih.gov/24121034). "See
Figure 2."

16 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

45. 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" (h
ttps://www.ncbi.nlm.nih.gov/pmc/articles/PMC4984655). Journal of Virology.
90 (16): 7415–28. doi:10.1128/JVI.00080-16 (https://doi.org/10.1128%2FJVI.0
0080-16). PMC 4984655 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4984
655). PMID 27279608 (https://pubmed.ncbi.nlm.nih.gov/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)."
46. 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 (https://doi.org/10.1007%2F978-1-4939-24
38-7_1). ISBN 978-1-4939-2438-7. PMC 4369385 (https://www.ncbi.nlm.nih.go
v/pmc/articles/PMC4369385). PMID 25720466 (https://pubmed.ncbi.nlm.nih.g
ov/25720466). "See section: Coronavirus Life Cycle—Assembly and Release"
47. 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 (https://doi.org/10.1016%2FS0065-352
7%2806%2966005-3). ISBN 978-0120398690. PMID 16877062 (https://pubme
d.ncbi.nlm.nih.gov/16877062). "Nevertheless, the interaction between S
protein and receptor remains the principal, if not sole, determinant of
coronavirus host species range and tissue tropism."
48. 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 (https://doi.org/10.1038%2Fs41579-018-011
8-9). PMID 30531947 (https://pubmed.ncbi.nlm.nih.gov/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)"
49. Li F, Li W, Farzan M, Harrison SC (September 2005). "Structure of SARS
coronavirus spike receptor-binding domain complexed with receptor" (https://
semanticscholar.org/paper/bbedaafec1ea70e9ae405d1f2ac4c143951630bc).
Science. 309 (5742): 1864–68. Bibcode:2005Sci...309.1864L (https://ui.adsab
s.harvard.edu/abs/2005Sci...309.1864L). doi:10.1126/science.1116480 (http
s://doi.org/10.1126%2Fscience.1116480). PMID 16166518 (https://pubmed.nc
bi.nlm.nih.gov/16166518).
50. Wertheim JO, Chu DK, Peiris JS, Kosakovsky Pond SL, Poon LL (June 2013). "A
case for the ancient origin of coronaviruses" (https://www.ncbi.nlm.nih.gov/p
mc/articles/PMC3676139). Journal of Virology. 87 (12): 7039–45.
doi:10.1128/JVI.03273-12 (https://doi.org/10.1128%2FJVI.03273-12).
PMC 3676139 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3676139).
PMID 23596293 (https://pubmed.ncbi.nlm.nih.gov/23596293).
"Alphacoronaviruses and betacoronaviruses are found exclusively in
mammals, whereas gammacoronaviruses and deltacoronaviruses primarily
infect birds."

17 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

51. Nextstrain, phylogenetic tree of Beta-CoV (https://nextstrain.org/groups/blab/

beta-cov)
52. Wertheim JO, Chu DK, Peiris JS, Kosakovsky Pond SL, Poon LL (June 2013). "A
case for the ancient origin of coronaviruses" (https://www.ncbi.nlm.nih.gov/p
mc/articles/PMC3676139). Journal of Virology. 87 (12): 7039–45.
doi:10.1128/JVI.03273-12 (https://doi.org/10.1128%2FJVI.03273-12).
PMC 3676139 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3676139).
PMID 23596293 (https://pubmed.ncbi.nlm.nih.gov/23596293).
53. 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" (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC3302495). Journal of Virology. 86 (7): 3995–4008.
doi:10.1128/JVI.06540-11 (https://doi.org/10.1128%2FJVI.06540-11).
PMC 3302495 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3302495).
PMID 22278237 (https://pubmed.ncbi.nlm.nih.gov/22278237).
54. Forni D, Cagliani R, Clerici M, Sironi M (January 2017). "Molecular Evolution of
Human Coronavirus Genomes" (https://www.ncbi.nlm.nih.gov/pmc/articles/PM
C7111218). Trends in Microbiology. 25 (1): 35–48.
doi:10.1016/j.tim.2016.09.001 (https://doi.org/10.1016%2Fj.tim.2016.09.001)
. PMC 7111218 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7111218).
PMID 27743750 (https://pubmed.ncbi.nlm.nih.gov/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]."
55. 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" (htt
ps://www.ncbi.nlm.nih.gov/pmc/articles/PMC3497669). Journal of Virology. 86
(23): 12816–25. doi:10.1128/JVI.00906-12 (https://doi.org/10.1128%2FJVI.009
06-12). PMC 3497669 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC349766
9). PMID 22993147 (https://pubmed.ncbi.nlm.nih.gov/22993147). "If these
predictions are correct, this observation suggests that HCoV-NL63 may have
originated from bats between 1190 and 1449 CE."
56. 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" (h
ttps://www.ncbi.nlm.nih.gov/pmc/articles/PMC2819850). Emerging Infectious
Diseases. 15 (9): 1377–84. doi:10.3201/eid1509.090224 (https://doi.org/10.3
201%2Feid1509.090224). PMC 2819850 (https://www.ncbi.nlm.nih.gov/pmc/a
rticles/PMC2819850). PMID 19788804 (https://pubmed.ncbi.nlm.nih.gov/1978
8804). "The most recent common ancestor of hCoV-229E and GhanaBt-
CoVGrp1 existed in ≈1686–1800 AD."
57. 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" (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC3528286). Viruses. 4 (12): 3689–700.
doi:10.3390/v4123689 (https://doi.org/10.3390%2Fv4123689). PMC 3528286
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3528286). PMID 23235471 (ht
tps://pubmed.ncbi.nlm.nih.gov/23235471).

18 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

58. Forni D, Cagliani R, Clerici M, Sironi M (January 2017). "Molecular Evolution of

Human Coronavirus Genomes" (https://www.ncbi.nlm.nih.gov/pmc/articles/PM
C7111218). Trends in Microbiology. 25 (1): 35–48.
doi:10.1016/j.tim.2016.09.001 (https://doi.org/10.1016%2Fj.tim.2016.09.001)
. PMC 7111218 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7111218).
PMID 27743750 (https://pubmed.ncbi.nlm.nih.gov/27743750).
59. 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" (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC3719811). Journal of Virology. 87 (15): 8638–50.
doi:10.1128/JVI.01055-13 (https://doi.org/10.1128%2FJVI.01055-13).
PMC 3719811 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3719811).
PMID 23720729 (https://pubmed.ncbi.nlm.nih.gov/23720729).
60. Vijaykrishna D, Smith GJ, Zhang JX, Peiris JS, Chen H, Guan Y (April 2007).
"Evolutionary insights into the ecology of coronaviruses" (https://www.ncbi.nl
m.nih.gov/pmc/articles/PMC1866124). Journal of Virology. 81 (8): 4012–20.
doi:10.1128/jvi.02605-06 (https://doi.org/10.1128%2Fjvi.02605-06).
PMC 1866124 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1866124).
PMID 17267506 (https://pubmed.ncbi.nlm.nih.gov/17267506).
61. 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 (https://doi.org/10.101
6%2Fj.meegid.2011.06.021). PMID 21763784 (https://pubmed.ncbi.nlm.nih.go
v/21763784).
62. Cui J, Han N, Streicker D, Li G, Tang X, Shi Z, et al. (October 2007).
"Evolutionary relationships between bat coronaviruses and their hosts" (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC2851503). Emerging Infectious
Diseases. 13 (10): 1526–32. doi:10.3201/eid1310.070448 (https://doi.org/10.
3201%2Feid1310.070448). PMC 2851503 (https://www.ncbi.nlm.nih.gov/pmc/
articles/PMC2851503). PMID 18258002 (https://pubmed.ncbi.nlm.nih.gov/182
58002).
63. 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" (https://www.ncbi.
nlm.nih.gov/pmc/articles/PMC4337523). Journal of Virology. 89 (6): 3076–92.
doi:10.1128/JVI.02420-14 (https://doi.org/10.1128%2FJVI.02420-14).
PMC 4337523 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4337523).
PMID 25552712 (https://pubmed.ncbi.nlm.nih.gov/25552712).
64. 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 (https://doi.org/10.1099%2Fvir.0.054940-0).
PMID 23804565 (https://pubmed.ncbi.nlm.nih.gov/23804565). "See Table 1"

19 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

65. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC544107). Journal of
Virology. 79 (3): 1595–604. doi:10.1128/jvi.79.3.1595-1604.2005 (https://doi.
org/10.1128%2Fjvi.79.3.1595-1604.2005). PMC 544107 (https://www.ncbi.nl
m.nih.gov/pmc/articles/PMC544107). PMID 15650185 (https://pubmed.ncbi.nl
m.nih.gov/15650185).
66. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC544107). Journal of
Virology. 79 (3): 1595–604. doi:10.1128/JVI.79.3.1595-1604.2005 (https://doi.
org/10.1128%2FJVI.79.3.1595-1604.2005). PMC 544107 (https://www.ncbi.nl
m.nih.gov/pmc/articles/PMC544107). PMID 15650185 (https://pubmed.ncbi.nl
m.nih.gov/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."
67. Corman VM, Muth D, Niemeyer D, Drosten C (2018). "Hosts and Sources of
Endemic Human Coronaviruses" (https://www.ncbi.nlm.nih.gov/pmc/articles/P
MC7112090). Advances in Virus Research. 100: 163–188.
doi:10.1016/bs.aivir.2018.01.001 (https://doi.org/10.1016%2Fbs.aivir.2018.0
1.001). PMC 7112090 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC711209
0). PMID 29551135 (https://pubmed.ncbi.nlm.nih.gov/29551135).
68. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC319
4943). Journal of Virology. 85 (21): 11325–37. doi:10.1128/JVI.05512-11 (http
s://doi.org/10.1128%2FJVI.05512-11). PMC 3194943 (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC3194943). PMID 21849456 (https://pubmed.ncbi.nlm.n
ih.gov/21849456).
69. Schaumburg CS, Held KS, Lane TE (May 2008). "Mouse hepatitis virus
infection of the CNS: a model for defense, disease, and repair" (https://www.n
cbi.nlm.nih.gov/pmc/articles/PMC5025298). Frontiers in Bioscience. 13:
4393–406. doi:10.2741/3012 (https://doi.org/10.2741%2F3012).
PMC 5025298 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5025298).
PMID 18508518 (https://pubmed.ncbi.nlm.nih.gov/18508518).
70. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC570073
9). Virology Journal. 14 (1): 230. doi:10.1186/s12985-017-0896-0 (https://doi.
org/10.1186%2Fs12985-017-0896-0). PMC 5700739 (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC5700739). PMID 29166910 (https://pubmed.ncbi.nlm.n
ih.gov/29166910).

20 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

71. 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 (https://doi.org/10.1055%2Fs-0028-1119
811). PMID 19199189 (https://pubmed.ncbi.nlm.nih.gov/19199189).
72. 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 (https://doi.org/10.1016%2Fbs.aivir.2018.0
1.001). ISBN 978-0-12-815201-0. PMID 29551135 (https://pubmed.ncbi.nlm.ni
h.gov/29551135).
73. Pelczar (2010). Microbiology: Application Based Approach (https://books.googl
e.com/books?id=xnClBCuo71IC&pg=PA656). p. 656.
ISBN 978-0-07-015147-5. Archived (https://web.archive.org/web/2016051613
4615/https://books.google.com/books?id=xnClBCuo71IC&pg=PA656) from the
original on 2016-05-16.
74. Russell La Fayette Cecil; Lee Goldman; Andrew I. Schafer (2012), Goldman's
Cecil Medicine, Expert Consult Premium Edition (https://books.google.com/bo
oks?id=Qd-vvNh0ee0C&pg=PA2103) (24 ed.), Elsevier Health Sciences,
pp. 2103–, ISBN 978-1-4377-1604-7, archived (https://web.archive.org/web/20
160504202334/https://books.google.com/books?id=Qd-vvNh0ee0C&pg=PA21
03) from the original on 2016-05-04
75. 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 (https://doi.org/10.1128%2FCMR.
00042-18). PMID 30541871 (https://pubmed.ncbi.nlm.nih.gov/30541871).
"See Figure 1."
76. 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 (https://doi.org/10.1093%2Finfdis%2Fjiaa161).
PMID 32246136 (https://pubmed.ncbi.nlm.nih.gov/32246136).
77. Abdul-Rasool S, Fielding BC (May 2010). "Understanding Human Coronavirus
HCoV-NL63" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2918871). The
Open Virology Journal. 4: 76–84. doi:10.2174/1874357901004010076 (https://
doi.org/10.2174%2F1874357901004010076). PMC 2918871 (https://www.ncb
i.nlm.nih.gov/pmc/articles/PMC2918871). PMID 20700397 (https://pubmed.nc
bi.nlm.nih.gov/20700397).
78. 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 (https://doi.org/10.1016%2FS0140-673
6%2820%2930185-9). PMID 31986257 (https://pubmed.ncbi.nlm.nih.gov/319
86257).

21 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

79. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2846944). BMC
Infectious Diseases. 10: 50. doi:10.1186/1471-2334-10-50 (https://doi.org/10.
1186%2F1471-2334-10-50). PMC 2846944 (https://www.ncbi.nlm.nih.gov/pmc
/articles/PMC2846944). PMID 20205928 (https://pubmed.ncbi.nlm.nih.gov/202
05928).
80. "Old age, sepsis tied to poor COVID-19 outcomes, death" (http://www.cidrap.u
mn.edu/news-perspective/2020/03/old-age-sepsis-tied-poor-covid-19-outcome
s-death). CIDRAP, University of Minnesota. Retrieved 2020-03-29.
81. 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
(https://doi.org/10.1093%2Faje%2Fkwh056). PMID 14742282 (https://pubme
d.ncbi.nlm.nih.gov/14742282).
82. "Summary of probable SARS cases with onset of illness from 1 November
2002 to 31 July 2003" (https://www.who.int/csr/sars/country/table2004_04_21/
en/). World Health Organization. April 2004.
83. "COVID-19 Dashboard by the Center for Systems Science and Engineering
(CSSE) at Johns Hopkins University (JHU)" (https://gisanddata.maps.arcgis.co
m/apps/opsdashboard/index.html#/bda7594740fd40299423467b48e9ecf6).
ArcGIS. Johns Hopkins University. Retrieved 2020-04-22.
84. "Report of the WHO-China Joint Mission on Coronavirus Disease 2019
(COVID-19)" (https://www.who.int/docs/default-source/coronaviruse/who-china
-joint-mission-on-covid-19-final-report.pdf) (PDF). World Health Organization.
February 2020.
85. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5840
604). The Korean Journal of Internal Medicine. 33 (2): 233–246.
doi:10.3904/kjim.2018.031 (https://doi.org/10.3904%2Fkjim.2018.031).
PMC 5840604 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5840604).
PMID 29506344 (https://pubmed.ncbi.nlm.nih.gov/29506344).
86. Ñamendys-Silva SA (March 2020). "Respiratory support for patients with
COVID-19 infection" (https://www.thelancet.com/journals/lanres/article/PIIS22
13-2600(20)30110-7/fulltext). The Lancet. Respiratory Medicine.
doi:10.1016/S2213-2600(20)30110-7 (https://doi.org/10.1016%2FS2213-260
0%2820%2930110-7). PMID 32145829 (https://pubmed.ncbi.nlm.nih.gov/321
45829).
87. Doucleef M (2012-09-26). "Scientists Go Deep On Genes Of SARS-Like Virus"
(https://www.npr.org/blogs/health/2012/09/25/161770135/scientists-go-deep-
on-genes-of-sars-like-virus). Associated Press. Archived (https://web.archive.or
g/web/20120927043755/http://www.npr.org/blogs/health/2012/09/25/161770
135/scientists-go-deep-on-genes-of-sars-like-virus) from the original on
2012-09-27. Retrieved 2012-09-27.

22 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

88. Falco M (2012-09-24). "New SARS-like virus poses medical mystery" (http://th
echart.blogs.cnn.com/2012/09/24/new-sars-like-virus-poses-medical-mystery
/?hpt=he_c2). CNN Health. Archived (https://web.archive.org/web/201311010
42029/http://thechart.blogs.cnn.com/2012/09/24/new-sars-like-virus-poses-m
edical-mystery/?hpt=he_c2) from the original on 2013-11-01. Retrieved
2013-03-16.
89. "New SARS-like virus found in Middle East" (http://www.aljazeera.com/news/m
iddleeast/2012/09/2012924182013530585.html). Al-Jazeera. 2012-09-24.
Archived (https://web.archive.org/web/20130309203607/http://www.aljazeer
a.com/news/middleeast/2012/09/2012924182013530585.html) from the
original on 2013-03-09. Retrieved 2013-03-16.
90. Kelland K (2012-09-28). "New virus not spreading easily between people:
WHO" (https://www.reuters.com/article/2012/09/28/us-virus-who-idUSBRE88R
0F220120928). Reuters. Archived (https://web.archive.org/web/20121124005
044/http://www.reuters.com/article/2012/09/28/us-virus-who-idUSBRE88R0F2
20120928) from the original on 2012-11-24. Retrieved 2013-03-16.
91. Nouveau coronavirus—Point de situation : Un nouveau cas d'infection
confirmé (http://www.social-sante.gouv.fr/actualite-presse,42/communiques,2
322/nouveau-coronavirus-point-de,15820.html) Archived (https://web.archive.
org/web/20130608140519/http://www.social-sante.gouv.fr/actualite-presse,42
/communiques,2322/nouveau-coronavirus-point-de,15820.html) 8 June 2013
at the Wayback Machine (Novel coronavirus—Status report: A new case of
confirmed infection) 12 May 2013, social-sante.gouv.fr
92. "MERS Transmission" (https://www.cdc.gov/coronavirus/mers/about/transmissi
on.html). Centers for Disease Control and Prevention (CDC). 2019-08-02.
Archived (https://web.archive.org/web/20191207073553/https://www.cdc.gov/
coronavirus/mers/about/transmission.html) from the original on 2019-12-07.
Retrieved 2019-12-10.
93. "Novel coronavirus infection" (http://www.who.int/csr/don/2013_05_22_ncov/e
n/index.html). World Health Association. 2013-05-22. Archived (https://web.ar
chive.org/web/20130607163823/http://www.who.int/csr/don/2013_05_22_nco
v/en/index.html) from the original on 2013-06-07. Retrieved 2013-05-23.
94. "MERS in the U.S." (https://www.cdc.gov/coronavirus/mers/us.html) Center for
Disease Control. 2019-08-02. Archived (https://web.archive.org/web/2019121
5030453/https://www.cdc.gov/coronavirus/mers/US.html) from the original on
2019-12-15. Retrieved 2019-12-10.
95. Sang-Hun C (2015-06-08). "MERS Virus's Path: One Man, Many South Korean
Hospitals" (https://www.nytimes.com/2015/06/09/world/asia/mers-viruss-path-
one-man-many-south-korean-hospitals.html). The New York Times. Archived
(https://web.archive.org/web/20170715170528/https://www.nytimes.com/201
5/06/09/world/asia/mers-viruss-path-one-man-many-south-korean-hospitals.ht
ml) from the original on 2017-07-15. Retrieved 2017-03-01.
96. "Middle East respiratory syndrome coronavirus (MERS-CoV)" (http://www.who.
int/emergencies/mers-cov/en/). WHO. Archived (https://web.archive.org/web/2
0191018010957/https://www.who.int/emergencies/mers-cov/en/) from the
original on 2019-10-18. Retrieved 2019-12-10.

23 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

97. 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" (https://www.nytimes.com/2020/01/29/opinion/coronavirus-outbrea
k.html). The New York Times. Retrieved 2020-01-30.
98. "WHO Statement Regarding Cluster of Pneumonia Cases in Wuhan, China" (ht
tps://www.who.int/china/news/detail/09-01-2020-who-statement-regarding-clu
ster-of-pneumonia-cases-in-wuhan-china). www.who.int. 2020-01-09. Archived
(https://web.archive.org/web/20200114133102/https://www.who.int/china/ne
ws/detail/09-01-2020-who-statement-regarding-cluster-of-pneumonia-cases-in
-wuhan-china) from the original on 2020-01-14. Retrieved 2020-01-10.
99. "Laboratory testing of human suspected cases of novel coronavirus (nCoV)
infection. Interim guidance, 10 January 2020" (https://apps.who.int/iris/bitstre
am/handle/10665/330374/WHO-2019-nCoV-laboratory-2020.1-eng.pdf) (PDF).
Archived (https://web.archive.org/web/20200120043516/https://apps.who.int/i
ris/bitstream/handle/10665/330374/WHO-2019-nCoV-laboratory-2020.1-eng.p
df) (PDF) from the original on 2020-01-20. Retrieved 2020-01-14.
100. "Novel Coronavirus 2019, Wuhan, China" (https://www.cdc.gov/coronavirus/20
19-ncov/index.html). www.cdc.gov (CDC). 2020-01-23. Archived (https://web.
archive.org/web/20200120144040/https://www.cdc.gov/coronavirus/2019-nco
v/index.html) from the original on 2020-01-20. Retrieved 2020-01-23.
101. "2019 Novel Coronavirus infection (Wuhan, China): Outbreak update" (https://
www.canada.ca/en/public-health/services/diseases/2019-novel-coronavirus-inf
ection.html). Canada.ca. 2020-01-21.
102. Cohen J (2020-01-26). "Wuhan seafood market may not be source of novel
virus spreading globally" (https://www.sciencemag.org/news/2020/01/wuhan-
seafood-market-may-not-be-source-novel-virus-spreading-globally).
ScienceMag American Association for the Advancement of Science. (AAAS) .
Archived (https://archive.today/20200127063253/https://www.sciencemag.or
g/news/2020/01/wuhan-seafood-market-may-not-be-source-novel-virus-sprea
ding-globally) from the original on 2020-01-27. Retrieved 2020-01-29.
103. Eschner K (2020-01-28). "We're still not sure where the COVID-19 really came
from" (https://www.popsci.com/story/health/wuhan-coronavirus-china-wet-ma
rket-wild-animal/). Popular Science. Archived (https://archive.today/20200130
003350/https://www.popsci.com/story/health/wuhan-coronavirus-china-wet-m
arket-wild-animal/) from the original on 2020-01-30. Retrieved 2020-01-30.
104. 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 (https://doi.org/10.1016%2Fj.ijid.2020.01.009).
PMID 31953166 (https://pubmed.ncbi.nlm.nih.gov/31953166).
105. Eschner K (2020-01-28). "We're still not sure where the COVID-19 really came
from" (https://www.popsci.com/story/health/wuhan-coronavirus-china-wet-ma
rket-wild-animal/). Popular Science. Archived (https://archive.today/20200130
003350/https://www.popsci.com/story/health/wuhan-coronavirus-china-wet-m
arket-wild-animal/) from the original on 2020-01-30. Retrieved 2020-01-30.
106. Murphy FA, Gibbs EP, Horzinek MC, Studdart MJ (1999). Veterinary Virology.
Boston: Academic Press. pp. 495–508. ISBN 978-0-12-511340-3.

24 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

107. Bande F, Arshad SS, Bejo MH, Moeini H, Omar AR (2015). "Progress and
challenges toward the development of vaccines against avian infectious
bronchitis" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4411447). Journal
of Immunology Research. 2015: 424860. doi:10.1155/2015/424860 (https://d
oi.org/10.1155%2F2015%2F424860). PMC 4411447 (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC4411447). PMID 25954763 (https://pubmed.ncbi.nlm.n
ih.gov/25954763).
108. Murray J (2014-04-16). "What's New With Ferret FIP-like Disease?" (http://ww
w.smallanimalchannel.com/ferrets/ferret-health/whats-new-with-ferret-fiplike-
disease.aspx) (xls). Archived (https://web.archive.org/web/20140424203951/
http://www.smallanimalchannel.com/ferrets/ferret-health/whats-new-with-ferr
et-fiplike-disease.aspx) from the original on 2014-04-24. Retrieved
2014-04-24.
109. Weiss SR, Navas-Martin S (December 2005). "Coronavirus pathogenesis and
the emerging pathogen severe acute respiratory syndrome coronavirus" (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC1306801). Microbiology and
Molecular Biology Reviews. 69 (4): 635–64.
doi:10.1128/MMBR.69.4.635-664.2005 (https://doi.org/10.1128%2FMMBR.69.
4.635-664.2005). PMC 1306801 (https://www.ncbi.nlm.nih.gov/pmc/articles/P
MC1306801). PMID 16339739 (https://pubmed.ncbi.nlm.nih.gov/16339739).
110. "Rat Coronavirus—an overview" (https://www.sciencedirect.com/topics/agricul
tural-and-biological-sciences/rat-coronavirus). www.ScienceDirect.com Topics.
111. 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 (http
s://ui.adsabs.harvard.edu/abs/2018Natur.556..255Z).
doi:10.1038/s41586-018-0010-9 (https://doi.org/10.1038%2Fs41586-018-001
0-9). PMID 29618817 (https://pubmed.ncbi.nlm.nih.gov/29618817).
112. Tirotta E, Carbajal KS, Schaumburg CS, Whitman L, Lane TE (July 2010). "Cell
replacement therapies to promote remyelination in a viral model of
demyelination" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2919340).
Journal of Neuroimmunology. 224 (1–2): 101–07.
doi:10.1016/j.jneuroim.2010.05.013 (https://doi.org/10.1016%2Fj.jneuroim.20
10.05.013). PMC 2919340 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC291
9340). PMID 20627412 (https://pubmed.ncbi.nlm.nih.gov/20627412).
113. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3111541). PLoS
Pathogens. 7 (6): e1002090. doi:10.1371/journal.ppat.1002090 (https://doi.or
g/10.1371%2Fjournal.ppat.1002090). PMC 3111541 (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC3111541). PMID 21695242 (https://pubmed.ncbi.nlm.n
ih.gov/21695242).
114. Cruz JL, Becares M, Sola I, Oliveros JC, Enjuanes L, Zúñiga S (September
2013). "Alphacoronavirus protein 7 modulates host innate immune response"
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3754097). Journal of Virology.
87 (17): 9754–67. doi:10.1128/JVI.01032-13 (https://doi.org/10.1128%2FJVI.0
1032-13). PMC 3754097 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3754
097). PMID 23824792 (https://pubmed.ncbi.nlm.nih.gov/23824792).

25 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

115. "Merck Veterinary Manual" (https://www.merckvetmanual.com/). Merck

Veterinary Manual. Archived (https://web.archive.org/web/20191213145951/h
ttps://www.merckvetmanual.com/) from the original on 2019-12-13. Retrieved
2020-01-24.
116. "Enteric Coronavirus" (http://dora.missouri.edu/rabbits/enteric-coronavirus/).
Diseases of Research Animals. Archived (https://web.archive.org/web/201907
01054046/http://dora.missouri.edu/rabbits/enteric-coronavirus/) from the
original on 2019-07-01. Retrieved 2020-01-24.
117. 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" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC701
1528). Veterinary Research. 51 (1): 10. doi:10.1186/s13567-020-0739-7 (http
s://doi.org/10.1186%2Fs13567-020-0739-7). PMC 7011528 (https://www.ncbi.
nlm.nih.gov/pmc/articles/PMC7011528). PMID 32041637 (https://pubmed.ncb
i.nlm.nih.gov/32041637).

Further reading
Alwan A, Mahjour J, Memish ZA (2013). "Novel coronavirus infection: time to
stay ahead of the curve" (http://www.emro.who.int/emhj-volume-19-2013/volu
me-19-supplement-1-coronavirus/volume-19-supplement-1-coronavirus.htm).
Eastern Mediterranean Health Journal = La Revue De Sante De La
Mediterranee Orientale = Al-Majallah Al-Sihhiyah Li-Sharq Al-Mutawassit . 19
Suppl 1: S3–4. doi:10.26719/2013.19.supp1.S3 (https://doi.org/10.26719%2F2
013.19.supp1.S3). PMID 23888787 (https://pubmed.ncbi.nlm.nih.gov/238887
87).
Laude H, Rasschaert D, Delmas B, Godet M, Gelfi J, Charley B (June 1990).
"Molecular biology of transmissible gastroenteritis virus". Veterinary
Microbiology. 23 (1–4): 147–54. doi:10.1016/0378-1135(90)90144-K (https://d
oi.org/10.1016%2F0378-1135%2890%2990144-K). PMID 2169670 (https://pu
bmed.ncbi.nlm.nih.gov/2169670).
Sola I, Alonso S, Zúñiga S, Balasch M, Plana-Durán J, Enjuanes L (April 2003).
"Engineering the transmissible gastroenteritis virus genome as an expression
vector inducing lactogenic immunity" (https://www.ncbi.nlm.nih.gov/pmc/artic
les/PMC150661). Journal of Virology. 77 (7): 4357–69.
doi:10.1128/JVI.77.7.4357-4369.2003 (https://doi.org/10.1128%2FJVI.77.7.435
7-4369.2003). PMC 150661 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1
50661). PMID 12634392 (https://pubmed.ncbi.nlm.nih.gov/12634392).
Tajima M (1970). "Morphology of transmissible gastroenteritis virus of pigs. A
possible member of coronaviruses. Brief report" (https://www.semanticschola
r.org/paper/fb11c63ac7ca2c5a32f08511c9167afb953dfc30). Archiv Fur Die
Gesamte Virusforschung. 29 (1): 105–08. doi:10.1007/BF01253886 (https://do
i.org/10.1007%2FBF01253886). PMC 7086923 (https://www.ncbi.nlm.nih.gov/
pmc/articles/PMC7086923). PMID 4195092 (https://pubmed.ncbi.nlm.nih.gov/
4195092).

Classification ICD-10: D
B97.2 (https://i

26 of 27 4/22/20, 5:00 PM
Coronavirus - Wikipedia https://en.wikipedia.org/wiki/Coronavirus

cd.who.int/brow
se10/2019/en#
/B97.2) ·
MeSH:
D017934 (http
s://www.nlm.ni
h.gov/cgi/mesh/
2015/MB_cgi?fi
eld=uid&term=
D017934)

Retrieved from "https://en.wikipedia.org/w/index.php?title=Coronavirus&oldid=952483263"

This page was last edited on 22 April 2020, at 13:29 (UTC).

Text is available under the Creative Commons Attribution-ShareAlike License; additional terms
may apply. By using this site, you agree to the Terms of Use and Privacy Policy. Wikipedia® is a
registered trademark of the Wikimedia Foundation, Inc., a non-profit organization.

27 of 27 4/22/20, 5:00 PM