Severe acute respiratory syndrome coronavirus

2
Severe acute respiratory syndrome coronavirus 2
(SARS‑CoV‑2),[2] is the coronavirus that causes COVID-19 Severe acute respiratory
(coronavirus disease 2019), the respiratory illness responsible for syndrome coronavirus 2
the ongoing COVID-19 pandemic.[3] The virus was previously
referred to by its provisional name, 2019 novel coronavirus
(2019-nCoV),[4][5][6][7] and has also been called human
coronavirus 2019 (HCoV-19 or hCoV-19).[8][9][10][11] First
identified in the city of Wuhan, Hubei, China, the World Health
Organization declared the outbreak a Public Health Emergency
of International Concern on 30 January 2020, and a pandemic
on 11 March 2020.[12][13] SARS‑CoV‑2 is a positive-sense
single-stranded RNA virus[14] that is contagious in humans.[15]
As described by the US National Institutes of Health, it is the
Colourised transmission electron
successor to SARS-CoV-1, the virus that caused the 2002–2004
SARS outbreak.[16] micrograph of SARS-CoV-2 virions
with visible coronae
SARS‑CoV‑2 is a virus of the species severe acute respiratory
syndrome–related coronavirus (SARSr-CoV).[2] It is believed to
have zoonotic origins and has close genetic similarity to bat
coronaviruses, suggesting it emerged from a bat-borne
virus.[9][17] Research is ongoing as to whether SARS‑CoV‑2
came directly from bats or indirectly through any intermediate
hosts.[18] The virus shows little genetic diversity, indicating that
the spillover event introducing SARS‑CoV‑2 to humans is likely
to have occurred in late 2019.[19]

Epidemiological studies estimate that, in December 2019 —

September 2020 period, each infection resulted in an average of
2.4 to 3.4 new ones when no members of the community are
Atomic model of the external
immune and no preventive measures are taken.[20] The virus
primarily spreads between people through close contact and via structure of the SARS-CoV-2 virion.
aerosols and respiratory droplets that are exhaled when talking, Each "ball" is an atom.[1]
breathing, or otherwise exhaling, as well as those produced from ● Blue: envelope
coughs or sneezes.[21][22] It mainly enters human cells by ● Turquoise: spike glycoprotein (S)
binding to angiotensin converting enzyme 2 (ACE2), a
membrane protein that regulates the renin-angiotensin ● Red: envelope proteins (E)
system.[23][24] ● Green: membrane proteins (M)
● Orange: glycan
Virus classification
Contents (unranked): Virus
Terminology Realm: Riboviria
Infection and transmission
 Asymptomatic transmission Kingdom: Orthornavirae
Reinfection
Phylum: Pisuviricota
Reservoir and origin
Class: Pisoniviricetes
Phylogenetics and taxonomy
Order: Nidovirales
Phylogenetic tree
Variants Family: Coronaviridae
Virology Genus: Betacoronavirus
Structure
Subgenus: Sarbecovirus
Genome
Replication cycle Species: Severe acute
respiratory
Treatment and drug development
syndrome–related
Epidemiology
coronavirus
See also
Virus: Severe acute
References
respiratory
Further reading
syndrome
External links
coronavirus 2
Notable variants
Terminology
B.1.1.7 (Alpha)
During the initial outbreak in B.1.351 (Beta)
Wuhan, China, various names were
used for the virus; some names used P.1 (Gamma)
by different sources included "the B.1.617.2 (Delta)
coronavirus" or "Wuhan
coronavirus".[25][26] In January Full list
2020, the World Health Organization Synonyms
recommended "2019 novel
coronavirus" (2019-nCov)[5][27] as 2019-nCoV
the provisional name for the virus.
Sign with provisional This was in accordance with WHO's
name "2019-nCoV" 2015 guidance[28] against using geographical locations, animal species, or
groups of people in disease and virus names.[29][30]

On 11 February 2020, the International Committee on Taxonomy of Viruses adopted the official name
"severe acute respiratory syndrome coronavirus 2" (SARS‑CoV‑2).[31] To avoid confusion with the
disease SARS, the WHO sometimes refers to SARS‑CoV‑2 as "the COVID-19 virus" in public health
communications[32][33] and the name HCoV-19 was included in some research articles.[8][9][10]

Infection and transmission

Human-to-human transmission of SARS‑CoV‑2 was confirmed on 20 January 2020 during the COVID-19
pandemic.[15][34][35][36] Transmission was initially assumed to occur primarily via respiratory droplets from
coughs and sneezes within a range of about 1.8 metres (6 ft).[37][38] Laser light scattering experiments
suggest that speaking is an additional mode of transmission[39][40] and a far-reaching[41] and under-
researched[42] one, indoors, with little air flow.[43][44] Other studies have suggested that the virus may be
airborne as well, with aerosols potentially being able to transmit the virus.[45][46][47] During human-to-
human transmission, between 200 and 800 infectious SARS‑CoV‑2 virions are thought to initiate a new
infection.[48][49][50] If confirmed, aerosol transmission has biosafety implications because a major concern
associated with the risk of working with emerging viruses in the laboratory is the generation of aerosols
from various laboratory activities which are not immediately recognizable and may affect other scientific
personnel.[51] Indirect contact via contaminated surfaces is another possible cause of infection.[52]
Preliminary research indicates that the virus may remain viable on plastic (polypropylene) and stainless steel
(AISI 304) for up to three days, but it does not survive on cardboard for more than one day or on copper
for more than four hours.[10] The virus is inactivated by soap, which destabilizes its lipid bilayer.[53][54]
Viral RNA has also been found in stool samples and semen from infected individuals.[55][56]

The degree to which the virus is infectious during the incubation period is uncertain, but research has
indicated that the pharynx reaches peak viral load approximately four days after infection[57][58] or in the
first week of symptoms and declines thereafter.[59] The duration of SARS-CoV-2 RNA shedding is
generally between 3 and 46 days after symptom onset.[60]

A study by a team of researchers from the University of North Carolina found that the nasal cavity is
seemingly the dominant initial site of infection, with subsequent aspiration-mediated virus-seeding into the
lungs in SARS‑CoV‑2 pathogenesis.[61] They found that there was an infection gradient from high in
proximal towards low in distal pulmonary epithelial cultures, with a focal infection in ciliated cells and type
2 pneumocytes in the airway and alveolar regions respectively.[61]

Studies have identified a range of animals—such as cats, ferrets, hamsters, non-human primates, minks, tree
shrews, raccoon dogs, fruit bats, and rabbits—that are susceptible and permissive to SARS-CoV-2
infection.[62][63][64] Some institutions have advised that those infected with SARS‑CoV‑2 restrict their
contact with animals.[65][66]

Asymptomatic transmission

On 1 February 2020, the World Health Organization (WHO) indicated that "transmission from
asymptomatic cases is likely not a major driver of transmission".[67] One meta-analysis found that 17% of
infections are asymptomatic, and asymptomatic individuals were 42% less likely to transmit the virus.[68]

However, an epidemiological model of the beginning of the outbreak in China suggested that "pre-
symptomatic shedding may be typical among documented infections" and that subclinical infections may
have been the source of a majority of infections.[69] That may explain how out of 217 on board a cruise
liner that docked at Montevideo, only 24 of 128 who tested positive for viral RNA showed symptoms.[70]
Similarly, a study of ninety-four patients hospitalized in January and February 2020 estimated patients shed
the most virus two to three days before symptoms appear and that "a substantial proportion of transmission
probably occurred before first symptoms in the index case".[71]

Reinfection

There is uncertainty about reinfection and long-term immunity.[72] It is not known how common reinfection
is, but reports have indicated that it is occurring with variable severity.[72]

The first reported case of reinfection was a 33-year-old man from Hong Kong who first tested positive on
26 March 2020, was discharged on 15 April 2020 after two negative tests, and tested positive again on 15
August 2020 (142 days later), which was confirmed by whole-genome sequencing showing that the viral
genomes between the episodes belong to different clades.[73] The findings had the implications that herd
immunity may not eliminate the virus if reinfection is not an uncommon occurrence and that vaccines may
not be able to provide lifelong protection against the virus.[73]

Another case study described a 25-year-old man from Nevada who tested positive for SARS‑CoV‑2 on 18
April 2020 and on 5 June 2020 (separated by two negative tests). Since genomic analyses showed
significant genetic differences between the SARS‑CoV‑2 variant sampled on those two dates, the case
study authors determined this was a reinfection.[74] The man's second infection was symptomatically more
severe than the first infection, but the mechanisms that could account for this are not known.[74]

Reservoir and origin

The first known infections from SARS‑CoV‑2 were
discovered in Wuhan, China.[17] The original source of
viral transmission to humans remains unclear, as does
whether the virus became pathogenic before or after the
spillover event.[9][19][75] Because many of the early
infectees were workers at the Huanan Seafood
Market,[76][77] it has been suggested that the virus might
Transmission of SARS-CoV-1 and
have originated from the market.[9][78] However, other SARS‑CoV‑2 from mammals as biological
research indicates that visitors may have introduced the carriers to humans
virus to the market, which then facilitated rapid
expansion of the infections.[19][79] A March 2021 WHO-
convened report stated that human spillover via an intermediate animal host was the most likely
explanation, with direct spillover from bats next most likely. Introduction through the food supply chain and
the Huanan Seafood Market was considered another possible, but less likely, explanation.[80] An analysis
in December 2021, however, said that the index case had been misidentified and that the preponderance of
early cases linked to the Huanan Market, specifically a section where raccoon dogs were caged, argued for
it being the source.[81]

For a virus recently acquired through a cross-species transmission, rapid evolution is expected.[82] The
mutation rate estimated from early cases of SARS-CoV-2 was of 6.54 × 10−4 per site per year.[80]
Coronaviruses in general have high genetic plasticity,[83] but SARS-CoV-2's viral evolution is slowed by
the RNA proofreading capability of its replication machinery.[84] For comparison, the viral mutation rate in
vivo of SARS-CoV-2 has been found to be lower than that of influenza.[85]

Research into the natural reservoir of the virus that caused the 2002–2004 SARS outbreak has resulted in
the discovery of many SARS-like bat coronaviruses, most originating in horseshoe bats. Phylogenetic
analysis indicates that samples taken from Rhinolophus sinicus show a resemblance of 80% to
SARS‑CoV‑2.[86][87][88] Phylogenetic analysis also indicates that a virus from Rhinolophus affinis,
collected in Yunnan province and designated RaTG13, has a 96.1% resemblance to SARS‑CoV‑2.[17][89]
This sequence was the closest known to SARS-CoV-2 at the time of its identification,[80] but it is not its
direct ancestor.[90] Other closely-related sequences were also identified in samples from local bat
populations.[91]

Bats are considered the most likely natural reservoir of SARS‑CoV‑2.[80][92] Differences between the bat
coronavirus and SARS‑CoV‑2 suggest that humans may have been infected via an intermediate host;[78]
although the source of introduction into humans remains unknown.[93][94]
 Although the role of pangolins as an intermediate host was initially
posited (a study published in July 2020 suggested that pangolins are an
intermediate host of SARS‑CoV‑2-like coronaviruses[95][96]),
subsequent studies have not substantiated their contribution to the
spillover.[80] Evidence against this hypothesis includes the fact that
pangolin virus samples are too distant to SARS-CoV-2: isolates obtained
from pangolins seized in Guangdong were only 92% identical in
sequence to the SARS‑CoV‑2 genome (matches above 90 percent may
Samples taken from
sound high, but in genomic terms it is a wide evolutionary gap[97]). In
Rhinolophus sinicus, a
species of horseshoe bats,
addition, despite similarities in a few critical amino acids,[98] pangolin
show an 80% resemblance to virus samples exhibit poor binding to the human ACE2 receptor.[99]
SARS‑CoV‑2.
Phylogenetics and taxonomy
SARS‑CoV‑2 belongs to the broad family of viruses known Genomic information
as coronaviruses.[26] It is a positive-sense single-stranded
RNA (+ssRNA) virus, with a single linear RNA segment.
Coronaviruses infect humans, other mammals, including
livestock and companion animals, and avian species.[100]
Human coronaviruses are capable of causing illnesses ranging
from the common cold to more severe diseases such as
Middle East respiratory syndrome (MERS, fatality rate Genomic organisation of isolate Wuhan-
~34%). SARS-CoV-2 is the seventh known coronavirus to Hu-1, the earliest sequenced sample of
infect people, after 229E, NL63, OC43, HKU1, MERS- SARS-CoV-2
CoV, and the original SARS-CoV.[101]
NCBI 86693 (https://www.ncbi.nl
Like the SARS-related coronavirus implicated in the 2003 genome ID m.nih.gov/genome/?term=
SARS outbreak, SARS‑CoV‑2 is a member of the subgenus 86693)
Sarbecovirus (beta-CoV lineage B).[102][103] Coronaviruses
Genome 29,903 bases
undergo frequent recombination.[104] The mechanism of
size
recombination in unsegmented RNA viruses such as SARS-
CoV-2 is generally by copy-choice replication, in which gene Year of 2020
material switches from one RNA template molecule to completion
another during replication.[105] SARS-CoV-2 RNA sequence Genome browser (https://genome.ucsc.e
is approximately 30,000 bases in length,[106] relatively long du/cgi-bin/hgTracks?db=wuhCor1)
for a coronavirus (which in turn carry the largest genomes
(UCSC)
among all RNA families)[107] Its genome consists nearly
entirely of protein-coding sequences, a trait shared with other
coronaviruses.[104]

A distinguishing feature of SARS‑CoV‑2 is its incorporation of a polybasic site cleaved by furin,[98] which
appears to be an important element enhancing its virulence.[108] It was suggested that the acquisition of the
furin-cleavage site in the SARS-CoV-2 S protein was essential for zoonotic transfer to humans.[109] The
furin protease recognizes the canonical peptide sequence RX[R/K]R↓X where the cleavage site is indicated
by a down arrow and X is any amino acid.[110][111] In SARS-CoV-2 the recognition site is formed by the
incorporated 12 codon nucleotide sequence CCT CGG CGG GCA which corresponds to the amino acid
sequence PRRA.[112] This sequence is upstream of an arginine and serine which forms the S1/S2 cleavage
site (PRRAR↓S) of the spike protein.[113] Although such sites are a common naturally-occurring feature of
other viruses within the Subfamily Orthocoronavirinae,[112] it appears in few other viruses from the Beta-
CoV genus,[114] and it is unique among members of its subgenus for such a site.[98] The furin cleavage site
PRRAR↓ is identical to that of the feline coronavirus, an alphacoronavirus 1 strain.[115]
Viral genetic sequence data can provide critical information about whether viruses separated by time and
space are likely to be epidemiologically linked.[116] With a sufficient number of sequenced genomes, it is
possible to reconstruct a phylogenetic tree of the mutation history of a family of viruses. By 12 January
2020, five genomes of SARS‑CoV‑2 had been isolated from Wuhan and reported by the Chinese Center
for Disease Control and Prevention (CCDC) and other institutions;[106][117] the number of genomes
increased to 42 by 30 January 2020.[118] A phylogenetic analysis of those samples showed they were
"highly related with at most seven mutations relative to a common ancestor", implying that the first human
infection occurred in November or December 2019.[118] Examination of the topology of the phylogenetic
tree at the start of the pandemic also found high similarities between human isolates.[119] As of
21 August 2021, 3,422 SARS‑CoV‑2 genomes, belonging to 19 strains, sampled on all continents except
Antarctica were publicly available.[120]

On 11 February 2020, the International Committee on Taxonomy of Viruses announced that according to
existing rules that compute hierarchical relationships among coronaviruses based on five conserved
sequences of nucleic acids, the differences between what was then called 2019-nCoV and the virus from
the 2003 SARS outbreak were insufficient to make them separate viral species. Therefore, they identified
2019-nCoV as a virus of Severe acute respiratory syndrome–related coronavirus.[121]

In July 2020, scientists reported that a more infectious SARS‑CoV‑2 variant with spike protein variant
G614 has replaced D614 as the dominant form in the pandemic.[122][123]

Coronavirus genomes and subgenomes encode six open reading frames (ORFs).[124] In October 2020,
researchers discovered a possible overlapping gene named ORF3d, in the SARS‑CoV‑2 genome. It is
unknown if the protein produced by ORF3d has any function, but it provokes a strong immune response.
ORF3d has been identified before, in a variant of coronavirus that infects pangolins.[125][126]

Phylogenetic tree

A phylogenetic tree based on whole-genome sequences of SARS-CoV-2 and related coronaviruses

is:[127][128]

SARS‑CoV‑2 related coronavirus

(Bat) Rc-o319, 81% to SARS-CoV-2, Rhinolophus cornutus,
Iwate, Japan [129]

Bat SL-ZXC21, 88% to SARS-CoV-2, Rhinolophus

pusillus, Zhoushan, Zhejiang [130]

Bat SL-ZC45, 88% to SARS-CoV-2, Rhinolophus

pusillus, Zhoushan, Zhejiang [130]

Pangolin SARSr-CoV-GX, 89% to SARS-CoV-2,

Manis javanica, Smuggled from Southeast Asia [131]

Pangolin SARSr-CoV-GD, 91% to SARS-CoV-2,

Manis javanica, Smuggled from Southeast Asia [132]
 Bat RshSTT182, 92.6% to SARS-CoV-2,
Rhinolophus shameli, Steung Treng,
Cambodia [133]

Bat RshSTT200, 92.6% to SARS-CoV-2,

Rhinolophus shameli, Steung Treng,
Cambodia [133]

(Bat) RacCS203, 91.5% to SARS-CoV-2,

Rhinolophus acuminatus, Chachoengsao,
Thailand [128]

(Bat) RmYN02, 93.3% to SARS-CoV-2,

Rhinolophus malayanus Mengla, Yunnan [134]

(Bat) RpYN06, 94.4% to SARS-CoV-2,

Rhinolophus pusillus, Xishuangbanna, Yunnan [127]

(Bat) RaTG13, 96.1% to SARS-CoV-2,

Rhinolophus affinis, Mojiang, Yunnan

SARS-CoV-2

SARS-CoV-1, 79% to SARS-CoV-2

Variants

There are many thousands of variants of SARS-CoV-2, which can be grouped into the much larger
clades.[135] Several different clade nomenclatures have been proposed. Nextstrain divides the variants into
five clades (19A, 19B, 20A, 20B, and 20C), while GISAID divides them into seven (L, O, V, S, G, GH,
and GR).[136]

Several notable variants of SARS-CoV-2 emerged in late 2020. The World Health Organization has
currently declared four variants of concern, which are as follows:[137]

Alpha: Lineage B.1.1.7 emerged in the United Kingdom in September 2020, with evidence
of increased transmissibility and virulence. Notable mutations include N501Y and P681H.
An E484K mutation in some lineage B.1.1.7 virions has been noted and is also tracked
by various public health agencies.
Beta: Lineage B.1.351 emerged in South Africa in May 2020, with evidence of increased
transmissibility and changes to antigenicity, with some public health officials raising alarms
about its impact on the efficacy of some vaccines. Notable mutations include K417N, E484K
and N501Y.
 Gamma: Lineage P.1 emerged in Brazil in November
2020, also with evidence of increased transmissibility
and virulence, alongside changes to antigenicity.
Similar concerns about vaccine efficacy have been
raised. Notable mutations also include K417N, E484K
and N501Y.
Delta: Lineage B.1.617.2 emerged in India in October
2020. There is also evidence of increased
transmissibility and changes to antigenicity.

Other notable variants include 6 other WHO-designated variants

under investigation and Cluster 5, which emerged among mink in
Denmark and resulted in a mink euthanasia campaign rendering it False-colour transmission electron
virtually extinct.[138] micrograph of a B.1.1.7 variant
coronavirus. The variant's increased
transmissibility is believed to be due
Virology to changes in the structure of the
spike proteins, shown here in green.

Structure

Each SARS-CoV-2 virion is 50–200 nanometres (2.0 × 10−6 –

7.9 × 10−6 in) in diameter.[77] Like other coronaviruses, SARS-
CoV-2 has four structural proteins, known as the S (spike), E
(envelope), M (membrane), and N (nucleocapsid) proteins; the N
protein holds the RNA genome, and the S, E, and M proteins
together create the viral envelope.[139] Coronavirus S proteins are
glycoproteins and also type I membrane proteins (membranes
containing a single transmembrane domain oriented on the
extracellular side).[109] They are divided into two functional parts
Structure of a SARSr-CoV virion
(S1 and S2).[100] In SARS-CoV-2, the spike protein, which has
been imaged at the atomic level using cryogenic electron
microscopy,[140][141] is the protein responsible for allowing the
virus to attach to and fuse with the membrane of a host cell;[139] specifically, its S1 subunit catalyzes
attachment, the S2 subunit fusion.[142]

Genome

SARS-CoV-2 has a linear, positive-sense, single-stranded RNA genome about 30,000 bases long.[100] Its
genome has a bias against cytosine (C) and guanine (G) nucleotides like other coronaviruses.[143] The
genome has the highest composition of U (32.2%), followed by A (29.9%), and a similar composition of G
(19.6%) and C (18.3%).[144] The nucleotide bias arises from the mutation of guanines and cytosines to
adenosines and uracils, respectively.[145] The mutation of CG dinucleotides is thought to arise to avoid the
zinc finger antiviral protein related defense mechanism of cells,[146] and to lower the energy to unbind the
genome during replication and translation (adenosine and uracil base pair via two hydrogen bonds, cytosine
and guanine via three).[145] The depletion of CG dinucleotides in its genome has led the virus to have a
noticeable codon usage bias. For instance, arginine's six different codons have a relative synonymous
codon usage of AGA (2.67), CGU (1.46), AGG (.81), CGC (.58), CGA (.29), and CGG (.19).[144] A
similar codon usage bias trend is seen in other SARS–related coronaviruses.[147]
Replication cycle

Virus infections start when viral particles bind to host surface cellular
receptors.[148] Protein modeling experiments on the spike protein of the
virus soon suggested that SARS‑CoV‑2 has sufficient affinity to the
receptor angiotensin converting enzyme 2 (ACE2) on human cells to use
them as a mechanism of cell entry.[149] By 22 January 2020, a group in
China working with the full virus genome and a group in the United States
using reverse genetics methods independently and experimentally
demonstrated that ACE2 could act as the receptor for
SARS‑CoV‑2.[17][150][151][152] Studies have shown that SARS‑CoV‑2
has a higher affinity to human ACE2 than the original SARS
virus.[140][153] SARS‑CoV‑2 may also use basigin to assist in cell
entry.[154]
SARS‑CoV‑2 spike
Initial spike protein priming by transmembrane protease, serine 2 homotrimer with one protein
(TMPRSS2) is essential for entry of SARS‑CoV‑2.[23] The host protein subunit highlighted. The
ACE2 binding domain is
neuropilin 1 (NRP1) may aid the virus in host cell entry using ACE2.[155]
magenta.
After a SARS‑CoV‑2 virion attaches to a target cell, the cell's TMPRSS2
cuts open the spike protein of the virus, exposing a fusion peptide in the S2
subunit, and the host receptor ACE2.[142] After fusion, an endosome
forms around the virion, separating it from the rest of the host cell. The virion escapes when the pH of the
endosome drops or when cathepsin, a host cysteine protease, cleaves it.[142] The virion then releases RNA
into the cell and forces the cell to produce and disseminate copies of the virus, which infect more cells.[156]

SARS‑CoV‑2 produces at least three virulence factors that promote shedding of new virions from host cells
and inhibit immune response.[139] Whether they include downregulation of ACE2, as seen in similar
coronaviruses, remains under investigation (as of May 2020).[157]

Digitally colourised scanning electron micrographs of SARS-CoV-2 virions (yellow) emerging from human cells
cultured in a laboratory

Treatment and drug development

Very few drugs are known to effectively inhibit SARS‑CoV‑2. Masitinib is a clinically safe drug and was
recently found to inhibit its main protease, 3CLpro and showed >200-fold reduction in viral titers in the
lungs and nose in mice. However, it is not approved for the treatment of COVID-19 in humans as of
August 2021.[158]
Epidemiology
Retrospective tests collected within the Chinese surveillance
system revealed no clear indication of substantial unrecognized
circulation of SARS‑CoV‑2 in Wuhan during the latter part of
2019.[80]

A meta-analysis from November 2020 estimated the basic

reproduction number ( ) of the virus to be between 2.39 and
3.44.[20] This means each infection from the virus is expected to
result in 2.39 to 3.44 new infections when no members of the
community are immune and no preventive measures are taken. The
reproduction number may be higher in densely populated
conditions such as those found on cruise ships.[159] Many forms of
preventive efforts may be employed in specific circumstances to Transmission electron micrograph of
SARS‑CoV‑2 virions (red) isolated
reduce the propagation of the virus.[124]
from a patient during the COVID-19
There have been about 96,000 confirmed cases of infection in pandemic
mainland China.[160] While the proportion of infections that result
in confirmed cases or progress to diagnosable disease remains
unclear,[161] one mathematical model estimated that 75,815 people were infected on 25 January 2020 in
Wuhan alone, at a time when the number of confirmed cases worldwide was only 2,015.[162] Before 24
February 2020, over 95% of all deaths from COVID-19 worldwide had occurred in Hubei province, where
Wuhan is located.[163][164] As of 23 November 2021, the percentage had decreased to 0.062%.[160]

As of 23 November 2021, there have been 258,249,487 total confirmed cases of SARS‑CoV‑2 infection in
the ongoing pandemic.[160] The total number of deaths attributed to the virus is 5,159,012.[160]

See also
3C-like protease (NS5)

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Further reading
Bar-On YM, Flamholz A, Phillips R, Milo R (April 2020). "SARS-CoV-2 (COVID-19) by the
numbers" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7224694). eLife. 9.
arXiv:2003.12886 (https://arxiv.org/abs/2003.12886). Bibcode:2020arXiv200312886B (http
s://ui.adsabs.harvard.edu/abs/2020arXiv200312886B). doi:10.7554/eLife.57309 (https://doi.
org/10.7554%2FeLife.57309). PMC 7224694 (https://www.ncbi.nlm.nih.gov/pmc/articles/PM
C7224694). PMID 32228860 (https://pubmed.ncbi.nlm.nih.gov/32228860).
Brüssow H (May 2020). "The Novel Coronavirus - A Snapshot of Current Knowledge" (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC7111068). Microbial Biotechnology. 13 (3): 607–
612. doi:10.1111/1751-7915.13557 (https://doi.org/10.1111%2F1751-7915.13557).
PMC 7111068 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7111068). PMID 32144890
(https://pubmed.ncbi.nlm.nih.gov/32144890).
Cascella M, Rajnik M, Aleem A, Dulebohn S, Di Napoli R (January 2020). "Features,
Evaluation and Treatment Coronavirus (COVID-19)" (https://www.ncbi.nlm.nih.gov/books/NB
 K554776/). StatPearls. PMID 32150360 (https://pubmed.ncbi.nlm.nih.gov/32150360).
Archived (https://web.archive.org/web/20200406135818/https://www.ncbi.nlm.nih.gov/books/
NBK554776/) from the original on 6 April 2020. Retrieved 4 April 2020.
Laboratory testing for coronavirus disease 2019 (COVID-19) in suspected human cases
(Report). World Health Organization. 2 March 2020. hdl:10665/331329 (https://hdl.handle.ne
t/10665%2F331329).
Zoumpourlis V, Goulielmaki M, Rizos E, Baliou S, Spandidos DA (October 2020). "
[Comment] The COVID‑19 pandemic as a scientific and social challenge in the 21st century"
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7453598). Molecular Medicine Reports
(Review). 22 (4): 3035–3048. doi:10.3892/mmr.2020.11393 (https://doi.org/10.3892%2Fmmr.
2020.11393). PMC 7453598 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7453598).
PMID 32945405 (https://pubmed.ncbi.nlm.nih.gov/32945405).

External links
"Coronavirus Disease 2019 (COVID-19)" (https://www.cdc.gov/coronavirus/2019-ncov/inde
x.html). Centers for Disease Control and Prevention (CDC). 11 February 2020.
"Coronavirus disease (COVID-19) Pandemic" (https://www.who.int/emergencies/diseases/n
ovel-coronavirus-2019). World Health Organization (WHO).
"SARS-CoV-2 (Severe acute respiratory syndrome coronavirus 2) Sequences" (https://www.
ncbi.nlm.nih.gov/genbank/sars-cov-2-seqs/). National Center for Biotechnology Information
(NCBI).
"COVID-19 Resource Centre" (https://www.thelancet.com/coronavirus). The Lancet.
"Coronavirus (Covid-19)" (https://www.nejm.org/coronavirus). The New England Journal of
Medicine.
"Covid-19: Novel Coronavirus Outbreak" (https://novel-coronavirus.onlinelibrary.wiley.com/).
Wiley.
"SARS-CoV-2" (https://www.viprbrc.org/brc/home.spg?decorator=corona_ncov). Virus
Pathogen Database and Analysis Resource.
"SARS-CoV-2 related protein structures" (http://www.rcsb.org/news?year=2020&article=5e3
c4bcba5007a04a313edcc). Protein Data Bank.
Classification ICD-10: U07.1 (httD
ps://icd.who.int/bro
wse10/2019/en#/U
07.1) · MeSH:
C000656484 (http
s://www.nlm.nih.go
v/cgi/mesh/2015/M
B_cgi?field=uid&ter
m=C000656484) ·
SNOMED CT:
840533007 (http://s
nomed.info/id/8405
33007)

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