Middle East respiratory syndrome coronavirus: transmission and phylogenetic evolution
Identifieur interne : 001417 ( Pmc/Corpus ); précédent : 001416; suivant : 001418Middle East respiratory syndrome coronavirus: transmission and phylogenetic evolution
Auteurs : Jaffar A. Al-Tawfiq ; Ziad A. MemishSource :
- Trends in Microbiology [ 0966-842X ] ; 2014.
Abstract
MERS-CoV continues to cause sporadic cases with amplification of cases in the healthcare setting. Camels have been linked as an intermediate host. Proper infection control measures are needed to prevent nosocomial outbreaks.
Url:
DOI: 10.1016/j.tim.2014.08.001
PubMed: 25178651
PubMed Central: 7133228
Links to Exploration step
PMC:7133228Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Middle East respiratory syndrome coronavirus: transmission and phylogenetic evolution</title><author><name sortKey="Al Tawfiq, Jaffar A" sort="Al Tawfiq, Jaffar A" uniqKey="Al Tawfiq J" first="Jaffar A." last="Al-Tawfiq">Jaffar A. Al-Tawfiq</name><affiliation><nlm:aff id="aff0005">Johns Hopkins Aramco Healthcare, Dhahran, Kingdom of Saudi Arabia</nlm:aff></affiliation><affiliation><nlm:aff id="aff0010">Indiana University School of Medicine, Indianapolis, IN, USA</nlm:aff></affiliation></author><author><name sortKey="Memish, Ziad A" sort="Memish, Ziad A" uniqKey="Memish Z" first="Ziad A." last="Memish">Ziad A. Memish</name><affiliation><nlm:aff id="aff0015">Ministry of Health, Riyadh, Kingdom of Saudi Arabia</nlm:aff></affiliation><affiliation><nlm:aff id="aff0020">College of Medicine, Alfaisal University, Riyadh, Kingdom of Saudi Arabia</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">25178651</idno><idno type="pmc">7133228</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7133228</idno><idno type="RBID">PMC:7133228</idno><idno type="doi">10.1016/j.tim.2014.08.001</idno><date when="2014">2014</date><idno type="wicri:Area/Pmc/Corpus">001417</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">001417</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Middle East respiratory syndrome coronavirus: transmission and phylogenetic evolution</title><author><name sortKey="Al Tawfiq, Jaffar A" sort="Al Tawfiq, Jaffar A" uniqKey="Al Tawfiq J" first="Jaffar A." last="Al-Tawfiq">Jaffar A. Al-Tawfiq</name><affiliation><nlm:aff id="aff0005">Johns Hopkins Aramco Healthcare, Dhahran, Kingdom of Saudi Arabia</nlm:aff></affiliation><affiliation><nlm:aff id="aff0010">Indiana University School of Medicine, Indianapolis, IN, USA</nlm:aff></affiliation></author><author><name sortKey="Memish, Ziad A" sort="Memish, Ziad A" uniqKey="Memish Z" first="Ziad A." last="Memish">Ziad A. Memish</name><affiliation><nlm:aff id="aff0015">Ministry of Health, Riyadh, Kingdom of Saudi Arabia</nlm:aff></affiliation><affiliation><nlm:aff id="aff0020">College of Medicine, Alfaisal University, Riyadh, Kingdom of Saudi Arabia</nlm:aff></affiliation></author></analytic><series><title level="j">Trends in Microbiology</title><idno type="ISSN">0966-842X</idno><idno type="eISSN">1878-4380</idno><imprint><date when="2014">2014</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><title>Highlights</title><p><list list-type="simple" id="lis0005"><list-item id="lsti0005"><label>•</label><p id="par0005">MERS-CoV continues to cause sporadic cases with amplification of cases in the healthcare setting.</p></list-item><list-item id="lsti0010"><label>•</label><p id="par0010">Camels have been linked as an intermediate host.</p></list-item><list-item id="lsti0015"><label>•</label><p id="par0015">Proper infection control measures are needed to prevent nosocomial outbreaks.</p></list-item></list></p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Zaki, A M" uniqKey="Zaki A">A.M. 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Trends Microbiol</journal-id><journal-id journal-id-type="iso-abbrev">Trends Microbiol</journal-id><journal-title-group><journal-title>Trends in Microbiology</journal-title></journal-title-group><issn pub-type="ppub">0966-842X</issn><issn pub-type="epub">1878-4380</issn><publisher><publisher-name>Elsevier Ltd.</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">25178651</article-id><article-id pub-id-type="pmc">7133228</article-id><article-id pub-id-type="publisher-id">S0966-842X(14)00157-7</article-id><article-id pub-id-type="doi">10.1016/j.tim.2014.08.001</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>Middle East respiratory syndrome coronavirus: transmission and phylogenetic evolution</article-title></title-group><contrib-group><contrib contrib-type="author" id="aut0005"><name><surname>Al-Tawfiq</surname><given-names>Jaffar A.</given-names></name><xref rid="aff0005" ref-type="aff">1</xref><xref rid="aff0010" ref-type="aff">2</xref></contrib><contrib contrib-type="author" id="aut0010"><name><surname>Memish</surname><given-names>Ziad A.</given-names></name><email>zmemish@yahoo.com</email><xref rid="aff0015" ref-type="aff">3</xref><xref rid="aff0020" ref-type="aff">4</xref></contrib></contrib-group><aff id="aff0005"><label>1</label>Johns Hopkins Aramco Healthcare, Dhahran, Kingdom of Saudi Arabia</aff><aff id="aff0010"><label>2</label>Indiana University School of Medicine, Indianapolis, IN, USA</aff><aff id="aff0015"><label>3</label>Ministry of Health, Riyadh, Kingdom of Saudi Arabia</aff><aff id="aff0020"><label>4</label>College of Medicine, Alfaisal University, Riyadh, Kingdom of Saudi Arabia</aff><pub-date pub-type="pmc-release"><day>29</day><month>8</month><year>2014</year></pub-date><pmc-comment> PMC Release delay is 0 months and 0 days and was based on .</pmc-comment>
<pub-date pub-type="ppub"><month>10</month><year>2014</year></pub-date><pub-date pub-type="epub"><day>29</day><month>8</month><year>2014</year></pub-date><volume>22</volume><issue>10</issue><fpage>573</fpage><lpage>579</lpage><permissions><copyright-statement>Copyright © 2014 Elsevier Ltd. All rights reserved.</copyright-statement><copyright-year>2014</copyright-year><copyright-holder>Elsevier Ltd</copyright-holder><license><license-p>Since January 2020 Elsevier has created a COVID-19 resource centre with free information in English and Mandarin on the novel coronavirus COVID-19. The COVID-19 resource centre is hosted on Elsevier Connect, the company's public news and information website. Elsevier hereby grants permission to make all its COVID-19-related research that is available on the COVID-19 resource centre - including this research content - immediately available in PubMed Central and other publicly funded repositories, such as the WHO COVID database with rights for unrestricted research re-use and analyses in any form or by any means with acknowledgement of the original source. These permissions are granted for free by Elsevier for as long as the COVID-19 resource centre remains active.</license-p></license></permissions><abstract abstract-type="author-highlights" id="abs0005"><title>Highlights</title><p><list list-type="simple" id="lis0005"><list-item id="lsti0005"><label>•</label><p id="par0005">MERS-CoV continues to cause sporadic cases with amplification of cases in the healthcare setting.</p></list-item><list-item id="lsti0010"><label>•</label><p id="par0010">Camels have been linked as an intermediate host.</p></list-item><list-item id="lsti0015"><label>•</label><p id="par0015">Proper infection control measures are needed to prevent nosocomial outbreaks.</p></list-item></list></p></abstract><abstract id="abs0010"><p>The Middle East respiratory syndrome coronavirus (MERS-CoV) was first described in 2012 and, subsequently, many cases were reported with a lower case fatality rate than initial cases. Humans can become infected within their communities and transmission can then be amplified in the healthcare setting. Contact investigation among cases shows a variable amount of spread among family members and healthcare workers. So far, circulating virus strains remain similar under continuous monitoring, with no genetic changes. Here, we discuss the transmission pattern, phylogenetic evolution, and pathogenesis of MERS-CoV infection.</p></abstract><kwd-group id="kwd0005"><title>Keywords</title><kwd>Middle East</kwd><kwd>MERS-CoV</kwd><kwd>coronavirus</kwd><kwd>pathogenesis</kwd></kwd-group></article-meta></front><body><sec id="sec0005"><title>Global spread of Middle East respiratory syndrome coronavirus</title><p id="par0020">The initial case of MERS-CoV was reported from a patient from Bisha who was subsequently admitted to a private hospital in Jeddah, the Kingdom of Saudi Arabia (KSA) in 2012 <xref rid="bib0005" ref-type="bibr">[1]</xref>. Given that the etiology of this severe community-acquired pneumonia became known as MERS-CoV, formerly novel coronavirus, a retrospective investigation of an outbreak of 13 cases of severe acute respiratory infection clustered in a hospital in Zarqa, Jordan was conducted. The investigation revealed two confirmed and 11 probable cases in March 2012 <xref rid="bib0010" ref-type="bibr">[2]</xref>. Since then, cases have been reported from KSA <xref rid="bib0005" ref-type="bibr">1</xref>, <xref rid="bib0015" ref-type="bibr">3</xref>, <xref rid="bib0020" ref-type="bibr">4</xref>, <xref rid="bib0025" ref-type="bibr">5</xref>, <xref rid="bib0030" ref-type="bibr">6</xref>, <xref rid="bib0035" ref-type="bibr">7</xref>, <xref rid="bib0040" ref-type="bibr">8</xref>, <xref rid="bib0045" ref-type="bibr">9</xref>, <xref rid="bib0050" ref-type="bibr">10</xref>, <xref rid="bib0055" ref-type="bibr">11</xref>, Jordan <xref rid="bib0010" ref-type="bibr">2</xref>, <xref rid="bib0060" ref-type="bibr">12</xref>, France <xref rid="bib0065" ref-type="bibr">13</xref>, <xref rid="bib0070" ref-type="bibr">14</xref>, United Arab Emirates (UAE) <xref rid="bib0065" ref-type="bibr">13</xref>, <xref rid="bib0070" ref-type="bibr">14</xref>, <xref rid="bib0075" ref-type="bibr">15</xref>, Qatar <xref rid="bib0015" ref-type="bibr">3</xref>, <xref rid="bib0080" ref-type="bibr">16</xref>, Tunisia <xref rid="bib0015" ref-type="bibr">[3]</xref> (<ext-link ext-link-type="uri" xlink:href="http://www.promedmail.org/direct.php%3Fid=1725864" id="intr0010">http://www.promedmail.org/direct.php?id=1725864</ext-link>), UK <xref rid="bib0085" ref-type="bibr">[17]</xref>, Italy <xref rid="bib0090" ref-type="bibr">[18]</xref>, Oman (<ext-link ext-link-type="uri" xlink:href="http://www.who.int/csr/don/2014_05_15_mers/en/" id="intr0015">http://www.who.int/csr/don/2014_05_15_mers/en/</ext-link>), Kuwait, Yemen, Germany, The Netherlands, Greece, Malaysia, Philippines, Iran, and Egypt (<ext-link ext-link-type="uri" xlink:href="http://www.cdc.gov/coronavirus/mers/index.html" id="intr0020">http://www.cdc.gov/coronavirus/mers/index.html</ext-link>). The total global cases as of 16 June 2014 were 701 laboratory-confirmed cases officially reported to the World Health Organization (WHO), including 249 (35.5%) deaths (<ext-link ext-link-type="uri" xlink:href="http://www.who.int/csr/don/2014_06_16_mers/en/" id="intr0025">http://www.who.int/csr/don/2014_06_16_mers/en/</ext-link>). The number of cases per reporting country is shown in <xref rid="tbl0005" ref-type="table">Table 1</xref>. In this review, we highlight the transmission, evolution, and pathogenesis of MERS-CoV infection.<table-wrap position="float" id="tbl0005"><label>Table 1</label><caption><p>Number of reported MES-CoV cases and number of deaths per country<xref rid="tblfn0005" ref-type="table-fn">a</xref></p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left">Country</th><th align="left">Cases</th><th align="left">Deaths</th></tr></thead><tbody><tr><td align="left">Saudi Arabia</td><td align="char">689</td><td align="char">283</td></tr><tr><td align="left">United Arab Emirates</td><td align="char">70</td><td align="char">9</td></tr><tr><td align="left">Qatar</td><td align="char">7</td><td align="char">4</td></tr><tr><td align="left">Jordan</td><td align="char">18</td><td align="char">5</td></tr><tr><td align="left">Oman</td><td align="char">2</td><td align="char">2</td></tr><tr><td align="left">Kuwait</td><td align="char">3</td><td align="char">1</td></tr><tr><td align="left">Egypt</td><td align="char">1</td><td align="char">0</td></tr><tr><td align="left">Yemen</td><td align="char">1</td><td align="char">1</td></tr><tr><td align="left">Lebanon</td><td align="char">1</td><td align="char">0</td></tr><tr><td align="left">Iran</td><td align="char">2</td><td align="char">1</td></tr><tr><td align="left">UK</td><td align="char">4</td><td align="char">3</td></tr><tr><td align="left">Germany</td><td align="char">2</td><td align="char">1</td></tr><tr><td align="left">France</td><td align="char">2</td><td align="char">1</td></tr><tr><td align="left">Italy</td><td align="char">1</td><td align="char">0</td></tr><tr><td align="left">Greece</td><td align="char">1</td><td align="char">0</td></tr><tr><td align="left">The Netherlands</td><td align="char">2</td><td align="char">0</td></tr><tr><td align="left">Malaysia</td><td align="char">1</td><td align="char">1</td></tr><tr><td align="left">Tunisia</td><td align="char">3</td><td align="char">1</td></tr><tr><td align="left">Algeria</td><td align="char">2</td><td align="char">0</td></tr><tr><td align="left">Philippines</td><td align="char">1</td><td align="char">0</td></tr><tr><td align="left">USA</td><td align="char">2</td><td align="char">0</td></tr><tr><td align="left">Total</td><td align="char">815</td><td align="char">313</td></tr></tbody></table><table-wrap-foot><fn id="tblfn0005"><label>a</label><p id="npar0005">As of 4 June 2014, according to European Center for Disease Prevention and Control (<ext-link ext-link-type="uri" xlink:href="http://www.ecdc.europa.eu/en/press/news/_layouts/forms/News_DispForm.aspx%3FList=8db7286c-fe2d-476c-9133-18ff4cb1b568%26ID=1016" id="intr0005">http://www.ecdc.europa.eu/en/press/news/_layouts/forms/News_DispForm.aspx?List=8db7286c-fe2d-476c-9133-18ff4cb1b568&ID=1016</ext-link>).</p></fn></table-wrap-foot></table-wrap></p></sec><sec id="sec0010"><title>Interhuman transmission of MERS-CoV and genetic analysis</title><p id="par0025">It is thought that most MERS-CoV cases are acquired through human-to-human transmission. An analysis of 144 confirmed and 17 probable cases showed that 95 cases (59%) were secondary cases with an epidemiologic link to other confirmed cases <xref rid="bib0015" ref-type="bibr">[3]</xref>. The acquisition of infection in these 95 cases occurred more frequently in healthcare settings (63.2%) and among family contacts (13.7%) <xref rid="bib0015" ref-type="bibr">[3]</xref>. In a recent update by the WHO, it was noted that the number of laboratory-confirmed MERS-CoV cases increased sharply at the beginning of mid-March 2014, in KSA and UAE, in relation to the occurrence of healthcare-associated outbreaks (<ext-link ext-link-type="uri" xlink:href="http://www.who.int/csr/don/2014_05_28_mers/en/" id="intr0030">http://www.who.int/csr/don/2014_05_28_mers/en/</ext-link>). The occurrences of healthcare-associated clusters were reported from KSA <xref rid="bib0015" ref-type="bibr">[3]</xref>, Jordan <xref rid="bib0010" ref-type="bibr">2</xref>, <xref rid="bib0060" ref-type="bibr">12</xref>, France <xref rid="bib0065" ref-type="bibr">[13]</xref>, UAE, and Qatar <xref rid="bib0015" ref-type="bibr">[3]</xref>. Household transmission and clusters were also reported from KSA <xref rid="bib0025" ref-type="bibr">5</xref>, <xref rid="bib0030" ref-type="bibr">6</xref> and Tunisia <xref rid="bib0015" ref-type="bibr">[3]</xref>, and transmission occurring in both healthcare and household settings was reported from UK <xref rid="bib0085" ref-type="bibr">[17]</xref> and Italy <xref rid="bib0090" ref-type="bibr">[18]</xref>.</p><p id="par0030">Another large healthcare-associated cluster reported from Al-Hasa, KSA involved 23 hemodialysis patients, family members, and healthcare workers <xref rid="bib0020" ref-type="bibr">[4]</xref>. It was evident epidemiologically that person-to-person transmission occurred in 21 (91.3%) of the cases <xref rid="bib0020" ref-type="bibr">[4]</xref>. Subsequent phylogenetic analysis of the virus isolates from the Al-Hasa outbreak revealed that there was more than one possibility of the MERS-CoV virus being introduced from the community <xref rid="bib0095" ref-type="bibr">[19]</xref>. The Al-Hasa MERS-CoV variants were not detected in any other part of KSA, and the Al-Hasa region has remained free of other virus variants <xref rid="bib0100" ref-type="bibr">[20]</xref>. The same analysis of 32 MERS-CoV isolates from May 2013 to September 2013 revealed four different MERS-CoV clades, namely: Hafr-Al_Batin_1 clade; Buraidah_1 clade; Riyadh_3 clade; and Al-Hasa clade <xref rid="bib0100" ref-type="bibr">[20]</xref>. The second three clades (i.e., not the Hafr-Al_Batin_1 clade) were no longer circulating or contributing to cases in KSA <xref rid="bib0100" ref-type="bibr">[20]</xref>. In addition, the Al-Hasa clade was confined to an area in the eastern region of KSA. The other three clades were detected in cases from many locations, suggesting a mobile human or nonhuman source. The duration of the existence of each clade was 98 days (range: 62–147 days) <xref rid="bib0100" ref-type="bibr">[20]</xref>. Recently, analysis of reported genetic data from a community cluster in the city of Hafr Al-Batin showed that at least two of the infected contacts could not have been directly infected from the index patient and, thus, an alternate source was a possibility <xref rid="bib0055" ref-type="bibr">[11]</xref>. Therefore, the reported cases seemed to originate in the community with nonsustained human-to-human transmission and subsequent amplification of cases in the healthcare setting. This is more evident with the current increase in the number of cases in the Jeddah region, where there was also amplification in the healthcare setting for these cases <xref rid="bib0105" ref-type="bibr">[21]</xref> (<ext-link ext-link-type="uri" xlink:href="http://www.moh.gov.sa/en/CoronaNew/PressReleases/Pages/default.aspx" id="intr0035">http://www.moh.gov.sa/en/CoronaNew/PressReleases/Pages/default.aspx</ext-link>). Thus, immediate isolation of these MERS-CoV cases in the healthcare setting would prevent subsequent transmission of the virus.</p><p id="par0035">It is estimated that the rate of molecular evolution of MERS-CoV is 1.12×10<sup>−3</sup> substitutions/site per year <xref rid="bib0100" ref-type="bibr">[20]</xref>. The spike protein is important in potentiating membrane fusion and coronavirus entry and a variation in this protein was documented in the main Saudi Arabian lineage and the Munich/Abu Dhabi/2013 cluster <xref rid="bib0095" ref-type="bibr">[19]</xref>. The genetic analysis of MERS-CoV, especially in the Al-Hasa outbreak and in the community from Hafr Al-Batin, showed multiple introduction of MERS-CoV, possibly from an animal source as well as limited human-to-human transmission, with different clades in circulation in different regions.</p></sec><sec id="sec0015"><title>Transmission among family and healthcare worker contacts</title><p id="par0040">Screening of healthcare workers and family contacts of presumed index cases have been extensively studied <xref rid="bib0010" ref-type="bibr">2</xref>, <xref rid="bib0035" ref-type="bibr">7</xref>, <xref rid="bib0070" ref-type="bibr">14</xref>, <xref rid="bib0080" ref-type="bibr">16</xref>, <xref rid="bib0085" ref-type="bibr">17</xref>, <xref rid="bib0110" ref-type="bibr">22</xref>, <xref rid="bib0115" ref-type="bibr">23</xref>. A summary of the contact investigation results is shown in <xref rid="tbl0010" ref-type="table">Table 2</xref><xref rid="bib0010" ref-type="bibr">2</xref>, <xref rid="bib0035" ref-type="bibr">7</xref>, <xref rid="bib0060" ref-type="bibr">12</xref>, <xref rid="bib0070" ref-type="bibr">14</xref>, <xref rid="bib0080" ref-type="bibr">16</xref>, <xref rid="bib0085" ref-type="bibr">17</xref>, <xref rid="bib0110" ref-type="bibr">22</xref>, <xref rid="bib0115" ref-type="bibr">23</xref>. In the UK, contacts of two index cases and an additional two (2.17%) secondary cases out of 92 contacts were identified, but none of the 43 contacts of the secondary cases were positive <xref rid="bib0085" ref-type="bibr">17</xref>, <xref rid="bib0110" ref-type="bibr">22</xref>. In France, one (0.8%) of 124 contacts of the index case and none of the 39 contacts of the secondary case were positive for MERS-CoV <xref rid="bib0070" ref-type="bibr">[14]</xref>. Using serology, investigation of 85 contacts of an index case failed to demonstrate any positive cases <xref rid="bib0080" ref-type="bibr">[16]</xref>, and the use of respiratory PCR and serology among 83 contacts of another case did not reveal any positive cases <xref rid="bib0115" ref-type="bibr">[23]</xref>. The largest screening of contacts is from KSA <xref rid="bib0035" ref-type="bibr">[7]</xref>. Of the 1695 healthcare contacts and 462 family contacts of confirmed MERS cases, a positive result was obtained from 19 (1.12%) healthcare workers and 14 (3.6%) cases of family contacts <xref rid="bib0035" ref-type="bibr">[7]</xref>. Thus, the rate of secondary transmission among family contact is higher than other contacts. Thus, close and prolonged contact with the index case seems to be a requirement for the spread of the disease. Another possible explanation is exposure of other family members to the same source of infection, as was evident from the Hafr Al-Batin community outbreak <xref rid="bib0055" ref-type="bibr">[11]</xref>.<table-wrap position="float" id="tbl0010"><label>Table 2</label><caption><p>Results of contact investigation</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left">Method</th><th align="left">Case</th><th align="left">Number of contacts</th><th align="left">Number (%) positive</th><th align="left">Country</th><th align="left">Refs</th></tr></thead><tbody><tr><td align="left">Respiratory PCR</td><td align="left">Outbreak</td><td align="char">13</td><td align="char">2 (15.4)</td><td align="left">Jordan</td><td align="left"><xref rid="bib0010" ref-type="bibr">[2]</xref></td></tr><tr><td align="left">Serology</td><td align="left">Outbreak</td><td align="char">124</td><td align="char">9 (7.5)</td><td align="left">Jordan</td><td align="left"><xref rid="bib0060" ref-type="bibr">[12]</xref></td></tr><tr><td align="left">Respiratory PCR</td><td align="left">Index</td><td align="char">92</td><td align="char">2 (2.2)</td><td align="left">UK</td><td align="left"><xref rid="bib0085" ref-type="bibr">17</xref>, <xref rid="bib0110" ref-type="bibr">22</xref></td></tr><tr><td align="left">Respiratory PCR</td><td align="left">Secondary</td><td align="char">43</td><td align="char">0 (0)</td><td align="left">UK</td><td align="left"><xref rid="bib0010" ref-type="bibr">2</xref>, <xref rid="bib0085" ref-type="bibr">17</xref></td></tr><tr><td align="left">Respiratory PCR</td><td align="left">Index</td><td align="char">124</td><td align="char">1 (0.8)</td><td align="left">France</td><td align="left"><xref rid="bib0070" ref-type="bibr">[14]</xref></td></tr><tr><td align="left">Respiratory PCR</td><td align="left">Secondary</td><td align="char">39</td><td align="char">0 (0)</td><td align="left">France</td><td align="left"><xref rid="bib0070" ref-type="bibr">[14]</xref></td></tr><tr><td align="left">Serology</td><td align="left">Index</td><td align="char">85</td><td align="char">0 (0)</td><td align="left">Germany</td><td align="left"><xref rid="bib0080" ref-type="bibr">[16]</xref></td></tr><tr><td align="left">Serology</td><td align="left">Secondary</td><td align="char">83</td><td align="char">0 (0)</td><td align="left">Germany</td><td align="left"><xref rid="bib0115" ref-type="bibr">[23]</xref></td></tr><tr><td align="left">Respiratory PCR</td><td align="left">Index secondary</td><td align="char">1695 (HCW)<xref rid="tblfn0010" ref-type="table-fn">a</xref>; 462 (family)</td><td align="char">19 (1.12); 14 (3.6)</td><td align="left">KSA</td><td align="left"><xref rid="bib0035" ref-type="bibr">[7]</xref></td></tr></tbody></table><table-wrap-foot><fn id="tblfn0010"><label>a</label><p id="npar0010">Abbreviation: HCW, healthcare workers.</p></fn></table-wrap-foot></table-wrap></p></sec><sec id="sec0020"><title>Zoonotic transmission of MERS-CoV</title><p id="par0045">The origin of the MERS-CoV and the intermediate host is currently not well known. CoVs are grouped into: <italic>Alphacoronavirus, Betacoronavirus</italic> (classified as clades 2a–2d), <italic>Gammacoronavirus</italic>, and <italic>Deltacoronavirus</italic>, and all known 2c bat CoVs originate from bats <xref rid="bib0120" ref-type="bibr">[24]</xref>. It was also thought that MERS-CoV originates from bats because MERS is classified in the C betacoronaviruses <xref rid="bib0125" ref-type="bibr">[25]</xref>. A fragment of CoV sequence (182 nucleotides) was recovered from <italic>Taphozous perforatus</italic> (Egyptian tomb bats) <xref rid="bib0125" ref-type="bibr">[25]</xref>. The bat was found living near the home and work location of a case of MERS-CoV infection in the city of Bisha in western KSA <xref rid="bib0130" ref-type="bibr">[26]</xref>. It was also found that DPP4 receptor is also the receptor for bats HKU4 virus <xref rid="bib0435" ref-type="bibr">[87]</xref>. Analysis of many human MERS-CoV genomes showed diversity within the isolates and suggested the possibility of multiple independent zoonotic events <xref rid="bib0075" ref-type="bibr">15</xref>, <xref rid="bib0095" ref-type="bibr">19</xref>. More recent studies point toward camels as possible reservoirs or intermediate hosts.</p><p id="par0050">The first study to point to camels as a possible source and reservoir was based on positive serological results from 50 serum samples from dromedary camels (<italic>Camelus dromedaries</italic>) from Oman that showed a high titer of neutralizing antibodies against MERS-CoV <xref rid="bib0135" ref-type="bibr">[27]</xref>. Subsequently, multiple studies showed positive antibodies in camels from many countries (<xref rid="tbl0015" ref-type="table">Table 3</xref>) <xref rid="bib0135" ref-type="bibr">27</xref>, <xref rid="bib0140" ref-type="bibr">28</xref>, <xref rid="bib0145" ref-type="bibr">29</xref>, <xref rid="bib0150" ref-type="bibr">30</xref>, <xref rid="bib0155" ref-type="bibr">31</xref>, <xref rid="bib0160" ref-type="bibr">32</xref>, <xref rid="bib0165" ref-type="bibr">33</xref>, <xref rid="bib0170" ref-type="bibr">34</xref>, <xref rid="bib0175" ref-type="bibr">35</xref>, <xref rid="bib0180" ref-type="bibr">36</xref>, <xref rid="bib0185" ref-type="bibr">37</xref>, including Spain (Canary Islands), Egypt, Qatar, KSA, UAE, and Jordan. The detection of MERS-CoV by PCR was also obtained from two out of nine (22.2%) camels from the farm of a patient in Jeddah, KSA <xref rid="bib0160" ref-type="bibr">[32]</xref>. Similarly, in an investigation of two human cases of MERS-CoV in Qatar, of the 14 tested dromedary camels, three tested positive by PCR for MERS-CoV <xref rid="bib0140" ref-type="bibr">[28]</xref>. In dromedary camels, there is more than one genomic variant compared with humans, who have one variant, and this suggests interspecies transmission of MERS-CoV of specific genotypes <xref rid="bib0190" ref-type="bibr">[38]</xref>. There was one amino acid change in the spike protein within the receptor-binding domain and other changes were outside the receptor-binding domain <xref rid="bib0190" ref-type="bibr">[38]</xref>. Despite the evidence linking camels to MERS-CoV, camel contact of patients was reported in only 4.3% of the total 161 cases and in 18.3% of the 49 cases with documented animal exposure <xref rid="bib0085" ref-type="bibr">[17]</xref>. More recently, MERS-CoV was isolated from the milk of five out of 12 camels, suggesting another mode of transmission <xref rid="bib0195" ref-type="bibr">[39]</xref>. However, it is unusual for milk consumption to cause respiratory illness because the pathogenesis of the initial events of MERS-CoV is thought to be upper respiratory tract infection with subsequent viremia rather than viremia causing respiratory infection. Interestingly, a recent study showed that camels may have acquired MERS-CoV from bats in sub-Saharan Africa, with subsequent importation of camels to the Arabian peninsula <xref rid="bib0200" ref-type="bibr">[40]</xref>. Identification of the major route of animal-to-human transmission is needed to understand the overall picture of persistence and evolution in the different hosts <xref rid="bib0205" ref-type="bibr">[41]</xref>.<table-wrap position="float" id="tbl0015"><label>Table 3</label><caption><p>Summary of different studies of MERS-CoV in camels</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left">Country of study</th><th align="left">Date of sampling</th><th align="left">Test</th><th align="left">Number tested</th><th align="left">Number positive</th><th align="left">% positive</th><th align="left">Refs</th></tr></thead><tbody><tr><td align="left">Oman</td><td align="left">2013</td><td align="left">Serology</td><td align="char">50</td><td align="char">50</td><td align="char">100</td><td align="left"><xref rid="bib0135" ref-type="bibr">[27]</xref></td></tr><tr><td align="left">Spain (Canary Islands)</td><td align="left">2012–2013</td><td align="left">Serology</td><td align="char">195</td><td align="char">15</td><td align="char">14</td><td align="left"><xref rid="bib0135" ref-type="bibr">[27]</xref></td></tr><tr><td rowspan="2" align="left">Qatar</td><td rowspan="2" align="left">17 October 2013</td><td align="left">Neutralization assay</td><td align="char">14</td><td align="char">14</td><td align="char">100</td><td align="left"><xref rid="bib0140" ref-type="bibr">[28]</xref></td></tr><tr><td align="left">PCR</td><td align="char">14</td><td align="char">3</td><td align="char">21</td><td align="left"><xref rid="bib0140" ref-type="bibr">[28]</xref></td></tr><tr><td align="left">Egypt</td><td align="left">June 2013</td><td align="left">Serology</td><td align="char">110</td><td align="char">103</td><td align="char">94</td><td align="left"><xref rid="bib0145" ref-type="bibr">[29]</xref></td></tr><tr><td align="left">Saudi Arabia (Al-Hasa & Riyadh)</td><td align="left">2010–2013</td><td align="left">Serology</td><td align="char">310</td><td align="char">280</td><td align="char">90</td><td align="left"><xref rid="bib0150" ref-type="bibr">[30]</xref></td></tr><tr><td rowspan="2" align="left">UAE</td><td align="left">2003</td><td align="left">Serology</td><td align="char">151</td><td align="char">151</td><td align="char">100</td><td align="left"><xref rid="bib0155" ref-type="bibr">[31]</xref></td></tr><tr><td align="left">2013</td><td align="left">Serology</td><td align="char">500</td><td align="char">481</td><td align="char">96</td><td align="left"><xref rid="bib0155" ref-type="bibr">[31]</xref></td></tr><tr><td align="left">KSA (Jeddah)</td><td align="left">2014</td><td align="left">PCR</td><td align="char">9</td><td align="char">2</td><td align="char">22.2</td><td align="left"><xref rid="bib0160" ref-type="bibr">[32]</xref></td></tr><tr><td rowspan="2" align="left">Egypt</td><td rowspan="2" align="left">June–December 2013</td><td align="left">Pseudoparticle neutralization assay</td><td align="char">52</td><td align="char">48</td><td align="char">92.3</td><td align="left"><xref rid="bib0165" ref-type="bibr">[33]</xref></td></tr><tr><td align="left">PCR</td><td align="char">110</td><td align="char">4</td><td align="char">3.6</td><td align="left"><xref rid="bib0165" ref-type="bibr">[33]</xref></td></tr><tr><td align="left">Oman</td><td align="left">December 2013</td><td align="left">PCR</td><td align="char">76</td><td align="char">5</td><td align="char">6.6</td><td align="left"><xref rid="bib0170" ref-type="bibr">[34]</xref></td></tr><tr><td align="left">UAE</td><td align="left">February–October 2005</td><td align="left">Neutralization test</td><td align="char">11</td><td align="char">9</td><td align="char">81.8</td><td align="left"><xref rid="bib0175" ref-type="bibr">[35]</xref></td></tr><tr><td align="left">USA, Canada</td><td align="left">2000–2001</td><td align="left">Neutralization test</td><td align="char">6</td><td align="char">0</td><td align="char">0</td><td align="left"><xref rid="bib0175" ref-type="bibr">[35]</xref></td></tr><tr><td align="left">Ethiopia</td><td align="left">2010–11</td><td align="left">Serology</td><td align="char">188</td><td align="char">181</td><td align="char">96</td><td align="left"><xref rid="bib0180" ref-type="bibr">[36]</xref></td></tr><tr><td align="left">Nigeria</td><td align="left">2010–2011</td><td align="left">Serology</td><td align="char">358</td><td align="char">336</td><td align="char">94</td><td align="left"><xref rid="bib0180" ref-type="bibr">[36]</xref></td></tr><tr><td align="left">Tunisia</td><td align="left">2010–2011</td><td align="left">Serology</td><td align="char">204</td><td align="char">99</td><td align="char">48.5</td><td align="left"><xref rid="bib0180" ref-type="bibr">[36]</xref></td></tr><tr><td align="left">Jordan</td><td align="left">June and September 2013</td><td align="left">Serology</td><td align="char">11</td><td align="char">11</td><td align="char">100</td><td align="left"><xref rid="bib0185" ref-type="bibr">[37]</xref></td></tr><tr><td align="left">Kenya</td><td align="left">1993–2013</td><td align="left">Serology</td><td align="char">774</td><td align="char">213</td><td align="char">27.5</td><td align="left"><xref rid="bib0440" ref-type="bibr">[88]</xref></td></tr><tr><td align="left">Egypt</td><td align="left">1997</td><td align="left">Serology</td><td align="char">43</td><td align="char">35</td><td align="char">81.4</td><td align="left"><xref rid="bib0445" ref-type="bibr">[89]</xref></td></tr><tr><td align="left">Sudan</td><td align="left">1983</td><td align="left">Serology</td><td align="char">60</td><td align="char">52</td><td align="char">86.7</td><td align="left"><xref rid="bib0445" ref-type="bibr">[89]</xref></td></tr><tr><td align="left">Somalia</td><td align="left">1983</td><td align="left">Serology</td><td align="char">25</td><td align="char">20</td><td align="char">80</td><td align="left"><xref rid="bib0445" ref-type="bibr">[89]</xref></td></tr><tr><td align="left">Somalia</td><td align="left">1984</td><td align="left">Serology</td><td align="char">61</td><td align="char">52</td><td align="char">85.2</td><td align="left"><xref rid="bib0445" ref-type="bibr">[89]</xref></td></tr></tbody></table></table-wrap></p></sec><sec id="sec0025"><title>Transmission mode of MERS-CoV</title><p id="par0055">Possible routes of transmission were elucidated from the largest nosocomial outbreak of MERS-CoV in the Al-Hasa region in May–June 2012 <xref rid="bib0020" ref-type="bibr">4</xref>, <xref rid="bib0210" ref-type="bibr">42</xref>, <xref rid="bib0215" ref-type="bibr">43</xref>, <xref rid="bib0220" ref-type="bibr">44</xref>. Respiratory droplet transmission was the most likely mode of transmission, with the possibility of direct or indirect contact and airborne transmission during aerosol-generating procedures as other possible routes of transmission <xref rid="bib0020" ref-type="bibr">4</xref>, <xref rid="bib0225" ref-type="bibr">45</xref>. Healthcare workers are advised to follow contact and droplet isolation precautions when dealing with patients with MERS-CoV with the addition of airborne infection isolation precautions during aerosol-generating procedures <xref rid="bib0225" ref-type="bibr">45</xref>, <xref rid="bib0230" ref-type="bibr">46</xref>. The only study describing infection among healthcare workers showed that many healthcare workers were involved in aerosol-generating procedures, such as intubation (71.4%), airway suctioning (57%), and sputum induction (28.6%) <xref rid="bib0235" ref-type="bibr">[47]</xref>. Further recommendations for the use of airborne infection isolation precautions were recently discussed in two recent publications <xref rid="bib0240" ref-type="bibr">[48]</xref> (<ext-link ext-link-type="uri" xlink:href="http://www.ecdc.europa.eu/en/press/news/_layouts/forms/News_DispForm.aspx%3FList=8db7286c-fe2d-476c-9133-18ff4cb1b568%26ID=1002" id="intr0040">http://www.ecdc.europa.eu/en/press/news/_layouts/forms/News_DispForm.aspx?List=8db7286c-fe2d-476c-9133-18ff4cb1b568&ID=1002</ext-link>). Airborne spread of severe acute respiratory syndrome (SARS)-CoV and MERS-CoV is thought to be predominantly in the form of large respiratory droplets expelled during coughing and sneezing and via contact with fomites <xref rid="bib0240" ref-type="bibr">[48]</xref>. A recent study showed that identical MERS-CoV RNA fragments were detected in an air sample collected from the barn of a camel that shared similar MERS-CoV with an infected person <xref rid="bib0245" ref-type="bibr">[49]</xref>. However, the evidence of aerosol dissemination was lacking.</p><p id="par0060">Coronaviruses are categorized based on the levels of oral–fecal and respiratory transmission, as indicated by protein intrinsic disorder predictions <xref rid="bib0250" ref-type="bibr">[50]</xref>. Based on these predictions, MERS-CoV has hard inner and outer shells that enable its persistence in the environment <xref rid="bib0250" ref-type="bibr">[50]</xref>. Thus, MERS-CoV is predicted to have the highest oral–fecal transmission rates and relatively low respiratory transmission rates <xref rid="bib0250" ref-type="bibr">[50]</xref>. In a patient with MERS-CoV, the virus was detected in stool samples obtained on days 12 and 16, with a viral load of up to 1031 RNA copies/g <xref rid="bib0075" ref-type="bibr">[15]</xref>. Patients with SARS commonly show high virus concentrations and prolonged virus excretion in stools, leading to the use of stool samples for routine virological diagnosis <xref rid="bib0255" ref-type="bibr">51</xref>, <xref rid="bib0260" ref-type="bibr">52</xref>, <xref rid="bib0265" ref-type="bibr">53</xref>. Diarrhea was reported to occur in patients with MERS-CoV <xref rid="bib0030" ref-type="bibr">6</xref>, <xref rid="bib0040" ref-type="bibr">8</xref>, <xref rid="bib0065" ref-type="bibr">13</xref>. An oral–fecal route of transmission of MERS-CoV was not documented in the clinical setting. MERS-CoV seems to be more stable at low temperatures and low humidity. MERS-CoV was recovered after 48 h at 20°C and 40% relative humidity (RH), and the virus was viable for 8 h at 30°C and 80% RH and for 24 h at 30°C and 30% RH <xref rid="bib0270" ref-type="bibr">[54]</xref>.</p><p id="par0065">In addition to the required respiratory or airborne infection isolation precautions, it is recommended to use contact isolation precautions <xref rid="bib0210" ref-type="bibr">42</xref>, <xref rid="bib0215" ref-type="bibr">43</xref>, <xref rid="bib0225" ref-type="bibr">45</xref>. In the SARS-CoV era, the amplification of infection in the healthcare setting is thought to be related to multiple factors, including overcrowding and reduced space between patient beds of less than 1 m; poor ventilation; use of nebulization; oxygen or non-invasive ventilation therapy; and resuscitation of patients <xref rid="bib0275" ref-type="bibr">55</xref>, <xref rid="bib0280" ref-type="bibr">56</xref>. The US and European Centers for Disease Control and Prevention (CDC) have called for the use of airborne infection isolation precautions when dealing with patients with MERS-CoV while they await further evidence to support their superiority over contact and droplet precautions (<ext-link ext-link-type="uri" xlink:href="http://www.ecdc.europa.eu/en/press/news/_layouts/forms/News_DispForm.aspx%3FList=8db7286c-fe2d-476c-9133-18ff4cb1b568%26ID=1002" id="intr0045">http://www.ecdc.europa.eu/en/press/news/_layouts/forms/News_DispForm.aspx?List=8db7286c-fe2d-476c-9133-18ff4cb1b568&ID=1002</ext-link>).</p></sec><sec id="sec0030"><title>MERS and SARS pathogenesis comparison</title><p id="par0070">Most of the data relating to the pathogenesis of SARS-CoV come from <italic>in vitro</italic> studies. The virus is thought to enter the cells through dipeptidyl peptidase 4 (DPP4), which is also known as CD26 <xref rid="bib0285" ref-type="bibr">[57]</xref>. DPP4 is found on the surfaces of nonciliated bronchial epithelial cells in humans <xref rid="bib0285" ref-type="bibr">[57]</xref>. The MERS-CoV spike protein interacts with cellular DPP4 and mediates viral attachment to the host cells <xref rid="bib0290" ref-type="bibr">[58]</xref>. In studies of the pathogenesis of MERS-CoV, the use of anti-DPP4 antibodies resulted in the inhibition of MERS-CoV infection of primary human bronchial epithelial cells <xref rid="bib0285" ref-type="bibr">[57]</xref>. Adenosine deaminase is a DPP4-binding protein that competes for virus binding and acts as a natural antagonist for MERS-CoV infection <xref rid="bib0295" ref-type="bibr">[59]</xref>. Thus, the presence of DPP4 dictates which species are infected with MERS-CoV and explains the difference in hosts between MERS-CoV and SARS-CoV. SARS-CoV-like viruses have been found in Himalayan palm civets (<italic>Paguma larvata</italic>), raccoon dogs (<italic>Nyctereutes procyonoides</italic>), and the Chinese ferret badger (<italic>Melogale moschata</italic>) <xref rid="bib0300" ref-type="bibr">[60]</xref>. The MERS-CoV was found to bind to DPP4 from various animals, such as camel, goat, cow, and sheep <xref rid="bib0305" ref-type="bibr">[61]</xref>. However, there was no serological evidence of MERS-CoV in these animals <xref rid="bib0185" ref-type="bibr">[37]</xref> and only camels were found to have MERS-CoV antibodies <xref rid="bib0135" ref-type="bibr">27</xref>, <xref rid="bib0140" ref-type="bibr">28</xref>, <xref rid="bib0145" ref-type="bibr">29</xref>, <xref rid="bib0150" ref-type="bibr">30</xref>, <xref rid="bib0155" ref-type="bibr">31</xref>, <xref rid="bib0160" ref-type="bibr">32</xref>, <xref rid="bib0165" ref-type="bibr">33</xref>, <xref rid="bib0170" ref-type="bibr">34</xref>, <xref rid="bib0175" ref-type="bibr">35</xref>, <xref rid="bib0180" ref-type="bibr">36</xref>, <xref rid="bib0185" ref-type="bibr">37</xref>.</p><p id="par0075">Similar to SARS-CoV, MERS-CoV papain-like proteases (PLpro) block the regulation of the cytokines CCL5, interferon β (IFN-β), and CXCL10 in stimulated cells <xref rid="bib0310" ref-type="bibr">[62]</xref>. Human, camel, and horse cells were more likely to be infected with MERS-CoV than were goat and bat cells, and the presence of a cell surface lung protease increased susceptibility to MERS-CoV <xref rid="bib0315" ref-type="bibr">[63]</xref>. MERS-CoV can infect lower respiratory, kidney, intestinal, and liver cells, as well as histiocytes <xref rid="bib0320" ref-type="bibr">[64]</xref>, and may infect nonhuman primate, porcine, bat, civet, and rabbit cell lines <xref rid="bib0320" ref-type="bibr">64</xref>, <xref rid="bib0325" ref-type="bibr">65</xref>. When human monocyte-derived macrophages (MDMs) were infected with either MERS-CoV or SARS-CoV, neither virus significantly stimulated the expression of antiviral cytokines (IFN-α and IFN-β) but both induced similar levels of tumor necrosis factor-α (TNF-α) and interleukin-6 (IL-6). However, MERS-CoV induced significantly higher levels of IL-12, IFN-γ and chemokines (IP-10/CXCL-10, MCP-1/CCL-2, MIP-1α/CCL-3, RANTES/CCL-5, and IL-8) <xref rid="bib0330" ref-type="bibr">[66]</xref>. The presence and activation of immune cell recruiting chemokines and immunostimulating cytokines were higher and more prolonged in cases of MERS-CoV, compared with SARS-CoV and this may explain the presence of more severe disease and a higher fatality rate in patients with MERS-CoV <xref rid="bib0330" ref-type="bibr">[66]</xref>. MERS-CoV uses a viral double-stranded RNA-binding protein (4a protein) to evade the innate host response <xref rid="bib0335" ref-type="bibr">[67]</xref>. The MERS-CoV 4a protein is an immunosuppressive factor that antagonizes type I interferon production <xref rid="bib0335" ref-type="bibr">[67]</xref>. Thus, MERS-CoV might cause severe disease in humans by counteracting host immunity and the sensing of virus invasion <xref rid="bib0335" ref-type="bibr">[67]</xref>. Understanding how MERS-CoV weakens virus sensing would be informative for the development of new strategies for therapy and vaccine.</p><p id="par0080">Recently, a recombinant protein containing the receptor-binding domain (RBD) of MERS-CoV spike (S) glycoprotein infused with Fc of human IgG (RBD-Fc) resulted in neutralizing antibodies in mice <xref rid="bib0340" ref-type="bibr">68</xref>, <xref rid="bib0345" ref-type="bibr">69</xref>. The use of the intranasal MERS-CoV RBD-Fc vaccination resulted in a similar systemic humoral immune responses to subcutaneous vaccination, and higher systemic cellular immune responses and local mucosal immune responses <xref rid="bib0350" ref-type="bibr">[70]</xref>. DPP4 in mice did not support MERS-CoV infection, but using mouse DPP4 as a scaffold showed that two critical amino acids (A288L and T330R) are important for the development of a mouse-adapted MERS-CoV strain for rapid assessment of therapy and vaccines <xref rid="bib0355" ref-type="bibr">[71]</xref>. Understanding the contribution of different antigenic components of the MERS-CoV and utilizing spike glycoprotein may result in the development of an effective vaccine.</p></sec><sec id="sec0035"><title>Variation in the MERS-CoV infection response</title><p id="par0085">There is wide variation in the host response and clinical outcome of MERS-CoV infection from asymptomatic, to mildly symptomatic to a fatal disease <xref rid="bib0035" ref-type="bibr">7</xref>, <xref rid="bib0040" ref-type="bibr">8</xref>. MERS-CoV infection was also found to have a male predominance and more specifically among those with underlying comorbid conditions. These differences in the susceptibility may be related to underlying genetic variation, environmental exposure, or demography <xref rid="bib0360" ref-type="bibr">[72]</xref>. A patient with a poor outcome did not promote type 1 IFN and IFN-α compared with a patient who survived <xref rid="bib0365" ref-type="bibr">[73]</xref>. IFN-α promotes antigen presentation in response to viruses, impairs the development of a robust antiviral response and, thus, decreases viral clearance <xref rid="bib0365" ref-type="bibr">[73]</xref>. Two human RBD-specific neutralizing monoclonal antibodies (MERS-4 and MERS-27) inhibited infection of live MERS-CoV with IC<sub>50</sub> (half-maximal inhibitory concentration) at nanomolar concentrations <xref rid="bib0370" ref-type="bibr">[74]</xref>. Given that MERS-CoV was initially limited to patients with chronic diseases <xref rid="bib0020" ref-type="bibr">[4]</xref>, causing severe disease in those with comorbid conditions and tends to affect certain individuals among family contacts <xref rid="bib0035" ref-type="bibr">[7]</xref>. Understanding the factors involved in the pathogenesis of the disease would facilitate the development of therapeutic options.</p></sec><sec id="sec0040"><title>Prospects for control of MERS-CoV</title><p id="par0090">The emergence of SARS in 2003 and the subsequent control of the virus shed light on important measures for the control of emerging viral respiratory infections. These measures include: quick case detection, isolation of suspected cases, effective contact tracing, and the practice of basic infection control measures <xref rid="bib0375" ref-type="bibr">[75]</xref> (<ext-link ext-link-type="uri" xlink:href="http://www.who.int/csr/sars/en/WHOconsensus.pdf" id="intr0050">http://www.who.int/csr/sars/en/WHOconsensus.pdf</ext-link>). In a mathematical model, a combination of household-based quarantine, isolation of cases outside the household, and targeted prophylactic use was shown to be effective in the prevention of viral transmission <xref rid="bib0380" ref-type="bibr">[76]</xref>. Delays of more than 3 days in contact tracing, case identification, and quarantine adversely affected the effectiveness of isolation <xref rid="bib0385" ref-type="bibr">[77]</xref>. One study estimated the effectiveness of the isolation of symptomatic patients and the tracing and quarantine of contacts <xref rid="bib0390" ref-type="bibr">[78]</xref>. The authors concluded that SARS could be controlled by effective isolation of symptomatic patients <xref rid="bib0390" ref-type="bibr">[78]</xref>. This finding is in contrast to the case of pandemic influenza because of the high level of presymptomatic transmission that does not occur with SARS <xref rid="bib0390" ref-type="bibr">[78]</xref>. In a review of the effectiveness of community quarantine, it was found that this is an effective preventive measure, but quantification of the effect was lacking <xref rid="bib0395" ref-type="bibr">79</xref>, <xref rid="bib0400" ref-type="bibr">80</xref>. The delay in the isolation of patients in the healthcare setting also contributed to the spread of SARS <xref rid="bib0395" ref-type="bibr">[79]</xref>. The Al-Hasa MERS-CoV outbreak and the recent increase of MERS-CoV cases in April–May 2014 also point to inadequate infection control measures <xref rid="bib0020" ref-type="bibr">[4]</xref> (<ext-link ext-link-type="uri" xlink:href="http://www.who.int/csr/disease/coronavirus_infections/MERS_CoV_RA_20140424.pdf%3Fua=1" id="intr0055">http://www.who.int/csr/disease/coronavirus_infections/MERS_CoV_RA_20140424.pdf?ua=1</ext-link>). Prompt identification of patients with SARS and the implementation of infection control measures resulted in the prevention of secondary transmission of SARS <xref rid="bib0405" ref-type="bibr">[81]</xref>. Different strategies may be used for the control of an outbreak in a healthcare setting. A hospital closure is needed to control nosocomial transmission if the outbreak is detected late, or closure early in an outbreak is sufficient to remove exposed persons to a designated location or keeping them in one place <xref rid="bib0410" ref-type="bibr">[82]</xref>. In a study of family members and other visiting patients with SARS, it was shown that the rate of SARS among visitors was lower when they used masks <xref rid="bib0415" ref-type="bibr">[83]</xref>. Thus, prompt identification of cases with subsequent proper isolation in the healthcare setting or community quarantine is a key measure to prevent the spread of emerging respiratory pathogens.</p></sec><sec id="sec0045"><title>Concluding remarks</title><p id="par0095">In conclusion, the number of MERS-CoV cases has recently increased due to multiple introductions into humans from the animal reservoir, with no long-term sustained human-to-human transmission. There is no human reservoir of cases with few or no symptoms <xref rid="bib0420" ref-type="bibr">[84]</xref>. Infection in the healthcare setting is an important risk factor for human-to-human transmission. It is not easy to identify patients with MERS-CoV during the early stages of the disease based on clinical presentation alone because patients may have mild or atypical symptoms <xref rid="bib0420" ref-type="bibr">[84]</xref>. In a univariate analysis, the presence of a normal white blood cell count (WBC) and interstitial infiltrate may be used as clues to the presence of MERS-CoV infection <xref rid="bib0050" ref-type="bibr">[10]</xref>. Thus, it is important to have a high index of suspicion and to isolate those patients early to avoid healthcare transmission. It is also important to have early diagnostic tests based on respiratory specimens, preferably lower respiratory tract specimen for rapid diagnosis <xref rid="bib0425" ref-type="bibr">[85]</xref>. The implications of MERS-CoV for travelers with comparison with SARS-CoV were recently reviewed <xref rid="bib0105" ref-type="bibr">21</xref>, <xref rid="bib0430" ref-type="bibr">86</xref>. The origin of SARS-COV and MERS-CoV is an animal source with subsequent human transmission <xref rid="bib0105" ref-type="bibr">21</xref>, <xref rid="bib0430" ref-type="bibr">86</xref>. Although SARS resulted in a widespread infection, MERS-CoV has caused limited travel-associated human cases with no major events related to the Hajj <xref rid="bib0430" ref-type="bibr">[86]</xref>. In regard to their clinical presentation, both SARS-CoV and MERS-CoV have significant overlap of signs and symptoms <xref rid="bib0105" ref-type="bibr">[21]</xref>. Further studies are needed to identify the presence of other animal reservoirs for MERS-CoV and to pinpoint the risk for the development of severe disease (<xref rid="tb0005" ref-type="boxed-text">Box 1</xref>).<boxed-text id="tb0005"><label>Box 1</label><caption><title>Outstanding questions</title></caption><p id="par0100"><list list-type="simple" id="lis0010"><list-item id="lsti0020"><label>•</label><p id="par0105">How is MERS-CoV transmitted to humans?</p></list-item><list-item id="lsti0025"><label>•</label><p id="par0110">What promotes interspecies transmission?</p></list-item><list-item id="lsti0030"><label>•</label><p id="par0115">Are camels the only animal host?</p></list-item><list-item id="lsti0035"><label>•</label><p id="par0120">What is the prevalence of immunity among the general population in the affected countries?</p></list-item><list-item id="lsti0040"><label>•</label><p id="par0125">Why is there enhanced disease severity in people with comorbidities or the elderly?</p></list-item><list-item id="lsti0045"><label>•</label><p id="par0130">What is the exact period of infectiousness?</p></list-item></list></p></boxed-text></p></sec></body><back><ref-list id="bibl0005"><title>References</title><ref id="bib0005"><label>1</label><element-citation publication-type="journal" id="sbref0005"><person-group person-group-type="author"><name><surname>Zaki</surname><given-names>A.M.</given-names></name></person-group><article-title>Isolation of a novel coronavirus from a man with pneumonia in Saudi Arabia</article-title><source>N. 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