MERS-CoV: Understanding the Latest Human Coronavirus Threat
Identifieur interne : 000804 ( Pmc/Corpus ); précédent : 000803; suivant : 000805MERS-CoV: Understanding the Latest Human Coronavirus Threat
Auteurs : Aasiyah Chafekar ; Burtram C. FieldingSource :
- Viruses [ 1999-4915 ] ; 2018.
Abstract
Human coronaviruses cause both upper and lower respiratory tract infections in humans. In 2012, a sixth human coronavirus (hCoV) was isolated from a patient presenting with severe respiratory illness. The 60-year-old man died as a result of renal and respiratory failure after admission to a hospital in Jeddah, Saudi Arabia. The aetiological agent was eventually identified as a coronavirus and designated Middle East respiratory syndrome coronavirus (MERS-CoV). MERS-CoV has now been reported in more than 27 countries across the Middle East, Europe, North Africa and Asia. As of July 2017, 2040 MERS-CoV laboratory confirmed cases, resulting in 712 deaths, were reported globally, with a majority of these cases from the Arabian Peninsula. This review summarises the current understanding of MERS-CoV, with special reference to the (i) genome structure; (ii) clinical features; (iii) diagnosis of infection; and (iv) treatment and vaccine development.
Url:
DOI: 10.3390/v10020093
PubMed: 29495250
PubMed Central: 5850400
Links to Exploration step
PMC:5850400Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">MERS-CoV: Understanding the Latest Human Coronavirus Threat</title><author><name sortKey="Chafekar, Aasiyah" sort="Chafekar, Aasiyah" uniqKey="Chafekar A" first="Aasiyah" last="Chafekar">Aasiyah Chafekar</name></author><author><name sortKey="Fielding, Burtram C" sort="Fielding, Burtram C" uniqKey="Fielding B" first="Burtram C." last="Fielding">Burtram C. Fielding</name></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">29495250</idno><idno type="pmc">5850400</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5850400</idno><idno type="RBID">PMC:5850400</idno><idno type="doi">10.3390/v10020093</idno><date when="2018">2018</date><idno type="wicri:Area/Pmc/Corpus">000804</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000804</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">MERS-CoV: Understanding the Latest Human Coronavirus Threat</title><author><name sortKey="Chafekar, Aasiyah" sort="Chafekar, Aasiyah" uniqKey="Chafekar A" first="Aasiyah" last="Chafekar">Aasiyah Chafekar</name></author><author><name sortKey="Fielding, Burtram C" sort="Fielding, Burtram C" uniqKey="Fielding B" first="Burtram C." last="Fielding">Burtram C. Fielding</name></author></analytic><series><title level="j">Viruses</title><idno type="eISSN">1999-4915</idno><imprint><date when="2018">2018</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Human coronaviruses cause both upper and lower respiratory tract infections in humans. In 2012, a sixth human coronavirus (hCoV) was isolated from a patient presenting with severe respiratory illness. The 60-year-old man died as a result of renal and respiratory failure after admission to a hospital in Jeddah, Saudi Arabia. The aetiological agent was eventually identified as a coronavirus and designated Middle East respiratory syndrome coronavirus (MERS-CoV). MERS-CoV has now been reported in more than 27 countries across the Middle East, Europe, North Africa and Asia. As of July 2017, 2040 MERS-CoV laboratory confirmed cases, resulting in 712 deaths, were reported globally, with a majority of these cases from the Arabian Peninsula. This review summarises the current understanding of MERS-CoV, with special reference to the (i) genome structure; (ii) clinical features; (iii) diagnosis of infection; and (iv) treatment and vaccine development.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Zaki, A M" uniqKey="Zaki A">A.M. Zaki</name></author><author><name sortKey="Van Boheemen, S" uniqKey="Van Boheemen S">S. van Boheemen</name></author><author><name sortKey="Bestebroer, T M" uniqKey="Bestebroer T">T.M. Bestebroer</name></author><author><name sortKey="Osterhaus, A D" uniqKey="Osterhaus A">A.D. Osterhaus</name></author><author><name sortKey="Fouchier, R A" uniqKey="Fouchier R">R.A. 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Viruses</journal-id><journal-id journal-id-type="iso-abbrev">Viruses</journal-id><journal-id journal-id-type="publisher-id">viruses</journal-id><journal-title-group><journal-title>Viruses</journal-title></journal-title-group><issn pub-type="epub">1999-4915</issn><publisher><publisher-name>MDPI</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">29495250</article-id><article-id pub-id-type="pmc">5850400</article-id><article-id pub-id-type="doi">10.3390/v10020093</article-id><article-id pub-id-type="publisher-id">viruses-10-00093</article-id><article-categories><subj-group subj-group-type="heading"><subject>Review</subject></subj-group></article-categories><title-group><article-title>MERS-CoV: Understanding the Latest Human Coronavirus Threat</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Chafekar</surname><given-names>Aasiyah</given-names></name></contrib><contrib contrib-type="author"><name><surname>Fielding</surname><given-names>Burtram C.</given-names></name><xref rid="c1-viruses-10-00093" ref-type="corresp">*</xref></contrib></contrib-group><aff id="af1-viruses-10-00093">Molecular Biology and Virology Research Laboratory, Department of Medical Biosciences, Faculty of Natural Sciences, University of the Western Cape, Private Bag X17, Robert Sobukwe Drive, Bellville 7535, South Africa;<email>achafekar@uwc.ac.za</email></aff><author-notes><corresp id="c1-viruses-10-00093"><label>*</label>Correspondence: <email>bfielding@uwc.ac.za</email>; Tel.: +27-21-959-3620</corresp></author-notes><pub-date pub-type="epub"><day>24</day><month>2</month><year>2018</year></pub-date><pub-date pub-type="collection"><month>2</month><year>2018</year></pub-date><volume>10</volume><issue>2</issue><elocation-id>93</elocation-id><history><date date-type="received"><day>29</day><month>11</month><year>2017</year></date><date date-type="accepted"><day>02</day><month>2</month><year>2018</year></date></history><permissions><copyright-statement>© 2018 by the authors.</copyright-statement><copyright-year>2018</copyright-year><license license-type="open-access"><license-p>Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (<ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">http://creativecommons.org/licenses/by/4.0/</ext-link>).</license-p></license></permissions><abstract><p>Human coronaviruses cause both upper and lower respiratory tract infections in humans. In 2012, a sixth human coronavirus (hCoV) was isolated from a patient presenting with severe respiratory illness. The 60-year-old man died as a result of renal and respiratory failure after admission to a hospital in Jeddah, Saudi Arabia. The aetiological agent was eventually identified as a coronavirus and designated Middle East respiratory syndrome coronavirus (MERS-CoV). MERS-CoV has now been reported in more than 27 countries across the Middle East, Europe, North Africa and Asia. As of July 2017, 2040 MERS-CoV laboratory confirmed cases, resulting in 712 deaths, were reported globally, with a majority of these cases from the Arabian Peninsula. This review summarises the current understanding of MERS-CoV, with special reference to the (i) genome structure; (ii) clinical features; (iii) diagnosis of infection; and (iv) treatment and vaccine development.</p></abstract><kwd-group><kwd>human coronavirus</kwd><kwd>MERS-CoV</kwd><kwd>clinical features</kwd><kwd>upper respiratory tract infections</kwd><kwd>lower respiratory tract infections</kwd><kwd>respiratory viruses</kwd></kwd-group></article-meta></front><body><sec id="sec1-viruses-10-00093"><title>1. Introduction</title><p>Given the diversity of animal coronaviruses, it was not surprising when another human coronavirus was isolated from a patient presenting with severe respiratory illness in June 2012. The 60-year-old man died as a result of renal and respiratory failure 11 days after admission to a hospital in Jeddah, Saudi Arabia [<xref rid="B1-viruses-10-00093" ref-type="bibr">1</xref>]. The novel etiological agent was subsequently named Middle East Respiratory syndrome coronavirus (MERS-CoV) [<xref rid="B2-viruses-10-00093" ref-type="bibr">2</xref>]. MERS-CoV is one of six known human coronaviruses that cause respiratory disease in humans and, with a mortality rate >35% [<xref rid="B3-viruses-10-00093" ref-type="bibr">3</xref>], it is the first highly pathogenic human coronavirus to emerge since the global scare caused by the severe acute respiratory syndrome coronavirus (SARS-CoV) in 2003.</p><p>With the Kingdom of Saudi Arabia, the focal point of an ongoing MERS-CoV outbreak, the large number of religious pilgrims congregating annually in Saudi Arabia drastically increases the potential for the uncontrolled global spread of MERS-CoV infections [<xref rid="B4-viruses-10-00093" ref-type="bibr">4</xref>]. In fact, infections have already been reported in more than 27 countries across the Middle East, Europe, North Africa and Asia [<xref rid="B5-viruses-10-00093" ref-type="bibr">5</xref>,<xref rid="B6-viruses-10-00093" ref-type="bibr">6</xref>,<xref rid="B7-viruses-10-00093" ref-type="bibr">7</xref>,<xref rid="B8-viruses-10-00093" ref-type="bibr">8</xref>].</p><p>This review focusses on the current information of MERS-CoV, with special reference to the genome structure, clinical features, diagnosis of infection and treatment and vaccine development. We also look at future prospects for MERS-CoV spread and prevention.</p></sec><sec id="sec2-viruses-10-00093"><title>2. Genome Structure and Gene Functions</title><p>MERS-CoV, a lineage C <italic>Betacoronavirus</italic> (βCoVs), has a positive-sense single-stranded RNA (ssRNA) genome about 30-kb in size [<xref rid="B9-viruses-10-00093" ref-type="bibr">9</xref>,<xref rid="B10-viruses-10-00093" ref-type="bibr">10</xref>]. As of 2016, phylogenetic analysis of MERS-CoV has been done on 182 full-length genomes or multiple concatenated genome fragments, including 94 from humans and 88 from dromedary camels [<xref rid="B11-viruses-10-00093" ref-type="bibr">11</xref>,<xref rid="B12-viruses-10-00093" ref-type="bibr">12</xref>]. The MERS-CoV genomes share more than 99% sequence identity, indicating a low mutation rate and low variance among the genomes. MERS-CoV genomes are roughly divided into two clades: clade A, which contains only a few strains, and clade B, to which most strains belong [<xref rid="B12-viruses-10-00093" ref-type="bibr">12</xref>].</p><p>As with other CoV genomes, the first 5′ two-thirds of the MERS-CoV genome consist of the replicase complex (ORF1a and ORF1b). The remaining 3′ one-third encodes the structural proteins spike (S), envelope (E), membrane (M), and nucleocapsid (N), as well as five accessory proteins (ORF3, ORF4a, ORF4b, ORF5 and ORF8b) that are not required for genome replication (<xref ref-type="fig" rid="viruses-10-00093-f001">Figure 1</xref>), but are likely involved in pathogenesis [<xref rid="B9-viruses-10-00093" ref-type="bibr">9</xref>,<xref rid="B13-viruses-10-00093" ref-type="bibr">13</xref>,<xref rid="B14-viruses-10-00093" ref-type="bibr">14</xref>,<xref rid="B15-viruses-10-00093" ref-type="bibr">15</xref>,<xref rid="B16-viruses-10-00093" ref-type="bibr">16</xref>,<xref rid="B17-viruses-10-00093" ref-type="bibr">17</xref>] . The flanking regions of the genome contain the 5′ and 3′ untranslated regions (UTR) [<xref rid="B13-viruses-10-00093" ref-type="bibr">13</xref>,<xref rid="B14-viruses-10-00093" ref-type="bibr">14</xref>]. Typical of the coronaviruses, the MERS-CoV accessory proteins do not share homology with any known host or virus protein, other than those of its closely related lineage C βCoVs [<xref rid="B12-viruses-10-00093" ref-type="bibr">12</xref>].</p><p>MERS-CoV structural and accessory protein-coding plasmids transiently transfected into cells showed that, while ORF4b was localised mostly in the nucleus, all of the other proteins (S, E, M, N, ORF3, ORF4a and ORF5) localised to the cytoplasm [<xref rid="B18-viruses-10-00093" ref-type="bibr">18</xref>]. Furthermore, studies with MERS-CoV deletion-mutants of ORFs 3 to 5 are attenuated for replication in human airway-derived (Calu-3) cells [<xref rid="B19-viruses-10-00093" ref-type="bibr">19</xref>], and deletion-mutants of ORFs 4a and 4b are attenuated for replication in hepatic carcinoma-derived (Huh-7) cells [<xref rid="B16-viruses-10-00093" ref-type="bibr">16</xref>,<xref rid="B20-viruses-10-00093" ref-type="bibr">20</xref>]. This clearly points to important putative roles for the MERS-CoV accessory proteins in viral replication, at least in an in vitro setting [<xref rid="B21-viruses-10-00093" ref-type="bibr">21</xref>].</p><p>The principal response of mammalian cells to viral infection is the activation of the type I interferon (IFN)-mediated innate immune response through the production of type I IFNs (IFN-α and IFN-β). On the other hand, evasion of host innate immunity through IFN antagonism is a critical component of viral pathogenesis and is mediated by virus-encoded IFN antagonist proteins. Each protein blocks one or more key signaling proteins in the IFN and NF-κB pathways to enhance viral replication and pathogenesis [<xref rid="B22-viruses-10-00093" ref-type="bibr">22</xref>,<xref rid="B23-viruses-10-00093" ref-type="bibr">23</xref>,<xref rid="B24-viruses-10-00093" ref-type="bibr">24</xref>,<xref rid="B25-viruses-10-00093" ref-type="bibr">25</xref>]. Coronaviruses have similarly evolved these mechanisms to impede or bypass the innate immunity of their hosts at various levels, which ultimately contribute to coronavirus virulence. Various coronavirus proteins have previously been implicated in the disruption of signal transduction events required for the IFN response [<xref rid="B26-viruses-10-00093" ref-type="bibr">26</xref>], often by interfering with the host’s type I interferon induction.</p><p>Evidence of MERS-CoV inducing type I IFN only weakly and late in infection (9–15) suggests that MERS-CoV has also evolved mechanisms to evade the host immune system. In fact, MERS-CoV M, ORF4a, ORF4b and ORF5 proteins are reported to be strong IFN antagonists [<xref rid="B18-viruses-10-00093" ref-type="bibr">18</xref>]. Further studies, using the transient overexpression of MERS-CoV accessory protein ORF4a, ORF4b, and ORF5, show that the MERS-CoV accessory proteins inhibit both type I IFN induction [<xref rid="B18-viruses-10-00093" ref-type="bibr">18</xref>,<xref rid="B27-viruses-10-00093" ref-type="bibr">27</xref>,<xref rid="B28-viruses-10-00093" ref-type="bibr">28</xref>] and NF-kappaB signaling pathways [<xref rid="B28-viruses-10-00093" ref-type="bibr">28</xref>]. MERS-CoV ORF4a, a double-stranded RNA (dsRNA) binding protein [<xref rid="B27-viruses-10-00093" ref-type="bibr">27</xref>], potentially acts as an antagonist of the antiviral activity of IFN via the inhibition of both the interferon production (IFN-β promoter activity, IRF-3/7 and NF-κB activation) and the ISRE promoter element signaling pathways [<xref rid="B18-viruses-10-00093" ref-type="bibr">18</xref>]. MERS-CoV ORF4b, on the other hand, is an enzyme in the 2H-phosphoesterase (2H-PE) family with phosphodiesterase (PDE) activity. Even though MERS-CoV ORF4b is detected primarily in the nucleus of both infected and transfected cells [<xref rid="B18-viruses-10-00093" ref-type="bibr">18</xref>,<xref rid="B27-viruses-10-00093" ref-type="bibr">27</xref>,<xref rid="B28-viruses-10-00093" ref-type="bibr">28</xref>], the expression levels of cytoplasmic MERS-CoV ORF4b are still sufficient to inhibit activation of RNase L, an interferon-induced potent antiviral activity [<xref rid="B18-viruses-10-00093" ref-type="bibr">18</xref>,<xref rid="B28-viruses-10-00093" ref-type="bibr">28</xref>]. MERS-CoV ORF4b is the first identified RNase L antagonist expressed by a human or bat coronavirus and provides a possible MERS-CoV mechanism for evasion of innate immunity by inhibiting the type I IFN and NF-kappaβ signaling pathways [<xref rid="B16-viruses-10-00093" ref-type="bibr">16</xref>,<xref rid="B28-viruses-10-00093" ref-type="bibr">28</xref>]. The MERS-CoV replicase proteins, including nsp1, nsp3 and nsp14, were also shown to interfere with the innate immune response signaling pathways through different mechanisms [<xref rid="B21-viruses-10-00093" ref-type="bibr">21</xref>,<xref rid="B29-viruses-10-00093" ref-type="bibr">29</xref>,<xref rid="B30-viruses-10-00093" ref-type="bibr">30</xref>]. Evidently, MERS-CoV has developed various mechanisms to evade the host immune system.</p></sec><sec id="sec3-viruses-10-00093"><title>3. Clinical Features</title><p>The median age of persons with laboratory-confirmed MERS-CoV infection is 49 years (range, <1–94 years); 65% of patients are males. The median time from illness onset to hospitalization is approximately four days, resulting in a median length of stay of 41 days [<xref rid="B31-viruses-10-00093" ref-type="bibr">31</xref>]. Currently, among all patients, the morbidity rate is approximately 36% [<xref rid="B3-viruses-10-00093" ref-type="bibr">3</xref>], with the median time from the onset of symptoms to death 11.5 days [<xref rid="B32-viruses-10-00093" ref-type="bibr">32</xref>]. Chest radiography and computed tomography findings are generally consistent with viral pneumonitis and acute respiratory distress syndrome [<xref rid="B33-viruses-10-00093" ref-type="bibr">33</xref>]. Laboratory findings include lymphopenia, thrombocytopenia and elevated lactate dehydrogenase levels [<xref rid="B1-viruses-10-00093" ref-type="bibr">1</xref>,<xref rid="B31-viruses-10-00093" ref-type="bibr">31</xref>,<xref rid="B34-viruses-10-00093" ref-type="bibr">34</xref>,<xref rid="B35-viruses-10-00093" ref-type="bibr">35</xref>,<xref rid="B36-viruses-10-00093" ref-type="bibr">36</xref>,<xref rid="B37-viruses-10-00093" ref-type="bibr">37</xref>,<xref rid="B38-viruses-10-00093" ref-type="bibr">38</xref>,<xref rid="B39-viruses-10-00093" ref-type="bibr">39</xref>], with some cases with a consumptive coagulopathy and elevations in creatinine, lactate dehydrogenase and liver enzymes [<xref rid="B31-viruses-10-00093" ref-type="bibr">31</xref>,<xref rid="B33-viruses-10-00093" ref-type="bibr">33</xref>,<xref rid="B40-viruses-10-00093" ref-type="bibr">40</xref>].</p><p>The clinical spectrum of MERS-CoV infection ranges from asymptomatic infection [<xref rid="B41-viruses-10-00093" ref-type="bibr">41</xref>,<xref rid="B42-viruses-10-00093" ref-type="bibr">42</xref>,<xref rid="B43-viruses-10-00093" ref-type="bibr">43</xref>] to rapidly progressive, acute respiratory distress syndrome, septic shock and multi-organ failure and death (see [<xref rid="B32-viruses-10-00093" ref-type="bibr">32</xref>,<xref rid="B44-viruses-10-00093" ref-type="bibr">44</xref>] for review of clinical spectrum). Initial symptoms are often nonspecific and patients report general malaise, including low grade fever, chills, headache, nonproductive cough, dyspnea, and myalgia [<xref rid="B38-viruses-10-00093" ref-type="bibr">38</xref>,<xref rid="B45-viruses-10-00093" ref-type="bibr">45</xref>]. Other symptoms can include sore throat and similar to SARS-CoV, MERS-CoV patients can also present with gastrointestinal symptoms such as anorexia, nausea and vomiting, abdominal pain and diarrhea [<xref rid="B46-viruses-10-00093" ref-type="bibr">46</xref>,<xref rid="B47-viruses-10-00093" ref-type="bibr">47</xref>,<xref rid="B48-viruses-10-00093" ref-type="bibr">48</xref>]. Atypical presentations, including mild respiratory illness without fever and diarrheal illness, preceding the development of pneumonia have been documented [<xref rid="B49-viruses-10-00093" ref-type="bibr">49</xref>]. Up to 50% of adult symptomatic patients require intensive care unit (ICU) treatment. These patients often show no sign of improvement and 40–70% typically require mechanical ventilation within the first week [<xref rid="B32-viruses-10-00093" ref-type="bibr">32</xref>,<xref rid="B41-viruses-10-00093" ref-type="bibr">41</xref>,<xref rid="B50-viruses-10-00093" ref-type="bibr">50</xref>]. Renal replacement therapy is required for between 22–70% of critically ill patients [<xref rid="B31-viruses-10-00093" ref-type="bibr">31</xref>,<xref rid="B34-viruses-10-00093" ref-type="bibr">34</xref>,<xref rid="B35-viruses-10-00093" ref-type="bibr">35</xref>,<xref rid="B40-viruses-10-00093" ref-type="bibr">40</xref>,<xref rid="B51-viruses-10-00093" ref-type="bibr">51</xref>], with the higher-end of the estimation possibly due to over-estimation as a result of hospital-acquired infections in patients with pre-existing renal disease [<xref rid="B32-viruses-10-00093" ref-type="bibr">32</xref>,<xref rid="B35-viruses-10-00093" ref-type="bibr">35</xref>].</p><p>MERS-CoV is linked with more severe disease in older people, people with weakened immune systems, and those with chronic diseases such as cancer, chronic lung disease and diabetes. The majority of patients who are hospitalized with MERS-CoV infection had chronic co-morbidities such as obesity, diabetes, hypertension, cardiovascular diseases or end-stage renal disease [<xref rid="B40-viruses-10-00093" ref-type="bibr">40</xref>,<xref rid="B52-viruses-10-00093" ref-type="bibr">52</xref>,<xref rid="B53-viruses-10-00093" ref-type="bibr">53</xref>,<xref rid="B54-viruses-10-00093" ref-type="bibr">54</xref>]. In fact, about 75% of patients testing positive for MERS-CoV have at least one co-morbid disease; fatal cases are more likely to have an underlying condition (86% among fatal cases vs. 42% among recovered or asymptomatic cases) [<xref rid="B33-viruses-10-00093" ref-type="bibr">33</xref>].</p><p>Interestingly, MERS-CoV cases have been reported mainly in adults [<xref rid="B55-viruses-10-00093" ref-type="bibr">55</xref>], with children rarely affected [<xref rid="B56-viruses-10-00093" ref-type="bibr">56</xref>,<xref rid="B57-viruses-10-00093" ref-type="bibr">57</xref>]. Even so, a recent case study of a MERS-CoV infected a 9-month-old child, newly diagnosed to have infantile nephrotic syndrome, showed complications that resulted in severe respiratory symptoms, multi-organ dysfunction and death [<xref rid="B58-viruses-10-00093" ref-type="bibr">58</xref>]. In another study of 11 pediatric cases that tested positive for MERS-CoV, the two symptomatic patients had Down’s syndrome and cystic fibrosis, respectively, indicating that severe disease could potentially occur in children with serious underlying conditions [<xref rid="B43-viruses-10-00093" ref-type="bibr">43</xref>]. Even with these reported pediatric cases, data on infection in children remain scarce, making it difficult to ascertain whether MERS-CoV is really a predominantly adult disease, or whether it often presents differently in children.</p><p>Simultaneous infection of the respiratory tract with at least two viruses is common in hospitalized patients, and although it is not clear whether these infections are more, or less, severe than single virus infections [<xref rid="B59-viruses-10-00093" ref-type="bibr">59</xref>], mixed clinical features are commonly observed [<xref rid="B60-viruses-10-00093" ref-type="bibr">60</xref>]; this makes clinical diagnosis unreliable and severely limit epidemiological studies of etiological agents. Similar to other respiratory viruses, MERS-CoV has been found in combination with a second respiratory virus, such as influenza A virus [<xref rid="B48-viruses-10-00093" ref-type="bibr">48</xref>,<xref rid="B61-viruses-10-00093" ref-type="bibr">61</xref>] respiratory syncytial virus, human parainfluenza virus 3 or human metapneumovirus [<xref rid="B62-viruses-10-00093" ref-type="bibr">62</xref>,<xref rid="B63-viruses-10-00093" ref-type="bibr">63</xref>,<xref rid="B64-viruses-10-00093" ref-type="bibr">64</xref>]. MERS-CoV infected patients requiring mechanical ventilation also exhibited a similar co-infection profile with nosocomial bacterial infections including, <italic>Klebsiella pneumoniae</italic>, <italic>Staphylococcus aureus</italic>, <italic>Acinetobacter species</italic> and <italic>Candida</italic> species [<xref rid="B47-viruses-10-00093" ref-type="bibr">47</xref>,<xref rid="B65-viruses-10-00093" ref-type="bibr">65</xref>]. Preceding or concurrent viral respiratory tract infections can predispose the host to secondary co-infections from other microorganisms throughout the airway. The mechanisms by which viruses promote these superinfections are diverse and replete [<xref rid="B66-viruses-10-00093" ref-type="bibr">66</xref>]. As yet, not much is known as to how MERS-CoV damages the airway and dysregulate the lung barrier function, which, in turn, supports the adherence and invasion of other pathogens into normally sterile sites within the respiratory tract.</p><p>Neuromuscular complications are not rare during MERS treatment, and could simply have been underdiagnosed previously [<xref rid="B67-viruses-10-00093" ref-type="bibr">67</xref>]. The first cases of severe neurological syndrome, characterized by varying degrees of disturbed consciousness, ataxia, focal motor deficit and bilateral hyper-intense lesions were reported from a retrospective study of patients in ICU [<xref rid="B68-viruses-10-00093" ref-type="bibr">68</xref>]. Another subsequent small retrospective study in Saudi Arabia reported that 25.7% of MERS patients developed confusion and 8.6% experienced some kind of seizure [<xref rid="B69-viruses-10-00093" ref-type="bibr">69</xref>]. To date, other cases with central nervous system involvement, including one case of intracerebral haemorrhage as a result of thrombocytopenia, disseminated intravascular coagulation and platelet dysfunction, one case of critical illness polyneuropathy [<xref rid="B70-viruses-10-00093" ref-type="bibr">70</xref>] and four cases that included Bickerstaff’s encephalitis overlapping with Guillain–Barre syndrome, intensive-care-unit-acquired weakness, or other toxic or infectious neuropathies [<xref rid="B67-viruses-10-00093" ref-type="bibr">67</xref>], have been reported. Neurological complications in the latter study did not appear concomitantly with respiratory symptoms, but were delayed by 2–3 weeks [<xref rid="B67-viruses-10-00093" ref-type="bibr">67</xref>].</p><p>MERS-CoV can be detected in respiratory tract secretions, with tracheal secretions and broncho-alveolar lavage specimens containing a higher viral load than nasopharyngeal swabs. The virus has also been detected in feces, serum and urine [<xref rid="B48-viruses-10-00093" ref-type="bibr">48</xref>,<xref rid="B71-viruses-10-00093" ref-type="bibr">71</xref>,<xref rid="B72-viruses-10-00093" ref-type="bibr">72</xref>,<xref rid="B73-viruses-10-00093" ref-type="bibr">73</xref>]. Virus excretion peaks approximately 10 days after the onset of symptoms [<xref rid="B48-viruses-10-00093" ref-type="bibr">48</xref>], but viable viruses can be shed through respiratory secretions for up to 25 days from clinically fully recovered patients. In the healthcare setting, MERS-CoV has been isolated from environmental objects such as bed sheets, bedrails, IV fluid hangers and X-ray devices [<xref rid="B7-viruses-10-00093" ref-type="bibr">7</xref>]. Another study also reported that MERS-CoV could survive for longer than two days at 20 °C and 40% relative humidity, confirming the risk of contact or fomite transmission in healthcare settings [<xref rid="B74-viruses-10-00093" ref-type="bibr">74</xref>]. Viral RNA, on the other hand, is detected for up to five days on environmental surfaces following the last positive PCR from patients’ respiratory samples; RNA was detected in samples from anterooms, medical devices and air-ventilating equipment [<xref rid="B7-viruses-10-00093" ref-type="bibr">7</xref>], but this is not necessarily indicative of viable virus.</p></sec><sec id="sec4-viruses-10-00093"><title>4. Diagnosis of Infection</title><p>With no specific, reliable antiviral drug or vaccine approved for clinical use in MERS-CoV infections, rapid diagnostic tests are required to manage outbreaks of this virus. The first probe and primer sets for MERS-CoV detection by real-time RT-PCR were developed shortly after the initial reports of the disease [<xref rid="B75-viruses-10-00093" ref-type="bibr">75</xref>,<xref rid="B76-viruses-10-00093" ref-type="bibr">76</xref>]. Other early diagnostic tools included virus culture in Vero and LLCMK2 cells [<xref rid="B1-viruses-10-00093" ref-type="bibr">1</xref>,<xref rid="B77-viruses-10-00093" ref-type="bibr">77</xref>], but isolation and identification of viruses in cell culture is a slow, specialized and insensitive method [<xref rid="B78-viruses-10-00093" ref-type="bibr">78</xref>].</p><p>Laboratory detection and confirmation of MERS-CoV infections has broadly included (i) molecular detection of MERS-CoV RNA; (ii) MERS-CoV antigen detection; or (iii) assays to identify a humoral response to prior MERS-CoV infection among humans [<xref rid="B79-viruses-10-00093" ref-type="bibr">79</xref>] (<xref ref-type="table" rid="viruses-10-00093-t001">Table 1</xref>). These assays have been used with varying degrees of success in terms of specificity, sensitivity, etc.</p><p>Currently, according to the WHO case definition, a positive real-time RT-PCR assay, targeting at least two different genomic regions, is used to confirm MERS-CoV infection (<uri xlink:href="http://www.who.int/csr/disease/coronavirus_infections/case_definition/en/index.html">http://www.who.int/csr/disease/coronavirus_infections/case_definition/en/index.html</uri>) [<xref rid="B80-viruses-10-00093" ref-type="bibr">80</xref>]. Of the different assay probes and primers sets used, those targeting ORF1a and upstream of the E gene show the highest sensitivity and remain the most widely used targets for MERS-CoV detection [<xref rid="B75-viruses-10-00093" ref-type="bibr">75</xref>,<xref rid="B81-viruses-10-00093" ref-type="bibr">81</xref>]. Additionally, a single positive assay result, confirmed by gene sequencing, can also be considered positive for MERS-CoV infection. A stumbling block here, though, is the fact that, when compared to real-time PCR, conventional RT-PCR typically generates lower quality sequence-ready templates [<xref rid="B80-viruses-10-00093" ref-type="bibr">80</xref>,<xref rid="B82-viruses-10-00093" ref-type="bibr">82</xref>,<xref rid="B83-viruses-10-00093" ref-type="bibr">83</xref>,<xref rid="B84-viruses-10-00093" ref-type="bibr">84</xref>,<xref rid="B85-viruses-10-00093" ref-type="bibr">85</xref>,<xref rid="B86-viruses-10-00093" ref-type="bibr">86</xref>], thereby limiting the usefulness of conventional RT-PCR in these single positive-sequencing assays.</p><p>Molecular tests can detect nucleic acids derived from MERS-CoV in clinical respiratory, serum, and stool specimens [<xref rid="B81-viruses-10-00093" ref-type="bibr">81</xref>,<xref rid="B87-viruses-10-00093" ref-type="bibr">87</xref>]. However, a major obstacle of conventional nucleic acid-based tests is that they require specialized molecular techniques and equipment, and are therefore not appropriate for point-of-care testing or bedside diagnosis. For this reason, for effective diagnosis and treatment of MERS-CoV infection, it is necessary to develop alternative methods that can be adapted to rapid and reliable clinical detection of MERS-CoV antigens. Here, the most appropriate tests would be assays detecting viral antigens or antibodies in the infected host [<xref rid="B87-viruses-10-00093" ref-type="bibr">87</xref>].</p></sec><sec id="sec5-viruses-10-00093"><title>5. Animal Models</title><p>Not only are laboratory animal species often used as models for human disease progression, they are also needed to study and evaluate novel therapies against emerging viruses [<xref rid="B105-viruses-10-00093" ref-type="bibr">105</xref>]. Studies have shown that rabbits [<xref rid="B106-viruses-10-00093" ref-type="bibr">106</xref>], ferrets [<xref rid="B107-viruses-10-00093" ref-type="bibr">107</xref>], Syrian hamsters [<xref rid="B108-viruses-10-00093" ref-type="bibr">108</xref>] and wild-type mice [<xref rid="B109-viruses-10-00093" ref-type="bibr">109</xref>] are not suitable as models of MERS-CoV infection. More recently, four transgenic mouse models for MERS-CoV infection have been developed. In the first, a modified adenovirus expressing human DPP4 (huDPP4) is introduced intranasally to mice, which results in the expression of huDPP4 in all cells of the lung, not just those that natively express DPP4. In this model, mice show transient human DPP4 expression and mild lung disease. A concern with this model is that cells constitutively expressing DPP4 will be infected and the role of a broader infection of all cell types may change pathogenesis [<xref rid="B110-viruses-10-00093" ref-type="bibr">110</xref>]. In the second model, a transgenic mouse was produced that expresses huDPP4 systemically. In this model, MERS-CoV infection leads to high levels of viral RNA and inflammation in the lungs, but, unfortunately, significant inflammation and viral RNA are also detected in the brains of infected mice, which represent a non-physiological expression pattern [<xref rid="B111-viruses-10-00093" ref-type="bibr">111</xref>]. In the third model, a novel transgenic humanized mouse model was generated by replacing the mouse DPP4 coding sequence with that encoding huDPP4, ensuring correct physiological expression of huDPP4. Mice in this model show lung pathology consistent with the radiographic findings of interstitial pneumonia and significant lung disease as seen in humans infected with MERS-CoV. This suggests that this mouse model recapitulates pathological sequelae that are seen in MERS-CoV infection of humans. Importantly, unlike what is seen in other mouse models of MERS-CoV infection, virus replication and pathology in the huDPP4 mice are localized in the lungs and no inflammation develops in the brain, ensuring a more physiological accurate model of the human disease [<xref rid="B112-viruses-10-00093" ref-type="bibr">112</xref>]. Finally, in 2016, Cockrell et al. generated a mouse model permissive for MERS-CoV infection, but with functional DPP4 immune function. Infecting this DDP4-chimeric mouse with a mouse-adapted strain of MERS-CoV, mimics MERS-CoV-induced respiratory disease without bystander neurologic disease [<xref rid="B113-viruses-10-00093" ref-type="bibr">113</xref>]. </p><p>Non-human primate models, including the rhesus macaque [<xref rid="B114-viruses-10-00093" ref-type="bibr">114</xref>,<xref rid="B115-viruses-10-00093" ref-type="bibr">115</xref>,<xref rid="B116-viruses-10-00093" ref-type="bibr">116</xref>] and common marmoset [<xref rid="B117-viruses-10-00093" ref-type="bibr">117</xref>] have also been reported as suitable animal models of MERS-CoV infection. Even though both species are susceptible to MERS-CoV infection, the extent of virus replication and severity of disease vary [<xref rid="B105-viruses-10-00093" ref-type="bibr">105</xref>]. Rhesus macaques infected with MERS-CoV via intra-tracheal inoculation show clinical signs of disease, virus replication, histological lesions and neutralizing antibody production, indicating that this monkey model is suitable for studies of MERS-CoV infection [<xref rid="B116-viruses-10-00093" ref-type="bibr">116</xref>]. On the other hand, the common marmoset reproduces several, but not all, features of MERS-CoV infection, and can potentially be used to evaluate novel therapies for human use [<xref rid="B105-viruses-10-00093" ref-type="bibr">105</xref>,<xref rid="B117-viruses-10-00093" ref-type="bibr">117</xref>].</p></sec><sec id="sec6-viruses-10-00093"><title>6. Treatment and Vaccine Development</title><p>When no vaccines or specific antiviral drugs are available during an outbreak, nonspecific therapeutic interventions are often introduced to prevent severe morbidity and mortality. However, for this to be done effectively, a basic understanding of the pathogenesis of the disease is required and interventions are implemented based on observations of the clinical course of disease and complications. Due to the nature of many diseases, however, it is often not possible to assess, or systematically compare, different therapeutic approaches during an outbreak [<xref rid="B118-viruses-10-00093" ref-type="bibr">118</xref>]. Similarly, in the case of MERS-CoV, it is necessary to monitor epidemic patterns and investigate the spread of infections to efficiently identify, control and prevent possible epidemics. For MERS-CoV infections, supportive care, which includes rest, fluids and analgesics are used, and mainly depends on the provision of organ support and management of complications [<xref rid="B49-viruses-10-00093" ref-type="bibr">49</xref>,<xref rid="B119-viruses-10-00093" ref-type="bibr">119</xref>,<xref rid="B120-viruses-10-00093" ref-type="bibr">120</xref>]. Broad-spectrum antimicrobials, antivirals [<xref rid="B121-viruses-10-00093" ref-type="bibr">121</xref>,<xref rid="B122-viruses-10-00093" ref-type="bibr">122</xref>], interferon-α2b (96) and antifungals can be used to minimize the risk of co-infection with opportunistic pathogens [<xref rid="B49-viruses-10-00093" ref-type="bibr">49</xref>,<xref rid="B119-viruses-10-00093" ref-type="bibr">119</xref>].</p><p>Interestingly, combination treatment with ribavirin and interferons inhibits MERS-CoV replication in vitro, and it was also shown to improve clinical outcomes in MERS-CoV-infected non-human primates. However, this treatment in the rhesus macaques was initiated very soon after viral challenge (~8 h), resulting in reduced disease severity in the rhesus macaque model. This appears to simulate mild-to-moderate human MERS-CoV cases, making it difficult to extrapolate the outcome of this early intervention in severe human cases. Even though the authors recommended that combined IFN-α2b and ribavirin therapy should be considered as an early intervention therapy for MERS-CoV [<xref rid="B115-viruses-10-00093" ref-type="bibr">115</xref>], we also need to keep in mind that, due to the limited effective therapeutic window of opportunity, broad spectrum antivirals might not be sufficient to treat severe MERS-CoV patients [<xref rid="B122-viruses-10-00093" ref-type="bibr">122</xref>].</p><p>Resveratrol has been shown to inhibit various human viruses in vivo and in vitro, including influenza virus, Epstein–Barr virus, herpes simplex virus, respiratory syncytial virus, HIV-1, varicella zoster virus, enterovirus 71, human metapneumovirus, human rhinovirus 16, polyomavirus and cytomegalovirus ([<xref rid="B123-viruses-10-00093" ref-type="bibr">123</xref>,<xref rid="B124-viruses-10-00093" ref-type="bibr">124</xref>] for review). The antiviral effects of resveratrol are mainly associated with the inhibition of viral replication, protein synthesis, gene expression, and/or nucleic acid synthesis [<xref rid="B123-viruses-10-00093" ref-type="bibr">123</xref>,<xref rid="B124-viruses-10-00093" ref-type="bibr">124</xref>,<xref rid="B125-viruses-10-00093" ref-type="bibr">125</xref>]. In an in vitro study, resveratrol was shown to significantly inhibit MERS-CoV infection, most likely due to the observed inhibition of MERS-CoV nucleocapsid (N) protein expression [<xref rid="B126-viruses-10-00093" ref-type="bibr">126</xref>], a multifunctional protein essential for CoV replication [<xref rid="B127-viruses-10-00093" ref-type="bibr">127</xref>]. Furthermore, resveratrol downregulated apoptosis induced by MERS-CoV, thereby prolonging cellular survival post-infection [<xref rid="B126-viruses-10-00093" ref-type="bibr">126</xref>]. Although the beneficial roles of resveratrol in several viral diseases have been well documented, adverse effects have been also been reported, including increasing viral RNA replication during Hep-C virus infection in vitro (OR6 cells) [<xref rid="B128-viruses-10-00093" ref-type="bibr">128</xref>], strong cytotoxicity in cultured cells [<xref rid="B129-viruses-10-00093" ref-type="bibr">129</xref>], as well as enhanced HBV transcription and replication in vitro and in vivo [<xref rid="B130-viruses-10-00093" ref-type="bibr">130</xref>]. Clearly, the antiviral potential of resveratrol in MERS-CoV infections needs to be studied more extensively, but, based on the various unintended negative effects, this needs to proceed with caution.</p><p>More recently, de Wilde et al. [<xref rid="B131-viruses-10-00093" ref-type="bibr">131</xref>] reported that in an in vitro test, low-micromolar concentrations of alisporivir, a non-immunosuppressive cyclosporin A-analog, inhibit the replication of four different coronaviruses, including MERS-CoV. In this study, ribavirin was found to further potentiate the antiviral effect of alisporivir in the in vitro infection models, which warrants the further exploration of cyclophilin inhibitors as potential host-directed, broad-spectrum inhibitors of coronavirus replication [<xref rid="B131-viruses-10-00093" ref-type="bibr">131</xref>].</p><p>3C-like protease (3CL<sup>pro</sup>)—analogous to picornavirus 3C protease (3C<sup>pro</sup>)—is functionally important in the CoV replication cycle [<xref rid="B132-viruses-10-00093" ref-type="bibr">132</xref>] and is thus regarded as a validated drug target. Peptidomimetic inhibitors of enterovirus 3C<sup>pro</sup> (6b, 6c and 6d) inhibited MERS-CoV 3CL<sup>pro</sup> and in MERS-CoV-infected cells, the inhibitors showed antiviral activity by downregulating viral protein production in cells, as well as reducing release of infectious viral particles into culture supernatants. These compounds exhibited good selectivity index and should be investigated further as, not only an inhibitor of MERS-CoV replication and infections, but also as broad-spectrum antiviral activity drugs against other CoVs and picornaviruses [<xref rid="B133-viruses-10-00093" ref-type="bibr">133</xref>]. Our laboratory has also previously screened the ZINC drugs-now library for candidates with potential anti-3CL<sup>pro</sup> activity with a consensus high-throughput pharmacophore modelling and molecular docking approach. Molecular dynamics was used to confirm results obtained from structure-based techniques, resulting in a highly defined hit-list of 19 compounds, which represent valuable scaffolds that could be used as a basis for future anti-coronaviral inhibitor discovery experiments [<xref rid="B50-viruses-10-00093" ref-type="bibr">50</xref>,<xref rid="B134-viruses-10-00093" ref-type="bibr">134</xref>]. Even with all of these potential anti-MERS-CoV candidates, no experimental interventions have demonstrated significant benefit in acutely ill patients in a consistent or controlled manner. Therefore, supportive management, adapted from guidelines developed for SARS-CoV, has thus far been the mainstay of MERS-CoV treatment [<xref rid="B135-viruses-10-00093" ref-type="bibr">135</xref>].</p><p>Because of the highly sophisticated immune evasion mechanisms of viral pathogens, human vaccine development remains a major challenge [<xref rid="B136-viruses-10-00093" ref-type="bibr">136</xref>]. In addition, the development of safe and effective coronavirus vaccines has been even more challenging, being curtailed by major obstacles, including (1) coronavirus immunity often wanes rapidly; (2) individuals needing to be protected include the elderly; and (3) vaccines may exacerbate rather than prevent coronavirus lung immunopathology [<xref rid="B137-viruses-10-00093" ref-type="bibr">137</xref>,<xref rid="B138-viruses-10-00093" ref-type="bibr">138</xref>]. Various vaccines against MERS-CoV have been designed, one of which are currently being tested in clinical trials (<xref ref-type="table" rid="viruses-10-00093-t002">Table 2</xref>). All of the MERS-CoV structural proteins could potentially induce neutralizing antibodies and protective responses. However, prior to identification of the major neutralizing antibody-inducing epitopes, inactivated viruses could be used in the production of first-generation vaccines; this is an easy first-response approach since it is relatively simple to produce whole killed virus particles [<xref rid="B139-viruses-10-00093" ref-type="bibr">139</xref>]. With the many safety concerns associated with the production of inactivated vaccines [<xref rid="B140-viruses-10-00093" ref-type="bibr">140</xref>,<xref rid="B141-viruses-10-00093" ref-type="bibr">141</xref>,<xref rid="B142-viruses-10-00093" ref-type="bibr">142</xref>], these types of vaccines must preferably be replaced by safer and more effective neutralizing epitope-based vaccines, as soon as the fragments containing the neutralizing epitopes are identified [<xref rid="B139-viruses-10-00093" ref-type="bibr">139</xref>]. Current MERS-CoV vaccines provide effective protection in a few animal models [<xref rid="B143-viruses-10-00093" ref-type="bibr">143</xref>,<xref rid="B144-viruses-10-00093" ref-type="bibr">144</xref>,<xref rid="B145-viruses-10-00093" ref-type="bibr">145</xref>,<xref rid="B146-viruses-10-00093" ref-type="bibr">146</xref>,<xref rid="B147-viruses-10-00093" ref-type="bibr">147</xref>].</p></sec><sec id="sec7-viruses-10-00093"><title>7. Future Perspectives</title><p>The emergence of Middle East respiratory syndrome (MERS) and the discovery of the MERS coronavirus (MERS-CoV) in 2012 suggest that another SARS-like epidemic is occurring. Unlike the severe acute respiratory syndrome (SARS) epidemic, which rapidly disappeared in less than one year, MERS has persisted for over three years. More than 2000 cases of MERS have been reported worldwide, and the disease carries a worryingly high fatality rate of >30% [<xref rid="B12-viruses-10-00093" ref-type="bibr">12</xref>]. While this number seems low, the virus remains a global threat due to its propensity to cause severe disease in patients with underlying medical conditions and its apparent ability to readily spread within hospital settings [<xref rid="B148-viruses-10-00093" ref-type="bibr">148</xref>]. In addition, the pattern of MERS-CoV lineages is more consistent with the movement of infected livestock or animal products [<xref rid="B171-viruses-10-00093" ref-type="bibr">171</xref>] and epidemiological evidence suggests that it is periodically introduced into human populations [<xref rid="B172-viruses-10-00093" ref-type="bibr">172</xref>,<xref rid="B173-viruses-10-00093" ref-type="bibr">173</xref>], which increases the risk for various future pandemics.</p><p>Even though the clinical outcomes of MERS-CoV infections are well documented, more comprehensive population-based studies are required to determine the involvement of MERS-CoV in other body systems. In addition, the continued development of technologies to routinely and accurately identify asymptomatic MERS-CoV infections will shed light on the true incidence of this virus in the human population. It would appear the MERS-CoV has been circulating in the human population for greater than one year without detection and suggests independent transmission from an unknown source. However, as discussed previously with regard to the emergence of severe acute respiratory syndrome coronavirus (SARS-CoV) in 2002, other evolutionary aspects, such as mutation rates and selection pressure, should be considered to understand the evolutionary dynamics of MERS-CoV [<xref rid="B174-viruses-10-00093" ref-type="bibr">174</xref>,<xref rid="B175-viruses-10-00093" ref-type="bibr">175</xref>,<xref rid="B176-viruses-10-00093" ref-type="bibr">176</xref>,<xref rid="B177-viruses-10-00093" ref-type="bibr">177</xref>]. Possibly different molecular clock rates of MERS-CoV in animal hosts and humans may also have to be taken into account. Similarly to the genomic evolution of influenza A viruses [<xref rid="B178-viruses-10-00093" ref-type="bibr">178</xref>], MERS-CoV might experience different evolutionary courses in different hosts. To better understand these dynamics, the chain of MERS-CoV zoonotic transmissions should be further clarified [<xref rid="B174-viruses-10-00093" ref-type="bibr">174</xref>].</p><p>As with other HCoVs, a detailed manipulation of the MERS-CoV genome to understand the role of the MERS-CoV viral genes in pathogenesis and replication, and for the subsequent development of MERS-CoV as a vaccine vector, is needed. The development of MERS-CoV full-length infectious clones [<xref rid="B19-viruses-10-00093" ref-type="bibr">19</xref>,<xref rid="B20-viruses-10-00093" ref-type="bibr">20</xref>,<xref rid="B179-viruses-10-00093" ref-type="bibr">179</xref>] already allows for the systematic experimental study of the roles of the various corresponding MERS-CoV proteins, which should lead to a better understanding of the role of the viral genes in infectivity and pathogenicity [<xref rid="B180-viruses-10-00093" ref-type="bibr">180</xref>]. This manipulation of the virus genome also provides a reverse genetics platform that could lead to the future development of MERS-CoV-based vector vaccines [<xref rid="B181-viruses-10-00093" ref-type="bibr">181</xref>].</p><p>As a result of the increase in MERS spread, the WHO (World Health Organisation) and CDC (Center for Disease Control) have released various case definitions to allow for the likelihood of a pandemic threat to be reduced. Fever, pneumonia, and acute respiratory distress syndrome with a history of travel to the Arab Peninsula are some of the symptoms that are used to diagnose a MERS-CoV infection. Due to the increase in nosocomial infections, health care workers are also advised to be aware of any upper respiratory tract infections and exposure to MERS-CoV-positive individuals [<xref rid="B182-viruses-10-00093" ref-type="bibr">182</xref>]. For the foreseeable future, important measures to prevent nosocomial outbreaks should include good compliance with appropriate personal protection equipment by health-care workers when managing patients with suspected and confirmed MERS-CoV infection, early diagnosis, prompt isolation of infected patients, and improvement of ventilation in health-care facilities [<xref rid="B183-viruses-10-00093" ref-type="bibr">183</xref>,<xref rid="B184-viruses-10-00093" ref-type="bibr">184</xref>].</p></sec></body><back><ack><title>Acknowledgments</title><p>Burtram C. Fielding receives funding from the National Research Foundation (NRF) (South Africa) and the University of the Western Cape Senate Research Fund. Any opinion, findings and conclusions or recommendations expressed in this material are those of the authors and, therefore, the NRF does not accept any liability in regard thereto.</p></ack><notes><title>Author Contributions</title><p>Burtram C. 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HCoVs genomes are 26 kb to 32 kb in size. At the 5′-end, overlapping reading frames 1a and 1b (blue) make up two-thirds of the genome. The remaining one third of the genome (expanded region) encodes for the structural (white) and accessory proteins (grey).</p></caption><graphic xlink:href="viruses-10-00093-g001"></graphic></fig><table-wrap id="viruses-10-00093-t001" orientation="portrait" position="float"><object-id pub-id-type="pii">viruses-10-00093-t001_Table 1</object-id><label>Table 1</label><caption><p>Detection methods of MERS-CoV.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Method Used for Detection</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">1 Sensitivity/2 Specificity/3 Viral Target Gene</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Reference</th></tr></thead><tbody><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">rtRT-PCR</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Sensitivity for upE is 3.4 copies per reaction (95% confidence interval (CI): 2.5–6.9 copies) or 291 copies/mL of sample. <break></break>2 No cross-reactivity was observed with coronaviruses OC43, NL63, 229E, SARS-CoV, nor with 92 clinical specimens containing common human respiratory viruses.<break></break>3 Targeting regions upstream of the E gene (upE) or within open reading frame (ORF) 1b, respectively. </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B75-viruses-10-00093" ref-type="bibr">75</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">qRT-PCR <sup>#</sup></td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Sensitivity to widely used upE gene as well as a ORF1a&b was introduced<break></break>2 No false-positive amplifications were obtained with other human coronaviruses or common respiratory viral pathogens or with 336 diverse clinical specimens from non-MERS-CoV cases; specimens from two confirmed MERS-CoV cases were positive with all assay signatures.<break></break>3 Two novel signatures used one that targets the MERS-CoV N gene in combination with the upE test. The other a positive test to add to an efficient MERS-CoV kit.</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B81-viruses-10-00093" ref-type="bibr">81</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RT-Sequence-Validated-LAMP Assays</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Could detect 0.02 to 0.2 plaque forming units (PFU) (5 to 50 PFU/mL) of MERS-CoV in infected cell culture supernatants. <break></break>2 Did not cross-react with common human respiratory pathogens.</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B88-viruses-10-00093" ref-type="bibr">88</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RT-LAMP</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Capable of detecting as few as 3.4 copies of MERS-CoV RNA; Assay exhibited sensitivity similar to that of MERS-CoV real-time RT-PCR. <break></break>2 No cross-reaction to other respiratory viruses. <break></break>3 Assay designed to amplify the MERS-CoV gene </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B80-viruses-10-00093" ref-type="bibr">80</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">rt-RPA </td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Highly sensitive, is able to detect 10 MERS-CoV RNA copies with a more rapid detection time than MERS-RT-PCR. <break></break>2 No cross-reaction to other respiratory viruses including HCoVs.<break></break>3 Assay designed to amplify the partial nucleocapsid gene of MERS-CoV</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B89-viruses-10-00093" ref-type="bibr">89</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">mAb Test </td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Rapid detection and cost effective ELISA<break></break>2 High specificity used to detect the MERS-CoV nucleocapsid protein </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B87-viruses-10-00093" ref-type="bibr">87</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Immuno-chromotagraphic tool</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Highly sensitive,<break></break>2 No cross reactivity with other respiratory pathogens observed in vitro and in silico <break></break>3 Detects recombinant MERS-CoV N protein </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B90-viruses-10-00093" ref-type="bibr">90</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Immunofluorescence Assay</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Highly sensitive, antigen based detection <break></break>2 Cross reactivity seen with convalescent SARS patient (sera) <break></break>3 Assay used both whole virus and S1 portion of the spike protein</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B91-viruses-10-00093" ref-type="bibr">91</xref>,<xref rid="B92-viruses-10-00093" ref-type="bibr">92</xref>,<xref rid="B93-viruses-10-00093" ref-type="bibr">93</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">ppNT Assay </td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Highly sensitive, more sensitive that MNT test <break></break>2 Lack of MERS neutralizing activity indicated high specificity by this assay. No cross reactivity seen with SARS-CoV<break></break>3 Assay was designed for two different genes used: a codon optimized spike gene and a HIV/MERS pseudoparticle was generated </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B94-viruses-10-00093" ref-type="bibr">94</xref>,<xref rid="B95-viruses-10-00093" ref-type="bibr">95</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MNT Test </td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Highly sensitive; less so than ppNT assay <break></break>2 Highly specific, as SARS-CoV antigen was not detected compared to MERS-CoV. <break></break>3 Test designed to detect IgG antibodies generated when using the RBD of the S1 subunit of the spike gene </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B94-viruses-10-00093" ref-type="bibr">94</xref>,<xref rid="B96-viruses-10-00093" ref-type="bibr">96</xref>,<xref rid="B97-viruses-10-00093" ref-type="bibr">97</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Protein Microarray</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Highly sensitive assay using protein microarray technology to detect IgG and IgM antibodies <break></break>2 No cross reactivity seen with sera of patients that had been exposed to four common HCoVs.<break></break>3 Assay designed to use the S1 receptor-binding subunit of the spike protein of MERS and SARS as antigens.</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B98-viruses-10-00093" ref-type="bibr">98</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">One pot RT-LAMP</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Capable of detecting four viral copies MERS within 60 min<break></break>2 No cross-reaction to the other acute respiratory disease viruses (influenza type A virus (H1N1 and H3N2), influenza type B virus, HCoV-229E, and human metapneumovirus)<break></break>3 Six sets of primers designed specifically to amplify the MERS-CoV genes</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B99-viruses-10-00093" ref-type="bibr">99</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RT-iiPCR assays</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Could detect 3.7 × 10<sup>−1</sup> plaque forming units (PFU) of MERS-CoV in infected cell culture supernatants and sputum samples.<break></break>2 Viral nucleic acids extracted from infected cultures that contained HCoV-229E, HCoV-OC43, FIPV, influenza type A and B virus strains yielded negative results, indicating no cross reactivity. <break></break>3 Targeting regions upstream of the E gene (upE) or within open reading frame (ORF) 1b</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B100-viruses-10-00093" ref-type="bibr">100</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Powerchek MERS Assay</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 95% limits of detection of assay for the upE and ORF1a were 16.2 copies/μL and 8.2 copies/μL, respectively.<break></break>2 No cross reactivity with other respiratory pathogens observed in vitro and in silico <break></break>3 Targeting regions upstream of the E gene (upE) or within open reading frame (ORF) 1b</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B101-viruses-10-00093" ref-type="bibr">101</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">acpcPNA-AgNP aggregation assay </td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Probe designed for targets makes this assay highly specific. Limit of detection found to be 1.53 nM<break></break>2 Cross reactivity with other CoVs was not evaluated<break></break>3 Synthetic oligonucleotides were designed to target MERS</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B102-viruses-10-00093" ref-type="bibr">102</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">mCoV-MS</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Highly sensitive, multiplex PCR based to target specific genes in HcoVs<break></break>2 Cross reactivity with other respiratory pathogens was not evaluated<break></break>3 Targeting regions upstream of the E gene (upE) or within open reading frame (ORF) 1b</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B103-viruses-10-00093" ref-type="bibr">103</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Duplex-RT-PCR method</td><td align="left" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1 Highly sensitive, simultaneous detection of MERS and SARS viruses. <break></break>2 Cross reactivity with other respiratory pathogens was not evaluated<break></break>3 Primers and probes that target the conserved spike S2 region of SARS-CoV, MERS-CoV, and their related bat CoVs were used</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B104-viruses-10-00093" ref-type="bibr">104</xref>]</td></tr></tbody></table><table-wrap-foot><fn><p><bold>rtRT-PCR:</bold> Real-time reverse transcription polymerase chain reaction; <bold>LAMP:</bold> Loop-mediated isothermal amplification; <bold>qRT-PCR:</bold> Quantitative real-time reverse transcription polymerase chain reaction; <bold>rtRPA:</bold> reverse transcription isothermal Recombinase Polymerase Amplification; <bold>mAb:</bold> monoclonal Antibody; <bold>ELISA:</bold> Enzyme linked immunoabsorbent assay; <bold>ppNT:</bold> pseudoparticle neutralisation; <bold>MNT:</bold> microneutralisation; <bold>RT-iiPCR:</bold> reverse transcription-insulated isothermal PCR; <bold>Powerchek:</bold> PowerChek MERS assay; Kogene Biotech, Korea; <bold>acpcPNA-AgNP:</bold> DNA detection based on pyrrolidinyl peptide nucleic acid induced silver nanoparticle (colorimetric assay); <bold>mCoV-MS:</bold> MassARRAY matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF MS) system; <bold>N</bold><bold>:</bold> Nucleocapsid; <sup>#</sup> FDA approved (RealStar MERS-CoV RT-PCR kit 1.0, Altona Diagnostics GmbH, Hamburg, Germany).</p></fn></table-wrap-foot></table-wrap><table-wrap id="viruses-10-00093-t002" orientation="portrait" position="float"><object-id pub-id-type="pii">viruses-10-00093-t002_Table 2</object-id><label>Table 2</label><caption><p>MERS-CoV vaccines developed (adapted from [<xref rid="B135-viruses-10-00093" ref-type="bibr">135</xref>,<xref rid="B148-viruses-10-00093" ref-type="bibr">148</xref>]).</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Vaccine Categories</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Target Antigen</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Immunization</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Animal Model</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Immunogenicity</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Stage of Development</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Reference</th></tr></thead><tbody><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Anti-MERS-CoV monoclonal antibodies</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Surface (S) glycoprotein </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Passive </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">marmosets</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Animals developed pneumonia, high viral titre detected in lungs</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical: in vivo, efficacy stage</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B149-viruses-10-00093" ref-type="bibr">149</xref>,<xref rid="B150-viruses-10-00093" ref-type="bibr">150</xref>,<xref rid="B151-viruses-10-00093" ref-type="bibr">151</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Human polyclonal anti-MERS-CoV antibodies</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus structural proteins</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Passive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Ad5-hDPP4-transduced mouse</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nab developed to reduce viral titres post exposure</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical: in vivo, efficacy stage</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B152-viruses-10-00093" ref-type="bibr">152</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Inactivated virion vaccines</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MERS-CoV</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Active</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">hDPP4-transgenic mice</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nab produced without adjuvant, T-cell response not done</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical: in vivo, efficacy stage</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B153-viruses-10-00093" ref-type="bibr">153</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Live attenuated vaccines (deleted E protein; mutated in nsp14)</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">rMERS-CoV-∆E</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Active </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Not tested</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Not indicated</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical development: in vitro</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B20-viruses-10-00093" ref-type="bibr">20</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Recombinant viral vectors (MVA, Adenovirus, Parainfluenza virus, Measles, Rabies)</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S and SolS proteins</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Active </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Ad/hDPP4-mice Camels</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nab in mice, antigen specific humoral and in some case T cell immune responses </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical: in vitro, efficacy stage</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B145-viruses-10-00093" ref-type="bibr">145</xref>,<xref rid="B154-viruses-10-00093" ref-type="bibr">154</xref>,<xref rid="B155-viruses-10-00093" ref-type="bibr">155</xref>,<xref rid="B156-viruses-10-00093" ref-type="bibr">156</xref>,<xref rid="B157-viruses-10-00093" ref-type="bibr">157</xref>,<xref rid="B158-viruses-10-00093" ref-type="bibr">158</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Replicon particles (e.g., Venezuelan (VRP-S)</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S protein </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Active </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Ad/hDPP4-mice mice</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nab produced, mice developed progressive pneumonia with virus replication detected in airways</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical: in vivo, efficacy stage</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B110-viruses-10-00093" ref-type="bibr">110</xref>,<xref rid="B159-viruses-10-00093" ref-type="bibr">159</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Subunit vaccines RBDs rRBDs RBDs-Fc rNTDs </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S/S1protein with various amino acid residues</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Active </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">-hDPP4-transgenic-Ad5-hDPP4 mice Rabbit NHPs</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">High mucosal and humoral immune response, strong Nab in mice and rabbits. Good T-cell response in mice. Tg-Mice protected from MERS-CoV</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical: in vitro, efficacy stage</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B147-viruses-10-00093" ref-type="bibr">147</xref>,<xref rid="B160-viruses-10-00093" ref-type="bibr">160</xref>,<xref rid="B161-viruses-10-00093" ref-type="bibr">161</xref>,<xref rid="B162-viruses-10-00093" ref-type="bibr">162</xref>,<xref rid="B163-viruses-10-00093" ref-type="bibr">163</xref>,<xref rid="B164-viruses-10-00093" ref-type="bibr">164</xref>,<xref rid="B165-viruses-10-00093" ref-type="bibr">165</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DNA vaccines</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S protein</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Active </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NHP:Rhesus Macaques Camels Mice</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Cellular immune response and Nab response in mice, NHPs and camels.</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Phase 1 clinical trials </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B166-viruses-10-00093" ref-type="bibr">166</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DNA prime/Protein-boost Vaccines</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S and S1 protein</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Active </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NHP:Rhesus Macaques Mice</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nab response seen in mice and NHPs</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical: in vitro, efficacy stage</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B167-viruses-10-00093" ref-type="bibr">167</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">VLPs</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S, M, E</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Active</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NHP:Rhesus Macaques</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus specific Nab and IgG antibody response against the RBD</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical: in vivo, efficacy stage</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B168-viruses-10-00093" ref-type="bibr">168</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nanoparticle vaccine</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">S protein</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Active</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Mice</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nab with the presence of adjuvant (M1 and Alum) </td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Preclinical: in vivo, efficacy stage</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B169-viruses-10-00093" ref-type="bibr">169</xref>,<xref rid="B170-viruses-10-00093" ref-type="bibr">170</xref>]</td></tr></tbody></table><table-wrap-foot><fn><p><bold>Ad:</bold> Adenovirus; <bold>Ad/hDPP4-mice:</bold> mice transduced with hDPP4 in an adenovirus vector; <bold>Alum</bold>: aluminum hydroxide (adjuvant); <bold>∆E:</bold> truncated envelope protein, <bold>hDPP4:</bold> human dipeptidyl peptidase 4; <bold>M1:</bold> matrix protein 1 (adjuvant); <bold>MERS-CoV:</bold> Middle East Respiratory Syndrome Coronavirus; <bold>M:</bold> membrane protein; <bold>MVA:</bold> modified vaccinia virus Ankara; <bold>N:</bold> nucleocapsid protein; <bold>Nab:</bold> neutralizing antibody; <bold>NHP:</bold> non-human primates; <bold>rMERS-CoV:</bold> recombinant Middle East respiratory syndrome coronavirus; <bold>rNTD:</bold> recombinant N-terminal domain; <bold>RBD:</bold> receptor-binding domain; <bold>rRBD:</bold> recombinant RBD; <bold>RBD-Fc:</bold> RBD fused to the human IgG antibody crytallizable fragment; S: spike protein; <bold>S1:</bold> S1 domain of the spike protein, <bold>SolS:</bold> spike protein lacking transmembrane domain; <bold>Tg-mice</bold>: transgenic mice; <bold>VRP:</bold> virus replicon particle; <bold>VLP’s:</bold> virus like particles.</p></fn></table-wrap-foot></table-wrap></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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