Evidence of the Recombinant Origin of a Bat Severe Acute Respiratory Syndrome (SARS)-Like Coronavirus and Its Implications on the Direct Ancestor of SARS Coronavirus▿
Identifieur interne : 000F63 ( Pmc/Checkpoint ); précédent : 000F62; suivant : 000F64Evidence of the Recombinant Origin of a Bat Severe Acute Respiratory Syndrome (SARS)-Like Coronavirus and Its Implications on the Direct Ancestor of SARS Coronavirus▿
Auteurs : Chung-Chau Hon ; Tsan-Yuk Lam ; Zheng-Li Shi ; Alexei J. Drummond ; Chi-Wai Yip ; Fanya Zeng ; Pui-Yi Lam ; Frederick Chi-Ching LeungSource :
- Journal of Virology [ 0022-538X ] ; 2007.
Abstract
Bats have been identified as the natural reservoir of severe acute respiratory syndrome (SARS)-like and SARS coronaviruses (SLCoV and SCoV). However, previous studies suggested that none of the currently sampled bat SLCoVs is the descendant of the direct ancestor of SCoV, based on their relatively distant phylogenetic relationship. In this study, evidence of the recombinant origin of the genome of a bat SLCoV is demonstrated. We identified a potential recombination breakpoint immediately after the consensus intergenic sequence between open reading frame 1 and the S coding region, suggesting the replication intermediates may participate in the recombination event, as previously speculated for other CoVs. Phylogenetic analysis of its parental regions suggests the presence of an uncharacterized SLCoV lineage that is phylogenetically closer to SCoVs than any of the currently sampled bat SLCoVs. Using various Bayesian molecular-clock models, interspecies transfer of this SLCoV lineage from bats to the amplifying host (e.g., civets) was estimated to have happened a median of 4.08 years before the SARS outbreak. Based on this relatively short window period, we speculate that this uncharacterized SLCoV lineage may contain the direct ancestor of SCoV. This study sheds light on the possible host bat species of the direct ancestor of SCoV, providing valuable information on the scope and focus of surveillance for the origin of SCoV.
Url:
DOI: 10.1128/JVI.01926-07
PubMed: 18057240
PubMed Central: 2258724
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Evidence of the Recombinant Origin of a Bat Severe Acute Respiratory Syndrome (SARS)-Like Coronavirus and Its Implications on the Direct Ancestor of SARS Coronavirus<xref ref-type="fn" rid="fn1">▿</xref> </title><author><name sortKey="Hon, Chung Chau" sort="Hon, Chung Chau" uniqKey="Hon C" first="Chung-Chau" last="Hon">Chung-Chau Hon</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Lam, Tsan Yuk" sort="Lam, Tsan Yuk" uniqKey="Lam T" first="Tsan-Yuk" last="Lam">Tsan-Yuk Lam</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Shi, Zheng Li" sort="Shi, Zheng Li" uniqKey="Shi Z" first="Zheng-Li" last="Shi">Zheng-Li Shi</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Drummond, Alexei J" sort="Drummond, Alexei J" uniqKey="Drummond A" first="Alexei J." last="Drummond">Alexei J. Drummond</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Yip, Chi Wai" sort="Yip, Chi Wai" uniqKey="Yip C" first="Chi-Wai" last="Yip">Chi-Wai Yip</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Zeng, Fanya" sort="Zeng, Fanya" uniqKey="Zeng F" first="Fanya" last="Zeng">Fanya Zeng</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Lam, Pui Yi" sort="Lam, Pui Yi" uniqKey="Lam P" first="Pui-Yi" last="Lam">Pui-Yi Lam</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Leung, Frederick Chi Ching" sort="Leung, Frederick Chi Ching" uniqKey="Leung F" first="Frederick Chi-Ching" last="Leung">Frederick Chi-Ching Leung</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">18057240</idno><idno type="pmc">2258724</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2258724</idno><idno type="RBID">PMC:2258724</idno><idno type="doi">10.1128/JVI.01926-07</idno><date when="2007">2007</date><idno type="wicri:Area/Pmc/Corpus">000726</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000726</idno><idno type="wicri:Area/Pmc/Curation">000726</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Curation">000726</idno><idno type="wicri:Area/Pmc/Checkpoint">000F63</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Checkpoint">000F63</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Evidence of the Recombinant Origin of a Bat Severe Acute Respiratory Syndrome (SARS)-Like Coronavirus and Its Implications on the Direct Ancestor of SARS Coronavirus<xref ref-type="fn" rid="fn1">▿</xref> </title><author><name sortKey="Hon, Chung Chau" sort="Hon, Chung Chau" uniqKey="Hon C" first="Chung-Chau" last="Hon">Chung-Chau Hon</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Lam, Tsan Yuk" sort="Lam, Tsan Yuk" uniqKey="Lam T" first="Tsan-Yuk" last="Lam">Tsan-Yuk Lam</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Shi, Zheng Li" sort="Shi, Zheng Li" uniqKey="Shi Z" first="Zheng-Li" last="Shi">Zheng-Li Shi</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Drummond, Alexei J" sort="Drummond, Alexei J" uniqKey="Drummond A" first="Alexei J." last="Drummond">Alexei J. Drummond</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Yip, Chi Wai" sort="Yip, Chi Wai" uniqKey="Yip C" first="Chi-Wai" last="Yip">Chi-Wai Yip</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Zeng, Fanya" sort="Zeng, Fanya" uniqKey="Zeng F" first="Fanya" last="Zeng">Fanya Zeng</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Lam, Pui Yi" sort="Lam, Pui Yi" uniqKey="Lam P" first="Pui-Yi" last="Lam">Pui-Yi Lam</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author><author><name sortKey="Leung, Frederick Chi Ching" sort="Leung, Frederick Chi Ching" uniqKey="Leung F" first="Frederick Chi-Ching" last="Leung">Frederick Chi-Ching Leung</name><affiliation><nlm:aff id="aff1"></nlm:aff></affiliation></author></analytic><series><title level="j">Journal of Virology</title><idno type="ISSN">0022-538X</idno><idno type="eISSN">1098-5514</idno><imprint><date when="2007">2007</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Bats have been identified as the natural reservoir of severe acute respiratory syndrome (SARS)-like and SARS coronaviruses (SLCoV and SCoV). However, previous studies suggested that none of the currently sampled bat SLCoVs is the descendant of the direct ancestor of SCoV, based on their relatively distant phylogenetic relationship. In this study, evidence of the recombinant origin of the genome of a bat SLCoV is demonstrated. We identified a potential recombination breakpoint immediately after the consensus intergenic sequence between open reading frame 1 and the S coding region, suggesting the replication intermediates may participate in the recombination event, as previously speculated for other CoVs. Phylogenetic analysis of its parental regions suggests the presence of an uncharacterized SLCoV lineage that is phylogenetically closer to SCoVs than any of the currently sampled bat SLCoVs. Using various Bayesian molecular-clock models, interspecies transfer of this SLCoV lineage from bats to the amplifying host (e.g., civets) was estimated to have happened a median of 4.08 years before the SARS outbreak. Based on this relatively short window period, we speculate that this uncharacterized SLCoV lineage may contain the direct ancestor of SCoV. This study sheds light on the possible host bat species of the direct ancestor of SCoV, providing valuable information on the scope and focus of surveillance for the origin of SCoV.</p></div></front></TEI><pmc article-type="research-article"><pmc-comment>The publisher of this article does not allow downloading of the full text in XML form.</pmc-comment>
<front><journal-meta><journal-id journal-id-type="nlm-ta">J Virol</journal-id><journal-title>Journal of Virology</journal-title><issn pub-type="ppub">0022-538X</issn><issn pub-type="epub">1098-5514</issn><publisher><publisher-name>American Society for Microbiology (ASM)</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">18057240</article-id><article-id pub-id-type="pmc">2258724</article-id><article-id pub-id-type="publisher-id">1926-07</article-id><article-id pub-id-type="doi">10.1128/JVI.01926-07</article-id><article-categories><subj-group subj-group-type="heading"><subject>Genetic Diversity and Evolution</subject></subj-group></article-categories><title-group><article-title>Evidence of the Recombinant Origin of a Bat Severe Acute Respiratory Syndrome (SARS)-Like Coronavirus and Its Implications on the Direct Ancestor of SARS Coronavirus<xref ref-type="fn" rid="fn1">▿</xref> </article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Hon</surname><given-names>Chung-Chau</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Lam</surname><given-names>Tsan-Yuk</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Shi</surname><given-names>Zheng-Li</given-names></name><xref ref-type="aff" rid="aff1">2</xref></contrib><contrib contrib-type="author"><name><surname>Drummond</surname><given-names>Alexei J.</given-names></name><xref ref-type="aff" rid="aff1">3</xref></contrib><contrib contrib-type="author"><name><surname>Yip</surname><given-names>Chi-Wai</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Zeng</surname><given-names>Fanya</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Lam</surname><given-names>Pui-Yi</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Leung</surname><given-names>Frederick Chi-Ching</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="corresp" rid="cor1">*</xref></contrib></contrib-group><aff id="aff1">School of Biological Sciences, The University of Hong Kong, Hong Kong, China,<label>1</label> State Key Laboratory of Virology, Wuhan Institute of Virology, Chinese Academy of Sciences, Wuhan, Hubei, China,<label>2</label> Bioinformatics Institute, University of Auckland, Auckland, New Zealand<label>3</label></aff><author-notes><fn id="cor1"><label>*</label><p>Corresponding author. Mailing address: 5N-12, Kadoorie Biological Science Building, The University of Hong Kong, Hong Kong, China. Phone: 852-2299 0825. Fax: 852-2857 4672. E-mail: <email>fcleung@hkucc.hku.hk</email></p></fn></author-notes><pub-date pub-type="ppub"><month>2</month><year>2008</year></pub-date><pub-date pub-type="epub"><day>5</day><month>12</month><year>2007</year></pub-date><volume>82</volume><issue>4</issue><fpage>1819</fpage><lpage>1826</lpage><history><date date-type="received"><day>3</day><month>9</month><year>2007</year></date><date date-type="accepted"><day>21</day><month>11</month><year>2007</year></date></history><permissions><copyright-statement>Copyright © 2008, American Society for Microbiology</copyright-statement></permissions><self-uri xlink:title="pdf" xlink:href="zjv00408001819.pdf"></self-uri><abstract><p>Bats have been identified as the natural reservoir of severe acute respiratory syndrome (SARS)-like and SARS coronaviruses (SLCoV and SCoV). However, previous studies suggested that none of the currently sampled bat SLCoVs is the descendant of the direct ancestor of SCoV, based on their relatively distant phylogenetic relationship. In this study, evidence of the recombinant origin of the genome of a bat SLCoV is demonstrated. We identified a potential recombination breakpoint immediately after the consensus intergenic sequence between open reading frame 1 and the S coding region, suggesting the replication intermediates may participate in the recombination event, as previously speculated for other CoVs. Phylogenetic analysis of its parental regions suggests the presence of an uncharacterized SLCoV lineage that is phylogenetically closer to SCoVs than any of the currently sampled bat SLCoVs. Using various Bayesian molecular-clock models, interspecies transfer of this SLCoV lineage from bats to the amplifying host (e.g., civets) was estimated to have happened a median of 4.08 years before the SARS outbreak. Based on this relatively short window period, we speculate that this uncharacterized SLCoV lineage may contain the direct ancestor of SCoV. This study sheds light on the possible host bat species of the direct ancestor of SCoV, providing valuable information on the scope and focus of surveillance for the origin of SCoV.</p></abstract></article-meta></front><floats-wrap><fig position="float" id="f1"><label>FIG. 1.</label><caption><p>Detection of recombination and estimation of a breakpoint within the genome of Rp3. A similarity plot (A) and a bootscan analysis (B) detected a single recombination breakpoint at around the ORF1b/S junction. Both analyses were performed with an F84 distance model, a window size of 1,500 bp, and a step size of 300 bp. The Hu-SCoV group includes strains Tor2 (AY274119), GD01 (AY278489), ZJ01 (AY297028), SZ3 (AY304486), GZ0402 (AY613947), and PC4 (AY613950). (C) Organization of essential ORFs of the SCoV genome and location of the estimated breakpoint. The blue and red horizontal arrows represent the essential ORFs from the major and minor parents, respectively. A sequence alignment of the ORF1b/S junction regions of Rp3, Tor2, and Rm1 is shown below. A consensus IGS and the coding regions of ORF1b and S are annotated above the alignment. The black vertical arrow below the alignment indicates the estimated breakpoint located immediately after the start codon of the S coding region.</p></caption><graphic xlink:href="zjv0040802380001"></graphic></fig><fig position="float" id="f2"><label>FIG. 2.</label><caption><p>Phylogenetic origins of the major and minor parental regions of Rp3. ML phylogenies were constructed from the concatenated sequences of the essential ORFs of the major (A) and minor (B) parental regions of selected CoVs. For the purposes of display, the phylogenies were midpoint rooted. The taxa were annotated according to their accession numbers and host species—civets (C), humans (H), or bats (B)—and strain names. The numbers on the left of the nodes refer to the BMCMC posterior probabilities. The percentages of support for all other internal nodes within the two lineages were omitted for simplicity. The recombinant strain Rp3, the most recent common ancestor of Hu-SCoVs (MRCA-Hu), and the divergence event between Hu-SCoVs and HB-SLCoVs (DIV-Hu/HB) are indicated. The scale bars are in units of nucleotide substitutions per site.</p></caption><graphic xlink:href="zjv0040802380002"></graphic></fig><fig position="float" id="f3"><label>FIG. 3.</label><caption><p>tMRCA-Hu estimated from the S1 data set. (A) tMRCA-Hu estimated from the S1 data set under various Bayesian clock models and the ML SRDT model. (B) Posterior MCMC samples (left <italic>y</italic> axis) of tMRCA-Hu estimated from the S1 data set under the UCED model and the lognormal distribution (right <italic>y</italic> axis) fitted using Easyfit. The values of the parameters for the lognormal distribution are as follows: σ = 0.56, μ = −1.00, and γ = 2.04.</p></caption><graphic xlink:href="zjv0040802380003"></graphic></fig><fig position="float" id="f4"><label>FIG. 4.</label><caption><p>Specification of an S1-derived lognormal tMRCA-Hu prior in the analysis of the ORF1 data set under the UCED model. (A) Prior and posterior distributions of tMRCA-Hu. (B) Effects of the tMRCA-Hu prior on the posterior distribution of tDIV-Hu/HB.</p></caption><graphic xlink:href="zjv0040802380004"></graphic></fig><fig position="float" id="f5"><label>FIG. 5.</label><caption><p>Estimation of the window period between the cross-species event and the onset of the 2003 SARS epidemic. This time-scaled phylogeny was summarized from all MCMC phylogenies of the ORF1 data set analyzed under the UCED model with the S1-derived tMRCA-Hu prior. The heights of the nodes are represented by the median of their estimates. The HPD of tMRCA-Hu and tDIV-Hu/HB are indicated by gray boxes at these nodes. The taxa were labeled in the same style as in Fig. <xref rid="f2" ref-type="fig">2</xref>, except their sampling dates were annotated.</p></caption><graphic xlink:href="zjv0040802380005"></graphic></fig><table-wrap position="float" id="t1"><label>TABLE 1.</label><caption><p>Details of the two data sets and results of the ML molecular-clock tests</p></caption><table frame="hsides" rules="groups"><thead><tr><th colspan="1" rowspan="1" align="center" valign="bottom">Data set</th><th colspan="1" rowspan="1" align="center" valign="bottom">Viral lineages included<xref ref-type="table-fn" rid="t1fn1"><italic>a</italic></xref></th><th colspan="1" rowspan="1" align="center" valign="bottom">No. of taxa<xref ref-type="table-fn" rid="t1fn2"><italic>b</italic></xref></th><th colspan="1" rowspan="1" align="center" valign="bottom">df<xref ref-type="table-fn" rid="t1fn3"><italic>c</italic></xref></th><th colspan="1" rowspan="1" align="center" valign="bottom">2Δ<xref ref-type="table-fn" rid="t1fn4"><italic>d</italic></xref></th><th colspan="1" rowspan="1" align="center" valign="bottom">LRT (<italic>P</italic>)<xref ref-type="table-fn" rid="t1fn5"><italic>e</italic></xref></th></tr></thead><tbody><tr><td colspan="1" rowspan="1" align="left" valign="top">S1</td><td colspan="1" rowspan="1" align="left" valign="top">Hu-SCoVs only</td><td colspan="1" rowspan="1" align="char" char="." valign="top">36</td><td colspan="1" rowspan="1" align="char" char="." valign="top">34</td><td colspan="1" rowspan="1" align="char" char="." valign="top">39.29</td><td colspan="1" rowspan="1" align="char" char="." valign="top">0.24</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top">ORF1</td><td colspan="1" rowspan="1" align="left" valign="top">All Hu-SCoVs and Bt-SLCoVs</td><td colspan="1" rowspan="1" align="char" char="." valign="top">24</td><td colspan="1" rowspan="1" align="char" char="." valign="top">22</td><td colspan="1" rowspan="1" align="char" char="." valign="top">87.55</td><td colspan="1" rowspan="1" align="char" char="." valign="top"><0.001</td></tr></tbody></table><table-wrap-foot><fn id="t1fn1"><label>a</label><p>S1 and ORF1 data sets were used for estimation of tMRCA-Hu and tDIV-Hu/HB, respectively.</p></fn><fn id="t1fn2"><label>b</label><p>Due to the relatively low variability among Hu-SCoV ORF1 sequences, highly similar Hu-SCoV ORF1 taxa with identical sampling dates were removed (<italic>n</italic> = 24 after removal).</p></fn><fn id="t1fn3"><label>c</label><p>df.refers to the degree of freedom in the LRT.</p></fn><fn id="t1fn4"><label>d</label><p>2Δ is twice the difference between the log likelihoods for the SRDT and DR models.</p></fn><fn id="t1fn5"><label>e</label><p>The SRDT model cannot be rejected if <italic>P</italic> is >0.05.</p></fn></table-wrap-foot></table-wrap><table-wrap position="float" id="t2"><label>TABLE 2.</label><caption><p>Performances of the Bayesian clock models</p></caption><table frame="hsides" rules="groups"><thead><tr><th colspan="1" rowspan="2" align="center" valign="middle">Parameter</th><th colspan="3" rowspan="1" align="center" valign="bottom">Value
<hr></hr></th></tr><tr><th colspan="1" rowspan="1" align="center" valign="bottom">Clock model<xref ref-type="table-fn" rid="t2fn3"><italic>c</italic></xref></th><th colspan="1" rowspan="1" align="center" valign="bottom">S1 data set</th><th colspan="1" rowspan="1" align="center" valign="bottom">ORF1 data set</th></tr></thead><tbody><tr><td colspan="1" rowspan="1" align="left" valign="top">Marginal likelihood<xref ref-type="table-fn" rid="t2fn1"><italic>a</italic></xref></td><td colspan="1" rowspan="1" align="left" valign="top">CLOC</td><td colspan="1" rowspan="1" align="char" char="." valign="top">−3,159.31</td><td colspan="1" rowspan="1" align="char" char="." valign="top">−52,478.47</td></tr><tr><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top">UCED</td><td colspan="1" rowspan="1" align="char" char="." valign="top">−3,155.66</td><td colspan="1" rowspan="1" align="char" char="." valign="top">−52,452.64</td></tr><tr><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top">UCLN</td><td colspan="1" rowspan="1" align="char" char="." valign="top">−3,157.55</td><td colspan="1" rowspan="1" align="char" char="." valign="top">−52,467.24</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top">BF<xref ref-type="table-fn" rid="t2fn2"><italic>b</italic></xref></td><td colspan="1" rowspan="1" align="left" valign="top">UCED vs. CLOC</td><td colspan="1" rowspan="1" align="char" char="." valign="top">3.65</td><td colspan="1" rowspan="1" align="char" char="." valign="top">25.82</td></tr><tr><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top">UCLN vs. CLOC</td><td colspan="1" rowspan="1" align="char" char="." valign="top">1.76</td><td colspan="1" rowspan="1" align="char" char="." valign="top">11.23</td></tr><tr><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top">UCED vs. UCLN</td><td colspan="1" rowspan="1" align="char" char="." valign="top">1.89</td><td colspan="1" rowspan="1" align="char" char="." valign="top">14.59</td></tr></tbody></table><table-wrap-foot><fn id="t2fn1"><label>a</label><p>The marginal likelihoods are presented in natural log scale.</p></fn><fn id="t2fn2"><label>b</label><p>The BFs are presented in natural log scale (i.e., ln BF); a ln BF of >2.99 is defined as a strong support for the favored model.</p></fn><fn id="t2fn3"><label>c</label><p>The clock models for the ORF1 data set refer to analyses without the S1-derived lognormal prior on tMRCA-Hu.</p></fn></table-wrap-foot></table-wrap><table-wrap position="float" id="t3"><label>TABLE 3.</label><caption><p>Estimates from the ORF1 data set under the Bayesian UCED model</p></caption><table frame="hsides" rules="groups"><thead><tr><th colspan="1" rowspan="1" align="center" valign="bottom">tMRCA-Hu prior</th><th colspan="1" rowspan="1" align="center" valign="bottom">tMRCA-Hu<xref ref-type="table-fn" rid="t3fn1"><italic>a</italic></xref></th><th colspan="1" rowspan="1" align="center" valign="bottom">tDIV-Hu/HB<xref ref-type="table-fn" rid="t3fn1"><italic>a</italic></xref></th><th colspan="1" rowspan="1" align="center" valign="bottom">Branch A<xref ref-type="table-fn" rid="t3fn2"><italic>b</italic></xref></th></tr></thead><tbody><tr><td colspan="1" rowspan="1" align="left" valign="top">With S1-derived prior</td><td colspan="1" rowspan="1" align="char" char="." valign="top">2002.63 (2002.14-2002.96)</td><td colspan="1" rowspan="1" align="char" char="." valign="top">1998.51 (1993.55-2001.32)</td><td colspan="1" rowspan="1" align="char" char="." valign="top">4.08 (1.45-8.84)</td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top">Without S1-derived prior</td><td colspan="1" rowspan="1" align="char" char="." valign="top">2002.40 (2000.69-2003.01)</td><td colspan="1" rowspan="1" align="char" char="." valign="top">1997.44 (1987.68-2001.49)</td><td colspan="1" rowspan="1" align="char" char="." valign="top">4.86 (1.37-13.47)</td></tr></tbody></table><table-wrap-foot><fn id="t3fn1"><label>a</label><p>Medians with HPD in parentheses.</p></fn><fn id="t3fn2"><label>b</label><p>Length of Branch A (Fig. <xref rid="f5" ref-type="fig">5</xref>) in years. Similar results were obtained under the UCLN model (data not shown for simplicity).</p></fn></table-wrap-foot></table-wrap></floats-wrap></pmc><affiliations><list></list><tree><noCountry><name sortKey="Drummond, Alexei J" sort="Drummond, Alexei J" uniqKey="Drummond A" first="Alexei J." last="Drummond">Alexei J. Drummond</name><name sortKey="Hon, Chung Chau" sort="Hon, Chung Chau" uniqKey="Hon C" first="Chung-Chau" last="Hon">Chung-Chau Hon</name><name sortKey="Lam, Pui Yi" sort="Lam, Pui Yi" uniqKey="Lam P" first="Pui-Yi" last="Lam">Pui-Yi Lam</name><name sortKey="Lam, Tsan Yuk" sort="Lam, Tsan Yuk" uniqKey="Lam T" first="Tsan-Yuk" last="Lam">Tsan-Yuk Lam</name><name sortKey="Leung, Frederick Chi Ching" sort="Leung, Frederick Chi Ching" uniqKey="Leung F" first="Frederick Chi-Ching" last="Leung">Frederick Chi-Ching Leung</name><name sortKey="Shi, Zheng Li" sort="Shi, Zheng Li" uniqKey="Shi Z" first="Zheng-Li" last="Shi">Zheng-Li Shi</name><name sortKey="Yip, Chi Wai" sort="Yip, Chi Wai" uniqKey="Yip C" first="Chi-Wai" last="Yip">Chi-Wai Yip</name><name sortKey="Zeng, Fanya" sort="Zeng, Fanya" uniqKey="Zeng F" first="Fanya" last="Zeng">Fanya Zeng</name></noCountry></tree></affiliations></record>Pour manipuler ce document sous Unix (Dilib)
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