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Diversity and Global Distribution of IncL/M Plasmids Enabling Horizontal Dissemination of β-Lactam Resistance Genes among the Enterobacteriaceae

Identifieur interne : 000088 ( Pmc/Checkpoint ); précédent : 000087; suivant : 000089

Diversity and Global Distribution of IncL/M Plasmids Enabling Horizontal Dissemination of β-Lactam Resistance Genes among the Enterobacteriaceae

Auteurs : Marcin Adamczuk [Pologne] ; Piotr Zaleski [Pologne] ; Lukasz Dziewit [Pologne] ; Renata Wolinowska [Pologne] ; Marta Nieckarz [Pologne] ; Pawel Wawrzyniak [Pologne] ; Piotr Kieryl [Pologne] ; Andrzej Plucienniczak [Pologne] ; Dariusz Bartosik [Pologne]

Source :

RBID : PMC:4510254

Abstract

Antibiotic resistance determinants are frequently associated with plasmids and other mobile genetic elements, which simplifies their horizontal transmission. Several groups of plasmids (including replicons of the IncL/M incompatibility group) were found to play an important role in the dissemination of resistance genes encoding β-lactamases. The IncL/M plasmids are large, broad host range, and self-transmissible replicons. We have identified and characterized two novel members of this group: pARM26 (isolated from bacteria inhabiting activated sludge from a wastewater treatment plant) and pIGT15 (originating from a clinical strain of Escherichia coli). This instigated a detailed comparative analysis of all available sequences of IncL/M plasmids encoding β-lactamases. The core genome of these plasmids is comprised of 20 genes with conserved synteny. Phylogenetic analyses of these core genes allowed clustering of the plasmids into four separate groups, which reflect their antibiotic resistance profiles. Examination of the biogeography of the IncL/M plasmids revealed that they are most frequently found in bacteria of the family Enterobacteriaceae originating from the Mediterranean region and Western Europe and that they are able to persist in various ecological niches even in the absence of direct antibiotic selection pressure.


Url:
DOI: 10.1155/2015/414681
PubMed: 26236726
PubMed Central: 4510254


Affiliations:


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PMC:4510254

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<p>Antibiotic resistance determinants are frequently associated with plasmids and other mobile genetic elements, which simplifies their horizontal transmission. Several groups of plasmids (including replicons of the IncL/M incompatibility group) were found to play an important role in the dissemination of resistance genes encoding
<italic>β</italic>
-lactamases. The IncL/M plasmids are large, broad host range, and self-transmissible replicons. We have identified and characterized two novel members of this group: pARM26 (isolated from bacteria inhabiting activated sludge from a wastewater treatment plant) and pIGT15 (originating from a clinical strain of
<italic>Escherichia coli</italic>
). This instigated a detailed comparative analysis of all available sequences of IncL/M plasmids encoding
<italic>β</italic>
-lactamases. The core genome of these plasmids is comprised of 20 genes with conserved synteny. Phylogenetic analyses of these core genes allowed clustering of the plasmids into four separate groups, which reflect their antibiotic resistance profiles. Examination of the biogeography of the IncL/M plasmids revealed that they are most frequently found in bacteria of the family Enterobacteriaceae originating from the Mediterranean region and Western Europe and that they are able to persist in various ecological niches even in the absence of direct antibiotic selection pressure.</p>
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</TEI>
<pmc article-type="research-article">
<pmc-dir>properties open_access</pmc-dir>
<front>
<journal-meta>
<journal-id journal-id-type="nlm-ta">Biomed Res Int</journal-id>
<journal-id journal-id-type="iso-abbrev">Biomed Res Int</journal-id>
<journal-id journal-id-type="publisher-id">BMRI</journal-id>
<journal-title-group>
<journal-title>BioMed Research International</journal-title>
</journal-title-group>
<issn pub-type="ppub">2314-6133</issn>
<issn pub-type="epub">2314-6141</issn>
<publisher>
<publisher-name>Hindawi Publishing Corporation</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="pmid">26236726</article-id>
<article-id pub-id-type="pmc">4510254</article-id>
<article-id pub-id-type="doi">10.1155/2015/414681</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Research Article</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Diversity and Global Distribution of IncL/M Plasmids Enabling Horizontal Dissemination of
<italic>β</italic>
-Lactam Resistance Genes among the Enterobacteriaceae</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Adamczuk</surname>
<given-names>Marcin</given-names>
</name>
<xref ref-type="aff" rid="I1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zaleski</surname>
<given-names>Piotr</given-names>
</name>
<xref ref-type="aff" rid="I2">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Dziewit</surname>
<given-names>Lukasz</given-names>
</name>
<xref ref-type="aff" rid="I1">
<sup>1</sup>
</xref>
<xref ref-type="corresp" rid="cor1">
<sup>*</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wolinowska</surname>
<given-names>Renata</given-names>
</name>
<xref ref-type="aff" rid="I3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Nieckarz</surname>
<given-names>Marta</given-names>
</name>
<xref ref-type="aff" rid="I1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wawrzyniak</surname>
<given-names>Pawel</given-names>
</name>
<xref ref-type="aff" rid="I1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="I2">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Kieryl</surname>
<given-names>Piotr</given-names>
</name>
<xref ref-type="aff" rid="I2">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Plucienniczak</surname>
<given-names>Andrzej</given-names>
</name>
<xref ref-type="aff" rid="I2">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Bartosik</surname>
<given-names>Dariusz</given-names>
</name>
<xref ref-type="aff" rid="I1">
<sup>1</sup>
</xref>
<xref ref-type="corresp" rid="cor2">
<sup>*</sup>
</xref>
</contrib>
</contrib-group>
<aff id="I1">
<sup>1</sup>
Department of Bacterial Genetics, Institute of Microbiology, Faculty of Biology, University of Warsaw, Miecznikowa 1, 02-096 Warsaw, Poland</aff>
<aff id="I2">
<sup>2</sup>
Institute of Biotechnology and Antibiotics, Staroscinska 5, 02-516 Warsaw, Poland</aff>
<aff id="I3">
<sup>3</sup>
Department of Pharmaceutical Microbiology, Medical University of Warsaw, Oczki 3, 02-007 Warsaw, Poland</aff>
<author-notes>
<corresp id="cor1">*Lukasz Dziewit:
<email>ldziewit@biol.uw.edu.pl</email>
and </corresp>
<corresp id="cor2">*Dariusz Bartosik:
<email>bartosik@biol.uw.edu.pl</email>
</corresp>
<fn fn-type="other">
<p>Academic Editor: Jorge F. Quarleri</p>
</fn>
</author-notes>
<pub-date pub-type="ppub">
<year>2015</year>
</pub-date>
<pub-date pub-type="epub">
<day>8</day>
<month>7</month>
<year>2015</year>
</pub-date>
<volume>2015</volume>
<elocation-id>414681</elocation-id>
<history>
<date date-type="received">
<day>23</day>
<month>12</month>
<year>2014</year>
</date>
<date date-type="accepted">
<day>22</day>
<month>6</month>
<year>2015</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright © 2015 Marcin Adamczuk et al.</copyright-statement>
<copyright-year>2015</copyright-year>
<license license-type="open-access">
<license-p>This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.</license-p>
</license>
</permissions>
<abstract>
<p>Antibiotic resistance determinants are frequently associated with plasmids and other mobile genetic elements, which simplifies their horizontal transmission. Several groups of plasmids (including replicons of the IncL/M incompatibility group) were found to play an important role in the dissemination of resistance genes encoding
<italic>β</italic>
-lactamases. The IncL/M plasmids are large, broad host range, and self-transmissible replicons. We have identified and characterized two novel members of this group: pARM26 (isolated from bacteria inhabiting activated sludge from a wastewater treatment plant) and pIGT15 (originating from a clinical strain of
<italic>Escherichia coli</italic>
). This instigated a detailed comparative analysis of all available sequences of IncL/M plasmids encoding
<italic>β</italic>
-lactamases. The core genome of these plasmids is comprised of 20 genes with conserved synteny. Phylogenetic analyses of these core genes allowed clustering of the plasmids into four separate groups, which reflect their antibiotic resistance profiles. Examination of the biogeography of the IncL/M plasmids revealed that they are most frequently found in bacteria of the family Enterobacteriaceae originating from the Mediterranean region and Western Europe and that they are able to persist in various ecological niches even in the absence of direct antibiotic selection pressure.</p>
</abstract>
</article-meta>
</front>
<floats-group>
<fig id="fig1" orientation="portrait" position="float">
<label>Figure 1</label>
<caption>
<p>Linear map showing the genetic structure of circular plasmids pACM1, pARM26, and pIGT15. Arrows indicate the direction of gene transcription. The gray-shaded area connects DNA regions of different plasmids with at least 99% nucleotide sequence identity. Yellow frames indicate DNA regions (I and II) of pARM26 not present within pIGT15. REP: replication system; TRA: conjugal transfer system; PAR: partitioning system; InA and InB: class 1 integrons.</p>
</caption>
<graphic xlink:href="BMRI2015-414681.001"></graphic>
</fig>
<fig id="fig2" orientation="portrait" position="float">
<label>Figure 2</label>
<caption>
<p>Comparative analyses of IncL/M plasmids. (a) Phylogenetic tree based on the 20 core genes of the IncL/M plasmids. (b) Comparative genomic analysis of the IncL/M plasmids (sequence annotations of several plasmids have been manually corrected; the distinguished genes are shown by arrows on the linear maps: red:
<italic> bla</italic>
genes; yellow: other AR genes). The gray-shaded areas connect DNA regions of different plasmids with at least 68% nucleotide sequence identity. (c–k) Comparison of the antibiotic resistance gene contents of the IncL/M plasmids: (c)
<italic>β</italic>
-lactam resistance genes; (d) aminoglycoside resistance genes; (e) macrolide resistance genes; (f) quinolone resistance genes; (g) trimethoprim resistance genes; (h) sulfonamide resistance genes; (i) bleomycin resistance genes; (j) chloramphenicol resistance genes; (k) tetracycline resistance genes.</p>
</caption>
<graphic xlink:href="BMRI2015-414681.002"></graphic>
</fig>
<fig id="fig3" orientation="portrait" position="float">
<label>Figure 3</label>
<caption>
<p>Geographical distribution of IncL/M plasmids.</p>
</caption>
<graphic xlink:href="BMRI2015-414681.003"></graphic>
</fig>
<table-wrap id="tab1" orientation="portrait" position="float">
<label>Table 1</label>
<caption>
<p>IncL/M plasmid sequences deposited in the GenBank database.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" rowspan="1" colspan="1">Plasmid name</th>
<th align="left" rowspan="1" colspan="1">Host strain</th>
<th align="left" rowspan="1" colspan="1">Strain origin</th>
<th align="center" rowspan="1" colspan="1">Size
<break></break>
(bp)</th>
<th align="center" rowspan="1" colspan="1">%
<break></break>
GC </th>
<th align="left" rowspan="1" colspan="1">Reference/GenBank accession number</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" rowspan="1" colspan="1">pACM1</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella oxytoca </italic>
</td>
<td align="left" rowspan="1" colspan="1">New York, USA; hospital patient</td>
<td align="center" rowspan="1" colspan="1">89,977</td>
<td align="center" rowspan="1" colspan="1">54.4</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B10" ref-type="bibr">10</xref>
]/NC_024997</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pARM26</td>
<td align="left" rowspan="1" colspan="1">unknown bacterium</td>
<td align="left" rowspan="1" colspan="1">Warsaw, Poland; wastewater treatment plant</td>
<td align="center" rowspan="1" colspan="1">86,948</td>
<td align="center" rowspan="1" colspan="1">54.4</td>
<td align="left" rowspan="1" colspan="1">this work/KP294350</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pCTX-M3</td>
<td align="left" rowspan="1" colspan="1">
<italic>Citrobacter freundii </italic>
2526</td>
<td align="left" rowspan="1" colspan="1">Warsaw, Poland; hospital patient</td>
<td align="center" rowspan="1" colspan="1">89,468</td>
<td align="center" rowspan="1" colspan="1">51.0</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B11" ref-type="bibr">11</xref>
]/NC_004464</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pCTX-M360</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella pneumoniae</italic>
0773</td>
<td align="left" rowspan="1" colspan="1">Guangzhou, China; hospital patient</td>
<td align="center" rowspan="1" colspan="1">68,018</td>
<td align="center" rowspan="1" colspan="1">51.4</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B12" ref-type="bibr">12</xref>
]/NC_011641</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pE71T</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella pneumoniae</italic>
E71T</td>
<td align="left" rowspan="1" colspan="1">Ireland; hospital patient</td>
<td align="center" rowspan="1" colspan="1">63,578</td>
<td align="center" rowspan="1" colspan="1">51.2</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B13" ref-type="bibr">13</xref>
]/NC_023027</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pEl1573</td>
<td align="left" rowspan="1" colspan="1">
<italic>Enterobacter cloacae</italic>
El1573</td>
<td align="left" rowspan="1" colspan="1">Sydney, Australia; hospital patient</td>
<td align="center" rowspan="1" colspan="1">87,731</td>
<td align="center" rowspan="1" colspan="1">52.8</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B14" ref-type="bibr">14</xref>
]/NC_019368</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pEL60</td>
<td align="left" rowspan="1" colspan="1">
<italic>Erwinia amylovora</italic>
LebB66</td>
<td align="left" rowspan="1" colspan="1">Lebanon; fruits</td>
<td align="center" rowspan="1" colspan="1">60,145</td>
<td align="center" rowspan="1" colspan="1">51.5</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B22" ref-type="bibr">22</xref>
]/NC_005246</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pENT-d0d</td>
<td align="left" rowspan="1" colspan="1">
<italic>Enterobacter cloacae</italic>
ECNIH5</td>
<td align="left" rowspan="1" colspan="1">Bethesda, USA; hospital sink drain</td>
<td align="center" rowspan="1" colspan="1">77,575</td>
<td align="center" rowspan="1" colspan="1">50.7</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B39" ref-type="bibr">26</xref>
]/CP009857</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pENT-e56</td>
<td align="left" rowspan="1" colspan="1">
<italic>Enterobacter cloacae</italic>
ECNIH4</td>
<td align="left" rowspan="1" colspan="1">Bethesda, USA; hospital sink drain</td>
<td align="center" rowspan="1" colspan="1">77,248</td>
<td align="center" rowspan="1" colspan="1">51.5</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B39" ref-type="bibr">26</xref>
]/CP009852</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pFOX-7a</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella pneumonia</italic>
e 45A03</td>
<td align="left" rowspan="1" colspan="1">Siena, Italy; hospital patient</td>
<td align="center" rowspan="1" colspan="1">90,439</td>
<td align="center" rowspan="1" colspan="1">53.4</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B15" ref-type="bibr">15</xref>
]/NC_025134</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pIGT15</td>
<td align="left" rowspan="1" colspan="1">
<italic>Escherichia coli</italic>
CZD1527</td>
<td align="left" rowspan="1" colspan="1">Warsaw, Poland; hospital patient</td>
<td align="center" rowspan="1" colspan="1">74,839</td>
<td align="center" rowspan="1" colspan="1">54.7</td>
<td align="left" rowspan="1" colspan="1">this work/KP294351</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pJEG011</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella pneumoniae</italic>
Kp002</td>
<td align="left" rowspan="1" colspan="1">Sydney, Australia; hospital patient</td>
<td align="center" rowspan="1" colspan="1">71,446</td>
<td align="center" rowspan="1" colspan="1">50.6</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B16" ref-type="bibr">16</xref>
]/NC_021078</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pKOI-34</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella oxytoca</italic>
MS5279</td>
<td align="left" rowspan="1" colspan="1">Japan; hospital patient</td>
<td align="center" rowspan="1" colspan="1">87,343</td>
<td align="center" rowspan="1" colspan="1">53.3</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B84" ref-type="bibr">27</xref>
]/AB715422</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pKPN-068</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella pneumoniae </italic>
KPNIH27</td>
<td align="left" rowspan="1" colspan="1">USA; hospital patient</td>
<td align="center" rowspan="1" colspan="1">80,411</td>
<td align="center" rowspan="1" colspan="1">51.3</td>
<td align="left" rowspan="1" colspan="1">—/CP007733</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pKPoxa-48N1</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella pneumoniae </italic>
KP1a</td>
<td align="left" rowspan="1" colspan="1">Nancy, France; hospital patient</td>
<td align="center" rowspan="1" colspan="1">62,592</td>
<td align="center" rowspan="1" colspan="1">51.1</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B17" ref-type="bibr">17</xref>
]/NC_021488</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pNDM-HK</td>
<td align="left" rowspan="1" colspan="1">
<italic>Escherichia coli</italic>
HK-01</td>
<td align="left" rowspan="1" colspan="1">Hong Kong, China; hospital patient</td>
<td align="center" rowspan="1" colspan="1">88,803</td>
<td align="center" rowspan="1" colspan="1">51.5</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B18" ref-type="bibr">18</xref>
]/NC_019063</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pNDM-OM</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella pneumoniae</italic>
601</td>
<td align="left" rowspan="1" colspan="1">Oman; hospital patient</td>
<td align="center" rowspan="1" colspan="1">87,185</td>
<td align="center" rowspan="1" colspan="1">51.5</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B19" ref-type="bibr">19</xref>
]/NC_019889</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pNE1280</td>
<td align="left" rowspan="1" colspan="1">
<italic>Enterobacter cloacae</italic>
1623</td>
<td align="left" rowspan="1" colspan="1">Omaha, USA; hospital patient</td>
<td align="center" rowspan="1" colspan="1">66,531</td>
<td align="center" rowspan="1" colspan="1">52.3</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B20" ref-type="bibr">20</xref>
]/NC_019346</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pOXA-48a</td>
<td align="left" rowspan="1" colspan="1">
<italic>Klebsiella pneumoniae </italic>
Kp11978</td>
<td align="left" rowspan="1" colspan="1">Turkey; hospital patient</td>
<td align="center" rowspan="1" colspan="1">61,881</td>
<td align="center" rowspan="1" colspan="1">51.1</td>
<td align="left" rowspan="1" colspan="1">[
<xref rid="B21" ref-type="bibr">21</xref>
]/NC_019154</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">R830b</td>
<td align="left" rowspan="1" colspan="1">
<italic>Serratia marcescens </italic>
</td>
<td align="left" rowspan="1" colspan="1">USA</td>
<td align="center" rowspan="1" colspan="1">81,793</td>
<td align="center" rowspan="1" colspan="1">53.2</td>
<td align="left" rowspan="1" colspan="1">—/NC_019344</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="tab2" orientation="portrait" position="float">
<label>Table 2</label>
<caption>
<p>Unique genes identified within IncL/M plasmids.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" rowspan="1" colspan="1">Plasmid name</th>
<th align="center" rowspan="1" colspan="1">Number of singletons</th>
<th align="left" rowspan="1" colspan="1">Singleton gene/locus tag [possible function]</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" rowspan="1" colspan="1">pACM1</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="left" rowspan="1" colspan="1">
<italic>dfrA1</italic>
/D733_p1025 [dihydrofolate reductase]</td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pARM26</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pCTX-M3</td>
<td align="center" rowspan="1" colspan="1">3</td>
<td align="left" rowspan="1" colspan="1">
<italic>dfrA12</italic>
/pCTX-M3_084 [dihydrofolate reductase];
<italic>orfF</italic>
/pCTX-M3_085 [integron gene cassette, hypothetical protein];
<italic>aadA2</italic>
/pCTX-M3_086 [streptomycin 3′-adenyltransferase]</td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pCTX-M360</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pE71T</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pEl1573</td>
<td align="center" rowspan="1" colspan="1">5</td>
<td align="left" rowspan="1" colspan="1">
<italic>mphA</italic>
/D727_p1024 [macrolide phosphorylase];
<italic>mrx</italic>
/D727_p1026 [protein of unknown function, required for MphA expression];
<italic>mphR(A)</italic>
/D727_p1027 [transcriptional regulator for macrolide phosphorylase gene
<italic>mph(A</italic>
)];
<italic>catB3</italic>
/D727_p1035 [chloramphenicol acetyltransferase];
<italic>bla</italic>
<sub>IMP-4</sub>
/D727_p1038 [IMP-4 metallo-
<italic>β</italic>
-lactamase]</td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pEL60</td>
<td align="center" rowspan="1" colspan="1">2</td>
<td align="left" rowspan="1" colspan="1">
<italic>orf8</italic>
/pEL60p08 [hypothetical protein];
<italic>kfrA</italic>
/pEL60p09 [plasmid maintenance protein KfrA]</td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pENT-d0d</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pENT-e56</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pFOX-7a</td>
<td align="center" rowspan="1" colspan="1">9</td>
<td align="left" rowspan="1" colspan="1">bla
<sub>FOX-7</sub>
/D647_p51042 [FOX-7
<italic>β</italic>
-lactamase];
<italic>mdrL</italic>
/D647_p51043 [major facilitator transporter];
<italic>lysR</italic>
/D647_p51044 [LysR family transcriptional regulator];
<italic> tnpA</italic>
/D647_p51045 [transposase of IS
<italic>1634</italic>
-family insertion sequence]; tnpA/D647_p51046 [transposase of IS
<italic>Apu1</italic>
(IS
<italic>4</italic>
family/IS
<italic>H8</italic>
group)]; D647_p51047 [hypothetical protein];
<italic>tnpA</italic>
/D647_p51048 [transposase of IS
<italic>Apu2</italic>
(IS
<italic>4</italic>
family/IS
<italic>H8</italic>
group)];
<italic>istA</italic>
/D647_p51052 [IstA protein of IS
<italic>1326</italic>
(IS
<italic>21</italic>
family)];
<italic>istB</italic>
/D647_p51053 [IstB protein of IS
<italic>1326</italic>
(IS
<italic>21</italic>
family)]</td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pIGT15</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pJEG011</td>
<td align="center" rowspan="1" colspan="1">5</td>
<td align="left" rowspan="1" colspan="1">
<italic>strA</italic>
/D647_p23045 [streptomycin phosphotransferase A];
<italic>strB</italic>
/D647_p23040 [streptomycin phosphotransferase B];
<italic>aphA6</italic>
/D647_p23043 [aminoglycoside 3′-phosphotransferase]; tnpA/D647_p23042 [transposase of IS
<italic>1</italic>
-family insertion sequence];
<italic>bla</italic>
<sub>CTX-M-14</sub>
/D647_p23049 [extended-spectrum
<italic>β</italic>
-lactamase]</td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pKOI-34</td>
<td align="center" rowspan="1" colspan="1">7</td>
<td align="left" rowspan="1" colspan="1">
<italic>arsR</italic>
[arsenical resistance operon repressor]; arsD [arsenical resistance operon transacting repressor];
<italic>arsA</italic>
[arsenical pump-driving ATPase];
<italic>arsB</italic>
[arsenic efflux pump protein];
<italic>arsC</italic>
[arsenate reductase];
<italic>ycgA</italic>
[arsenical pump-driving ATPase];
<italic>bla</italic>
<sub>IMP-34</sub>
[IMP-34 metallo-
<italic>β</italic>
-lactamase]</td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pKPN-068</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pKPoxa-48N1</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pNDM-HK</td>
<td align="center" rowspan="1" colspan="1">1</td>
<td align="left" rowspan="1" colspan="1">
<italic>insL</italic>
/D616_p59011 [transposase of IS
<italic>4</italic>
family/IS
<italic>231</italic>
group insertion sequence]</td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pNDM-OM</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pNE1280</td>
<td align="center" rowspan="1" colspan="1">8</td>
<td align="left" rowspan="1" colspan="1">
<italic>istB</italic>
/KPC4_0043 [IstB protein of IS
<italic>21</italic>
-family insertion sequence];
<italic>bla</italic>
<sub>KPC-4</sub>
/KPC4_0045 [
<italic>β</italic>
-lactamase];
<italic>tnpA</italic>
/KPC4_0047 [transposase]; KPC4_0049 [hypothetical protein]; KPC4_0051 [hypothetical protein]; KPC4_0053 [hypothetical protein];
<italic>rom</italic>
/KPC4_0055 [Rop family protein];
<italic>exc1</italic>
/KPC4_0057 [exclusion protein 1]</td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">pOXA-48a</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1"></td>
</tr>
<tr>
<td align="center" colspan="3" rowspan="1">
<hr></hr>
</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">R830b</td>
<td align="center" rowspan="1" colspan="1">15</td>
<td align="left" rowspan="1" colspan="1">
<italic>tniQ</italic>
/D739_p1005 [transposase helper protein TniQ]; D739_p1006 [resolvase/integrase]; D739_p1010 [organomercurial lyase]; D739_p1020 [putative reverse transcriptase]; D739_p1024 [hypothetical protein]; D739_p1025 [Il-IS_2, transposase]; D739_p1026 [lipoprotein signal peptidase]; D739_p1027 [cobalt-zinc-cadmium resistance protein CzcD]; D739_p1028 [Cd(II)/Pb(II)-responsive transcriptional regulator]; D739_p1029 [hypothetical protein]; D739_p1107 [resolvase]; D739_p1108 [S-formylglutathione hydrolase]; D739_p1109 [glyoxalase]; D739_p1110 [S-(hydroxymethyl)glutathione dehydrogenase]; D739_p1111 [uncharacterized protein YaiN in formaldehyde detoxification operon]</td>
</tr>
</tbody>
</table>
</table-wrap>
</floats-group>
</pmc>
<affiliations>
<list>
<country>
<li>Pologne</li>
</country>
</list>
<tree>
<country name="Pologne">
<noRegion>
<name sortKey="Adamczuk, Marcin" sort="Adamczuk, Marcin" uniqKey="Adamczuk M" first="Marcin" last="Adamczuk">Marcin Adamczuk</name>
</noRegion>
<name sortKey="Bartosik, Dariusz" sort="Bartosik, Dariusz" uniqKey="Bartosik D" first="Dariusz" last="Bartosik">Dariusz Bartosik</name>
<name sortKey="Dziewit, Lukasz" sort="Dziewit, Lukasz" uniqKey="Dziewit L" first="Lukasz" last="Dziewit">Lukasz Dziewit</name>
<name sortKey="Kieryl, Piotr" sort="Kieryl, Piotr" uniqKey="Kieryl P" first="Piotr" last="Kieryl">Piotr Kieryl</name>
<name sortKey="Nieckarz, Marta" sort="Nieckarz, Marta" uniqKey="Nieckarz M" first="Marta" last="Nieckarz">Marta Nieckarz</name>
<name sortKey="Plucienniczak, Andrzej" sort="Plucienniczak, Andrzej" uniqKey="Plucienniczak A" first="Andrzej" last="Plucienniczak">Andrzej Plucienniczak</name>
<name sortKey="Wawrzyniak, Pawel" sort="Wawrzyniak, Pawel" uniqKey="Wawrzyniak P" first="Pawel" last="Wawrzyniak">Pawel Wawrzyniak</name>
<name sortKey="Wawrzyniak, Pawel" sort="Wawrzyniak, Pawel" uniqKey="Wawrzyniak P" first="Pawel" last="Wawrzyniak">Pawel Wawrzyniak</name>
<name sortKey="Wolinowska, Renata" sort="Wolinowska, Renata" uniqKey="Wolinowska R" first="Renata" last="Wolinowska">Renata Wolinowska</name>
<name sortKey="Zaleski, Piotr" sort="Zaleski, Piotr" uniqKey="Zaleski P" first="Piotr" last="Zaleski">Piotr Zaleski</name>
</country>
</tree>
</affiliations>
</record>

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