<?xml version="1.0" encoding="UTF-8"?><?xml-stylesheet type="text/xsl" href="static/style.xsl"?><OAI-PMH xmlns="http://www.openarchives.org/OAI/2.0/" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xsi:schemaLocation="http://www.openarchives.org/OAI/2.0/ http://www.openarchives.org/OAI/2.0/OAI-PMH.xsd"><responseDate>2026-06-27T23:05:09Z</responseDate><request verb="GetRecord" identifier="oai:docta.ucm.es:20.500.14352/94237" metadataPrefix="mets">https://docta.ucm.es/rest/oai/request</request><GetRecord><record><header><identifier>oai:docta.ucm.es:20.500.14352/94237</identifier><datestamp>2025-03-18T16:05:32Z</datestamp><setSpec>com_20.500.14352_14</setSpec><setSpec>col_20.500.14352_15</setSpec></header><metadata><mets xmlns="http://www.loc.gov/METS/" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:doc="http://www.lyncode.com/xoai" ID="&#xa;&#x9;&#x9;&#x9;&#x9;DSpace_ITEM_20.500.14352-94237" TYPE="DSpace ITEM" PROFILE="DSpace METS SIP Profile 1.0" xsi:schemaLocation="http://www.loc.gov/METS/ http://www.loc.gov/standards/mets/mets.xsd" OBJID="&#xa;&#x9;&#x9;&#x9;&#x9;hdl:20.500.14352/94237">
   <metsHdr CREATEDATE="2026-06-28T01:05:09Z">
      <agent ROLE="CUSTODIAN" TYPE="ORGANIZATION">
         <name>Docta Complutense</name>
      </agent>
   </metsHdr>
   <dmdSec ID="DMD_20.500.14352_94237">
      <mdWrap MDTYPE="MODS">
         <xmlData xmlns:mods="http://www.loc.gov/mods/v3" xsi:schemaLocation="http://www.loc.gov/mods/v3 http://www.loc.gov/standards/mods/v3/mods-3-1.xsd">
            <mods:mods xsi:schemaLocation="http://www.loc.gov/mods/v3 http://www.loc.gov/standards/mods/v3/mods-3-1.xsd">
               <mods:name>
                  <mods:role>
                     <mods:roleTerm type="text">author</mods:roleTerm>
                  </mods:role>
                  <mods:namePart>Escudero García-Calderón, José Antonio</mods:namePart>
               </mods:name>
               <mods:name>
                  <mods:role>
                     <mods:roleTerm type="text">author</mods:roleTerm>
                  </mods:role>
                  <mods:namePart>Rodríguez-Beltrán, Jerónimo</mods:namePart>
               </mods:name>
               <mods:name>
                  <mods:role>
                     <mods:roleTerm type="text">author</mods:roleTerm>
                  </mods:role>
                  <mods:namePart>Hernández-Beltrán, J. Carlos R.</mods:namePart>
               </mods:name>
               <mods:name>
                  <mods:role>
                     <mods:roleTerm type="text">author</mods:roleTerm>
                  </mods:role>
                  <mods:namePart>De la Fuente, Javier</mods:namePart>
               </mods:name>
               <mods:name>
                  <mods:role>
                     <mods:roleTerm type="text">author</mods:roleTerm>
                  </mods:role>
                  <mods:namePart>Fuentes-Hernández, Ayari</mods:namePart>
               </mods:name>
               <mods:name>
                  <mods:role>
                     <mods:roleTerm type="text">author</mods:roleTerm>
                  </mods:role>
                  <mods:namePart>MacLean, R. Craig</mods:namePart>
               </mods:name>
               <mods:name>
                  <mods:role>
                     <mods:roleTerm type="text">author</mods:roleTerm>
                  </mods:role>
                  <mods:namePart>Peña-Miller, Rafael</mods:namePart>
               </mods:name>
               <mods:name>
                  <mods:role>
                     <mods:roleTerm type="text">author</mods:roleTerm>
                  </mods:role>
                  <mods:namePart>San Millan, Álvaro</mods:namePart>
               </mods:name>
               <mods:extension>
                  <mods:dateAccessioned encoding="iso8601">2024-01-22T08:44:04Z</mods:dateAccessioned>
               </mods:extension>
               <mods:extension>
                  <mods:dateAvailable encoding="iso8601">2024-01-22T08:44:04Z</mods:dateAvailable>
               </mods:extension>
               <mods:originInfo>
                  <mods:dateIssued encoding="iso8601">2018-04-09</mods:dateIssued>
               </mods:originInfo>
               <mods:identifier type="issn">2397-334X</mods:identifier>
               <mods:identifier type="doi">10.1038/s41559-018-0529-z</mods:identifier>
               <mods:identifier type="uri">https://hdl.handle.net/20.500.14352/94237</mods:identifier>
               <mods:identifier type="officialurl">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6055991/</mods:identifier>
               <mods:abstract>Understanding the mechanisms governing innovation is a central element of evolutionary theory. Novel traits usually arise through mutations in existing genes, but trade-offs between new and ancestral protein functions are pervasive and constrain the evolution of innovation. Classical models posit that evolutionary innovation circumvents the constraints imposed by trade-offs through genetic amplifications, which provide functional redundancy. Bacterial multicopy plasmids provide a paradigmatic example of genetic amplification, yet their role in evolutionary innovation remains largely unexplored. Here, we reconstructed the evolution of a new trait encoded in a multicopy plasmid using TEM-1 β-lactamase as a model system. Through a combination of theory and experimentation, we show that multicopy plasmids promote the coexistence of ancestral and novel traits for dozens of generations, allowing bacteria to escape the evolutionary constraints imposed by trade-offs. Our results suggest that multicopy plasmids are excellent platforms for evolutionary innovation, contributing to explain their extreme abundance in bacteria.</mods:abstract>
               <mods:language>
                  <mods:languageTerm authority="rfc3066">eng</mods:languageTerm>
               </mods:language>
               <mods:accessCondition type="useAndReproduction">Attribution 4.0 International</mods:accessCondition>
               <mods:titleInfo>
                  <mods:title>Multicopy plasmids allow bacteria to escape from fitness trade-offs during evolutionary innovation</mods:title>
               </mods:titleInfo>
               <mods:genre>journal article</mods:genre>
            </mods:mods>
         </xmlData>
      </mdWrap>
   </dmdSec>
   <amdSec ID="TMD_20.500.14352_94237">
      <rightsMD ID="RIG_20.500.14352_94237">
         <mdWrap MIMETYPE="text/plain" MDTYPE="OTHER" OTHERMDTYPE="DSpaceDepositLicense">
            <binData>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</binData>
         </mdWrap>
      </rightsMD>
   </amdSec>
   <amdSec ID="FO_20.500.14352_94237_1">
      <techMD ID="TECH_O_20.500.14352_94237_1">
         <mdWrap MDTYPE="PREMIS">
            <xmlData xmlns:premis="http://www.loc.gov/standards/premis" xsi:schemaLocation="http://www.loc.gov/standards/premis http://www.loc.gov/standards/premis/PREMIS-v1-0.xsd">
               <premis:premis>
                  <premis:object>
                     <premis:objectIdentifier>
                        <premis:objectIdentifierType>URL</premis:objectIdentifierType>
                        <premis:objectIdentifierValue>https://docta.ucm.es/bitstreams/428b56fb-6eb8-4fd2-94b4-2fbf0da13cb0/download</premis:objectIdentifierValue>
                     </premis:objectIdentifier>
                     <premis:objectCategory>File</premis:objectCategory>
                     <premis:objectCharacteristics>
                        <premis:fixity>
                           <premis:messageDigestAlgorithm>MD5</premis:messageDigestAlgorithm>
                           <premis:messageDigest>4fc64b806c9a385f02da230214b52f1c</premis:messageDigest>
                        </premis:fixity>
                        <premis:size>2119047</premis:size>
                        <premis:format>
                           <premis:formatDesignation>
                              <premis:formatName>application/pdf</premis:formatName>
                           </premis:formatDesignation>
                        </premis:format>
                     </premis:objectCharacteristics>
                     <premis:originalName>emss-76535.pdf</premis:originalName>
                  </premis:object>
               </premis:premis>
            </xmlData>
         </mdWrap>
      </techMD>
   </amdSec>
   <amdSec ID="FT_20.500.14352_94237_5">
      <techMD ID="TECH_T_20.500.14352_94237_5">
         <mdWrap MDTYPE="PREMIS">
            <xmlData xmlns:premis="http://www.loc.gov/standards/premis" xsi:schemaLocation="http://www.loc.gov/standards/premis http://www.loc.gov/standards/premis/PREMIS-v1-0.xsd">
               <premis:premis>
                  <premis:object>
                     <premis:objectIdentifier>
                        <premis:objectIdentifierType>URL</premis:objectIdentifierType>
                        <premis:objectIdentifierValue>https://docta.ucm.es/bitstreams/68255c3d-395f-4d58-bed4-6fb0d0d0be92/download</premis:objectIdentifierValue>
                     </premis:objectIdentifier>
                     <premis:objectCategory>File</premis:objectCategory>
                     <premis:objectCharacteristics>
                        <premis:fixity>
                           <premis:messageDigestAlgorithm>MD5</premis:messageDigestAlgorithm>
                           <premis:messageDigest>1b43684650fcc9ba481d52a71112ae83</premis:messageDigest>
                        </premis:fixity>
                        <premis:size>71146</premis:size>
                        <premis:format>
                           <premis:formatDesignation>
                              <premis:formatName>text/plain</premis:formatName>
                           </premis:formatDesignation>
                        </premis:format>
                     </premis:objectCharacteristics>
                     <premis:originalName>emss-76535.pdf.txt</premis:originalName>
                  </premis:object>
               </premis:premis>
            </xmlData>
         </mdWrap>
      </techMD>
   </amdSec>
   <fileSec>
      <fileGrp USE="ORIGINAL">
         <file ID="BITSTREAM_ORIGINAL_20.500.14352_94237_1" MIMETYPE="application/pdf" SEQ="1" SIZE="2119047" CHECKSUM="4fc64b806c9a385f02da230214b52f1c" CHECKSUMTYPE="MD5" ADMID="FO_20.500.14352_94237_1" GROUPID="GROUP_BITSTREAM_20.500.14352_94237_1">
            <FLocat LOCTYPE="URL" xlink:type="simple" xlink:href="https://docta.ucm.es/bitstreams/428b56fb-6eb8-4fd2-94b4-2fbf0da13cb0/download"/>
         </file>
      </fileGrp>
      <fileGrp USE="TEXT">
         <file ID="BITSTREAM_TEXT_20.500.14352_94237_5" MIMETYPE="text/plain" SEQ="5" SIZE="71146" CHECKSUM="1b43684650fcc9ba481d52a71112ae83" CHECKSUMTYPE="MD5" ADMID="FT_20.500.14352_94237_5" GROUPID="GROUP_BITSTREAM_20.500.14352_94237_5">
            <FLocat LOCTYPE="URL" xlink:type="simple" xlink:href="https://docta.ucm.es/bitstreams/68255c3d-395f-4d58-bed4-6fb0d0d0be92/download"/>
         </file>
      </fileGrp>
   </fileSec>
   <structMap LABEL="DSpace Object" TYPE="LOGICAL">
      <div TYPE="DSpace Object Contents" ADMID="DMD_20.500.14352_94237">
         <div TYPE="DSpace BITSTREAM">
            <fptr FILEID="BITSTREAM_ORIGINAL_20.500.14352_94237_1"/>
         </div>
      </div>
   </structMap>
</mets></metadata></record></GetRecord></OAI-PMH>