The Human Antimicrobial Protein Bactericidal/Permeability-Increasing Protein (BPI) Inhibits the Infectivity of Influenza A Virus
Identifieur interne : 000A70 ( Pmc/Curation ); précédent : 000A69; suivant : 000A71The Human Antimicrobial Protein Bactericidal/Permeability-Increasing Protein (BPI) Inhibits the Infectivity of Influenza A Virus
Auteurs : Olaf Pinkenburg [Allemagne] ; Torben Meyer [Allemagne] ; Norbert Bannert [Allemagne] ; Steven Norley [Allemagne] ; Kathrin Bolte [Allemagne] ; Volker Czudai-Matwich [Allemagne] ; Susanne Herold [Allemagne] ; André Gessner [Allemagne] ; Markus Schnare [Allemagne]Source :
- PLoS ONE [ 1932-6203 ] ; 2016.
Abstract
In addition to their well-known antibacterial activity some antimicrobial peptides and proteins (AMPs) display also antiviral effects. A 27
Url:
DOI: 10.1371/journal.pone.0156929
PubMed: 27273104
PubMed Central: 4894568
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">The Human Antimicrobial Protein Bactericidal/Permeability-Increasing Protein (BPI) Inhibits the Infectivity of Influenza A Virus</title><author><name sortKey="Pinkenburg, Olaf" sort="Pinkenburg, Olaf" uniqKey="Pinkenburg O" first="Olaf" last="Pinkenburg">Olaf Pinkenburg</name><affiliation wicri:level="1"><nlm:aff id="aff001"><addr-line>Institute for Immunology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Immunology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author><author><name sortKey="Meyer, Torben" sort="Meyer, Torben" uniqKey="Meyer T" first="Torben" last="Meyer">Torben Meyer</name><affiliation wicri:level="1"><nlm:aff id="aff001"><addr-line>Institute for Immunology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Immunology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author><author><name sortKey="Bannert, Norbert" sort="Bannert, Norbert" uniqKey="Bannert N" first="Norbert" last="Bannert">Norbert Bannert</name><affiliation wicri:level="1"><nlm:aff id="aff002"><addr-line>Department for HIV and other Retroviruses, Robert Koch Institute, Berlin, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Department for HIV and other Retroviruses, Robert Koch Institute, Berlin</wicri:regionArea></affiliation></author><author><name sortKey="Norley, Steven" sort="Norley, Steven" uniqKey="Norley S" first="Steven" last="Norley">Steven Norley</name><affiliation wicri:level="1"><nlm:aff id="aff002"><addr-line>Department for HIV and other Retroviruses, Robert Koch Institute, Berlin, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Department for HIV and other Retroviruses, Robert Koch Institute, Berlin</wicri:regionArea></affiliation></author><author><name sortKey="Bolte, Kathrin" sort="Bolte, Kathrin" uniqKey="Bolte K" first="Kathrin" last="Bolte">Kathrin Bolte</name><affiliation wicri:level="1"><nlm:aff id="aff003"><addr-line>Laboratory for Cell Biology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Laboratory for Cell Biology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author><author><name sortKey="Czudai Matwich, Volker" sort="Czudai Matwich, Volker" uniqKey="Czudai Matwich V" first="Volker" last="Czudai-Matwich">Volker Czudai-Matwich</name><affiliation wicri:level="1"><nlm:aff id="aff004"><addr-line>Institute for Virology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Virology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author><author><name sortKey="Herold, Susanne" sort="Herold, Susanne" uniqKey="Herold S" first="Susanne" last="Herold">Susanne Herold</name><affiliation wicri:level="1"><nlm:aff id="aff005"><addr-line>Department of Internal Medicine II, University of Giessen Lung Center and German Center for Lung Research, Giessen, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Department of Internal Medicine II, University of Giessen Lung Center and German Center for Lung Research, Giessen</wicri:regionArea></affiliation></author><author><name sortKey="Gessner, Andre" sort="Gessner, Andre" uniqKey="Gessner A" first="André" last="Gessner">André Gessner</name><affiliation wicri:level="1"><nlm:aff id="aff006"><addr-line>Institute for Clinical Microbiology and Hygiene, University Regensburg, Regensburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Clinical Microbiology and Hygiene, University Regensburg, Regensburg</wicri:regionArea></affiliation></author><author><name sortKey="Schnare, Markus" sort="Schnare, Markus" uniqKey="Schnare M" first="Markus" last="Schnare">Markus Schnare</name><affiliation wicri:level="1"><nlm:aff id="aff001"><addr-line>Institute for Immunology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Immunology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">27273104</idno><idno type="pmc">4894568</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4894568</idno><idno type="RBID">PMC:4894568</idno><idno type="doi">10.1371/journal.pone.0156929</idno><date when="2016">2016</date><idno type="wicri:Area/Pmc/Corpus">000A70</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000A70</idno><idno type="wicri:Area/Pmc/Curation">000A70</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Curation">000A70</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">The Human Antimicrobial Protein Bactericidal/Permeability-Increasing Protein (BPI) Inhibits the Infectivity of Influenza A Virus</title><author><name sortKey="Pinkenburg, Olaf" sort="Pinkenburg, Olaf" uniqKey="Pinkenburg O" first="Olaf" last="Pinkenburg">Olaf Pinkenburg</name><affiliation wicri:level="1"><nlm:aff id="aff001"><addr-line>Institute for Immunology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Immunology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author><author><name sortKey="Meyer, Torben" sort="Meyer, Torben" uniqKey="Meyer T" first="Torben" last="Meyer">Torben Meyer</name><affiliation wicri:level="1"><nlm:aff id="aff001"><addr-line>Institute for Immunology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Immunology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author><author><name sortKey="Bannert, Norbert" sort="Bannert, Norbert" uniqKey="Bannert N" first="Norbert" last="Bannert">Norbert Bannert</name><affiliation wicri:level="1"><nlm:aff id="aff002"><addr-line>Department for HIV and other Retroviruses, Robert Koch Institute, Berlin, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Department for HIV and other Retroviruses, Robert Koch Institute, Berlin</wicri:regionArea></affiliation></author><author><name sortKey="Norley, Steven" sort="Norley, Steven" uniqKey="Norley S" first="Steven" last="Norley">Steven Norley</name><affiliation wicri:level="1"><nlm:aff id="aff002"><addr-line>Department for HIV and other Retroviruses, Robert Koch Institute, Berlin, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Department for HIV and other Retroviruses, Robert Koch Institute, Berlin</wicri:regionArea></affiliation></author><author><name sortKey="Bolte, Kathrin" sort="Bolte, Kathrin" uniqKey="Bolte K" first="Kathrin" last="Bolte">Kathrin Bolte</name><affiliation wicri:level="1"><nlm:aff id="aff003"><addr-line>Laboratory for Cell Biology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Laboratory for Cell Biology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author><author><name sortKey="Czudai Matwich, Volker" sort="Czudai Matwich, Volker" uniqKey="Czudai Matwich V" first="Volker" last="Czudai-Matwich">Volker Czudai-Matwich</name><affiliation wicri:level="1"><nlm:aff id="aff004"><addr-line>Institute for Virology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Virology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author><author><name sortKey="Herold, Susanne" sort="Herold, Susanne" uniqKey="Herold S" first="Susanne" last="Herold">Susanne Herold</name><affiliation wicri:level="1"><nlm:aff id="aff005"><addr-line>Department of Internal Medicine II, University of Giessen Lung Center and German Center for Lung Research, Giessen, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Department of Internal Medicine II, University of Giessen Lung Center and German Center for Lung Research, Giessen</wicri:regionArea></affiliation></author><author><name sortKey="Gessner, Andre" sort="Gessner, Andre" uniqKey="Gessner A" first="André" last="Gessner">André Gessner</name><affiliation wicri:level="1"><nlm:aff id="aff006"><addr-line>Institute for Clinical Microbiology and Hygiene, University Regensburg, Regensburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Clinical Microbiology and Hygiene, University Regensburg, Regensburg</wicri:regionArea></affiliation></author><author><name sortKey="Schnare, Markus" sort="Schnare, Markus" uniqKey="Schnare M" first="Markus" last="Schnare">Markus Schnare</name><affiliation wicri:level="1"><nlm:aff id="aff001"><addr-line>Institute for Immunology, Philipps-University of Marburg, Marburg, Germany</addr-line></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea>Institute for Immunology, Philipps-University of Marburg, Marburg</wicri:regionArea></affiliation></author></analytic><series><title level="j">PLoS ONE</title><idno type="eISSN">1932-6203</idno><imprint><date when="2016">2016</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>In addition to their well-known antibacterial activity some antimicrobial peptides and proteins (AMPs) display also antiviral effects. A 27 <monospace>aa</monospace> peptide from the N-terminal part of human bactericidal/permeability-increasing protein (BPI) previously shown to harbour antibacterial activity inhibits the infectivity of multiple Influenza A virus strains (H1N1, H3N2 and H5N1) the causing agent of the Influenza pneumonia. In contrast, the homologous murine BPI-peptide did not show activity against Influenza A virus. In addition human BPI-peptide inhibits the activation of immune cells mediated by Influenza A virus. By changing the human BPI-peptide to the sequence of the mouse homologous peptide the antiviral activity was completely abolished. Furthermore, the human BPI-peptide also inhibited the pathogenicity of the Vesicular Stomatitis Virus but failed to interfere with HIV and measles virus. Electron microscopy indicate that the human BPI-peptide interferes with the virus envelope and at high concentrations was able to destroy the particles completely.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Molinari, Na" uniqKey="Molinari N">NA Molinari</name></author><author><name sortKey="Ortega Sanchez, Ir" uniqKey="Ortega Sanchez I">IR Ortega-Sanchez</name></author><author><name sortKey="Messonnier, Ml" uniqKey="Messonnier M">ML Messonnier</name></author><author><name sortKey="Thompson, Ww" uniqKey="Thompson W">WW Thompson</name></author><author><name sortKey="Wortley, Pm" uniqKey="Wortley P">PM Wortley</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Johnson, Np" uniqKey="Johnson N">NP Johnson</name></author><author><name sortKey="Mueller, J" uniqKey="Mueller J">J Mueller</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Medina, Ra" uniqKey="Medina 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Types</subject><subj-group><subject>Animal Cells</subject><subj-group><subject>Blood Cells</subject><subj-group><subject>White Blood Cells</subject><subj-group><subject>Granulocytes</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and Life Sciences</subject><subj-group><subject>Cell Biology</subject><subj-group><subject>Cellular Types</subject><subj-group><subject>Animal Cells</subject><subj-group><subject>Immune Cells</subject><subj-group><subject>White Blood Cells</subject><subj-group><subject>Granulocytes</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and Life Sciences</subject><subj-group><subject>Immunology</subject><subj-group><subject>Immune Cells</subject><subj-group><subject>White Blood 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Control</subject><subj-group><subject>Antimicrobials</subject></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Research and Analysis Methods</subject><subj-group><subject>Immunologic Techniques</subject><subj-group><subject>Immunoassays</subject><subj-group><subject>Enzyme-Linked Immunoassays</subject></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and life sciences</subject><subj-group><subject>Organisms</subject><subj-group><subject>Viruses</subject><subj-group><subject>RNA viruses</subject><subj-group><subject>Rhabdoviruses</subject><subj-group><subject>Vesicular Stomatitis Virus</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and Life Sciences</subject><subj-group><subject>Microbiology</subject><subj-group><subject>Medical Microbiology</subject><subj-group><subject>Microbial Pathogens</subject><subj-group><subject>Viral Pathogens</subject><subj-group><subject>Vesicular Stomatitis Virus</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Medicine and Health Sciences</subject><subj-group><subject>Pathology and Laboratory Medicine</subject><subj-group><subject>Pathogens</subject><subj-group><subject>Microbial Pathogens</subject><subj-group><subject>Viral Pathogens</subject><subj-group><subject>Vesicular Stomatitis Virus</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and Life Sciences</subject><subj-group><subject>Organisms</subject><subj-group><subject>Viruses</subject><subj-group><subject>Viral Pathogens</subject><subj-group><subject>Vesicular Stomatitis Virus</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and Life Sciences</subject><subj-group><subject>Physiology</subject><subj-group><subject>Immune Physiology</subject><subj-group><subject>Antibodies</subject><subj-group><subject>Monoclonal Antibodies</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Medicine and Health Sciences</subject><subj-group><subject>Physiology</subject><subj-group><subject>Immune Physiology</subject><subj-group><subject>Antibodies</subject><subj-group><subject>Monoclonal Antibodies</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and Life Sciences</subject><subj-group><subject>Immunology</subject><subj-group><subject>Immune System Proteins</subject><subj-group><subject>Antibodies</subject><subj-group><subject>Monoclonal Antibodies</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Medicine and Health Sciences</subject><subj-group><subject>Immunology</subject><subj-group><subject>Immune System Proteins</subject><subj-group><subject>Antibodies</subject><subj-group><subject>Monoclonal Antibodies</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3"><subject>Biology and Life Sciences</subject><subj-group><subject>Biochemistry</subject><subj-group><subject>Proteins</subject><subj-group><subject>Immune System Proteins</subject><subj-group><subject>Antibodies</subject><subj-group><subject>Monoclonal Antibodies</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group></article-categories><title-group><article-title>The Human Antimicrobial Protein Bactericidal/Permeability-Increasing Protein (BPI) Inhibits the Infectivity of Influenza A Virus</article-title><alt-title alt-title-type="running-head">BPI and Influenza</alt-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Pinkenburg</surname><given-names>Olaf</given-names></name><xref ref-type="aff" rid="aff001"><sup>1</sup></xref></contrib><contrib contrib-type="author"><name><surname>Meyer</surname><given-names>Torben</given-names></name><xref ref-type="aff" rid="aff001"><sup>1</sup></xref></contrib><contrib contrib-type="author"><name><surname>Bannert</surname><given-names>Norbert</given-names></name><xref ref-type="aff" rid="aff002"><sup>2</sup></xref></contrib><contrib contrib-type="author"><name><surname>Norley</surname><given-names>Steven</given-names></name><xref ref-type="aff" rid="aff002"><sup>2</sup></xref></contrib><contrib contrib-type="author"><name><surname>Bolte</surname><given-names>Kathrin</given-names></name><xref ref-type="aff" rid="aff003"><sup>3</sup></xref></contrib><contrib contrib-type="author"><name><surname>Czudai-Matwich</surname><given-names>Volker</given-names></name><xref ref-type="aff" rid="aff004"><sup>4</sup></xref></contrib><contrib contrib-type="author"><name><surname>Herold</surname><given-names>Susanne</given-names></name><xref ref-type="aff" rid="aff005"><sup>5</sup></xref></contrib><contrib contrib-type="author"><name><surname>Gessner</surname><given-names>André</given-names></name><xref ref-type="aff" rid="aff006"><sup>6</sup></xref></contrib><contrib contrib-type="author"><name><surname>Schnare</surname><given-names>Markus</given-names></name><xref ref-type="aff" rid="aff001"><sup>1</sup></xref><xref ref-type="corresp" rid="cor001">*</xref></contrib></contrib-group><aff id="aff001"><label>1</label><addr-line>Institute for Immunology, Philipps-University of Marburg, Marburg, Germany</addr-line></aff><aff id="aff002"><label>2</label><addr-line>Department for HIV and other Retroviruses, Robert Koch Institute, Berlin, Germany</addr-line></aff><aff id="aff003"><label>3</label><addr-line>Laboratory for Cell Biology, Philipps-University of Marburg, Marburg, Germany</addr-line></aff><aff id="aff004"><label>4</label><addr-line>Institute for Virology, Philipps-University of Marburg, Marburg, Germany</addr-line></aff><aff id="aff005"><label>5</label><addr-line>Department of Internal Medicine II, University of Giessen Lung Center and German Center for Lung Research, Giessen, Germany</addr-line></aff><aff id="aff006"><label>6</label><addr-line>Institute for Clinical Microbiology and Hygiene, University Regensburg, Regensburg, Germany</addr-line></aff><contrib-group><contrib contrib-type="editor"><name><surname>Pöhlmann</surname><given-names>Stefan</given-names></name><role>Editor</role><xref ref-type="aff" rid="edit1"></xref></contrib></contrib-group><aff id="edit1"><addr-line>German Primate Center, GERMANY</addr-line></aff><author-notes><fn fn-type="COI-statement" id="coi001"><p><bold>Competing Interests: </bold>The authors have declared that no competing interests exist.</p></fn><fn fn-type="con" id="contrib001"><p>Conceived and designed the experiments: MS OP KB. Performed the experiments: OP TM SN KB MS VCM. Analyzed the data: OP NB KB MS. Contributed reagents/materials/analysis tools: SH AG. Wrote the paper: MS.</p></fn><corresp id="cor001">* E-mail: <email>markus.schnare@staff.uni-marburg.de</email></corresp></author-notes><pub-date pub-type="epub"><day>6</day><month>6</month><year>2016</year></pub-date><pub-date pub-type="collection"><year>2016</year></pub-date><volume>11</volume><issue>6</issue><elocation-id>e0156929</elocation-id><history><date date-type="received"><day>21</day><month>3</month><year>2016</year></date><date date-type="accepted"><day>20</day><month>5</month><year>2016</year></date></history><permissions><copyright-statement>© 2016 Pinkenburg et al</copyright-statement><copyright-year>2016</copyright-year><copyright-holder>Pinkenburg et al</copyright-holder><license xlink:href="http://creativecommons.org/licenses/by/4.0/"><license-p>This is an open access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License</ext-link>, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p></license></permissions><self-uri content-type="pdf" xlink:href="pone.0156929.pdf"></self-uri><abstract><p>In addition to their well-known antibacterial activity some antimicrobial peptides and proteins (AMPs) display also antiviral effects. A 27 <monospace>aa</monospace> peptide from the N-terminal part of human bactericidal/permeability-increasing protein (BPI) previously shown to harbour antibacterial activity inhibits the infectivity of multiple Influenza A virus strains (H1N1, H3N2 and H5N1) the causing agent of the Influenza pneumonia. In contrast, the homologous murine BPI-peptide did not show activity against Influenza A virus. In addition human BPI-peptide inhibits the activation of immune cells mediated by Influenza A virus. By changing the human BPI-peptide to the sequence of the mouse homologous peptide the antiviral activity was completely abolished. Furthermore, the human BPI-peptide also inhibited the pathogenicity of the Vesicular Stomatitis Virus but failed to interfere with HIV and measles virus. Electron microscopy indicate that the human BPI-peptide interferes with the virus envelope and at high concentrations was able to destroy the particles completely.</p></abstract><funding-group><award-group id="award001"><funding-source><institution>DFG</institution></funding-source><award-id>SCHN635/4-1</award-id><principal-award-recipient><name><surname>Schnare</surname><given-names>Markus</given-names></name></principal-award-recipient></award-group><award-group id="award002"><funding-source><institution>von-Behring-Röntgen Stiftung Giessen and Marburg</institution></funding-source><award-id>61-0053</award-id><principal-award-recipient><name><surname>Schnare</surname><given-names>Markus</given-names></name></principal-award-recipient></award-group><funding-statement>This work was supported in part by DFG grant SCHN635/4-1 (<ext-link ext-link-type="uri" xlink:href="http://www.dfg.de">http://www.dfg.de</ext-link>), von-Behring-Röntgen Stiftung Giessen and Marburg (grant number 61-0053 to MS)(<ext-link ext-link-type="uri" xlink:href="http://www.br-stiftung.de">http://www.br-stiftung.de</ext-link>). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.</funding-statement></funding-group><counts><fig-count count="6"></fig-count><table-count count="1"></table-count><page-count count="18"></page-count></counts><custom-meta-group><custom-meta id="data-availability"><meta-name>Data Availability</meta-name><meta-value>All relevant data are within the paper and its Supporting Information files.</meta-value></custom-meta></custom-meta-group></article-meta><notes><title>Data Availability</title><p>All relevant data are within the paper and its Supporting Information files.</p></notes></front></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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