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                  <publisher>Nature Publishing Group</publisher>
                  <biblScope unit="volume">520</biblScope>
                  <biblScope unit="issue">7546</biblScope>
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                  <date type="datePub">2015-04-09</date>
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              <p>The highly complex structure of the human brain is strongly shaped by genetic influences. Subcortical brain regions form circuits with cortical areas to coordinate movement, learning, memory and motivation, and altered circuits can lead to abnormal behaviour and disease. To investigate how common genetic variants affect the structure of these brain regions, here we conduct genome-wide association studies of the volumes of seven subcortical regions and the intracranial volume derived from magnetic resonance images of 30,717 individuals from 50 cohorts. We identify five novel genetic variants influencing the volumes of the putamen and caudate nucleus. We also find stronger evidence for three loci with previously established influences on hippocampal volume and intracranial volume. These variants show specific volumetric effects on brain structures rather than global effects across structures. The strongest effects were found for the putamen, where a novel intergenic locus with replicable influence on volume (rs945270; P = 1.08 × 10(-33); 0.52% variance explained) showed evidence of altering the expression of the KTN1 gene in both brain and blood tissue. Variants influencing putamen volume clustered near developmental genes that regulate apoptosis, axon guidance and vesicle transport. Identification of these genetic variants provides insight into the causes of variability in human brain development, and may help to determine mechanisms of neuropsychiatric dysfunction.</p>
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          <orgName>Centre National de la Recherche Scientifique</orgName>
          <orgName type="acronym">CNRS</orgName>
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          <desc>
            <address>
              <country key="FR"/>
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        <org type="institution" xml:id="struct-92114" status="OLD">
          <idno type="ROR">https://ror.org/01x3gbx83</idno>
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          <orgName type="acronym">INRA</orgName>
          <date type="start">1946-05-18</date>
          <date type="end">2019-12-31</date>
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        </org>
        <org type="institution" xml:id="struct-134741" status="VALID">
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          <orgName>McGill University = Université McGill [Montréal, Canada]</orgName>
          <desc>
            <address>
              <addrLine>845, rue Sherbrooke O. Montréal (Québec) Canada H3A 0G4</addrLine>
              <country key="CA"/>
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            <ref type="url">http://www.mcgill.ca/</ref>
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        </org>
        <org type="institution" xml:id="struct-469153" status="VALID">
          <idno type="ROR">https://ror.org/05s92vm98</idno>
          <orgName>Xidian University</orgName>
          <date type="start">2016-10-21</date>
          <desc>
            <address>
              <addrLine>Xi'an</addrLine>
              <country key="CN"/>
            </address>
            <ref type="url">https://en.xidian.edu.cn/</ref>
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        </org>
        <org type="institution" xml:id="struct-64587" status="VALID">
          <idno type="ROR">https://ror.org/02fa3aq29</idno>
          <orgName>McMaster University [Hamilton, Ontario]</orgName>
          <desc>
            <address>
              <addrLine>1280 Main Street WestHamilton, Ontario  L8S 4L8</addrLine>
              <country key="CA"/>
            </address>
            <ref type="url">http://www.mcmaster.ca/</ref>
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        <org type="institution" xml:id="struct-444582" status="VALID">
          <orgName>Royal Netherlands Academy of Arts and Sciences</orgName>
          <orgName type="acronym">KNAW</orgName>
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            <address>
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        <org type="institution" xml:id="struct-300728" status="VALID">
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          <idno type="ROR">https://ror.org/03zga2b32</idno>
          <orgName>University of Bergen</orgName>
          <orgName type="acronym">UiB</orgName>
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            <address>
              <addrLine>Universitetet i Bergen | Postboks 7800, NO-5020 Bergen</addrLine>
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            <ref type="url">https://www.uib.no/en</ref>
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          <idno type="ROR">https://ror.org/02kzqn938</idno>
          <idno type="Wikidata">Q3551621</idno>
          <orgName>Université de Lille</orgName>
          <desc>
            <address>
              <addrLine>EPE Université de Lille. -- 42 rue Paul Duez, 59000 Lille</addrLine>
              <country key="FR"/>
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            <ref type="url">https://www.univ-lille.fr/</ref>
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        <org type="institution" xml:id="struct-352867" status="VALID">
          <orgName>University Hospital Mannheim | Universitätsmedizin Mannheim</orgName>
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            <address>
              <country key="DE"/>
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            <ref type="url">https://www.umm.de/en</ref>
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        </org>
        <org type="laboratory" xml:id="struct-120233" status="VALID">
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          <orgName>Max Planck Institute for Psycholinguistics</orgName>
          <desc>
            <address>
              <addrLine>Wundtlaan 1 6525 XD Nijmegen The Netherlands</addrLine>
              <country key="NL"/>
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            <ref type="url">http://www.mpi.nl/</ref>
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        <org type="institution" xml:id="struct-5247" status="VALID">
          <orgName>Max-Planck-Gesellschaft</orgName>
          <desc>
            <address>
              <addrLine>Jägerstrasse, 10-11, D-10117 Berlin</addrLine>
              <country key="DE"/>
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            <ref type="url">https://www.mpg.de/en</ref>
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        </org>
        <org type="institution" xml:id="struct-309442" status="VALID">
          <idno type="ROR">https://ror.org/002pd6e78</idno>
          <orgName>Massachusetts General Hospital [Boston]</orgName>
          <desc>
            <address>
              <addrLine>55 Fruit StreetBoston, MA 02114</addrLine>
              <country key="US"/>
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            <ref type="url">http://www.massgeneral.org/</ref>
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        <org type="institution" xml:id="struct-300875" status="VALID">
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          <idno type="ROR">https://ror.org/02jx3x895</idno>
          <orgName>University College London [UCL]</orgName>
          <orgName type="acronym">UCL</orgName>
          <desc>
            <address>
              <addrLine>Gower Street, London WC1E 6BT</addrLine>
              <country key="GB"/>
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            <ref type="url">http://www.ucl.ac.uk/</ref>
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        <org type="regroupinstitution" xml:id="struct-566099" status="VALID">
          <orgName>Indiana University System</orgName>
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            <ref type="url">https://www.iu.edu/</ref>
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        </org>
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          <idno type="ROR">https://ror.org/016476m91</idno>
          <orgName>University of Aberdeen</orgName>
          <desc>
            <address>
              <addrLine>King's College, Aberdeen AB24 3FX</addrLine>
              <country key="GB"/>
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            <ref type="url">http://www.abdn.ac.uk/</ref>
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          <orgName>Medstar Research Institute</orgName>
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        <org type="institution" xml:id="struct-63690" status="VALID">
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          <orgName>Universität Bonn = University of Bonn</orgName>
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            <address>
              <addrLine>D-53012 Bonn</addrLine>
              <country key="DE"/>
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            <ref type="url">https://www.uni-bonn.de/</ref>
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        <org type="institution" xml:id="struct-19908" status="VALID">
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          <idno type="ROR">https://ror.org/01aj84f44</idno>
          <orgName>Aarhus University [Aarhus]</orgName>
          <desc>
            <address>
              <addrLine>Nordre Ringgade 1 DK-8000 Aarhus C</addrLine>
              <country key="DK"/>
            </address>
            <ref type="url">http://www.au.dk/en/</ref>
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        </org>
        <org type="institution" xml:id="struct-410432" status="VALID">
          <orgName>VU University Medical Center [Amsterdam]</orgName>
          <desc>
            <address>
              <addrLine>De Boelelaan 1118, 1081 HZ Amsterdam</addrLine>
              <country key="NL"/>
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            <ref type="url">http://www.vumc.com/</ref>
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        </org>
        <org type="institution" xml:id="struct-69530" status="VALID">
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          <idno type="ROR">https://ror.org/041kmwe10</idno>
          <orgName>Imperial College London</orgName>
          <date type="start">1907-01-01</date>
          <desc>
            <address>
              <addrLine>South Kensington Campus, London SW7 2AZ</addrLine>
              <country key="GB"/>
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            <ref type="url">http://www.imperial.ac.uk/</ref>
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        </org>
        <org type="regrouplaboratory" xml:id="struct-306744" status="INCOMING">
          <orgName>School of public health</orgName>
          <desc>
            <address>
              <country key="CN"/>
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          </desc>
          <listRelation>
            <relation active="#struct-93487" type="direct"/>
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        </org>
        <org type="institution" xml:id="struct-93487" status="VALID">
          <idno type="ROR">https://ror.org/02zhqgq86</idno>
          <orgName>The University of Hong Kong</orgName>
          <orgName type="acronym">HKU</orgName>
          <desc>
            <address>
              <addrLine>Pokfulam, Hong Kong</addrLine>
              <country key="HK"/>
            </address>
            <ref type="url">http://www.hku.hk/</ref>
          </desc>
        </org>
        <org type="institution" xml:id="struct-350388" status="INCOMING">
          <orgName>MRC-HPA Centre for Environment and Health</orgName>
          <desc>
            <address>
              <country key="GB"/>
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          </desc>
        </org>
        <org type="institution" xml:id="struct-93591" status="OLD">
          <idno type="ROR">https://ror.org/02en5vm52</idno>
          <orgName>Université Pierre et Marie Curie - Paris 6</orgName>
          <orgName type="acronym">UPMC</orgName>
          <date type="end">2017-12-31</date>
          <desc>
            <address>
              <addrLine>4 place Jussieu - 75005 Paris</addrLine>
              <country key="FR"/>
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            <ref type="url">http://www.upmc.fr/</ref>
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        <org type="institution" xml:id="struct-341038" status="VALID">
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          <idno type="ISNI">000000010626358X</idno>
          <idno type="ROR">https://ror.org/02cte4b68</idno>
          <orgName>Institut de Chimie - CNRS Chimie</orgName>
          <orgName type="acronym">INC-CNRS</orgName>
          <date type="start">2009-10-01</date>
          <desc>
            <address>
              <addrLine>CENTRE NATIONAL DE LA RECHERCHE SCIENTIFIQUE3 rue Michel-Ange75794 PARIS CEDEX 16 - France</addrLine>
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            <ref type="url">http://www.cnrs.fr/inc</ref>
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        <org type="institution" xml:id="struct-300563" status="VALID">
          <idno type="IdRef">026591790</idno>
          <idno type="ROR">https://ror.org/05f0yaq80</idno>
          <orgName>Stockholm University</orgName>
          <desc>
            <address>
              <addrLine>SE-106 91 Stockholm</addrLine>
              <country key="SE"/>
            </address>
            <ref type="url">http://www.su.se/english/</ref>
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        </org>
        <org type="institution" xml:id="struct-302344" status="VALID">
          <idno type="ROR">https://ror.org/056d84691</idno>
          <orgName>Karolinska Institutet = Karolinska Institute [Stockholm]</orgName>
          <desc>
            <address>
              <addrLine>Solnavägen 1, 171 77 SolnaStockholm</addrLine>
              <country key="SE"/>
            </address>
            <ref type="url">http://ki.se</ref>
          </desc>
        </org>
        <org type="regrouplaboratory" xml:id="struct-439680" status="VALID">
          <orgName>School of Medicine [Univ California San Diego]</orgName>
          <orgName type="acronym">UC San Diego</orgName>
          <desc>
            <address>
              <addrLine>University of California, San Diego | 9500 Gilman Dr, La Jolla, CA 92093</addrLine>
              <country key="US"/>
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            <ref type="url">https://medschool.ucsd.edu/</ref>
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          <idno type="ROR">https://ror.org/0168r3w48</idno>
          <orgName>University of California [San Diego]</orgName>
          <orgName type="acronym">UC San Diego</orgName>
          <desc>
            <address>
              <addrLine>UCSD, 9500 Gilman Dr., La Jolla, CA 92093 USA</addrLine>
              <country key="US"/>
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            <ref type="url">https://ucsd.edu/</ref>
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        <org type="institution" xml:id="struct-250752" status="VALID">
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          <idno type="ROR">https://ror.org/00rqy9422</idno>
          <idno type="Wikidata">Q866012</idno>
          <orgName>The University of Queensland</orgName>
          <orgName type="acronym">UQ [All campuses : Brisbane, Dutton Park Gatton, Herston, St Lucia and other locations]</orgName>
          <date type="start">1910-01-01</date>
          <desc>
            <address>
              <addrLine>Brisbane, St Lucia, QLD 4072</addrLine>
              <country key="AU"/>
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            <ref type="url">http://www.uq.edu.au/</ref>
          </desc>
        </org>
        <org type="regrouplaboratory" xml:id="struct-1003312" status="VALID">
          <orgName>Faculty of Social Sciences [Oslo]</orgName>
          <desc>
            <address>
              <addrLine>P.O box 1084 Blindern 0317 OSLO Norway</addrLine>
              <country key="NO"/>
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            <ref type="url">https://www.sv.uio.no/english/</ref>
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        <org type="institution" xml:id="struct-330962" status="INCOMING">
          <orgName>Deutsche Bundesbank</orgName>
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            <address>
              <country key="DE"/>
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        </org>
        <org type="institution" xml:id="struct-81095" status="VALID">
          <idno type="ROR">https://ror.org/01hxy9878</idno>
          <orgName>Royal College of Surgeons in Ireland</orgName>
          <orgName type="acronym">RCSI</orgName>
          <desc>
            <address>
              <addrLine>Royal College of Surgeons Ireland . 123 St Stephens Green . Dublin 2</addrLine>
              <country key="IE"/>
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            <ref type="url">http://www.rcsi.ie/</ref>
          </desc>
        </org>
        <org type="institution" xml:id="struct-337730" status="INCOMING">
          <orgName>Clinical And Experimental Epilepsy</orgName>
          <desc>
            <address>
              <country key="GB"/>
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          </desc>
        </org>
        <org type="institution" xml:id="struct-51214" status="VALID">
          <orgName>Yale School of Medicine [New Haven, Connecticut]</orgName>
          <orgName type="acronym">YSM</orgName>
          <desc>
            <address>
              <addrLine>Yale University School of Medicine - 333 Cedar Street - New Haven, CT 06510</addrLine>
              <country key="US"/>
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          </desc>
        </org>
        <org type="institution" xml:id="struct-339695" status="INCOMING">
          <orgName>Statistical Genetics Group</orgName>
          <desc>
            <address>
              <country key="DE"/>
            </address>
          </desc>
        </org>
        <org type="institution" xml:id="struct-424745" status="INCOMING">
          <orgName>WuXi AppTec, Co., Ltd</orgName>
          <desc>
            <address>
              <country key="CN"/>
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          </desc>
        </org>
        <org type="institution" xml:id="struct-1067803" status="VALID">
          <idno type="IdRef">154882658</idno>
          <idno type="ISNI">0000 0001 2191 4301</idno>
          <idno type="ROR">https://ror.org/05n0wgt02</idno>
          <idno type="Wikidata">Q6856599</idno>
          <orgName>King Faisal Specialist Hospital and Resarch Centre [Riyadh]</orgName>
          <orgName type="acronym">KFSHRC</orgName>
          <date type="start">1976-01-01</date>
          <desc>
            <address>
              <addrLine>Riyadh, Saudi Arabia</addrLine>
              <country key="SA"/>
            </address>
            <ref type="url">https://www.kfshrc.edu.sa/en/home</ref>
          </desc>
        </org>
        <org type="institution" xml:id="struct-346818" status="INCOMING">
          <orgName>CXR Biosciences Ltd</orgName>
          <desc>
            <address>
              <country key="FR"/>
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        </org>
        <org type="institution" xml:id="struct-300845" status="VALID">
          <idno type="ROR">https://ror.org/03h2bxq36</idno>
          <orgName>University of Dundee</orgName>
          <date type="start">2017-02-16</date>
          <desc>
            <address>
              <addrLine>University of DundeeNethergateDundeeScotland, UKDD1 4HN</addrLine>
              <country key="GB"/>
            </address>
            <ref type="url">https://www.dundee.ac.uk/</ref>
          </desc>
        </org>
        <org type="institution" xml:id="struct-305993" status="INCOMING">
          <orgName>Southwest Foundation for Biomedical Research</orgName>
          <desc>
            <address>
              <country key="US"/>
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          </desc>
        </org>
        <org type="institution" xml:id="struct-332777" status="INCOMING">
          <orgName>Department of Clinical Genetics</orgName>
          <desc>
            <address>
              <country key="AU"/>
            </address>
          </desc>
        </org>
        <org type="institution" xml:id="struct-351482" status="INCOMING">
          <orgName>Tehran University of Medical Siences</orgName>
          <desc>
            <address>
              <country key="IR"/>
            </address>
          </desc>
        </org>
        <org type="institution" xml:id="struct-343597" status="INCOMING">
          <orgName>Research Center Juelich</orgName>
          <desc>
            <address>
              <country key="DE"/>
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        </org>
        <org type="institution" xml:id="struct-149966" status="VALID">
          <idno type="ROR">https://ror.org/02s6k3f65</idno>
          <orgName>Université de Bâle = University of Basel = Basel Universität</orgName>
          <orgName type="acronym">Unibas</orgName>
          <desc>
            <address>
              <addrLine>Petersplatz 1, P. O. Box4001 Basel</addrLine>
              <country key="CH"/>
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            <ref type="url">https://www.unibas.ch/en.html</ref>
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        <org type="institution" xml:id="struct-333570" status="INCOMING">
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          <desc>
            <address>
              <country key="IE"/>
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        <org type="institution" xml:id="struct-237843" status="VALID">
          <idno type="IdRef">027706338</idno>
          <idno type="ROR">https://ror.org/013meh722</idno>
          <orgName>University of Cambridge [Cambridge, UK]</orgName>
          <orgName type="acronym">CAM</orgName>
          <desc>
            <address>
              <addrLine>The Old Schools, Trinity Lane, Cambridge CB2 1TN</addrLine>
              <country key="GB"/>
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            <ref type="url">http://www.cam.ac.uk/</ref>
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        <org type="institution" xml:id="struct-300572" status="VALID">
          <idno type="ROR">https://ror.org/04pp8hn57</idno>
          <orgName>Universiteit Utrecht / Utrecht University [Utrecht]</orgName>
          <desc>
            <address>
              <addrLine>Heidelberglaan 8, 3584 CS Utrecht</addrLine>
              <country key="NL"/>
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            <ref type="url">http://www.uu.nl/en</ref>
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        <org type="institution" xml:id="struct-341991" status="INCOMING">
          <orgName>Department of Genomics</orgName>
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              <country key="DE"/>
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        <org type="institution" xml:id="struct-346668" status="INCOMING">
          <orgName>Public Health Genomics Unit</orgName>
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              <country key="FI"/>
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        <org type="institution" xml:id="struct-162742" status="VALID">
          <idno type="ROR">https://ror.org/03cv38k47</idno>
          <orgName>University Medical Center Groningen [Groningen]</orgName>
          <orgName type="acronym">UMCG</orgName>
          <desc>
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              <addrLine>PO box 30.0019700 RB Groningen</addrLine>
              <country key="NL"/>
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            <ref type="url">https://www.umcg.nl/EN/</ref>
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        <org type="institution" xml:id="struct-347316" status="INCOMING">
          <orgName>Neuronal Plasticity / Mouse Behaviour</orgName>
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              <country key="DE"/>
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        <org type="institution" xml:id="struct-462070" status="VALID">
          <idno type="IdRef">235605212</idno>
          <idno type="ROR">https://ror.org/00r1edq15</idno>
          <orgName>Universität Greifswald = University of Greifswald</orgName>
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              <addrLine>Ernst-Moritz-Arndt- University of Greifswald, Domstraße 11, 17489 Greifswald</addrLine>
              <country key="DE"/>
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            <ref type="url">https://www.uni-greifswald.de/en/</ref>
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        <org type="institution" xml:id="struct-310910" status="VALID">
          <orgName>Mayo Clinic</orgName>
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        <org type="institution" xml:id="struct-302612" status="VALID">
          <idno type="ROR">https://ror.org/052gg0110</idno>
          <orgName>University of Oxford</orgName>
          <desc>
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              <addrLine>Wellington Square, Oxford OX1 2JD</addrLine>
              <country key="GB"/>
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            <ref type="url">http://www.ox.ac.uk/</ref>
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        <org type="institution" xml:id="struct-322053" status="INCOMING">
          <orgName>FMRIB Centre</orgName>
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              <country key="FR"/>
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        <org type="institution" xml:id="struct-331576" status="VALID">
          <orgName>John Radcliffe Hospital [Oxford University Hospital]</orgName>
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              <country key="US"/>
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          <orgName>University of Maryland [Baltimore County]</orgName>
          <orgName type="acronym">UMBC</orgName>
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            <address>
              <addrLine>1000 Hilltop Circle • Baltimore, MD 21250</addrLine>
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            <relation active="#struct-61400" type="direct"/>
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          <orgName>The Hong Kong Polytechnic University [Hong Kong]</orgName>
          <orgName type="acronym">POLYU</orgName>
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              <addrLine>Hung Hom, Kowloon, Hong Kong</addrLine>
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        <org type="institution" xml:id="struct-333179" status="INCOMING">
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        <org type="institution" xml:id="struct-468263" status="VALID">
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          <orgName>Oslo University Hospital [Oslo]</orgName>
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            <address>
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              <addrLine>Coventry CV4 7AL</addrLine>
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            <address>
              <addrLine>1-1 Yamadaoka, Suita,Osaka 565-0871 / Formerly known as Osaka University</addrLine>
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          <orgName>Martin-Luther-Universität Halle Wittenberg - Martin-Luther-University Halle Wittenberg</orgName>
          <orgName type="acronym">MLU</orgName>
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            <address>
              <addrLine>06108 Halle (Saale), Allemagne</addrLine>
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            <address>
              <addrLine>Place Eugène Bataillon - 34095 Montpellier cedex 5</addrLine>
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        <org type="institution" xml:id="struct-301759" status="VALID">
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        </org>
        <org type="institution" xml:id="struct-300713" status="VALID">
          <orgName>Macquarie University</orgName>
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            <address>
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            <ref type="url">http://www.mq.edu.au</ref>
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        </org>
        <org type="regrouplaboratory" xml:id="struct-343852" status="INCOMING">
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          <orgName>New Mexico Consortium</orgName>
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