Genetic variants linked to education predict longevity
Identifieur interne : 003E04 ( Ncbi/Merge ); précédent : 003E03; suivant : 003E05Genetic variants linked to education predict longevity
Auteurs : Riccardo E. Marioni [Royaume-Uni, Australie] ; Stuart J. Ritchie [Royaume-Uni] ; Peter K. Joshi [Royaume-Uni] ; Saskia P. Hagenaars [Royaume-Uni] ; Aysu Okbay [Pays-Bas] ; Krista Fischer [Estonie] ; Mark J. Adams [Royaume-Uni] ; W. David Hill [Royaume-Uni] ; Gail Davies [Royaume-Uni] ; Reka Nagy [Royaume-Uni] ; Carmen Amador [Royaume-Uni] ; Kristi L Ll [Estonie] ; Andres Metspalu [Estonie] ; David C. Liewald [Royaume-Uni] ; Archie Campbell [Royaume-Uni] ; James F. Wilson [Royaume-Uni] ; Caroline Hayward [Royaume-Uni] ; T Nu Esko [Estonie] ; David J. Porteous [Royaume-Uni] ; Catharine R. Gale [Royaume-Uni] ; Ian J. Deary [Royaume-Uni]Source :
- Proceedings of the National Academy of Sciences of the United States of America [ 0027-8424 ] ; 2016.
Abstract
Individuals with more education tend to live longer. Genetic variants have been discovered that predict educational attainment. We tested whether a “polygenic score” based on these genetic variants could make predictions about people’s lifespan. We used data from three cohort studies (including >130,000 participants) to examine the link between offspring polygenic score for education and parental longevity. Across the studies, we found that participants with more education-linked genetic variants had longer-living parents; compared with those with the lowest genetic education scores, those with the highest scores had parents who lived on average 6 months longer. This finding suggests the hypothesis that part of the ultimate explanation for the extended longevity of better-educated people is an underlying, quantifiable, genetic propensity.
Url:
DOI: 10.1073/pnas.1605334113
PubMed: 27799538
PubMed Central: 5127357
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<affiliation wicri:level="1"><nlm:aff id="aff199">Institute of Mathematical Statistics,<institution>University of Tartu</institution>
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<affiliation><nlm:aff id="aff200"><institution>Broad Institute</institution>
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<author><name sortKey="Porteous, David J" sort="Porteous, David J" uniqKey="Porteous D" first="David J." last="Porteous">David J. Porteous</name>
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<affiliation wicri:level="1"><nlm:aff id="aff191">Department of Psychology,<institution>University of Edinburgh</institution>
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<author><name sortKey="Hagenaars, Saskia P" sort="Hagenaars, Saskia P" uniqKey="Hagenaars S" first="Saskia P." last="Hagenaars">Saskia P. Hagenaars</name>
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<country xml:lang="fr">Royaume-Uni</country>
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, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
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, Edinburgh EH8 9YL,<country>United Kingdom</country>
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</author>
<author><name sortKey="Okbay, Aysu" sort="Okbay, Aysu" uniqKey="Okbay A" first="Aysu" last="Okbay">Aysu Okbay</name>
<affiliation wicri:level="1"><nlm:aff id="aff194">Department of Applied Economics, Erasmus School of Economics,<institution>Erasmus University</institution>
, 3062 PA Rotterdam,<country>The Netherlands</country>
;</nlm:aff>
<country xml:lang="fr">Pays-Bas</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff195">Department of Epidemiology,<institution>Erasmus Medical Center</institution>
, 3015 CE Rotterdam,<country>The Netherlands</country>
;</nlm:aff>
<country xml:lang="fr">Pays-Bas</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff196"><institution>Erasmus University Rotterdam Institute for Behavior and Biology</institution>
, Rotterdam 3062 PA,<country>The Netherlands</country>
;</nlm:aff>
<country xml:lang="fr">Pays-Bas</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Fischer, Krista" sort="Fischer, Krista" uniqKey="Fischer K" first="Krista" last="Fischer">Krista Fischer</name>
<affiliation wicri:level="1"><nlm:aff id="aff197">Estonian Genome Center,<institution>University of Tartu</institution>
, Tartu 51010,<country>Estonia</country>
;</nlm:aff>
<country xml:lang="fr">Estonie</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Adams, Mark J" sort="Adams, Mark J" uniqKey="Adams M" first="Mark J." last="Adams">Mark J. Adams</name>
<affiliation wicri:level="1"><nlm:aff id="aff193">Division of Psychiatry,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9YL,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Hill, W David" sort="Hill, W David" uniqKey="Hill W" first="W. David" last="Hill">W. David Hill</name>
<affiliation wicri:level="1"><nlm:aff id="aff188">Centre for Cognitive Ageing and Cognitive Epidemiology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff191">Department of Psychology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Davies, Gail" sort="Davies, Gail" uniqKey="Davies G" first="Gail" last="Davies">Gail Davies</name>
<affiliation wicri:level="1"><nlm:aff id="aff188">Centre for Cognitive Ageing and Cognitive Epidemiology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff191">Department of Psychology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Nagy, Reka" sort="Nagy, Reka" uniqKey="Nagy R" first="Reka" last="Nagy">Reka Nagy</name>
<affiliation wicri:level="1"><nlm:aff id="aff198">Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine,<institution>University of Edinburgh</institution>
, Edinburgh EH4 2XU,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Amador, Carmen" sort="Amador, Carmen" uniqKey="Amador C" first="Carmen" last="Amador">Carmen Amador</name>
<affiliation wicri:level="1"><nlm:aff id="aff198">Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine,<institution>University of Edinburgh</institution>
, Edinburgh EH4 2XU,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="L Ll, Kristi" sort="L Ll, Kristi" uniqKey="L Ll K" first="Kristi" last="L Ll">Kristi L Ll</name>
<affiliation wicri:level="1"><nlm:aff id="aff197">Estonian Genome Center,<institution>University of Tartu</institution>
, Tartu 51010,<country>Estonia</country>
;</nlm:aff>
<country xml:lang="fr">Estonie</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff199">Institute of Mathematical Statistics,<institution>University of Tartu</institution>
, Tartu 50409,<country>Estonia</country>
;</nlm:aff>
<country xml:lang="fr">Estonie</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Metspalu, Andres" sort="Metspalu, Andres" uniqKey="Metspalu A" first="Andres" last="Metspalu">Andres Metspalu</name>
<affiliation wicri:level="1"><nlm:aff id="aff197">Estonian Genome Center,<institution>University of Tartu</institution>
, Tartu 51010,<country>Estonia</country>
;</nlm:aff>
<country xml:lang="fr">Estonie</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Liewald, David C" sort="Liewald, David C" uniqKey="Liewald D" first="David C." last="Liewald">David C. Liewald</name>
<affiliation wicri:level="1"><nlm:aff id="aff188">Centre for Cognitive Ageing and Cognitive Epidemiology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff191">Department of Psychology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Campbell, Archie" sort="Campbell, Archie" uniqKey="Campbell A" first="Archie" last="Campbell">Archie Campbell</name>
<affiliation wicri:level="1"><nlm:aff id="aff189">Medical Genetics Section, Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine,<institution>University of Edinburgh</institution>
, Edinburgh EH4 2XU,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Wilson, James F" sort="Wilson, James F" uniqKey="Wilson J" first="James F." last="Wilson">James F. Wilson</name>
<affiliation wicri:level="1"><nlm:aff id="aff192">Usher Institute for Population Health Sciences and Informatics,<institution>University of Edinburgh</institution>
, Edinburgh EH16 4UX,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff198">Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine,<institution>University of Edinburgh</institution>
, Edinburgh EH4 2XU,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Hayward, Caroline" sort="Hayward, Caroline" uniqKey="Hayward C" first="Caroline" last="Hayward">Caroline Hayward</name>
<affiliation wicri:level="1"><nlm:aff id="aff198">Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine,<institution>University of Edinburgh</institution>
, Edinburgh EH4 2XU,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Esko, T Nu" sort="Esko, T Nu" uniqKey="Esko T" first="T Nu" last="Esko">T Nu Esko</name>
<affiliation wicri:level="1"><nlm:aff id="aff197">Estonian Genome Center,<institution>University of Tartu</institution>
, Tartu 51010,<country>Estonia</country>
;</nlm:aff>
<country xml:lang="fr">Estonie</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff199">Institute of Mathematical Statistics,<institution>University of Tartu</institution>
, Tartu 50409,<country>Estonia</country>
;</nlm:aff>
<country xml:lang="fr">Estonie</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation><nlm:aff id="aff200"><institution>Broad Institute</institution>
, Cambridge,<addr-line>MA</addr-line>
02142;</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="aff201">Department of Endocrinology,<institution>Children’s Hospital Boston</institution>
, Boston,<addr-line>MA</addr-line>
02115;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Porteous, David J" sort="Porteous, David J" uniqKey="Porteous D" first="David J." last="Porteous">David J. Porteous</name>
<affiliation wicri:level="1"><nlm:aff id="aff188">Centre for Cognitive Ageing and Cognitive Epidemiology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff189">Medical Genetics Section, Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine,<institution>University of Edinburgh</institution>
, Edinburgh EH4 2XU,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Gale, Catharine R" sort="Gale, Catharine R" uniqKey="Gale C" first="Catharine R." last="Gale">Catharine R. Gale</name>
<affiliation wicri:level="1"><nlm:aff id="aff188">Centre for Cognitive Ageing and Cognitive Epidemiology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff191">Department of Psychology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff202">Medical Research Council Lifecourse Epidemiology Unit,<institution>University of Southampton</institution>
, Southampton SO17 1BJ,<country>United Kingdom</country>
</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Deary, Ian J" sort="Deary, Ian J" uniqKey="Deary I" first="Ian J." last="Deary">Ian J. Deary</name>
<affiliation wicri:level="1"><nlm:aff id="aff188">Centre for Cognitive Ageing and Cognitive Epidemiology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="aff191">Department of Psychology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</nlm:aff>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
</author>
</analytic>
<series><title level="j">Proceedings of the National Academy of Sciences of the United States of America</title>
<idno type="ISSN">0027-8424</idno>
<idno type="eISSN">1091-6490</idno>
<imprint><date when="2016">2016</date>
</imprint>
</series>
</biblStruct>
</sourceDesc>
</fileDesc>
<profileDesc><textClass></textClass>
</profileDesc>
</teiHeader>
<front><div type="abstract" xml:lang="en"><title>Significance</title>
<p>Individuals with more education tend to live longer. Genetic variants have been discovered that predict educational attainment. We tested whether a “polygenic score” based on these genetic variants could make predictions about people’s lifespan. We used data from three cohort studies (including >130,000 participants) to examine the link between offspring polygenic score for education and parental longevity. Across the studies, we found that participants with more education-linked genetic variants had longer-living parents; compared with those with the lowest genetic education scores, those with the highest scores had parents who lived on average 6 months longer. This finding suggests the hypothesis that part of the ultimate explanation for the extended longevity of better-educated people is an underlying, quantifiable, genetic propensity.</p>
</div>
</front>
</TEI>
<pmc article-type="research-article"><pmc-comment>The publisher of this article does not allow downloading of the full text in XML form.</pmc-comment>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Proc Natl Acad Sci U S A</journal-id>
<journal-id journal-id-type="iso-abbrev">Proc. Natl. Acad. Sci. U.S.A</journal-id>
<journal-id journal-id-type="hwp">pnas</journal-id>
<journal-id journal-id-type="pmc">pnas</journal-id>
<journal-id journal-id-type="publisher-id">PNAS</journal-id>
<journal-title-group><journal-title>Proceedings of the National Academy of Sciences of the United States of America</journal-title>
</journal-title-group>
<issn pub-type="ppub">0027-8424</issn>
<issn pub-type="epub">1091-6490</issn>
<publisher><publisher-name>National Academy of Sciences</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">27799538</article-id>
<article-id pub-id-type="pmc">5127357</article-id>
<article-id pub-id-type="publisher-id">201605334</article-id>
<article-id pub-id-type="doi">10.1073/pnas.1605334113</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Social Sciences</subject>
<subj-group><subject>Social Sciences</subject>
</subj-group>
</subj-group>
<subj-group subj-group-type="heading"><subject>Biological Sciences</subject>
<subj-group><subject>Genetics</subject>
</subj-group>
</subj-group>
<series-title>From the Cover</series-title>
</article-categories>
<title-group><article-title>Genetic variants linked to education predict longevity</article-title>
<alt-title alt-title-type="short">Education-linked variants and longevity</alt-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Marioni</surname>
<given-names>Riccardo E.</given-names>
</name>
<xref ref-type="aff" rid="aff188"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff189"><sup>b</sup>
</xref>
<xref ref-type="aff" rid="aff190"><sup>c</sup>
</xref>
<xref ref-type="author-notes" rid="fn1"><sup>1</sup>
</xref>
<xref ref-type="corresp" rid="cor1"><sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Ritchie</surname>
<given-names>Stuart J.</given-names>
</name>
<xref ref-type="aff" rid="aff188"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff191"><sup>d</sup>
</xref>
<xref ref-type="author-notes" rid="fn1"><sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Joshi</surname>
<given-names>Peter K.</given-names>
</name>
<xref ref-type="aff" rid="aff192"><sup>e</sup>
</xref>
<xref ref-type="author-notes" rid="fn1"><sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Hagenaars</surname>
<given-names>Saskia P.</given-names>
</name>
<xref ref-type="aff" rid="aff188"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff191"><sup>d</sup>
</xref>
<xref ref-type="aff" rid="aff193"><sup>f</sup>
</xref>
</contrib>
<contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="false">http://orcid.org/0000-0002-5170-7781</contrib-id>
<name><surname>Okbay</surname>
<given-names>Aysu</given-names>
</name>
<xref ref-type="aff" rid="aff194"><sup>g</sup>
</xref>
<xref ref-type="aff" rid="aff195"><sup>h</sup>
</xref>
<xref ref-type="aff" rid="aff196"><sup>i</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Fischer</surname>
<given-names>Krista</given-names>
</name>
<xref ref-type="aff" rid="aff197"><sup>j</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Adams</surname>
<given-names>Mark J.</given-names>
</name>
<xref ref-type="aff" rid="aff193"><sup>f</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Hill</surname>
<given-names>W. David</given-names>
</name>
<xref ref-type="aff" rid="aff188"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff191"><sup>d</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Davies</surname>
<given-names>Gail</given-names>
</name>
<xref ref-type="aff" rid="aff188"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff191"><sup>d</sup>
</xref>
</contrib>
<contrib contrib-type="author"><collab>Social Science Genetic Association Consortium</collab>
<xref ref-type="fn" rid="fn2"><sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Nagy</surname>
<given-names>Reka</given-names>
</name>
<xref ref-type="aff" rid="aff198"><sup>k</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Amador</surname>
<given-names>Carmen</given-names>
</name>
<xref ref-type="aff" rid="aff198"><sup>k</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Läll</surname>
<given-names>Kristi</given-names>
</name>
<xref ref-type="aff" rid="aff197"><sup>j</sup>
</xref>
<xref ref-type="aff" rid="aff199"><sup>l</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Metspalu</surname>
<given-names>Andres</given-names>
</name>
<xref ref-type="aff" rid="aff197"><sup>j</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Liewald</surname>
<given-names>David C.</given-names>
</name>
<xref ref-type="aff" rid="aff188"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff191"><sup>d</sup>
</xref>
</contrib>
<contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="false">http://orcid.org/0000-0003-0198-5078</contrib-id>
<name><surname>Campbell</surname>
<given-names>Archie</given-names>
</name>
<xref ref-type="aff" rid="aff189"><sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wilson</surname>
<given-names>James F.</given-names>
</name>
<xref ref-type="aff" rid="aff192"><sup>e</sup>
</xref>
<xref ref-type="aff" rid="aff198"><sup>k</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Hayward</surname>
<given-names>Caroline</given-names>
</name>
<xref ref-type="aff" rid="aff198"><sup>k</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Esko</surname>
<given-names>Tõnu</given-names>
</name>
<xref ref-type="aff" rid="aff197"><sup>j</sup>
</xref>
<xref ref-type="aff" rid="aff199"><sup>l</sup>
</xref>
<xref ref-type="aff" rid="aff200"><sup>m</sup>
</xref>
<xref ref-type="aff" rid="aff201"><sup>n</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Porteous</surname>
<given-names>David J.</given-names>
</name>
<xref ref-type="aff" rid="aff188"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff189"><sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Gale</surname>
<given-names>Catharine R.</given-names>
</name>
<xref ref-type="aff" rid="aff188"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff191"><sup>d</sup>
</xref>
<xref ref-type="aff" rid="aff202"><sup>o</sup>
</xref>
<xref ref-type="author-notes" rid="fn3"><sup>4</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Deary</surname>
<given-names>Ian J.</given-names>
</name>
<xref ref-type="aff" rid="aff188"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff191"><sup>d</sup>
</xref>
<xref ref-type="author-notes" rid="fn3"><sup>4</sup>
</xref>
</contrib>
<aff id="aff188"><sup>a</sup>
Centre for Cognitive Ageing and Cognitive Epidemiology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</aff>
<aff id="aff189"><sup>b</sup>
Medical Genetics Section, Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine,<institution>University of Edinburgh</institution>
, Edinburgh EH4 2XU,<country>United Kingdom</country>
;</aff>
<aff id="aff190"><sup>c</sup>
Institute for Molecular Bioscience,<institution>University of Queensland</institution>
, Brisbane, QLD 4072,<country>Australia</country>
;</aff>
<aff id="aff191"><sup>d</sup>
Department of Psychology,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9JZ,<country>United Kingdom</country>
;</aff>
<aff id="aff192"><sup>e</sup>
Usher Institute for Population Health Sciences and Informatics,<institution>University of Edinburgh</institution>
, Edinburgh EH16 4UX,<country>United Kingdom</country>
;</aff>
<aff id="aff193"><sup>f</sup>
Division of Psychiatry,<institution>University of Edinburgh</institution>
, Edinburgh EH8 9YL,<country>United Kingdom</country>
;</aff>
<aff id="aff194"><sup>g</sup>
Department of Applied Economics, Erasmus School of Economics,<institution>Erasmus University</institution>
, 3062 PA Rotterdam,<country>The Netherlands</country>
;</aff>
<aff id="aff195"><sup>h</sup>
Department of Epidemiology,<institution>Erasmus Medical Center</institution>
, 3015 CE Rotterdam,<country>The Netherlands</country>
;</aff>
<aff id="aff196"><sup>i</sup>
<institution>Erasmus University Rotterdam Institute for Behavior and Biology</institution>
, Rotterdam 3062 PA,<country>The Netherlands</country>
;</aff>
<aff id="aff197"><sup>j</sup>
Estonian Genome Center,<institution>University of Tartu</institution>
, Tartu 51010,<country>Estonia</country>
;</aff>
<aff id="aff198"><sup>k</sup>
Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine,<institution>University of Edinburgh</institution>
, Edinburgh EH4 2XU,<country>United Kingdom</country>
;</aff>
<aff id="aff199"><sup>l</sup>
Institute of Mathematical Statistics,<institution>University of Tartu</institution>
, Tartu 50409,<country>Estonia</country>
;</aff>
<aff id="aff200"><sup>m</sup>
<institution>Broad Institute</institution>
, Cambridge,<addr-line>MA</addr-line>
02142;</aff>
<aff id="aff201"><sup>n</sup>
Department of Endocrinology,<institution>Children’s Hospital Boston</institution>
, Boston,<addr-line>MA</addr-line>
02115;</aff>
<aff id="aff202"><sup>o</sup>
Medical Research Council Lifecourse Epidemiology Unit,<institution>University of Southampton</institution>
, Southampton SO17 1BJ,<country>United Kingdom</country>
</aff>
</contrib-group>
<contrib-group><contrib contrib-type="collab"><name><surname>Okbay</surname>
<given-names>Aysu</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Beauchamp</surname>
<given-names>Jonathan P.</given-names>
</name>
<xref ref-type="aff" rid="aff8"><sup>8</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Fontana</surname>
<given-names>Mark A.</given-names>
</name>
<xref ref-type="aff" rid="aff4"><sup>4</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Lee</surname>
<given-names>James J.</given-names>
</name>
<xref ref-type="aff" rid="aff5"><sup>5</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Pers</surname>
<given-names>Tune H.</given-names>
</name>
<xref ref-type="aff" rid="aff6"><sup>6</sup>
</xref>
<xref ref-type="aff" rid="aff7"><sup>7</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Rietveld</surname>
<given-names>Cornelius A.</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Turley</surname>
<given-names>Patrick</given-names>
</name>
<xref ref-type="aff" rid="aff8"><sup>8</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Chen181</surname>
<given-names>Guo-Bo</given-names>
</name>
</contrib>
<contrib contrib-type="collab"><name><surname>Emilsson</surname>
<given-names>Valur</given-names>
</name>
<xref ref-type="aff" rid="aff9"><sup>9</sup>
</xref>
<xref ref-type="aff" rid="aff10"><sup>10</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Meddens</surname>
<given-names>S. Fleur W.</given-names>
</name>
<xref ref-type="aff" rid="aff43"><sup>43</sup>
</xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup>
</xref>
<xref ref-type="aff" rid="aff11"><sup>11</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Oskarsson</surname>
<given-names>Sven</given-names>
</name>
<xref ref-type="aff" rid="aff12"><sup>12</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Pickrell</surname>
<given-names>Joseph K.</given-names>
</name>
<xref ref-type="aff" rid="aff13"><sup>13</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Thom</surname>
<given-names>Kevin</given-names>
</name>
<xref ref-type="aff" rid="aff14"><sup>14</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Timshel</surname>
<given-names>Pascal</given-names>
</name>
<xref ref-type="aff" rid="aff15"><sup>15</sup>
</xref>
<xref ref-type="aff" rid="aff16"><sup>16</sup>
</xref>
<xref ref-type="aff" rid="aff7"><sup>7</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>de Vlaming</surname>
<given-names>Ronald</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Abdellaoui</surname>
<given-names>Abdel</given-names>
</name>
<xref ref-type="aff" rid="aff17"><sup>17</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Ahluwalia</surname>
<given-names>Tarunveer S.</given-names>
</name>
<xref ref-type="aff" rid="aff18"><sup>18</sup>
</xref>
<xref ref-type="aff" rid="aff15"><sup>15</sup>
</xref>
<xref ref-type="aff" rid="aff19"><sup>19</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Bacelis</surname>
<given-names>Jonas</given-names>
</name>
<xref ref-type="aff" rid="aff20"><sup>20</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Baumbach</surname>
<given-names>Clemens</given-names>
</name>
<xref ref-type="aff" rid="aff21"><sup>21</sup>
</xref>
<xref ref-type="aff" rid="aff22"><sup>22</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Bjornsdottir</surname>
<given-names>Gyda</given-names>
</name>
<xref ref-type="aff" rid="aff93"><sup>93</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Brandsma</surname>
<given-names>Johannes H.</given-names>
</name>
<xref ref-type="aff" rid="aff23"><sup>23</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Concas</surname>
<given-names>Maria Pina</given-names>
</name>
<xref ref-type="aff" rid="aff24"><sup>24</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Derringer</surname>
<given-names>Jaime</given-names>
</name>
<xref ref-type="aff" rid="aff25"><sup>25</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Furlotte</surname>
<given-names>Nicholas A.</given-names>
</name>
<xref ref-type="aff" rid="aff26"><sup>26</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Galesloot</surname>
<given-names>Tessel E.</given-names>
</name>
<xref ref-type="aff" rid="aff27"><sup>27</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Girotto</surname>
<given-names>Giorgia</given-names>
</name>
<xref ref-type="aff" rid="aff28"><sup>28</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Gupta</surname>
<given-names>Richa</given-names>
</name>
<xref ref-type="aff" rid="aff29"><sup>29</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Hall</surname>
<given-names>Leanne M.</given-names>
</name>
<xref ref-type="aff" rid="aff30"><sup>30</sup>
</xref>
<xref ref-type="aff" rid="aff31"><sup>31</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Harris</surname>
<given-names>Sarah E.</given-names>
</name>
<xref ref-type="aff" rid="aff32"><sup>32</sup>
</xref>
<xref ref-type="aff" rid="aff33"><sup>33</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Hofer</surname>
<given-names>Edith</given-names>
</name>
<xref ref-type="aff" rid="aff34"><sup>34</sup>
</xref>
<xref ref-type="aff" rid="aff35"><sup>35</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Horikoshi</surname>
<given-names>Momoko</given-names>
</name>
<xref ref-type="aff" rid="aff36"><sup>36</sup>
</xref>
<xref ref-type="aff" rid="aff37"><sup>37</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Huffman</surname>
<given-names>Jennifer E.</given-names>
</name>
<xref ref-type="aff" rid="aff38"><sup>38</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kaasik</surname>
<given-names>Kadri</given-names>
</name>
<xref ref-type="aff" rid="aff39"><sup>39</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kalafati</surname>
<given-names>Ioanna P.</given-names>
</name>
<xref ref-type="aff" rid="aff40"><sup>40</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Karlsson</surname>
<given-names>Robert</given-names>
</name>
<xref ref-type="aff" rid="aff41"><sup>41</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kong</surname>
<given-names>Augustine</given-names>
</name>
<xref ref-type="aff" rid="aff28"><sup>28</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Lahti</surname>
<given-names>Jari</given-names>
</name>
<xref ref-type="aff" rid="aff39"><sup>39</sup>
</xref>
<xref ref-type="aff" rid="aff42"><sup>42</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>van der Lee</surname>
<given-names>Sven J.</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>de Leeuw</surname>
<given-names>Christiaan</given-names>
</name>
<xref ref-type="aff" rid="aff43"><sup>43</sup>
</xref>
<xref ref-type="aff" rid="aff44"><sup>44</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Lind</surname>
<given-names>Penelope A.</given-names>
</name>
<xref ref-type="aff" rid="aff45"><sup>45</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Lindgren</surname>
<given-names>Karl-Oskar</given-names>
</name>
<xref ref-type="aff" rid="aff12"><sup>12</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Liu</surname>
<given-names>Tian</given-names>
</name>
<xref ref-type="aff" rid="aff46"><sup>46</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Mangino</surname>
<given-names>Massimo</given-names>
</name>
<xref ref-type="aff" rid="aff47"><sup>47</sup>
</xref>
<xref ref-type="aff" rid="aff48"><sup>48</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Marten</surname>
<given-names>Jonathan</given-names>
</name>
<xref ref-type="aff" rid="aff38"><sup>38</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Mihailov</surname>
<given-names>Evelin</given-names>
</name>
<xref ref-type="aff" rid="aff109"><sup>109</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Miller</surname>
<given-names>Michael B.</given-names>
</name>
<xref ref-type="aff" rid="aff5"><sup>5</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>van der Most</surname>
<given-names>Peter J.</given-names>
</name>
<xref ref-type="aff" rid="aff49"><sup>49</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Oldmeadow</surname>
<given-names>Christopher</given-names>
</name>
<xref ref-type="aff" rid="aff50"><sup>50</sup>
</xref>
<xref ref-type="aff" rid="aff51"><sup>51</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Payton</surname>
<given-names>Antony</given-names>
</name>
<xref ref-type="aff" rid="aff52"><sup>52</sup>
</xref>
<xref ref-type="aff" rid="aff53"><sup>53</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Pervjakova</surname>
<given-names>Natalia</given-names>
</name>
<xref ref-type="aff" rid="aff54"><sup>54</sup>
</xref>
<xref ref-type="aff" rid="aff109"><sup>109</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Peyrot</surname>
<given-names>Wouter J.</given-names>
</name>
<xref ref-type="aff" rid="aff55"><sup>55</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Qian</surname>
<given-names>Yong</given-names>
</name>
<xref ref-type="aff" rid="aff56"><sup>56</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Raitakari</surname>
<given-names>Olli</given-names>
</name>
<xref ref-type="aff" rid="aff57"><sup>57</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Rueedi</surname>
<given-names>Rico</given-names>
</name>
<xref ref-type="aff" rid="aff58"><sup>58</sup>
</xref>
<xref ref-type="aff" rid="aff59"><sup>59</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Salvi</surname>
<given-names>Erika</given-names>
</name>
<xref ref-type="aff" rid="aff60"><sup>60</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Schmidt</surname>
<given-names>Börge</given-names>
</name>
<xref ref-type="aff" rid="aff61"><sup>61</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Schraut</surname>
<given-names>Katharina E.</given-names>
</name>
<xref ref-type="aff" rid="aff62"><sup>62</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Shi</surname>
<given-names>Jianxin</given-names>
</name>
<xref ref-type="aff" rid="aff63"><sup>63</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Smith</surname>
<given-names>Albert V.</given-names>
</name>
<xref ref-type="aff" rid="aff64"><sup>64</sup>
</xref>
<xref ref-type="aff" rid="aff65"><sup>65</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Poot</surname>
<given-names>Raymond A.</given-names>
</name>
<xref ref-type="aff" rid="aff23"><sup>23</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>St Pourcain</surname>
<given-names>Beate</given-names>
</name>
<xref ref-type="aff" rid="aff66"><sup>66</sup>
</xref>
<xref ref-type="aff" rid="aff67"><sup>67</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Teumer</surname>
<given-names>Alexander</given-names>
</name>
<xref ref-type="aff" rid="aff68"><sup>68</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Thorleifsson</surname>
<given-names>Gudmar</given-names>
</name>
<xref ref-type="aff" rid="aff28"><sup>28</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Verweij</surname>
<given-names>Niek</given-names>
</name>
<xref ref-type="aff" rid="aff69"><sup>69</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Vuckovic</surname>
<given-names>Dragana</given-names>
</name>
<xref ref-type="aff" rid="aff28"><sup>28</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Wellmann</surname>
<given-names>Juergen</given-names>
</name>
<xref ref-type="aff" rid="aff70"><sup>70</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Westra</surname>
<given-names>Harm-Jan</given-names>
</name>
<xref ref-type="aff" rid="aff71"><sup>71</sup>
</xref>
<xref ref-type="aff" rid="aff72"><sup>72</sup>
</xref>
<xref ref-type="aff" rid="aff7"><sup>7</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Yang</surname>
<given-names>Jingyun</given-names>
</name>
<xref ref-type="aff" rid="aff73"><sup>73</sup>
</xref>
<xref ref-type="aff" rid="aff74"><sup>74</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Zhao</surname>
<given-names>Wei</given-names>
</name>
<xref ref-type="aff" rid="aff75"><sup>75</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Zhu</surname>
<given-names>Zhihong</given-names>
</name>
<xref ref-type="aff" rid="aff181"><sup>181</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Alizadeh</surname>
<given-names>Behrooz Z.</given-names>
</name>
<xref ref-type="aff" rid="aff49"><sup>49</sup>
</xref>
<xref ref-type="aff" rid="aff76"><sup>76</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Amin</surname>
<given-names>Najaf</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Bakshi</surname>
<given-names>Andrew</given-names>
</name>
<xref ref-type="aff" rid="aff181"><sup>181</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Baumeister</surname>
<given-names>Sebastian E.</given-names>
</name>
<xref ref-type="aff" rid="aff67"><sup>67</sup>
</xref>
<xref ref-type="aff" rid="aff77"><sup>77</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Biino</surname>
<given-names>Ginevra</given-names>
</name>
<xref ref-type="aff" rid="aff78"><sup>78</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Bønnelykke</surname>
<given-names>Klaus</given-names>
</name>
<xref ref-type="aff" rid="aff18"><sup>18</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Boyle</surname>
<given-names>Patricia A.</given-names>
</name>
<xref ref-type="aff" rid="aff72"><sup>72</sup>
</xref>
<xref ref-type="aff" rid="aff79"><sup>79</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Campbell</surname>
<given-names>Harry</given-names>
</name>
<xref ref-type="aff" rid="aff61"><sup>61</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Cappuccio</surname>
<given-names>Francesco P.</given-names>
</name>
<xref ref-type="aff" rid="aff80"><sup>80</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Davies</surname>
<given-names>Gail</given-names>
</name>
<xref ref-type="aff" rid="aff32"><sup>32</sup>
</xref>
<xref ref-type="aff" rid="aff81"><sup>81</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>De Neve</surname>
<given-names>Jan-Emmanuel</given-names>
</name>
<xref ref-type="aff" rid="aff82"><sup>82</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Deloukas</surname>
<given-names>Panos</given-names>
</name>
<xref ref-type="aff" rid="aff83"><sup>83</sup>
</xref>
<xref ref-type="aff" rid="aff84"><sup>84</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Demuth</surname>
<given-names>Ilja</given-names>
</name>
<xref ref-type="aff" rid="aff85"><sup>85</sup>
</xref>
<xref ref-type="aff" rid="aff86"><sup>86</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Ding</surname>
<given-names>Jun</given-names>
</name>
<xref ref-type="aff" rid="aff56"><sup>56</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Eibich</surname>
<given-names>Peter</given-names>
</name>
<xref ref-type="aff" rid="aff87"><sup>87</sup>
</xref>
<xref ref-type="aff" rid="aff88"><sup>88</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Eisele</surname>
<given-names>Lewin</given-names>
</name>
<xref ref-type="aff" rid="aff60"><sup>60</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Eklund</surname>
<given-names>Niina</given-names>
</name>
<xref ref-type="aff" rid="aff53"><sup>53</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Evans</surname>
<given-names>David M.</given-names>
</name>
<xref ref-type="aff" rid="aff66"><sup>66</sup>
</xref>
<xref ref-type="aff" rid="aff182"><sup>182</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Faul</surname>
<given-names>Jessica D.</given-names>
</name>
<xref ref-type="aff" rid="aff89"><sup>89</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Feitosa</surname>
<given-names>Mary F.</given-names>
</name>
<xref ref-type="aff" rid="aff90"><sup>90</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Forstner</surname>
<given-names>Andreas J.</given-names>
</name>
<xref ref-type="aff" rid="aff91"><sup>91</sup>
</xref>
<xref ref-type="aff" rid="aff92"><sup>92</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Gandin</surname>
<given-names>Ilaria</given-names>
</name>
<xref ref-type="aff" rid="aff28"><sup>28</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Gunnarsson</surname>
<given-names>Bjarni</given-names>
</name>
<xref ref-type="aff" rid="aff93"><sup>93</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Halldórsson</surname>
<given-names>Bjarni V.</given-names>
</name>
<xref ref-type="aff" rid="aff28"><sup>28</sup>
</xref>
<xref ref-type="aff" rid="aff94"><sup>94</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Harris</surname>
<given-names>Tamara B.</given-names>
</name>
<xref ref-type="aff" rid="aff95"><sup>95</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Heath</surname>
<given-names>Andrew C.</given-names>
</name>
<xref ref-type="aff" rid="aff96"><sup>96</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Hocking</surname>
<given-names>Lynne J.</given-names>
</name>
<xref ref-type="aff" rid="aff97"><sup>97</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Holliday</surname>
<given-names>Elizabeth G.</given-names>
</name>
<xref ref-type="aff" rid="aff50"><sup>50</sup>
</xref>
<xref ref-type="aff" rid="aff51"><sup>51</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Homuth</surname>
<given-names>Georg</given-names>
</name>
<xref ref-type="aff" rid="aff98"><sup>98</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Horan</surname>
<given-names>Michael A.</given-names>
</name>
<xref ref-type="aff" rid="aff99"><sup>99</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Hottenga</surname>
<given-names>Jouke-Jan</given-names>
</name>
<xref ref-type="aff" rid="aff17"><sup>17</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>de Jager</surname>
<given-names>Philip L.</given-names>
</name>
<xref ref-type="aff" rid="aff100"><sup>100</sup>
</xref>
<xref ref-type="aff" rid="aff101"><sup>101</sup>
</xref>
<xref ref-type="aff" rid="aff7"><sup>7</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Joshi</surname>
<given-names>Peter K.</given-names>
</name>
<xref ref-type="aff" rid="aff62"><sup>62</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Jugessur</surname>
<given-names>Astanand</given-names>
</name>
<xref ref-type="aff" rid="aff102"><sup>102</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kaakinen</surname>
<given-names>Marika A.</given-names>
</name>
<xref ref-type="aff" rid="aff103"><sup>103</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kähönen</surname>
<given-names>Mika</given-names>
</name>
<xref ref-type="aff" rid="aff104"><sup>104</sup>
</xref>
<xref ref-type="aff" rid="aff105"><sup>105</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kanoni</surname>
<given-names>Stavroula</given-names>
</name>
<xref ref-type="aff" rid="aff82"><sup>82</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Keltigangas-Järvinen</surname>
<given-names>Liisa</given-names>
</name>
<xref ref-type="aff" rid="aff39"><sup>39</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kiemeney</surname>
<given-names>Lambertus A.L.M.</given-names>
</name>
<xref ref-type="aff" rid="aff27"><sup>27</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kolcic</surname>
<given-names>Ivana</given-names>
</name>
<xref ref-type="aff" rid="aff106"><sup>106</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Koskinen</surname>
<given-names>Seppo</given-names>
</name>
<xref ref-type="aff" rid="aff54"><sup>54</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kraja</surname>
<given-names>Aldi T.</given-names>
</name>
<xref ref-type="aff" rid="aff90"><sup>90</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kroh</surname>
<given-names>Martin</given-names>
</name>
<xref ref-type="aff" rid="aff87"><sup>87</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kutalik</surname>
<given-names>Zoltan</given-names>
</name>
<xref ref-type="aff" rid="aff107"><sup>107</sup>
</xref>
<xref ref-type="aff" rid="aff57"><sup>57</sup>
</xref>
<xref ref-type="aff" rid="aff59"><sup>59</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Latvala</surname>
<given-names>Antti</given-names>
</name>
<xref ref-type="aff" rid="aff29"><sup>29</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Launer</surname>
<given-names>Lenore J.</given-names>
</name>
<xref ref-type="aff" rid="aff108"><sup>108</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Lebreton</surname>
<given-names>Maël P.</given-names>
</name>
<xref ref-type="aff" rid="aff11"><sup>11</sup>
</xref>
<xref ref-type="aff" rid="aff109"><sup>109</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Levinson</surname>
<given-names>Douglas F.</given-names>
</name>
<xref ref-type="aff" rid="aff110"><sup>110</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Lichtenstein</surname>
<given-names>Paul</given-names>
</name>
<xref ref-type="aff" rid="aff41"><sup>41</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Lichtner</surname>
<given-names>Peter</given-names>
</name>
<xref ref-type="aff" rid="aff116"><sup>116</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Liewald</surname>
<given-names>David C.M.</given-names>
</name>
<xref ref-type="aff" rid="aff32"><sup>32</sup>
</xref>
<xref ref-type="aff" rid="aff81"><sup>81</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><collab>LifeLines Cohort Study</collab>
<xref ref-type="aff" rid="aff111"><sup>111</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Loukola</surname>
<given-names>Anu</given-names>
</name>
<xref ref-type="aff" rid="aff29"><sup>29</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Madden</surname>
<given-names>Pamela A.</given-names>
</name>
<xref ref-type="aff" rid="aff96"><sup>96</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Mägi</surname>
<given-names>Reedik</given-names>
</name>
<xref ref-type="aff" rid="aff112"><sup>112</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Mäki-Opas</surname>
<given-names>Tomi</given-names>
</name>
<xref ref-type="aff" rid="aff54"><sup>54</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Marioni</surname>
<given-names>Riccardo E.</given-names>
</name>
<xref ref-type="aff" rid="aff32"><sup>32</sup>
</xref>
<xref ref-type="aff" rid="aff113"><sup>113</sup>
</xref>
<xref ref-type="aff" rid="aff181"><sup>181</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Marques-Vidal</surname>
<given-names>Pedro</given-names>
</name>
<xref ref-type="aff" rid="aff114"><sup>114</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Meddens</surname>
<given-names>Gerardus A.</given-names>
</name>
<xref ref-type="aff" rid="aff115"><sup>115</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>McMahon</surname>
<given-names>George</given-names>
</name>
<xref ref-type="aff" rid="aff66"><sup>66</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Meisinger</surname>
<given-names>Christa</given-names>
</name>
<xref ref-type="aff" rid="aff22"><sup>22</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Meitinger</surname>
<given-names>Thomas</given-names>
</name>
<xref ref-type="aff" rid="aff116"><sup>116</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Milaneschi</surname>
<given-names>Yusplitri</given-names>
</name>
<xref ref-type="aff" rid="aff55"><sup>55</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Milani</surname>
<given-names>Lili</given-names>
</name>
<xref ref-type="aff" rid="aff112"><sup>112</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Montgomery</surname>
<given-names>Grant W.</given-names>
</name>
<xref ref-type="aff" rid="aff117"><sup>117</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Myhre</surname>
<given-names>Ronny</given-names>
</name>
<xref ref-type="aff" rid="aff102"><sup>102</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Nelson</surname>
<given-names>Christopher P.</given-names>
</name>
<xref ref-type="aff" rid="aff30"><sup>30</sup>
</xref>
<xref ref-type="aff" rid="aff31"><sup>31</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Nyholt</surname>
<given-names>Dale R.</given-names>
</name>
<xref ref-type="aff" rid="aff118"><sup>118</sup>
</xref>
<xref ref-type="aff" rid="aff117"><sup>117</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Ollier</surname>
<given-names>William E.R.</given-names>
</name>
<xref ref-type="aff" rid="aff52"><sup>52</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Palotie</surname>
<given-names>Aarno</given-names>
</name>
<xref ref-type="aff" rid="aff119"><sup>119</sup>
</xref>
<xref ref-type="aff" rid="aff7"><sup>7</sup>
</xref>
<xref ref-type="aff" rid="aff120"><sup>120</sup>
</xref>
<xref ref-type="aff" rid="aff121"><sup>121</sup>
</xref>
<xref ref-type="aff" rid="aff122"><sup>122</sup>
</xref>
<xref ref-type="aff" rid="aff123"><sup>123</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Paternoster</surname>
<given-names>Lavinia</given-names>
</name>
<xref ref-type="aff" rid="aff66"><sup>66</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Pedersen</surname>
<given-names>Nancy L.</given-names>
</name>
<xref ref-type="aff" rid="aff41"><sup>41</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Petrovic</surname>
<given-names>Katja E.</given-names>
</name>
<xref ref-type="aff" rid="aff34"><sup>34</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Porteous</surname>
<given-names>David J.</given-names>
</name>
<xref ref-type="aff" rid="aff33"><sup>33</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Räikkönen</surname>
<given-names>Katri</given-names>
</name>
<xref ref-type="aff" rid="aff39"><sup>39</sup>
</xref>
<xref ref-type="aff" rid="aff42"><sup>42</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Ring</surname>
<given-names>Susan M.</given-names>
</name>
<xref ref-type="aff" rid="aff66"><sup>66</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Robino</surname>
<given-names>Antonietta</given-names>
</name>
<xref ref-type="aff" rid="aff124"><sup>124</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Rostapshova</surname>
<given-names>Olga</given-names>
</name>
<xref ref-type="aff" rid="aff125"><sup>125</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Rudan</surname>
<given-names>Igor</given-names>
</name>
<xref ref-type="aff" rid="aff62"><sup>62</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Rustichini</surname>
<given-names>Aldo</given-names>
</name>
<xref ref-type="aff" rid="aff126"><sup>126</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Salomaa</surname>
<given-names>Veikko</given-names>
</name>
<xref ref-type="aff" rid="aff54"><sup>54</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Sanders</surname>
<given-names>Alan R.</given-names>
</name>
<xref ref-type="aff" rid="aff127"><sup>127</sup>
</xref>
<xref ref-type="aff" rid="aff128"><sup>128</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Sarin</surname>
<given-names>Antti-Pekka</given-names>
</name>
<xref ref-type="aff" rid="aff122"><sup>122</sup>
</xref>
<xref ref-type="aff" rid="aff129"><sup>129</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Schmidt</surname>
<given-names>Helena</given-names>
</name>
<xref ref-type="aff" rid="aff130"><sup>130</sup>
</xref>
<xref ref-type="aff" rid="aff34"><sup>34</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Scott</surname>
<given-names>Rodney J.</given-names>
</name>
<xref ref-type="aff" rid="aff131"><sup>131</sup>
</xref>
<xref ref-type="aff" rid="aff51"><sup>51</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Smith</surname>
<given-names>Blair H.</given-names>
</name>
<xref ref-type="aff" rid="aff132"><sup>132</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Smith</surname>
<given-names>Jennifer A.</given-names>
</name>
<xref ref-type="aff" rid="aff75"><sup>75</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Staessen</surname>
<given-names>Jan A.</given-names>
</name>
<xref ref-type="aff" rid="aff133"><sup>133</sup>
</xref>
<xref ref-type="aff" rid="aff134"><sup>134</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Steinhagen-Thiessen</surname>
<given-names>Elisabeth</given-names>
</name>
<xref ref-type="aff" rid="aff85"><sup>85</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Strauch</surname>
<given-names>Konstantin</given-names>
</name>
<xref ref-type="aff" rid="aff135"><sup>135</sup>
</xref>
<xref ref-type="aff" rid="aff136"><sup>136</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Terracciano</surname>
<given-names>Antonio</given-names>
</name>
<xref ref-type="aff" rid="aff137"><sup>137</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Tobin</surname>
<given-names>Martin D.</given-names>
</name>
<xref ref-type="aff" rid="aff138"><sup>138</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Ulivi</surname>
<given-names>Sheila</given-names>
</name>
<xref ref-type="aff" rid="aff124"><sup>124</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Vaccargiu</surname>
<given-names>Simona</given-names>
</name>
<xref ref-type="aff" rid="aff24"><sup>24</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Quaye</surname>
<given-names>Lydia</given-names>
</name>
<xref ref-type="aff" rid="aff47"><sup>47</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>van Rooij</surname>
<given-names>Frank J.A.</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff139"><sup>139</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Venturini</surname>
<given-names>Cristina</given-names>
</name>
<xref ref-type="aff" rid="aff47"><sup>47</sup>
</xref>
<xref ref-type="aff" rid="aff48"><sup>48</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Vinkhuyzen</surname>
<given-names>Anna A.E.</given-names>
</name>
<xref ref-type="aff" rid="aff181"><sup>181</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Völker</surname>
<given-names>Uwe</given-names>
</name>
<xref ref-type="aff" rid="aff98"><sup>98</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Völzke</surname>
<given-names>Henry</given-names>
</name>
<xref ref-type="aff" rid="aff68"><sup>68</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Vonk</surname>
<given-names>Judith M.</given-names>
</name>
<xref ref-type="aff" rid="aff49"><sup>49</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Vozzi</surname>
<given-names>Diego</given-names>
</name>
<xref ref-type="aff" rid="aff124"><sup>124</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Waage</surname>
<given-names>Johannes</given-names>
</name>
<xref ref-type="aff" rid="aff18"><sup>18</sup>
</xref>
<xref ref-type="aff" rid="aff19"><sup>19</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Ware</surname>
<given-names>Erin B.</given-names>
</name>
<xref ref-type="aff" rid="aff75"><sup>75</sup>
</xref>
<xref ref-type="aff" rid="aff140"><sup>140</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Willemsen</surname>
<given-names>Gonneke</given-names>
</name>
<xref ref-type="aff" rid="aff17"><sup>17</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Attia</surname>
<given-names>John R.</given-names>
</name>
<xref ref-type="aff" rid="aff50"><sup>50</sup>
</xref>
<xref ref-type="aff" rid="aff51"><sup>51</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Bennett</surname>
<given-names>David A.</given-names>
</name>
<xref ref-type="aff" rid="aff73"><sup>73</sup>
</xref>
<xref ref-type="aff" rid="aff74"><sup>74</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Berger</surname>
<given-names>Klaus</given-names>
</name>
<xref ref-type="aff" rid="aff69"><sup>69</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Bertram</surname>
<given-names>Lars</given-names>
</name>
<xref ref-type="aff" rid="aff141"><sup>141</sup>
</xref>
<xref ref-type="aff" rid="aff142"><sup>142</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Bisgaard</surname>
<given-names>Hans</given-names>
</name>
<xref ref-type="aff" rid="aff18"><sup>18</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Boomsma</surname>
<given-names>Dorret I.</given-names>
</name>
<xref ref-type="aff" rid="aff17"><sup>17</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Borecki</surname>
<given-names>Ingrid B.</given-names>
</name>
<xref ref-type="aff" rid="aff90"><sup>90</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Bultmann</surname>
<given-names>Ute</given-names>
</name>
<xref ref-type="aff" rid="aff143"><sup>143</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Chabris</surname>
<given-names>Christopher F.</given-names>
</name>
<xref ref-type="aff" rid="aff144"><sup>144</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Cucca</surname>
<given-names>Francesco</given-names>
</name>
<xref ref-type="aff" rid="aff145"><sup>145</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Cusi</surname>
<given-names>Daniele</given-names>
</name>
<xref ref-type="aff" rid="aff60"><sup>60</sup>
</xref>
<xref ref-type="aff" rid="aff146"><sup>146</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Deary</surname>
<given-names>Ian J.</given-names>
</name>
<xref ref-type="aff" rid="aff32"><sup>32</sup>
</xref>
<xref ref-type="aff" rid="aff81"><sup>81</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Dedoussis</surname>
<given-names>George V.</given-names>
</name>
<xref ref-type="aff" rid="aff40"><sup>40</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>van Duijn</surname>
<given-names>Cornelia M.</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Eriksson</surname>
<given-names>Johan G.</given-names>
</name>
<xref ref-type="aff" rid="aff147"><sup>147</sup>
</xref>
<xref ref-type="aff" rid="aff42"><sup>42</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Franke</surname>
<given-names>Barbara</given-names>
</name>
<xref ref-type="aff" rid="aff148"><sup>148</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Franke</surname>
<given-names>Lude</given-names>
</name>
<xref ref-type="aff" rid="aff153"><sup>153</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Gasparini</surname>
<given-names>Paolo</given-names>
</name>
<xref ref-type="aff" rid="aff28"><sup>28</sup>
</xref>
<xref ref-type="aff" rid="aff124"><sup>124</sup>
</xref>
<xref ref-type="aff" rid="aff149"><sup>149</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Gejman</surname>
<given-names>Pablo V.</given-names>
</name>
<xref ref-type="aff" rid="aff127"><sup>127</sup>
</xref>
<xref ref-type="aff" rid="aff128"><sup>128</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Gieger</surname>
<given-names>Christian</given-names>
</name>
<xref ref-type="aff" rid="aff21"><sup>21</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Grabe</surname>
<given-names>Hans-Jörgen</given-names>
</name>
<xref ref-type="aff" rid="aff150"><sup>150</sup>
</xref>
<xref ref-type="aff" rid="aff151"><sup>151</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Gratten</surname>
<given-names>Jacob</given-names>
</name>
<xref ref-type="aff" rid="aff181"><sup>181</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Groenen</surname>
<given-names>Patrick J.F.</given-names>
</name>
<xref ref-type="aff" rid="aff152"><sup>152</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Gudnason</surname>
<given-names>Vilmundur</given-names>
</name>
<xref ref-type="aff" rid="aff9"><sup>9</sup>
</xref>
<xref ref-type="aff" rid="aff65"><sup>65</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>van der Harst</surname>
<given-names>Pim</given-names>
</name>
<xref ref-type="aff" rid="aff69"><sup>69</sup>
</xref>
<xref ref-type="aff" rid="aff153"><sup>153</sup>
</xref>
<xref ref-type="aff" rid="aff154"><sup>154</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Hayward</surname>
<given-names>Caroline</given-names>
</name>
<xref ref-type="aff" rid="aff38"><sup>38</sup>
</xref>
<xref ref-type="aff" rid="aff155"><sup>155</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Hinds</surname>
<given-names>David A.</given-names>
</name>
<xref ref-type="aff" rid="aff26"><sup>26</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Hoffmann</surname>
<given-names>Wolfgang</given-names>
</name>
<xref ref-type="aff" rid="aff68"><sup>68</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Hypponen</surname>
<given-names>Elina</given-names>
</name>
<xref ref-type="aff" rid="aff156"><sup>156</sup>
</xref>
<xref ref-type="aff" rid="aff157"><sup>157</sup>
</xref>
<xref ref-type="aff" rid="aff158"><sup>158</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Iacono</surname>
<given-names>William G.</given-names>
</name>
<xref ref-type="aff" rid="aff5"><sup>5</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Jacobsson</surname>
<given-names>Bo</given-names>
</name>
<xref ref-type="aff" rid="aff20"><sup>20</sup>
</xref>
<xref ref-type="aff" rid="aff102"><sup>102</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Järvelin</surname>
<given-names>Marjo- Riitta</given-names>
</name>
<xref ref-type="aff" rid="aff159"><sup>159</sup>
</xref>
<xref ref-type="aff" rid="aff160"><sup>160</sup>
</xref>
<xref ref-type="aff" rid="aff161"><sup>161</sup>
</xref>
<xref ref-type="aff" rid="aff162"><sup>162</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Jöckel</surname>
<given-names>Karl-Heinz</given-names>
</name>
<xref ref-type="aff" rid="aff61"><sup>61</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kaprio</surname>
<given-names>Jaakko</given-names>
</name>
<xref ref-type="aff" rid="aff29"><sup>29</sup>
</xref>
<xref ref-type="aff" rid="aff122"><sup>122</sup>
</xref>
<xref ref-type="aff" rid="aff54"><sup>54</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Kardia</surname>
<given-names>Sharon L.R.</given-names>
</name>
<xref ref-type="aff" rid="aff75"><sup>75</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Lehtimäki</surname>
<given-names>Terho</given-names>
</name>
<xref ref-type="aff" rid="aff163"><sup>163</sup>
</xref>
<xref ref-type="aff" rid="aff164"><sup>164</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Lehrer</surname>
<given-names>Steven F.</given-names>
</name>
<xref ref-type="aff" rid="aff165"><sup>165</sup>
</xref>
<xref ref-type="aff" rid="aff166"><sup>166</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Magnusson</surname>
<given-names>Patrik K.E.</given-names>
</name>
<xref ref-type="aff" rid="aff41"><sup>41</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Martin</surname>
<given-names>Nicholas G.</given-names>
</name>
<xref ref-type="aff" rid="aff167"><sup>167</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>McGue</surname>
<given-names>Matt</given-names>
</name>
<xref ref-type="aff" rid="aff5"><sup>5</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Metspalu</surname>
<given-names>Andres</given-names>
</name>
<xref ref-type="aff" rid="aff112"><sup>112</sup>
</xref>
<xref ref-type="aff" rid="aff168"><sup>168</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Pendleton</surname>
<given-names>Neil</given-names>
</name>
<xref ref-type="aff" rid="aff169"><sup>169</sup>
</xref>
<xref ref-type="aff" rid="aff170"><sup>170</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Penninx</surname>
<given-names>Brenda W.J.H.</given-names>
</name>
<xref ref-type="aff" rid="aff55"><sup>55</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Perola</surname>
<given-names>Markus</given-names>
</name>
<xref ref-type="aff" rid="aff54"><sup>54</sup>
</xref>
<xref ref-type="aff" rid="aff112"><sup>112</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Pirastu</surname>
<given-names>Nicola</given-names>
</name>
<xref ref-type="aff" rid="aff28"><sup>28</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Pirastu</surname>
<given-names>Mario</given-names>
</name>
<xref ref-type="aff" rid="aff24"><sup>24</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Polasek</surname>
<given-names>Ozren</given-names>
</name>
<xref ref-type="aff" rid="aff171"><sup>171</sup>
</xref>
<xref ref-type="aff" rid="aff62"><sup>62</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Posthuma</surname>
<given-names>Danielle</given-names>
</name>
<xref ref-type="aff" rid="aff43"><sup>43</sup>
</xref>
<xref ref-type="aff" rid="aff172"><sup>172</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Power</surname>
<given-names>Christine</given-names>
</name>
<xref ref-type="aff" rid="aff158"><sup>158</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Province</surname>
<given-names>Michael A.</given-names>
</name>
<xref ref-type="aff" rid="aff90"><sup>90</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Samani</surname>
<given-names>Nilesh J.</given-names>
</name>
<xref ref-type="aff" rid="aff30"><sup>30</sup>
</xref>
<xref ref-type="aff" rid="aff31"><sup>31</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Schlessinger</surname>
<given-names>David</given-names>
</name>
<xref ref-type="aff" rid="aff56"><sup>56</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Schmidt</surname>
<given-names>Reinhold</given-names>
</name>
<xref ref-type="aff" rid="aff34"><sup>34</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Sørensen</surname>
<given-names>Thorkild I.A.</given-names>
</name>
<xref ref-type="aff" rid="aff173"><sup>173</sup>
</xref>
<xref ref-type="aff" rid="aff15"><sup>15</sup>
</xref>
<xref ref-type="aff" rid="aff66"><sup>66</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Spector</surname>
<given-names>Tim D.</given-names>
</name>
<xref ref-type="aff" rid="aff47"><sup>47</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Stefansson</surname>
<given-names>Kari</given-names>
</name>
<xref ref-type="aff" rid="aff93"><sup>93</sup>
</xref>
<xref ref-type="aff" rid="aff65"><sup>65</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Thorsteinsdottir</surname>
<given-names>Unnur</given-names>
</name>
<xref ref-type="aff" rid="aff93"><sup>93</sup>
</xref>
<xref ref-type="aff" rid="aff65"><sup>65</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Thurik</surname>
<given-names>A. Roy</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff174"><sup>174</sup>
</xref>
<xref ref-type="aff" rid="aff175"><sup>175</sup>
</xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Timpson</surname>
<given-names>Nicholas J.</given-names>
</name>
<xref ref-type="aff" rid="aff66"><sup>66</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Tiemeier</surname>
<given-names>Henning</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff176"><sup>176</sup>
</xref>
<xref ref-type="aff" rid="aff177"><sup>177</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Tung</surname>
<given-names>Joyce Y.</given-names>
</name>
<xref ref-type="aff" rid="aff26"><sup>26</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Uitterlinden</surname>
<given-names>André G.</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Vitart</surname>
<given-names>Veronique</given-names>
</name>
<xref ref-type="aff" rid="aff38"><sup>38</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Vollenweider</surname>
<given-names>Peter</given-names>
</name>
<xref ref-type="aff" rid="aff114"><sup>114</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Weir</surname>
<given-names>David R.</given-names>
</name>
<xref ref-type="aff" rid="aff88"><sup>88</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Wilson</surname>
<given-names>James F.</given-names>
</name>
<xref ref-type="aff" rid="aff62"><sup>62</sup>
</xref>
<xref ref-type="aff" rid="aff38"><sup>38</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Wright</surname>
<given-names>Alan F.</given-names>
</name>
<xref ref-type="aff" rid="aff38"><sup>38</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Conley</surname>
<given-names>Dalton C.</given-names>
</name>
<xref ref-type="aff" rid="aff178"><sup>178</sup>
</xref>
<xref ref-type="aff" rid="aff179"><sup>179</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Krueger</surname>
<given-names>Robert F.</given-names>
</name>
<xref ref-type="aff" rid="aff5"><sup>5</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Smith</surname>
<given-names>George Davey</given-names>
</name>
<xref ref-type="aff" rid="aff66"><sup>66</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Hofman</surname>
<given-names>Albert</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Laibson</surname>
<given-names>David I.</given-names>
</name>
<xref ref-type="aff" rid="aff8"><sup>8</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Medland</surname>
<given-names>Sarah E.</given-names>
</name>
<xref ref-type="aff" rid="aff45"><sup>45</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Meyer</surname>
<given-names>Michelle N.</given-names>
</name>
<xref ref-type="aff" rid="aff180"><sup>180</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Yang</surname>
<given-names>Jian</given-names>
</name>
<xref ref-type="aff" rid="aff181"><sup>181</sup>
</xref>
<xref ref-type="aff" rid="aff182"><sup>182</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Johannesson</surname>
<given-names>Magnus</given-names>
</name>
<xref ref-type="aff" rid="aff183"><sup>183</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Visscher</surname>
<given-names>Peter M.</given-names>
</name>
<xref ref-type="aff" rid="aff181"><sup>181</sup>
</xref>
<xref ref-type="aff" rid="aff182"><sup>182</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Esko</surname>
<given-names>Tõnu</given-names>
</name>
<xref ref-type="aff" rid="aff112"><sup>112</sup>
</xref>
<xref ref-type="aff" rid="aff6"><sup>6</sup>
</xref>
<xref ref-type="aff" rid="aff184"><sup>184</sup>
</xref>
<xref ref-type="aff" rid="aff7"><sup>7</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Koellinger</surname>
<given-names>Philipp D.</given-names>
</name>
<xref ref-type="aff" rid="aff43"><sup>43</sup>
</xref>
<xref ref-type="aff" rid="aff11"><sup>11</sup>
</xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Cesarini</surname>
<given-names>David</given-names>
</name>
<xref ref-type="aff" rid="aff14"><sup>14</sup>
</xref>
<xref ref-type="aff" rid="aff185"><sup>185</sup>
</xref>
</contrib>
<contrib contrib-type="collab"><name><surname>Benjamin</surname>
<given-names>Daniel J.</given-names>
</name>
<xref ref-type="aff" rid="aff186"><sup>186</sup>
</xref>
<xref ref-type="aff" rid="aff187"><sup>187</sup>
</xref>
</contrib>
<aff id="aff1"><sup>1</sup>
Department of Applied Economics, Erasmus School of Economics, Erasmus University Rotterdam, 3062 PA, Rotterdam, The Netherlands</aff>
<aff id="aff2"><sup>2</sup>
Department of Epidemiology, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands</aff>
<aff id="aff3"><sup>3</sup>
Erasmus University Rotterdam Institute for Behavior and Biology, Rotterdam 3062 PA, The Netherlands</aff>
<aff id="aff4"><sup>4</sup>
Department of Economics, University of Michigan, Ann Arbor, MI 48109- 1220, USA</aff>
<aff id="aff5"><sup>5</sup>
Department of Psychology, University of Minnesota Twin Cities, Minneapolis, MN 55455, USA</aff>
<aff id="aff6"><sup>6</sup>
Division of Endocrinology and Center for Basic and Translational Obesity Research, Boston Children’s Hospital, Boston, MA 2116, USA</aff>
<aff id="aff7"><sup>7</sup>
Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA 02142, USA</aff>
<aff id="aff8"><sup>8</sup>
Department of Economics, Harvard University, Cambridge, MA 02138, USA</aff>
<aff id="aff9"><sup>9</sup>
Icelandic Heart Association, Kopavogur, 201, Iceland</aff>
<aff id="aff10"><sup>10</sup>
Faculty of Pharmaceutical Sciences, University of Iceland, 107 Reykjavík, Iceland</aff>
<aff id="aff11"><sup>11</sup>
Amsterdam Business School, University of Amsterdam, Amsterdam, 1018 TV, The Netherlands</aff>
<aff id="aff12"><sup>12</sup>
Department of Government, Uppsala University, Uppsala, 751 20, Sweden</aff>
<aff id="aff13"><sup>13</sup>
New York Genome Center, New York, NY 10013, USA</aff>
<aff id="aff14"><sup>14</sup>
Department of Economics, New York University, New York, NY 10012, USA</aff>
<aff id="aff15"><sup>15</sup>
The Novo Nordisk Foundation Center for Basic Metabolic Research, Section of Metabolic Genetics, University of Copenhagen, Faculty of Health and Medical Sciences, København, 2100, Denmark</aff>
<aff id="aff16"><sup>16</sup>
Center for Biological Sequence Analysis, Department of Systems Biology, Technical University of Denmark, Lyngby, 2800, Denmark</aff>
<aff id="aff17"><sup>17</sup>
Department of Biological Psychology, VU University Amsterdam, Amsterdam, 1081 BT, The Netherlands</aff>
<aff id="aff18"><sup>18</sup>
Copenhagen Prospective Studies on Asthma in Childhood, Faculty of Health and Medical Sciences, University of Copenhagen, Gentofte, 2820, Denmark</aff>
<aff id="aff19"><sup>19</sup>
Steno Diabetes Center, 2820 Gentofte, Denmark</aff>
<aff id="aff20"><sup>20</sup>
Department of Obstetrics and Gynecology, Institute of Clinical Sciences, Sahlgrenska Academy, Gothenburg, SE 416 85, Sweden</aff>
<aff id="aff21"><sup>21</sup>
Research Unit of Molecular Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, 85764, Germany</aff>
<aff id="aff22"><sup>22</sup>
Institute of Epidemiology II, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, 85764, Germany</aff>
<aff id="aff23"><sup>23</sup>
Department of Cell Biology, Erasmus Medical Center Rotterdam, 3015 CN, The Netherlands</aff>
<aff id="aff24"><sup>24</sup>
Istituto di Ricerca Genetica e Biomedica U.O.S. di Sassari, National Research Council of Italy, Sassari, 07100, Italy</aff>
<aff id="aff25"><sup>25</sup>
Psychology, University of Illinois, IL 61820, Champaign, USA</aff>
<aff id="aff26"><sup>26</sup>
23andMe, Inc., Mountain View, CA 94041, USA</aff>
<aff id="aff27"><sup>27</sup>
Radboud Institute for Health Sciences, Radboud University Medical Center, Nijmegen, 6500 HB, The Netherlands</aff>
<aff id="aff28"><sup>28</sup>
Department of Medical, Surgical and Health Sciences, University of Trieste, Trieste, 34100, Italy</aff>
<aff id="aff29"><sup>29</sup>
Department of Public Health, University of Helsinki, Helsinki, FI-00014, Finland</aff>
<aff id="aff30"><sup>30</sup>
Department of Cardiovascular Sciences, University of Leicester, Leicester, LE3 9QP, UK</aff>
<aff id="aff31"><sup>31</sup>
NIHR Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester, LE3 9QP, UK</aff>
<aff id="aff32"><sup>32</sup>
Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, EH8 9JZ, UK</aff>
<aff id="aff33"><sup>33</sup>
Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, EH4 2XU, UK</aff>
<aff id="aff34"><sup>34</sup>
Department of Neurology, General Hospital and Medical University Graz, Graz, 8036, Austria</aff>
<aff id="aff35"><sup>35</sup>
Institute for Medical Informatics, Statistics and Documentation, General Hospital and Medical University Graz, Graz, 8036, Austria</aff>
<aff id="aff36"><sup>36</sup>
Oxford Centre for Diabetes, Endocrinology & Metabolism, University of Oxford, Oxford, OX3 7LE, UK</aff>
<aff id="aff37"><sup>37</sup>
Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, OX3 7BN, UK</aff>
<aff id="aff38"><sup>38</sup>
MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, EH4 2XU, UK</aff>
<aff id="aff39"><sup>39</sup>
Institute of Behavioural Sciences, University of Helsinki, Helsinki, FI-00014, Finland</aff>
<aff id="aff40"><sup>40</sup>
Nutrition and Dietetics, Health Science and Education, Harokopio University, Athens, 17671, Greece</aff>
<aff id="aff41"><sup>41</sup>
Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, 171 77, Sweden</aff>
<aff id="aff42"><sup>42</sup>
Folkhälsan Research Centre, Helsingfors, FI-00014, Finland</aff>
<aff id="aff43"><sup>43</sup>
Department of Complex Trait Genetics, VU University, Center for Neurogenomics and Cognitive Research, Amsterdam, 1081 HV, The Netherlands</aff>
<aff id="aff44"><sup>44</sup>
Institute for Computing and Information Sciences, Radboud University Nijmegen, Nijmegen, 6525 EC, The Netherlands</aff>
<aff id="aff45"><sup>45</sup>
Quantitative Genetics, QIMR Berghofer Medical Research Institute, Brisbane, QLD 4029, Australia</aff>
<aff id="aff46"><sup>46</sup>
Lifespan Psychology, Max Planck Institute for Human Development, Berlin, 14195, Germany</aff>
<aff id="aff47"><sup>47</sup>
Department of Twin Research and Genetic Epidemiology, King's College London, London, SE1 7EH, UK</aff>
<aff id="aff48"><sup>48</sup>
NIHR Biomedical Research Centre, Guy’s and St. Thomas’ Foundation Trust, London, SE1 7EH, UK</aff>
<aff id="aff49"><sup>49</sup>
Department of Epidemiology, University of Groningen, University Medical Center Groningen, Groningen, 9700 RB, The Netherlands</aff>
<aff id="aff50"><sup>50</sup>
Public Health Stream, Hunter Medical Research Institute, New Lambton, NSW 2305, Australia</aff>
<aff id="aff51"><sup>51</sup>
Faculty of Health and Medicine, University of Newcastle, Newcastle, NSW 2300, Australia</aff>
<aff id="aff52"><sup>52</sup>
Centre for Integrated Genomic Medical Research, Institute of Population Health, The University of Manchester, Manchester, M13 9PT, UK</aff>
<aff id="aff53"><sup>53</sup>
School of Psychological Sciences, The University of Manchester, Manchester, M13 9PL, UK</aff>
<aff id="aff54"><sup>54</sup>
Department of Health, THL-National Institute for Health and Welfare, Helsinki, FI-00271, Finland</aff>
<aff id="aff55"><sup>55</sup>
Psychiatry, VU University Medical Center & GGZ inGeest, Amsterdam, 1081 HL, The Netherlands</aff>
<aff id="aff56"><sup>56</sup>
Laboratory of Genetics, National Institute on Aging, Baltimore, MD 21224, USA</aff>
<aff id="aff57"><sup>57</sup>
Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, 20521, Finland</aff>
<aff id="aff58"><sup>58</sup>
Department of Medical Genetics, University of Lausanne, Lausanne, 1005, Switzerland</aff>
<aff id="aff59"><sup>59</sup>
Swiss Institute of Bioinformatics, Lausanne, 1015, Switzerland</aff>
<aff id="aff60"><sup>60</sup>
Department Of Health Sciences, University of Milan, Milano, 20142, Italy</aff>
<aff id="aff61"><sup>61</sup>
Institute for Medical Informatics, Biometry and Epidemiology, University Hospital of Essen, Essen, 45147, Germany</aff>
<aff id="aff62"><sup>62</sup>
Centre for Global Health Research, The Usher Institute for Population Health Sciences and Informatics, University of Edinburgh, Edinburgh, EH8 9AG, UK</aff>
<aff id="aff63"><sup>63</sup>
Division of Cancer Epidemiology and Genetics, National Cancer Institute, Bethesda, MD 20892-9780, USA</aff>
<aff id="aff64"><sup>64</sup>
Icelandic Heart Association, Kopavogur, 201, Iceland</aff>
<aff id="aff65"><sup>65</sup>
Faculty of Medicine, University of Iceland, Reykjavik, 101, Iceland</aff>
<aff id="aff66"><sup>66</sup>
MRC Integrative Epidemiology Unit, University of Bristol, Bristol, BS8 2BN, UK</aff>
<aff id="aff67"><sup>67</sup>
School of Oral and Dental Sciences, University of Bristol, Bristol, BS1 2LY, UK</aff>
<aff id="aff68"><sup>68</sup>
Institute for Community Medicine, University Medicine Greifswald, Greifswald, 17475, Germany</aff>
<aff id="aff69"><sup>69</sup>
Department of Cardiology, University Medical Center Groningen, University of Groningen, Groningen, 9700 RB, The Netherlands</aff>
<aff id="aff70"><sup>70</sup>
Institute of Epidemiology and Social Medicine, University of Muenster, Muenster, 48149, Germany</aff>
<aff id="aff71"><sup>71</sup>
Divisions of Genetics and Rheumatology, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, MA 02115, Boston, USA</aff>
<aff id="aff72"><sup>72</sup>
Partners Center for Personalized Genetic Medicine, Boston, MA 02115, USA</aff>
<aff id="aff73"><sup>73</sup>
Rush Alzheimer’s Disease Center, Rush University Medical Center, Chicago, IL 60612, USA</aff>
<aff id="aff74"><sup>74</sup>
Department of Neurological Sciences, Rush University Medical Center, Chicago, IL 60612, USA</aff>
<aff id="aff75"><sup>75</sup>
Department of Epidemiology, University of Michigan, Ann Arbor, MI 48109, USA</aff>
<aff id="aff76"><sup>76</sup>
Department of Gastroenterology and Hepatology, University of Groningen, University Medical Center Groningen, Groningen, 9713 GZ, The Netherlands</aff>
<aff id="aff77"><sup>77</sup>
Institute of Epidemiology and Preventive Medicine, University of Regensburg, Regensburg, D-93053, Germany</aff>
<aff id="aff78"><sup>78</sup>
Institute of Molecular Genetics, National Research Council of Italy, Pavia, 27100, Italy</aff>
<aff id="aff79"><sup>79</sup>
Department of Behavioral Sciences, Rush University Medical Center, Chicago, IL 60612, USA</aff>
<aff id="aff80"><sup>80</sup>
Warwick Medical School, University of Warwick, Coventry, CV4 7AL, UK</aff>
<aff id="aff81"><sup>81</sup>
Department of Psychology, University of Edinburgh, Edinburgh, EH8 9JZ, UK</aff>
<aff id="aff82"><sup>82</sup>
Saïd Business School, University of Oxford, Oxford, OX1 1HP, UK</aff>
<aff id="aff83"><sup>83</sup>
William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, EC1M 6BQ, UK</aff>
<aff id="aff84"><sup>84</sup>
Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, Jeddah, 21589, Saudi Arabia</aff>
<aff id="aff85"><sup>85</sup>
The Berlin Aging Study II; Research Group on Geriatrics, Charité – Universitätsmedizin Berlin, Germany, Berlin, 13347, Germany</aff>
<aff id="aff86"><sup>86</sup>
Institute of Medical and Human Genetics, Charité-Universitätsmedizin, Berlin, Berlin, 13353, Germany</aff>
<aff id="aff87"><sup>87</sup>
German Socio- Economic Panel Study, DIW Berlin, Berlin, 10117, Germany</aff>
<aff id="aff88"><sup>88</sup>
Health Economics Research Centre, Nuffield Department of Population Health, University of Oxford, Oxford, OX3 7LF, UK</aff>
<aff id="aff89"><sup>89</sup>
Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, MI 48109, USA</aff>
<aff id="aff90"><sup>90</sup>
Department of Genetics, Division of Statistical Genomics, Washington University School of Medicine, St. Louis, MO 63018, USA</aff>
<aff id="aff91"><sup>91</sup>
Institute of Human Genetics, University of Bonn, Bonn, 53127, Germany</aff>
<aff id="aff92"><sup>92</sup>
Department of Genomics, Life and Brain Center, University of Bonn, Bonn, 53127, Germany</aff>
<aff id="aff93"><sup>93</sup>
deCODE Genetics/Amgen Inc., Reykjavik, IS-101, Iceland</aff>
<aff id="aff94"><sup>94</sup>
Institute of Biomedical and Neural Engineering, School of Science and Engineering, Reykjavik University, Reykjavik 101, Iceland</aff>
<aff id="aff95"><sup>95</sup>
Laboratory of Epidemiology, Demography, National Institute on Aging, National Institutes of Health, Bethesda, MD 20892-9205, United States</aff>
<aff id="aff96"><sup>96</sup>
Department of Psychiatry, Washington University School of Medicine, St. Louis, MO 63110, USA</aff>
<aff id="aff97"><sup>97</sup>
Division of Applied Health Sciences, University of Aberdeen, Aberdeen, AB25 2ZD, UK</aff>
<aff id="aff98"><sup>98</sup>
Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, Greifswald, 17475, Germany</aff>
<aff id="aff99"><sup>99</sup>
Manchester Medical School, The University of Manchester, Manchester, 9PT, UK</aff>
<aff id="aff100"><sup>100</sup>
Program in Translational NeuroPsychiatric Genomics, Departments of Neurology & Psychiatry, Brigham and Women’s Hospital, Boston, MA 02115, USA</aff>
<aff id="aff101"><sup>101</sup>
Harvard Medical School, Boston, MA 02115, USA</aff>
<aff id="aff102"><sup>102</sup>
Department of Genes and Environment, Norwegian Institute of Public Health, Oslo, N-0403, Norway</aff>
<aff id="aff103"><sup>103</sup>
Department of Genomics of Common Disease, Imperial College London, London, W12 0NN, UK</aff>
<aff id="aff104"><sup>104</sup>
Department of Clinical Physiology, Tampere University Hospital, Tampere, 33521, Finland</aff>
<aff id="aff105"><sup>105</sup>
Department of Clinical Physiology, University of Tampere, School of Medicine, Tampere, 33014, Finland</aff>
<aff id="aff106"><sup>106</sup>
Public Health, Medical School, University of Split, 21000 Split, Croatia</aff>
<aff id="aff107"><sup>107</sup>
Institute of Social and Preventive Medicine, Lausanne University Hospital (CHUV), Lausanne, 1010, Switzerland</aff>
<aff id="aff108"><sup>108</sup>
Neuroepidemiology Section, National Institute on Aging, National Institutes of Health, Bethesda, MD 20892-9205, USA</aff>
<aff id="aff109"><sup>109</sup>
Amsterdam Brain and Cognition Center, University of Amsterdam, 1018 XA, Amsterdam, The Netherlands</aff>
<aff id="aff110"><sup>110</sup>
Department of Psychiatry and Behavioral Sciences, Stanford University, Stanford, CA 94305-5797, USA</aff>
<aff id="aff111"><sup>111</sup>
LifeLines Cohort Study, University of Groningen, University Medical Center Groningen, Groningen, 9713 BZ, The Netherlands</aff>
<aff id="aff112"><sup>112</sup>
Estonian Genome Center, University of Tartu, Tartu, 51010, Estonia</aff>
<aff id="aff113"><sup>113</sup>
Medical Genetics Section, Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, EH4 2XU, UK</aff>
<aff id="aff114"><sup>114</sup>
Department of Internal Medicine, Internal Medicine, Lausanne University Hospital (CHUV), Lausanne, 1011, Switzerland</aff>
<aff id="aff115"><sup>115</sup>
Tema BV, 2131 HE Hoofddorp, The Netherlands</aff>
<aff id="aff116"><sup>116</sup>
Institute of Human Genetics, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, 85764, Germany</aff>
<aff id="aff117"><sup>117</sup>
Molecular Epidemiology, QIMR Berghofer Medical Research Institute, Brisbane, QLD 4029, Australia</aff>
<aff id="aff118"><sup>118</sup>
Institute of Health and Biomedical Innovation, Queensland Institute of Technology, Brisbane, QLD 4059, Australia</aff>
<aff id="aff119"><sup>119</sup>
Analytic and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital, Boston, MA 02114, USA</aff>
<aff id="aff120"><sup>120</sup>
The Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, MA 02142, USA.</aff>
<aff id="aff121"><sup>121</sup>
Psychiatric & Neurodevelopmental Genetics Unit, Department of Psychiatry, Massachusetts General Hospital, Boston, MA 02114, USA</aff>
<aff id="aff122"><sup>122</sup>
Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, 00014, Finland</aff>
<aff id="aff123"><sup>123</sup>
Department of Neurology, Massachusetts General Hospital, Boston, MA 02114, USA</aff>
<aff id="aff124"><sup>124</sup>
Medical Genetics, Institute for Maternal and Child Health IRCCS “Burlo Garofolo”, Trieste, 34100, Italy</aff>
<aff id="aff125"><sup>125</sup>
Harvard Kennedy School, Harvard University, Cambridge, MA 02139</aff>
<aff id="aff126"><sup>126</sup>
Department of Economics, University of Minnesota Twin Cities, Minneapolis, MN 55455, USA</aff>
<aff id="aff127"><sup>127</sup>
Department of Psychiatry and Behavioral Sciences, NorthShore University HealthSystem, Evanston, IL 60201-3137, USA</aff>
<aff id="aff128"><sup>128</sup>
Department of Psychiatry and Behavioral Neuroscience, University of Chicago, Chicago, IL 60637, USA</aff>
<aff id="aff129"><sup>129</sup>
Public Health Genomics Unit, National Institute for Health and Welfare, Helsinki 00300, Finland</aff>
<aff id="aff130"><sup>130</sup>
Research Unit for Genetic Epidemiology, Institute of Molecular Biology and Biochemistry, Center of Molecular Medicine, General Hospital and Medical University, Graz, Graz, 8010, Austria</aff>
<aff id="aff131"><sup>131</sup>
Information Based Medicine Stream, Hunter Medical Research Institute, New Lambton, NSW 2305, Australia</aff>
<aff id="aff132"><sup>132</sup>
Medical Research Institute, University of Dundee, Dundee, DD1 9SY, UK</aff>
<aff id="aff133"><sup>133</sup>
Research Unit Hypertension and Cardiovascular Epidemiology, Department of Cardiovascular Science, University of Leuven, Leuven, 3000, Belgium</aff>
<aff id="aff134"><sup>134</sup>
R&D VitaK Group, Maastricht University, Maastricht, 6229 EV, The Netherlands</aff>
<aff id="aff135"><sup>135</sup>
Institute of Genetic Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, 85764, Germany</aff>
<aff id="aff136"><sup>136</sup>
Institute of Medical Informatics, Biometry and Epidemiology, Chair of Genetic Epidemiology, Ludwig Maximilians-Universität, Munich, 81377, Germany</aff>
<aff id="aff137"><sup>137</sup>
Department of Geriatrics, Florida State University College of Medicine, Tallahassee, FL 32306, USA</aff>
<aff id="aff138"><sup>138</sup>
Department of Health Sciences and Genetics, University of Leicester, Leicester, LE1 7RH, UK</aff>
<aff id="aff139"><sup>139</sup>
Department of Internal Medicine, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands</aff>
<aff id="aff140"><sup>140</sup>
Research Center for Group Dynamics, Institute for Social Research, University of Michigan, Ann Arbor, MI 48104, USA</aff>
<aff id="aff141"><sup>141</sup>
Platform for Genome Analytics, Institutes of Neurogenetics & Integrative and Experimental Genomics, University of Lübeck, Lübeck, 23562, Germany</aff>
<aff id="aff142"><sup>142</sup>
Neuroepidemiology and Ageing Research Unit, School of Public Health, Faculty of Medicine, The Imperial College of Science, Technology and Medicine, London SW7 2AZ, UK</aff>
<aff id="aff143"><sup>143</sup>
Department of Health Sciences, Community & Occupational Medicine, University of Groningen, University Medical Center Groningen, Groningen, 9713 AV, The Netherlands</aff>
<aff id="aff144"><sup>144</sup>
Department of Psychology, Union College, Schenectady, NY 12308, USA</aff>
<aff id="aff145"><sup>145</sup>
Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, 9042, Italy</aff>
<aff id="aff146"><sup>146</sup>
Institute of Biomedical Technologies, Italian National Research Council, Segrate (Milano), 20090, Italy</aff>
<aff id="aff147"><sup>147</sup>
Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, 00014, Finland</aff>
<aff id="aff148"><sup>148</sup>
Departments of Human Genetics and Psychiatry, Donders Centre for Neuroscience, Nijmegen, 6500 HB, The Netherlands</aff>
<aff id="aff149"><sup>149</sup>
Sidra, Experimental Genetics Division, Sidra, Doha 26999, Qatar</aff>
<aff id="aff150"><sup>150</sup>
Department of Psychiatry and Psychotherapy, University Medicine Greifswald, Greifswald, 17475, Germany</aff>
<aff id="aff151"><sup>151</sup>
Department of Psychiatry and Psychotherapy, HELIOS-Hospital Stralsund, Stralsund, 18437, Germany</aff>
<aff id="aff152"><sup>152</sup>
Econometric Institute, Erasmus School of Economics, Erasmus University Rotterdam, Rotterdam, 3062 PA, The Netherlands</aff>
<aff id="aff153"><sup>153</sup>
Department of Genetics, University Medical Center Groningen, University of Groningen, Groningen. 9700 RB, The Netherlands</aff>
<aff id="aff154"><sup>154</sup>
Durrer Center for Cardiogenetic Research, ICIN-Netherlands Heart Institute, Utrecht, 1105 AZ, The Netherlands</aff>
<aff id="aff155"><sup>155</sup>
Generation Scotland, Centre for Genomics and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, EH4 2XU, UK</aff>
<aff id="aff156"><sup>156</sup>
School of Population Health and Sansom Institute, University of South Australia, SA5000, Adelaide, Australia</aff>
<aff id="aff157"><sup>157</sup>
South Australian Health and Medical Research Institute, Adelaide, SA5000, Australia</aff>
<aff id="aff158"><sup>158</sup>
Population, Policy and Practice, UCL Institute of Child Health, London, WC1N 1EH, UK</aff>
<aff id="aff159"><sup>159</sup>
Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment & Health, School of Public Health, Imperial College London, London, W2 1PG, UK</aff>
<aff id="aff160"><sup>160</sup>
Center for Life Course Epidemiology, Faculty of Medicine, University of Oulu, Oulu, FI-90014 , Finland</aff>
<aff id="aff161"><sup>161</sup>
Unit of Primary Care, Oulu University Hospital, Oulu, 90029 OYS, Finland</aff>
<aff id="aff162"><sup>162</sup>
Biocenter Oulu, University of Oulu, FI-90014 Oulu, Finland</aff>
<aff id="aff163"><sup>163</sup>
Fimlab Laboratories, Tampere, 33520, Finland</aff>
<aff id="aff164"><sup>164</sup>
Department of Clinical Chemistry, University of Tampere, School of Medicine, Tampere, 33014, Finland</aff>
<aff id="aff165"><sup>165</sup>
Economics, NYU Shanghai, 200122, Pudong, China</aff>
<aff id="aff166"><sup>166</sup>
Policy Studies, Queen's University, Kingston, K7L3N6, Canada</aff>
<aff id="aff167"><sup>167</sup>
Genetic Epidemiology, QIMR Berghofer Medical Research Institute, Brisbane, QLD 4029, Australia</aff>
<aff id="aff168"><sup>168</sup>
Institute of Molecular and Cell Biology, University of Tartu, Tartu, 51010, Estonia</aff>
<aff id="aff169"><sup>169</sup>
Centre for Clinical and Cognitive Neuroscience, Institute Brain Behaviour and Mental Health, Salford Royal Hospital, Manchester, M6 8HD, UK</aff>
<aff id="aff170"><sup>170</sup>
Manchester Institute Collaborative Research in Ageing, University of Manchester, Manchester, M13 9PL, UK</aff>
<aff id="aff171"><sup>171</sup>
Faculty of Medicine, University of Split, Croatia, Split 21000, Croatia</aff>
<aff id="aff172"><sup>172</sup>
Department of Clinical Genetics, VU Medical Centre, Amsterdam, 1081 HV, The Netherlands</aff>
<aff id="aff173"><sup>173</sup>
Institute of Preventive Medicine, Frederiksberg Hospital, The Capital Region, Frederiksberg, 2000, Denmark</aff>
<aff id="aff174"><sup>174</sup>
Montpellier Business School, Montpellier, 34080, France</aff>
<aff id="aff175"><sup>175</sup>
Panteia, Zoetermeer, 2715 CA, The Netherlands</aff>
<aff id="aff176"><sup>176</sup>
Department of Psychiatry, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands</aff>
<aff id="aff177"><sup>177</sup>
Department of Child and Adolescent Psychiatry, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands</aff>
<aff id="aff178"><sup>178</sup>
Department of Sociology, New York University, New York, NY 10012, USA</aff>
<aff id="aff179"><sup>179</sup>
School of Medicine, New York University, NY 10016, New York, USA</aff>
<aff id="aff180"><sup>180</sup>
Bioethics Program, Union Graduate College - Icahn School of Medicine at Mount Sinai, Schenectady, NY 12308, USA</aff>
<aff id="aff181"><sup>181</sup>
Queensland Brain Institute, The University of Queensland, Brisbane, QLD 4072, Australia</aff>
<aff id="aff182"><sup>182</sup>
The University of Queensland Diamantina Institute, The Translation Research Institute, Brisbane, QLD 4102, Australia</aff>
<aff id="aff183"><sup>183</sup>
Department of Economics, Stockholm School of Economics, Stockholm, 113 83, Sweden</aff>
<aff id="aff184"><sup>184</sup>
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA</aff>
<aff id="aff185"><sup>185</sup>
Research Institute for Industrial Economics, Stockholm, 10215, Sweden</aff>
<aff id="aff186"><sup>186</sup>
Department of Economics, Cornell University, Ithaca, NY 14853, USA</aff>
<aff id="aff187"><sup>187</sup>
Center for Economic and Social Research, University of Southern California, Los Angeles, CA 90089-3332, USA</aff>
</contrib-group>
<author-notes><corresp id="cor1"><sup>2</sup>
To whom correspondence should be addressed. Email: <email>Riccardo.Marioni@ed.ac.uk</email>
.</corresp>
<fn fn-type="edited-by"><p>Edited by Kenneth W. Wachter, University of California, Berkeley, CA, and approved September 20, 2016 (received for review April 4, 2016)</p>
</fn>
<fn fn-type="con"><p>Author contributions: R.E.M., S.J.R., P.K.J., S.S.G.A.C., K.L., A.M., A.C., J.F.W., C.H., T.E., D.J.P., C.R.G., and I.J.D. designed research; R.E.M., S.J.R., and P.K.J. performed research; R.E.M., S.J.R., P.K.J., S.P.H., A.O., K.F., M.J.A., W.D.H., G.D., S.S.G.A.C., R.N., C.A., D.C.L., A.C., and C.H. analyzed data; R.E.M., S.J.R., and P.K.J. wrote the paper; and J.F.W., C.H., T.E., D.J.P., and C.R.G. managed the study.</p>
</fn>
<fn fn-type="equal" id="fn1"><p><sup>1</sup>
R.E.M., S.J.R., and P.K.J. contributed equally to this work.</p>
</fn>
<fn fn-type="equal" id="fn3"><p><sup>4</sup>
C.R.G. and I.J.D. contributed equally to this work.</p>
</fn>
</author-notes>
<pub-date pub-type="ppub"><day>22</day>
<month>11</month>
<year>2016</year>
</pub-date>
<pub-date pub-type="epub"><day>31</day>
<month>10</month>
<year>2016</year>
</pub-date>
<pub-date pub-type="pmc-release"><day>31</day>
<month>10</month>
<year>2016</year>
</pub-date>
<pmc-comment> PMC Release delay is 0 months and 0 days and was based on the . </pmc-comment>
<volume>113</volume>
<issue>47</issue>
<fpage>13366</fpage>
<lpage>13371</lpage>
<permissions><license license-type="open-access"><license-p>Freely available online through the PNAS open access option.</license-p>
</license>
</permissions>
<self-uri xlink:title="pdf" xlink:href="pnas.201605334.pdf"></self-uri>
<abstract abstract-type="executive-summary"><title>Significance</title>
<p>Individuals with more education tend to live longer. Genetic variants have been discovered that predict educational attainment. We tested whether a “polygenic score” based on these genetic variants could make predictions about people’s lifespan. We used data from three cohort studies (including >130,000 participants) to examine the link between offspring polygenic score for education and parental longevity. Across the studies, we found that participants with more education-linked genetic variants had longer-living parents; compared with those with the lowest genetic education scores, those with the highest scores had parents who lived on average 6 months longer. This finding suggests the hypothesis that part of the ultimate explanation for the extended longevity of better-educated people is an underlying, quantifiable, genetic propensity.</p>
</abstract>
<abstract><p>Educational attainment is associated with many health outcomes, including longevity. It is also known to be substantially heritable. Here, we used data from three large genetic epidemiology cohort studies (Generation Scotland, <italic>n =</italic>
∼17,000; UK Biobank, <italic>n</italic>
= ∼115,000; and the Estonian Biobank, <italic>n</italic>
= ∼6,000) to test whether education-linked genetic variants can predict lifespan length. We did so by using cohort members’ polygenic profile score for education to predict their parents’ longevity. Across the three cohorts, meta-analysis showed that a 1 SD higher polygenic education score was associated with ∼2.7% lower mortality risk for both mothers (total <italic>n</italic>
<sub>deaths</sub>
= 79,702) and ∼2.4% lower risk for fathers (total <italic>n</italic>
<sub>deaths</sub>
= 97,630). On average, the parents of offspring in the upper third of the polygenic score distribution lived 0.55 y longer compared with those of offspring in the lower third. Overall, these results indicate that the genetic contributions to educational attainment are useful in the prediction of human longevity.</p>
</abstract>
<kwd-group><kwd>genetics</kwd>
<kwd>education</kwd>
<kwd>longevity</kwd>
<kwd>prediction</kwd>
<kwd>polygenic score</kwd>
</kwd-group>
<funding-group><award-group id="gs1"><funding-source id="sp1">Chief Scientist Office (CSO)<named-content content-type="funder-id">501100000589</named-content>
</funding-source>
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</front>
</pmc>
<affiliations><list><country><li>Australie</li>
<li>Estonie</li>
<li>Pays-Bas</li>
<li>Royaume-Uni</li>
</country>
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<name sortKey="Adams, Mark J" sort="Adams, Mark J" uniqKey="Adams M" first="Mark J." last="Adams">Mark J. Adams</name>
<name sortKey="Amador, Carmen" sort="Amador, Carmen" uniqKey="Amador C" first="Carmen" last="Amador">Carmen Amador</name>
<name sortKey="Campbell, Archie" sort="Campbell, Archie" uniqKey="Campbell A" first="Archie" last="Campbell">Archie Campbell</name>
<name sortKey="Davies, Gail" sort="Davies, Gail" uniqKey="Davies G" first="Gail" last="Davies">Gail Davies</name>
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<country name="Australie"><noRegion><name sortKey="Marioni, Riccardo E" sort="Marioni, Riccardo E" uniqKey="Marioni R" first="Riccardo E." last="Marioni">Riccardo E. Marioni</name>
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