Sequence variants at CHRNB3–CHRNA6 and CYP2A6 affect smoking behavior

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They report three loci associated to cigarettes smoked per day. Smoking is a common risk factor for many diseases1. We conducted genome-wide association meta-analyses for the number of cigarettes smoked per day (CPD) in smokers (n = 31,266) and smoking initiation (n = 46,481) using samples from the ENGAGE Consortium. In a second stage, we tested selected SNPs with in silico replication in the Tobacco and Genetics (TAG) and Glaxo Smith Kline (Ox-GSK) consortia cohorts (n = 45,691 smokers) and assessed some of those in a third sample of European ancestry (n = 9,040). Variants in three genomic regions associated with CPD (P < 5 × 10−8), including previously identified SNPs at 15q25 represented by rs1051730[A] (effect size = 0.80 CPD, P = 2.4 × 10−69), and SNPs at 19q13 and 8p11, represented by rs4105144[C] (effect size = 0.39 CPD, P = 2.2 × 10−12) and rs6474412-T (effect size = 0.29 CPD, P = 1.4 × 10−8), respectively. Among the genes at the two newly associated loci are genes encoding nicotine-metabolizing enzymes (CYP2A6 and CYP2B6) and nicotinic acetylcholine receptor subunits (CHRNB3 and CHRNA6), all of which have been highlighted in previous studies of smoking and nicotine dependence2,3,4. Nominal associations with lung cancer were observed at both 8p11 (rs6474412[T], odds ratio (OR) = 1.09, P = 0.04) and 19q13 (rs4105144[C], OR = 1.12, P = 0.0006).","datePublished":"2010-04-25T00:00:00Z","dateModified":"2010-04-25T00:00:00Z","pageStart":"448","pageEnd":"453","sameAs":"https://doi.org/10.1038/ng.573","keywords":["Addiction","Genome-wide association studies","Biomedicine","general","Human Genetics","Cancer Research","Agriculture","Gene Function","Animal Genetics and Genomics"],"image":["https://media.springernature.com/lw1200/springer-static/image/art%3A10.1038%2Fng.573/MediaObjects/41588_2010_Article_BFng573_Fig1_HTML.jpg"],"isPartOf":{"name":"Nature Genetics","issn":["1546-1718","1061-4036"],"volumeNumber":"42","@type":["Periodical","PublicationVolume"]},"publisher":{"name":"Nature Publishing Group US","logo":{"url":"https://www.springernature.com/app-sn/public/images/logo-springernature.png","@type":"ImageObject"},"@type":"Organization"},"author":[{"name":"Thorgeir E 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c-article-author-list--long" data-test="authors-list" data-component-authors-activator="authors-list"><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Thorgeir_E-Thorgeirsson-Aff1-Aff2" data-author-popup="auth-Thorgeir_E-Thorgeirsson-Aff1-Aff2" data-author-search="Thorgeirsson, Thorgeir E">Thorgeir E Thorgeirsson</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff2">2</a></sup>, </li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Daniel_F-Gudbjartsson-Aff1" data-author-popup="auth-Daniel_F-Gudbjartsson-Aff1" data-author-search="Gudbjartsson, Daniel F">Daniel F Gudbjartsson</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Ida-Surakka-Aff3-Aff4" data-author-popup="auth-Ida-Surakka-Aff3-Aff4" data-author-search="Surakka, Ida">Ida Surakka</a><sup class="u-js-hide"><a href="#Aff3">3</a>,<a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jacqueline_M-Vink-Aff5" data-author-popup="auth-Jacqueline_M-Vink-Aff5" data-author-search="Vink, Jacqueline M">Jacqueline M Vink</a><sup class="u-js-hide"><a href="#Aff5">5</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Najaf-Amin-Aff6" data-author-popup="auth-Najaf-Amin-Aff6" data-author-search="Amin, Najaf">Najaf Amin</a><sup class="u-js-hide"><a 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data-author-popup="auth-Thorunn-Rafnar-Aff1" data-author-search="Rafnar, Thorunn">Thorunn Rafnar</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-T_nu-Esko-Aff7-Aff8" data-author-popup="auth-T_nu-Esko-Aff7-Aff8" data-author-search="Esko, Tõnu">Tõnu Esko</a><sup class="u-js-hide"><a href="#Aff7">7</a>,<a href="#Aff8">8</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Stefan-Walter-Aff6" data-author-popup="auth-Stefan-Walter-Aff6" data-author-search="Walter, Stefan">Stefan Walter</a><sup class="u-js-hide"><a href="#Aff6">6</a></sup>, </li><li class="c-article-author-list__item 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data-author-search="Mangino, Massimo">Massimo Mangino</a><sup class="u-js-hide"><a href="#Aff10">10</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Inga-Prokopenko-Aff11-Aff12" data-author-popup="auth-Inga-Prokopenko-Aff11-Aff12" data-author-search="Prokopenko, Inga">Inga Prokopenko</a><sup class="u-js-hide"><a href="#Aff11">11</a>,<a href="#Aff12">12</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Reedik-M_gi-Aff11-Aff12" data-author-popup="auth-Reedik-M_gi-Aff11-Aff12" data-author-search="Mägi, Reedik">Reedik Mägi</a><sup class="u-js-hide"><a href="#Aff11">11</a>,<a href="#Aff12">12</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Kaisu-Keskitalo-Aff13" data-author-popup="auth-Kaisu-Keskitalo-Aff13" data-author-search="Keskitalo, Kaisu">Kaisu Keskitalo</a><sup class="u-js-hide"><a href="#Aff13">13</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Iris_H-Gudjonsdottir-Aff1" data-author-popup="auth-Iris_H-Gudjonsdottir-Aff1" data-author-search="Gudjonsdottir, Iris H">Iris H Gudjonsdottir</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Solveig-Gretarsdottir-Aff1" data-author-popup="auth-Solveig-Gretarsdottir-Aff1" data-author-search="Gretarsdottir, Solveig">Solveig Gretarsdottir</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Hreinn-Stefansson-Aff1" data-author-popup="auth-Hreinn-Stefansson-Aff1" data-author-search="Stefansson, Hreinn">Hreinn Stefansson</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-John_R-Thompson-Aff14" data-author-popup="auth-John_R-Thompson-Aff14" data-author-search="Thompson, John R">John R Thompson</a><sup class="u-js-hide"><a href="#Aff14">14</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Yurii_S-Aulchenko-Aff6" data-author-popup="auth-Yurii_S-Aulchenko-Aff6" data-author-search="Aulchenko, Yurii S">Yurii S Aulchenko</a><sup class="u-js-hide"><a href="#Aff6">6</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Mari-Nelis-Aff7-Aff8" data-author-popup="auth-Mari-Nelis-Aff7-Aff8" data-author-search="Nelis, Mari">Mari Nelis</a><sup class="u-js-hide"><a href="#Aff7">7</a>,<a href="#Aff8">8</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Katja_K-Aben-Aff15-Aff16" data-author-popup="auth-Katja_K-Aben-Aff15-Aff16" data-author-search="Aben, Katja K">Katja K Aben</a><sup class="u-js-hide"><a href="#Aff15">15</a>,<a href="#Aff16">16</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Martin-den_Heijer-Aff15-Aff17" data-author-popup="auth-Martin-den_Heijer-Aff15-Aff17" data-author-search="den Heijer, Martin">Martin den Heijer</a><sup class="u-js-hide"><a href="#Aff15">15</a>,<a href="#Aff17">17</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Asger-Dirksen-Aff18" data-author-popup="auth-Asger-Dirksen-Aff18" data-author-search="Dirksen, Asger">Asger Dirksen</a><sup class="u-js-hide"><a href="#Aff18">18</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Haseem-Ashraf-Aff18" data-author-popup="auth-Haseem-Ashraf-Aff18" data-author-search="Ashraf, Haseem">Haseem Ashraf</a><sup class="u-js-hide"><a href="#Aff18">18</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Nicole-Soranzo-Aff10-Aff19" data-author-popup="auth-Nicole-Soranzo-Aff10-Aff19" data-author-search="Soranzo, Nicole">Nicole Soranzo</a><sup class="u-js-hide"><a href="#Aff10">10</a>,<a href="#Aff19">19</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Ana_M-Valdes-Aff10" data-author-popup="auth-Ana_M-Valdes-Aff10" data-author-search="Valdes, Ana M">Ana M Valdes</a><sup class="u-js-hide"><a href="#Aff10">10</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Claire-Steves-Aff10" data-author-popup="auth-Claire-Steves-Aff10" data-author-search="Steves, Claire">Claire Steves</a><sup class="u-js-hide"><a href="#Aff10">10</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Andr__G-Uitterlinden-Aff6-Aff20" data-author-popup="auth-Andr__G-Uitterlinden-Aff6-Aff20" data-author-search="Uitterlinden, André G">André G Uitterlinden</a><sup class="u-js-hide"><a href="#Aff6">6</a>,<a href="#Aff20">20</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Albert-Hofman-Aff6" data-author-popup="auth-Albert-Hofman-Aff6" data-author-search="Hofman, Albert">Albert Hofman</a><sup class="u-js-hide"><a href="#Aff6">6</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Anke-T_njes-Aff21-Aff22" data-author-popup="auth-Anke-T_njes-Aff21-Aff22" data-author-search="Tönjes, Anke">Anke Tönjes</a><sup class="u-js-hide"><a href="#Aff21">21</a>,<a href="#Aff22">22</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Peter-Kovacs-Aff23" data-author-popup="auth-Peter-Kovacs-Aff23" data-author-search="Kovacs, Peter">Peter Kovacs</a><sup class="u-js-hide"><a href="#Aff23">23</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jouke_Jan-Hottenga-Aff5" data-author-popup="auth-Jouke_Jan-Hottenga-Aff5" data-author-search="Hottenga, Jouke Jan">Jouke Jan Hottenga</a><sup class="u-js-hide"><a href="#Aff5">5</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Gonneke-Willemsen-Aff5" data-author-popup="auth-Gonneke-Willemsen-Aff5" data-author-search="Willemsen, Gonneke">Gonneke Willemsen</a><sup class="u-js-hide"><a href="#Aff5">5</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Nicole-Vogelzangs-Aff24" data-author-popup="auth-Nicole-Vogelzangs-Aff24" data-author-search="Vogelzangs, Nicole">Nicole Vogelzangs</a><sup class="u-js-hide"><a href="#Aff24">24</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Angela-D_ring-Aff9" data-author-popup="auth-Angela-D_ring-Aff9" data-author-search="Döring, Angela">Angela Döring</a><sup class="u-js-hide"><a href="#Aff9">9</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Norbert-Dahmen-Aff25" data-author-popup="auth-Norbert-Dahmen-Aff25" data-author-search="Dahmen, Norbert">Norbert Dahmen</a><sup class="u-js-hide"><a href="#Aff25">25</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Barbara-Nitz-Aff9" data-author-popup="auth-Barbara-Nitz-Aff9" data-author-search="Nitz, Barbara">Barbara Nitz</a><sup class="u-js-hide"><a href="#Aff9">9</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Michele_L-Pergadia-Aff26" data-author-popup="auth-Michele_L-Pergadia-Aff26" data-author-search="Pergadia, Michele L">Michele L Pergadia</a><sup class="u-js-hide"><a href="#Aff26">26</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Berta-Saez-Aff27" data-author-popup="auth-Berta-Saez-Aff27" data-author-search="Saez, Berta">Berta Saez</a><sup class="u-js-hide"><a href="#Aff27">27</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Veronica-De_Diego-Aff28" data-author-popup="auth-Veronica-De_Diego-Aff28" data-author-search="De Diego, Veronica">Veronica De Diego</a><sup class="u-js-hide"><a href="#Aff28">28</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Victoria-Lezcano-Aff28" data-author-popup="auth-Victoria-Lezcano-Aff28" data-author-search="Lezcano, Victoria">Victoria Lezcano</a><sup class="u-js-hide"><a href="#Aff28">28</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Maria_D-Garcia_Prats-Aff29" data-author-popup="auth-Maria_D-Garcia_Prats-Aff29" data-author-search="Garcia-Prats, Maria D">Maria D Garcia-Prats</a><sup class="u-js-hide"><a href="#Aff29">29</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Samuli-Ripatti-Aff3-Aff4" data-author-popup="auth-Samuli-Ripatti-Aff3-Aff4" data-author-search="Ripatti, Samuli">Samuli Ripatti</a><sup class="u-js-hide"><a href="#Aff3">3</a>,<a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Markus-Perola-Aff4" data-author-popup="auth-Markus-Perola-Aff4" data-author-search="Perola, Markus">Markus Perola</a><sup class="u-js-hide"><a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Johannes-Kettunen-Aff19" data-author-popup="auth-Johannes-Kettunen-Aff19" data-author-search="Kettunen, Johannes">Johannes Kettunen</a><sup class="u-js-hide"><a href="#Aff19">19</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Anna_Liisa-Hartikainen-Aff30" data-author-popup="auth-Anna_Liisa-Hartikainen-Aff30" data-author-search="Hartikainen, Anna-Liisa">Anna-Liisa Hartikainen</a><sup class="u-js-hide"><a href="#Aff30">30</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Anneli-Pouta-Aff31" data-author-popup="auth-Anneli-Pouta-Aff31" data-author-search="Pouta, Anneli">Anneli Pouta</a><sup class="u-js-hide"><a href="#Aff31">31</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jaana-Laitinen-Aff32" data-author-popup="auth-Jaana-Laitinen-Aff32" data-author-search="Laitinen, Jaana">Jaana Laitinen</a><sup class="u-js-hide"><a href="#Aff32">32</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Matti-Isohanni-Aff30" data-author-popup="auth-Matti-Isohanni-Aff30" data-author-search="Isohanni, Matti">Matti Isohanni</a><sup class="u-js-hide"><a href="#Aff30">30</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Shen-Huei_Yi-Aff3-Aff4" data-author-popup="auth-Shen-Huei_Yi-Aff3-Aff4" data-author-search="Huei-Yi, Shen">Shen Huei-Yi</a><sup class="u-js-hide"><a href="#Aff3">3</a>,<a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Maxine-Allen-Aff11" data-author-popup="auth-Maxine-Allen-Aff11" data-author-search="Allen, Maxine">Maxine Allen</a><sup class="u-js-hide"><a href="#Aff11">11</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Maria-Krestyaninova-Aff33" data-author-popup="auth-Maria-Krestyaninova-Aff33" data-author-search="Krestyaninova, Maria">Maria Krestyaninova</a><sup class="u-js-hide"><a href="#Aff33">33</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Alistair_S-Hall-Aff34" data-author-popup="auth-Alistair_S-Hall-Aff34" data-author-search="Hall, Alistair S">Alistair S Hall</a><sup class="u-js-hide"><a href="#Aff34">34</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Gregory_T-Jones-Aff35" data-author-popup="auth-Gregory_T-Jones-Aff35" data-author-search="Jones, Gregory T">Gregory T Jones</a><sup class="u-js-hide"><a href="#Aff35">35</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Andre_M-van_Rij-Aff35" data-author-popup="auth-Andre_M-van_Rij-Aff35" data-author-search="van Rij, Andre M">Andre M van Rij</a><sup class="u-js-hide"><a href="#Aff35">35</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Thomas-Mueller-Aff36" data-author-popup="auth-Thomas-Mueller-Aff36" data-author-search="Mueller, Thomas">Thomas Mueller</a><sup class="u-js-hide"><a href="#Aff36">36</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Benjamin-Dieplinger-Aff36" data-author-popup="auth-Benjamin-Dieplinger-Aff36" data-author-search="Dieplinger, Benjamin">Benjamin Dieplinger</a><sup class="u-js-hide"><a href="#Aff36">36</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Meinhard-Haltmayer-Aff36" data-author-popup="auth-Meinhard-Haltmayer-Aff36" data-author-search="Haltmayer, Meinhard">Meinhard Haltmayer</a><sup class="u-js-hide"><a href="#Aff36">36</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Steinn-Jonsson-Aff37" data-author-popup="auth-Steinn-Jonsson-Aff37" data-author-search="Jonsson, Steinn">Steinn Jonsson</a><sup class="u-js-hide"><a href="#Aff37">37</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Stefan_E-Matthiasson-Aff38" data-author-popup="auth-Stefan_E-Matthiasson-Aff38" data-author-search="Matthiasson, Stefan E">Stefan E Matthiasson</a><sup class="u-js-hide"><a href="#Aff38">38</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Hogni-Oskarsson-Aff39" data-author-popup="auth-Hogni-Oskarsson-Aff39" data-author-search="Oskarsson, Hogni">Hogni Oskarsson</a><sup class="u-js-hide"><a href="#Aff39">39</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Thorarinn-Tyrfingsson-Aff40" data-author-popup="auth-Thorarinn-Tyrfingsson-Aff40" data-author-search="Tyrfingsson, Thorarinn">Thorarinn Tyrfingsson</a><sup class="u-js-hide"><a href="#Aff40">40</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Lambertus_A-Kiemeney-Aff15-Aff16-Aff41" data-author-popup="auth-Lambertus_A-Kiemeney-Aff15-Aff16-Aff41" data-author-search="Kiemeney, Lambertus A">Lambertus A Kiemeney</a><sup class="u-js-hide"><a href="#Aff15">15</a>,<a href="#Aff16">16</a>,<a href="#Aff41">41</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jose_I-Mayordomo-Aff42" data-author-popup="auth-Jose_I-Mayordomo-Aff42" data-author-search="Mayordomo, Jose I">Jose I Mayordomo</a><sup class="u-js-hide"><a href="#Aff42">42</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jes_S-Lindholt-Aff43" data-author-popup="auth-Jes_S-Lindholt-Aff43" data-author-search="Lindholt, Jes S">Jes S Lindholt</a><sup class="u-js-hide"><a href="#Aff43">43</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jesper_Holst-Pedersen-Aff44" data-author-popup="auth-Jesper_Holst-Pedersen-Aff44" data-author-search="Pedersen, Jesper Holst">Jesper Holst Pedersen</a><sup class="u-js-hide"><a href="#Aff44">44</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Wilbur_A-Franklin-Aff45" data-author-popup="auth-Wilbur_A-Franklin-Aff45" data-author-search="Franklin, Wilbur A">Wilbur A Franklin</a><sup class="u-js-hide"><a href="#Aff45">45</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Holly-Wolf-Aff46" data-author-popup="auth-Holly-Wolf-Aff46" data-author-search="Wolf, Holly">Holly Wolf</a><sup class="u-js-hide"><a href="#Aff46">46</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Grant_W-Montgomery-Aff47" data-author-popup="auth-Grant_W-Montgomery-Aff47" data-author-search="Montgomery, Grant W">Grant W Montgomery</a><sup class="u-js-hide"><a href="#Aff47">47</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Andrew_C-Heath-Aff26" data-author-popup="auth-Andrew_C-Heath-Aff26" data-author-search="Heath, Andrew C">Andrew C Heath</a><sup class="u-js-hide"><a href="#Aff26">26</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Nicholas_G-Martin-Aff47" data-author-popup="auth-Nicholas_G-Martin-Aff47" data-author-search="Martin, Nicholas G">Nicholas G Martin</a><sup class="u-js-hide"><a href="#Aff47">47</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Pamela_A_F-Madden-Aff26" data-author-popup="auth-Pamela_A_F-Madden-Aff26" data-author-search="Madden, Pamela A F">Pamela A F Madden</a><sup class="u-js-hide"><a href="#Aff26">26</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Ina-Giegling-Aff48" data-author-popup="auth-Ina-Giegling-Aff48" data-author-search="Giegling, Ina">Ina Giegling</a><sup class="u-js-hide"><a href="#Aff48">48</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Dan-Rujescu-Aff48" data-author-popup="auth-Dan-Rujescu-Aff48" data-author-search="Rujescu, Dan">Dan Rujescu</a><sup class="u-js-hide"><a href="#Aff48">48</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Marjo_Riitta-J_rvelin-Aff31-Aff49-Aff50-Aff51" data-author-popup="auth-Marjo_Riitta-J_rvelin-Aff31-Aff49-Aff50-Aff51" data-author-search="Järvelin, Marjo-Riitta">Marjo-Riitta Järvelin</a><sup class="u-js-hide"><a href="#Aff31">31</a>,<a href="#Aff49">49</a>,<a href="#Aff50">50</a>,<a href="#Aff51">51</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Veikko-Salomaa-Aff4" data-author-popup="auth-Veikko-Salomaa-Aff4" data-author-search="Salomaa, Veikko">Veikko Salomaa</a><sup class="u-js-hide"><a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Michael-Stumvoll-Aff21" data-author-popup="auth-Michael-Stumvoll-Aff21" data-author-search="Stumvoll, Michael">Michael Stumvoll</a><sup class="u-js-hide"><a href="#Aff21">21</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Tim_D-Spector-Aff10" data-author-popup="auth-Tim_D-Spector-Aff10" data-author-search="Spector, Tim D">Tim D Spector</a><sup class="u-js-hide"><a href="#Aff10">10</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-H_Erich-Wichmann-Aff9-Aff52-Aff53" data-author-popup="auth-H_Erich-Wichmann-Aff9-Aff52-Aff53" data-author-search="Wichmann, H-Erich">H-Erich Wichmann</a><sup class="u-js-hide"><a href="#Aff9">9</a>,<a href="#Aff52">52</a>,<a href="#Aff53">53</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Andres-Metspalu-Aff7-Aff8" data-author-popup="auth-Andres-Metspalu-Aff7-Aff8" data-author-search="Metspalu, Andres">Andres Metspalu</a><sup class="u-js-hide"><a href="#Aff7">7</a>,<a href="#Aff8">8</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Nilesh_J-Samani-Aff54" data-author-popup="auth-Nilesh_J-Samani-Aff54" data-author-search="Samani, Nilesh J">Nilesh J Samani</a><sup class="u-js-hide"><a href="#Aff54">54</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Brenda_W-Penninx-Aff24" data-author-popup="auth-Brenda_W-Penninx-Aff24" data-author-search="Penninx, Brenda W">Brenda W Penninx</a><sup class="u-js-hide"><a href="#Aff24">24</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Ben_A-Oostra-Aff55" data-author-popup="auth-Ben_A-Oostra-Aff55" data-author-search="Oostra, Ben A">Ben A Oostra</a><sup class="u-js-hide"><a href="#Aff55">55</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Dorret_I-Boomsma-Aff5" data-author-popup="auth-Dorret_I-Boomsma-Aff5" data-author-search="Boomsma, Dorret I">Dorret I Boomsma</a><sup class="u-js-hide"><a href="#Aff5">5</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Henning-Tiemeier-Aff6" data-author-popup="auth-Henning-Tiemeier-Aff6" data-author-search="Tiemeier, Henning">Henning Tiemeier</a><sup class="u-js-hide"><a href="#Aff6">6</a></sup>, </li><li 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c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jeffrey_R-Gulcher-Aff1" data-author-popup="auth-Jeffrey_R-Gulcher-Aff1" data-author-search="Gulcher, Jeffrey R">Jeffrey R Gulcher</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item"><a data-test="author-name" data-author-popup="group-1" href="#group-1">The ENGAGE Consortium</a>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Mark_I-McCarthy-Aff11-Aff12" data-author-popup="auth-Mark_I-McCarthy-Aff11-Aff12" data-author-search="McCarthy, Mark I">Mark I McCarthy</a><sup class="u-js-hide"><a href="#Aff11">11</a>,<a href="#Aff12">12</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Leena-Peltonen-Aff3-Aff19" data-author-popup="auth-Leena-Peltonen-Aff3-Aff19" data-author-search="Peltonen, Leena">Leena Peltonen</a><sup class="u-js-hide"><a href="#Aff3">3</a>,<a href="#Aff19">19</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Unnur-Thorsteinsdottir-Aff1-Aff38" data-author-popup="auth-Unnur-Thorsteinsdottir-Aff1-Aff38" data-author-search="Thorsteinsdottir, Unnur" data-corresp-id="c1">Unnur Thorsteinsdottir<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff38">38</a></sup> & </li><li class="c-article-author-list__show-more" aria-label="Show all 95 authors for this article" title="Show all 95 authors for this article">…</li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Kari-Stefansson-Aff1-Aff38" data-author-popup="auth-Kari-Stefansson-Aff1-Aff38" data-author-search="Stefansson, Kari" data-corresp-id="c2">Kari Stefansson<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff38">38</a></sup> </li></ul><button aria-expanded="false" class="c-article-author-list__button"><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-down-medium"></use></svg><span>Show authors</span></button> <p class="c-article-info-details" data-container-section="info"> <a data-test="journal-link" href="/ng" data-track="click" data-track-action="journal homepage" data-track-category="article body" data-track-label="link"><i data-test="journal-title">Nature Genetics</i></a> <b data-test="journal-volume"><span class="u-visually-hidden">volume</span> 42</b>, <span class="u-visually-hidden">pages </span>448–453 (<span data-test="article-publication-year">2010</span>)<a href="#citeas" class="c-article-info-details__cite-as u-hide-print" data-track="click" data-track-action="cite this article" data-track-label="link">Cite this article</a> </p> <div class="c-article-metrics-bar__wrapper u-clear-both"> <ul class="c-article-metrics-bar u-list-reset"> <li class=" c-article-metrics-bar__item" data-test="access-count"> <p class="c-article-metrics-bar__count">3833 <span class="c-article-metrics-bar__label">Accesses</span></p> </li> <li class="c-article-metrics-bar__item" data-test="citation-count"> <p class="c-article-metrics-bar__count">540 <span class="c-article-metrics-bar__label">Citations</span></p> </li> <li class="c-article-metrics-bar__item" data-test="altmetric-score"> <p class="c-article-metrics-bar__count">10 <span class="c-article-metrics-bar__label">Altmetric</span></p> </li> <li class="c-article-metrics-bar__item"> <p class="c-article-metrics-bar__details"><a href="/articles/ng.573/metrics" data-track="click" data-track-action="view metrics" data-track-label="link" rel="nofollow">Metrics <span class="u-visually-hidden">details</span></a></p> </li> </ul> </div> </header> <div class="u-js-hide" data-component="article-subject-links"> <h3 class="c-article__sub-heading">Subjects</h3> <ul class="c-article-subject-list"> <li class="c-article-subject-list__subject"><a href="/subjects/addiction" data-track="click" data-track-action="view subject" data-track-label="link">Addiction</a></li><li class="c-article-subject-list__subject"><a href="/subjects/genome-wide-association-studies" data-track="click" data-track-action="view subject" data-track-label="link">Genome-wide association studies</a></li> </ul> </div> </div> <div class="c-article-body"> <section aria-labelledby="Abs2" data-title="Abstract" lang="en"><div class="c-article-section" id="Abs2-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Abs2">Abstract</h2><div class="c-article-section__content" id="Abs2-content"><p>Smoking is a common risk factor for many diseases<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="WHO Report on the Global Tobacco Epidemic, 2008. 8 
 http://tobaccofreecenter.org/mpower-2008
 
 (2008)." href="/articles/ng.573#ref-CR1" id="ref-link-section-d159083860e2301">1</a></sup>. We conducted genome-wide association meta-analyses for the number of cigarettes smoked per day (CPD) in smokers (<i>n</i> = 31,266) and smoking initiation (<i>n</i> = 46,481) using samples from the ENGAGE Consortium. In a second stage, we tested selected SNPs with <i>in silico</i> replication in the Tobacco and Genetics (TAG) and Glaxo Smith Kline (Ox-GSK) consortia cohorts (<i>n</i> = 45,691 smokers) and assessed some of those in a third sample of European ancestry (<i>n</i> = 9,040). Variants in three genomic regions associated with CPD (<i>P</i> < 5 × 10<sup>−8</sup>), including previously identified SNPs at 15q25 represented by rs1051730[A] (effect size = 0.80 CPD, <i>P</i> = 2.4 × 10<sup>−69</sup>), and SNPs at 19q13 and 8p11, represented by rs4105144[C] (effect size = 0.39 CPD, <i>P</i> = 2.2 × 10<sup>−12</sup>) and rs6474412-T (effect size = 0.29 CPD, <i>P</i> = 1.4 × 10<sup>−8</sup>), respectively. Among the genes at the two newly associated loci are genes encoding nicotine-metabolizing enzymes (<i>CYP2A6</i> and <i>CYP2B6</i>) and nicotinic acetylcholine receptor subunits (<i>CHRNB3</i> and <i>CHRNA6</i>), all of which have been highlighted in previous studies of smoking and nicotine dependence<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Bierut, L.J. et al. Novel genes identified in a high-density genome wide association study for nicotine dependence. Hum. Mol. Genet. 16, 24–35 (2007)." href="/articles/ng.573#ref-CR2" id="ref-link-section-d159083860e2355">2</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 3" title="Saccone, S.F. et al. Cholinergic nicotinic receptor genes implicated in a nicotine dependence association study targeting 348 candidate genes with 3713 SNPs. Hum. Mol. Genet. 16, 36–49 (2007)." href="/articles/ng.573#ref-CR3" id="ref-link-section-d159083860e2358">3</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 4" title="Ray, R., Tyndale, R.F. & Lerman, C. Nicotine dependence pharmacogenetics: role of genetic variation in nicotine-metabolizing enzymes. J. Neurogenet. 23, 252–261 (2009)." href="/articles/ng.573#ref-CR4" id="ref-link-section-d159083860e2361">4</a></sup>. Nominal associations with lung cancer were observed at both 8p11 (rs6474412[T], odds ratio (OR) = 1.09, <i>P</i> = 0.04) and 19q13 (rs4105144[C], OR = 1.12, <i>P</i> = 0.0006).</p></div></div></section> <noscript> <div class="c-nature-box c-nature-box--side " data-component="entitlement-box"> <div class="js-access-button"> <a href="https://wayf.springernature.com?redirect_uri=https%3A%2F%2Fwww.nature.com%2Farticles%2Fng.573" class="c-article__button" data-test="ra21"> <svg class="u-icon" width="18" height="18" aria-hidden="true" focusable="false"><use href="#icon-institution"></use></svg> <span class="c-article__button-text">Access through your institution</span> </a> </div> <div class="js-buy-button"> <a href="#access-options" class="c-article__button c-article__button--inverted" data-test="ra21"> <span>Buy or subscribe</span> </a> </div> </div> </noscript> <div class="js-context-bar-sticky-point-mobile" data-track-context="article body"> </div> <div class="c-article-access-provider 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This work was supported in part by the US National Institutes of Health (R01-DA017932) and the European Commission's Sixth Framework Program, Integrated Project GENADDICT (LSHM-CT-2004-005166). The ENGAGE smoking consortium was formed through a component of the Integrated Project ENGAGE, supported by the European Commission's Seventh Framework Program, grant agreement HEALTH-F4-2007- 201413. ENGAGE projects have benefited from the SIMBioMS platform<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 28" title="Krestyaninova, M. et al. A System for Information Management in BioMedical Studies–SIMBioMS. Bioinformatics 25, 2768–2769 (2009)." href="/articles/ng.573#ref-CR28" id="ref-link-section-d159083860e5682">28</a></sup>, which has greatly facilitated data exchange and annotation. Further acknowledgments are listed in the <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/ng.573#MOESM34">Supplementary Note</a> online.</p></div></div></section><section aria-labelledby="author-information" data-title="Author information"><div class="c-article-section" id="author-information-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="author-information">Author information</h2><div class="c-article-section__content" id="author-information-content"><h3 class="c-article__sub-heading" id="affiliations">Authors and Affiliations</h3><ol class="c-article-author-affiliation__list"><li id="Aff1"><p class="c-article-author-affiliation__address">deCODE Genetics, Reykjavik, Iceland</p><p class="c-article-author-affiliation__authors-list">Thorgeir E Thorgeirsson, Daniel F Gudbjartsson, Frank Geller, Patrick Sulem, Thorunn Rafnar, Iris H Gudjonsdottir, Solveig Gretarsdottir, Hreinn Stefansson, Jeffrey R Gulcher, Unnur Thorsteinsdottir & Kari Stefansson</p></li><li id="Aff2"><p class="c-article-author-affiliation__address">Center for Biomolecular Science and Engineering, Jack Baskin School of Engineering, University of California, Santa Cruz, California, USA</p><p class="c-article-author-affiliation__authors-list">Thorgeir E Thorgeirsson</p></li><li id="Aff3"><p class="c-article-author-affiliation__address">Institute for Molecular Medicine (FIMM), University of Helsinki, Finland</p><p class="c-article-author-affiliation__authors-list">Ida Surakka, Samuli Ripatti, Shen Huei-Yi, Jaakko Kaprio & Leena Peltonen</p></li><li id="Aff4"><p class="c-article-author-affiliation__address">National Institute for Health and Welfare, Helsinki, Finland</p><p class="c-article-author-affiliation__authors-list">Ida Surakka, Samuli Ripatti, Markus Perola, Shen Huei-Yi & Veikko Salomaa</p></li><li id="Aff5"><p class="c-article-author-affiliation__address">Department of Biological Psychology, Vrije Universiteit (VU) Amsterdam, Amsterdam, The Netherlands</p><p class="c-article-author-affiliation__authors-list">Jacqueline M Vink, Jouke Jan Hottenga, Gonneke Willemsen & Dorret I Boomsma</p></li><li id="Aff6"><p class="c-article-author-affiliation__address">Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands</p><p class="c-article-author-affiliation__authors-list">Najaf Amin, Stefan Walter, Yurii S Aulchenko, André G Uitterlinden, Albert Hofman, Henning Tiemeier & Cornelia M van Duijn</p></li><li id="Aff7"><p class="c-article-author-affiliation__address">Estonian Genome Project of University of Tartu, Tartu, Estonia</p><p class="c-article-author-affiliation__authors-list">Tõnu Esko, Mari Nelis & Andres Metspalu</p></li><li id="Aff8"><p class="c-article-author-affiliation__address">Estonian Biocentre, Tartu, Estonia</p><p class="c-article-author-affiliation__authors-list">Tõnu Esko, Mari Nelis & Andres Metspalu</p></li><li id="Aff9"><p class="c-article-author-affiliation__address">Institute of Epidemiology, Helmholtz Zentrum München, Neuherberg, Germany., </p><p class="c-article-author-affiliation__authors-list">Christian Gieger, Rajesh Rawal, Angela Döring, Barbara Nitz & H-Erich Wichmann</p></li><li id="Aff10"><p class="c-article-author-affiliation__address">Department of Twin Research and Genetic Epidemiology, King's College London, St. Thomas' Hospital Campus, London, UK</p><p class="c-article-author-affiliation__authors-list">Massimo Mangino, Nicole Soranzo, Ana M Valdes, Claire Steves & Tim D Spector</p></li><li id="Aff11"><p class="c-article-author-affiliation__address">Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford, UK</p><p class="c-article-author-affiliation__authors-list">Inga Prokopenko, Reedik Mägi, Maxine Allen & Mark I McCarthy</p></li><li id="Aff12"><p class="c-article-author-affiliation__address">Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK</p><p class="c-article-author-affiliation__authors-list">Inga Prokopenko, Reedik Mägi & Mark I McCarthy</p></li><li id="Aff13"><p class="c-article-author-affiliation__address">Department of Public Health, University of Helsinki, Helsinki, Finland</p><p class="c-article-author-affiliation__authors-list">Kaisu Keskitalo & Jaakko Kaprio</p></li><li id="Aff14"><p class="c-article-author-affiliation__address">Department of Health Sciences and Genetics, University of Leicester, Leicester, UK</p><p class="c-article-author-affiliation__authors-list">John R Thompson</p></li><li id="Aff15"><p class="c-article-author-affiliation__address">Department of Epidemiology, Radboud University Nijmegen Medical Centre, Biostatistics and Health Technology Assessment, Nijmegen, The Netherlands</p><p class="c-article-author-affiliation__authors-list">Katja K Aben, Martin den Heijer & Lambertus A Kiemeney</p></li><li id="Aff16"><p class="c-article-author-affiliation__address">Comprehensive Cancer Centre East, Nijmegen, The Netherlands</p><p class="c-article-author-affiliation__authors-list">Katja K Aben & Lambertus A Kiemeney</p></li><li id="Aff17"><p class="c-article-author-affiliation__address">Department of Endocrinology, Radboud University Nijmegen Medical Centre, Nijmegen, The Netherlands</p><p class="c-article-author-affiliation__authors-list">Martin den Heijer</p></li><li id="Aff18"><p class="c-article-author-affiliation__address">Department of Pulmonary Medicine, Gentofte University Hospital, Hellerup, Denmark</p><p class="c-article-author-affiliation__authors-list">Asger Dirksen & Haseem Ashraf</p></li><li id="Aff19"><p class="c-article-author-affiliation__address">Wellcome Trust Sanger Institute, Hinxton, UK</p><p class="c-article-author-affiliation__authors-list">Nicole Soranzo, Johannes Kettunen & Leena Peltonen</p></li><li id="Aff20"><p class="c-article-author-affiliation__address">Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands</p><p class="c-article-author-affiliation__authors-list">André G Uitterlinden</p></li><li id="Aff21"><p class="c-article-author-affiliation__address">Department of Medicine, University of Leipzig, Leipzig, Germany</p><p class="c-article-author-affiliation__authors-list">Anke Tönjes & Michael Stumvoll</p></li><li id="Aff22"><p class="c-article-author-affiliation__address">Coordination Centre for Clinical Trials, University of Leipzig, Leipzig, Germany</p><p class="c-article-author-affiliation__authors-list">Anke Tönjes</p></li><li id="Aff23"><p class="c-article-author-affiliation__address">Interdisciplinary Centre for Clinical Research, University of Leipzig, Leipzig, Germany</p><p class="c-article-author-affiliation__authors-list">Peter Kovacs</p></li><li id="Aff24"><p class="c-article-author-affiliation__address">Department of Psychiatry, VU University Medical Center, Amsterdam, The Netherlands</p><p class="c-article-author-affiliation__authors-list">Nicole Vogelzangs & Brenda W Penninx</p></li><li id="Aff25"><p class="c-article-author-affiliation__address">Department of Psychiatry, University of Mainz, Mainz, Germany</p><p class="c-article-author-affiliation__authors-list">Norbert Dahmen</p></li><li id="Aff26"><p class="c-article-author-affiliation__address">Department of Psychiatry, Washington University School of Medicine, St. Louis, Missouri, USA., </p><p class="c-article-author-affiliation__authors-list">Michele L Pergadia, Andrew C Heath & Pamela A F Madden</p></li><li id="Aff27"><p class="c-article-author-affiliation__address">Instituto de Nanotecnología de Aragón, Zaragoza, Spain</p><p class="c-article-author-affiliation__authors-list">Berta Saez</p></li><li id="Aff28"><p class="c-article-author-affiliation__address">Division of Dermatology, Hospital San Pedro, Logroño, Spain</p><p class="c-article-author-affiliation__authors-list">Veronica De Diego & Victoria Lezcano</p></li><li id="Aff29"><p class="c-article-author-affiliation__address">Division of Pathology, San Jorge Hospital, Huesca, Spain</p><p class="c-article-author-affiliation__authors-list">Maria D Garcia-Prats</p></li><li id="Aff30"><p class="c-article-author-affiliation__address">Institute of Clinical Medicine, University of Oulu, Oulu, Finland</p><p class="c-article-author-affiliation__authors-list">Anna-Liisa Hartikainen & Matti Isohanni</p></li><li id="Aff31"><p class="c-article-author-affiliation__address">Lifecourse and Service Department, National Institute of Health and Welfare, Oulu, Finland</p><p class="c-article-author-affiliation__authors-list">Anneli Pouta & Marjo-Riitta Järvelin</p></li><li id="Aff32"><p class="c-article-author-affiliation__address">Finnish Institute of Occupational Health, Oulu, Finland</p><p class="c-article-author-affiliation__authors-list">Jaana Laitinen</p></li><li id="Aff33"><p class="c-article-author-affiliation__address">European Bioinformatics Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</p><p class="c-article-author-affiliation__authors-list">Maria Krestyaninova</p></li><li id="Aff34"><p class="c-article-author-affiliation__address">Multidisciplinary Cardiovascular Research Centre (MCRC), Leeds Institute of Genetics, Health and Therapeutics (LIGHT), University of Leeds, Leeds, UK</p><p class="c-article-author-affiliation__authors-list">Alistair S Hall</p></li><li id="Aff35"><p class="c-article-author-affiliation__address">Vascular Research Group, Otago Medical School, Dunedin, New Zealand</p><p class="c-article-author-affiliation__authors-list">Gregory T Jones & Andre M van Rij</p></li><li id="Aff36"><p class="c-article-author-affiliation__address">Department of Laboratory Medicine, Konventhospital Barmherzige Brueder Linz, Linz, Austria</p><p class="c-article-author-affiliation__authors-list">Thomas Mueller, Benjamin Dieplinger & Meinhard Haltmayer</p></li><li id="Aff37"><p class="c-article-author-affiliation__address">Department of Medicine, Landspitali University Hospital, Reykjavik, Iceland</p><p class="c-article-author-affiliation__authors-list">Steinn Jonsson</p></li><li id="Aff38"><p class="c-article-author-affiliation__address">University of Iceland, Faculty of Medicine, Reykjavik, Iceland</p><p class="c-article-author-affiliation__authors-list">Stefan E Matthiasson, Unnur Thorsteinsdottir & Kari Stefansson</p></li><li id="Aff39"><p class="c-article-author-affiliation__address">Therapeia, Reykjavik, Iceland</p><p class="c-article-author-affiliation__authors-list">Hogni Oskarsson</p></li><li id="Aff40"><p class="c-article-author-affiliation__address">National Center of Addiction Medicine, Vogur Hospital, Reykjavik, Iceland</p><p class="c-article-author-affiliation__authors-list">Thorarinn Tyrfingsson</p></li><li id="Aff41"><p class="c-article-author-affiliation__address">Department of Urology, Radboud University Nijmegen Medical Centre, Nijmegen, The Netherlands</p><p class="c-article-author-affiliation__authors-list">Lambertus A Kiemeney</p></li><li id="Aff42"><p class="c-article-author-affiliation__address">University of Zaragoza, Zaragoza, Spain</p><p class="c-article-author-affiliation__authors-list">Jose I Mayordomo</p></li><li id="Aff43"><p class="c-article-author-affiliation__address">Vascular Research Unit, Viborg Hospital, Viborg, Denmark</p><p class="c-article-author-affiliation__authors-list">Jes S Lindholt</p></li><li id="Aff44"><p class="c-article-author-affiliation__address">Department of Cardiothoracic Surgery, Rigshospitalet, University of Copenhagen, Copenhagen, Denmark</p><p class="c-article-author-affiliation__authors-list">Jesper Holst Pedersen</p></li><li id="Aff45"><p class="c-article-author-affiliation__address">Department of Pathology, University of Colorado Denver, Aurora, Colorado, USA., </p><p class="c-article-author-affiliation__authors-list">Wilbur A Franklin</p></li><li id="Aff46"><p class="c-article-author-affiliation__address">Community and Behavioral Health, Colorado School of Public Health, University of Colorado Denver, Aurora, Colorado, USA., </p><p class="c-article-author-affiliation__authors-list">Holly Wolf</p></li><li id="Aff47"><p class="c-article-author-affiliation__address">Queensland Institute of Medical Research, Queensland, Australia</p><p class="c-article-author-affiliation__authors-list">Grant W Montgomery & Nicholas G Martin</p></li><li id="Aff48"><p class="c-article-author-affiliation__address">Department of Psychiatry, University of Munich (LMU), Munich, Germany</p><p class="c-article-author-affiliation__authors-list">Ina Giegling & Dan Rujescu</p></li><li id="Aff49"><p class="c-article-author-affiliation__address">Department of Epidemiology and Public Health, Imperial College, Faculty of Medicine, London, UK</p><p class="c-article-author-affiliation__authors-list">Marjo-Riitta Järvelin</p></li><li id="Aff50"><p class="c-article-author-affiliation__address">Institute of Health Sciences, University of Oulu, Oulu, Finland</p><p class="c-article-author-affiliation__authors-list">Marjo-Riitta Järvelin</p></li><li id="Aff51"><p class="c-article-author-affiliation__address">Biocenter Oulu, University of Oulu, Oulu, Finland</p><p class="c-article-author-affiliation__authors-list">Marjo-Riitta Järvelin</p></li><li id="Aff52"><p class="c-article-author-affiliation__address">Institute of Medical Informatics, Biometry and Epidemiology, Ludwig-Maximilians-Universität, Munich, Germany</p><p class="c-article-author-affiliation__authors-list">H-Erich Wichmann</p></li><li id="Aff53"><p class="c-article-author-affiliation__address">Klinikum Grosshadern, Munich, Germany</p><p class="c-article-author-affiliation__authors-list">H-Erich Wichmann</p></li><li id="Aff54"><p class="c-article-author-affiliation__address">Department of Cardiovascular Sciences, University of Leicester, Clinical Sciences Wing, Glenfield Hospital, Leicester, UK</p><p class="c-article-author-affiliation__authors-list">Nilesh J Samani</p></li><li id="Aff55"><p class="c-article-author-affiliation__address">Department of Clinical Genetics, Erasmus University Medical Center, Rotterdam, The Netherlands</p><p class="c-article-author-affiliation__authors-list">Ben A Oostra</p></li><li id="Aff56"><p class="c-article-author-affiliation__address">Department of Mental Health and Alcohol Abuse Services, National Institute for Health and Welfare, Helsinki, Finland</p><p class="c-article-author-affiliation__authors-list">Jaakko Kaprio</p></li></ol><div class="u-js-hide u-hide-print" data-test="author-info"><span class="c-article__sub-heading">Authors</span><ol class="c-article-authors-search u-list-reset"><li id="auth-Thorgeir_E-Thorgeirsson-Aff1-Aff2"><span class="c-article-authors-search__title u-h3 js-search-name">Thorgeir E Thorgeirsson</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item 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class="c-article__sub-heading" id="contributions">Contributions</h3><p>The study was designed by and the results interpreted by T.E.T., D.F.G., F.G., J.R.G., U.T., K.S., L.P. and M.I.M. The meta-analysis was performed by D.F.G. and F.G., and D.F.G., F.G., I.S., J.M.V., P.S., N.A., T.E., S.W., C.G., R.R., M.M., I.P., R.M., J. Kettunen, Y.S.A., N.S. and J.J.H. were responsible for data analysis in each of the ENGAGE samples. Stage 3 and smoking-related disease samples were coordinated by I.H.G., H.S., S.G. and T.R. Those responsible for case and control ascertainment, recruitment and phenotypic information and project management at the study sites are: J.R.T., W.A.F., H.W., G.W.M., A.C.H., N.G.M., P.A.F.M., K.K.A., M.d.H., L.A.K., G.T.J., A.M.v.R., T.M., B.D., M.H., S.J., T.R., S.E.M., S.G., A.M.V., C.S., A.G.U., A.H., A.T., P.K., G.W., N.V., A. Dirksen, N.D., B.N., M.L.P., B.S., S.R., M.P., J. Kettunen, A.-L.H., A.P., J.L., M.I., A.S.H., T.E.T., H.O., T.T., V.D.D., V.L., M.D.G.-P., J.I.M., A. Döring, H.A., J.S.L., J.H.P., I.G., D.R., M.-R.J., V.S., M.S., T.D.S., H.-E.W., A.M., M.N., N.J.S., B.W.P., B.A.O., D.I.B., H.T., C.M.v.D., J. Kaprio, J.R.G., M.I.M., L.P., U.T. and K.S. Data submission coordination was provided by S.H.-Y., M.A. and M.K. Authors T.E.T., D.F.G. and U.T. wrote the first draft of the paper. All authors contributed to the final version of the paper.</p><h3 class="c-article__sub-heading" id="corresponding-author">Corresponding authors</h3><p id="corresponding-author-list">Correspondence to <a id="corresp-c1" href="mailto:unnur.thorsteinsdottir@decode.is">Unnur Thorsteinsdottir</a> or <a id="corresp-c2" href="mailto:kstefans@decode.is">Kari Stefansson</a>.</p></div></div></section><section data-title="Ethics declarations"><div class="c-article-section" id="ethics-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="ethics">Ethics declarations</h2><div class="c-article-section__content" id="ethics-content"> <h3 class="c-article__sub-heading">Competing interests</h3> <p>Authors whose affiliations are listed as deCODE genetics are employees of deCODE genetics, a biotechnology company.</p> </div></div></section><section data-title="Additional information"><div class="c-article-section" id="additional-information-section"><h2 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Surakka, I. <i>et al.</i> Sequence variants at <i>CHRNB3</i>–<i>CHRNA6</i> and <i>CYP2A6</i> affect smoking behavior. <i>Nat Genet</i> <b>42</b>, 448–453 (2010). https://doi.org/10.1038/ng.573</p><p class="c-bibliographic-information__download-citation u-hide-print"><a data-test="citation-link" data-track="click" data-track-action="download article citation" data-track-label="link" data-track-external="" rel="nofollow" href="https://citation-needed.springer.com/v2/references/10.1038/ng.573?format=refman&flavour=citation">Download citation<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-download-medium"></use></svg></a></p><ul class="c-bibliographic-information__list" data-test="publication-history"><li class="c-bibliographic-information__list-item"><p>Received<span class="u-hide">: </span><span class="c-bibliographic-information__value"><time datetime="2009-10-21">21 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Sequence variants at CHRNB3–CHRNA6 and CYP2A6 affect smoking behavior | Nature Genetics