**SCARF:** We thank all participants in this study. The study was funded by the Foundation for Strategic Research, the Swedish Heart-Lung Foundation, the Swedish Research Council (8691, 12660, 20653), the European Commission (LSHM-CT-2007-037273), the Knut and Alice Wallenberg Foundation, the Torsten and Ragnar Söderberg Foundation, the Strategic Cardiovascular and Diabetes Programmes of Karolinska Institutet and the Stockholm County Council, and the Stockholm County Council (560183). Bengt Sennblad acknowledges funding from the Magnus Bergvall Foundation and the Foundation for Old Servants. Mattias Frånberg acknowledges funding from the Swedish e-Science Research Center (SeRC).

**SCES:** The Singapore Chinese Eye Study (SCES) was supported by the National Medical Research Council (NMRC), Singapore (grants 0796/2003, IRG07nov013, IRG09nov014, NMRC 1176/2008, STaR/0003/2008, CG/SERI/2010) and Biomedical Research Council (BMRC), Singapore (08/1/35/19/550 and 09/1/35/19/616).

**TEENAGE (TEENS of Attica: Genes and Environment):** This research has been co-financed by the European Union (European Social Fund—ESF) and Greek national funds through the Operational Program 'Education and Lifelong Learning' of the National Strategic Reference Framework (NSRF)—Research Funding Program: Heracleitus II. Investing in knowledge society through the European Social Fund. This work was funded by the Wellcome Trust (098051).

We thank all study participants and their families as well as all volunteers for their contribution in this study. We thank the Sample Management and Genotyping Facilities staff at the Wellcome Trust Sanger Institute for sample preparation, quality control and genotyping.

**Uppsala Longitudinal Study of Adult Men (ULSAM):** The exome chip genotyping and data analyses were supported by Uppsala University, Knut och Alice Wallenberg Foundation, European Research Council, Swedish Diabetes Foundation (grant no. 2013-024), Swedish Research Council (grant no. 2012-1397), and Swedish Heart-Lung Foundation (20120197). C.M.L. is supported by a Wellcome Trust Research Career Development Fellowship (086596/Z/08/Z).

**INGI-VB:** The Val Borbera study (INGI-VB) thanks the inhabitants of the Val Borbera for participating in the study, the local administrations and the ASL-Novi Ligure for support and Fiammetta Viganò for technical help. We also thank Professor Clara Camaschella, Professor Federico Caligaris-Cappio and the MDs of the Medicine Dept. of the San Raffaele Hospital for help with clinical data collection. The study was supported by funds from Fondazione Compagnia di San Paolo-Torino, Fondazione Cariplo-Milano, Italian Ministry of Health Progetto Finalizzato 2007 and 2012, Italian Ministry of Health Progetto CCM 2010, and PRIN 2009.

**WGHS:** The WGHS is supported by HL043851 and HL080467 from the National Heart, Lung, and Blood Institute and CA047988 from the National Cancer Institute, the Donald W. Reynolds Foundation and the Fondation Leducq, with collaborative scientific support and funding for genotyping provided by Amgen.

## Author contributions

**Writing group:** J.W., A.Y.C., S.M.W., S.W., H.Y., J.A.B., M.D., M.-F.H., S.R., K.F., L.L., B.H., R.A., J.B.B., M.S., J.C.F., J.D., J.B.M., J.I.R., R.A.S., M.O.G.

**Project, design, management and coordination:** J.D., B.M.P., D.S.S., J.B.M., J.I.R., R.A.S., M.O.G.

**Cohort PI:** R.A., A.C., Y.L., D.M.B., L.A.C., G.G., T.J., E.I., A.J.K., C.L., R.A.M., J.M.N., W.H.-H.S., D.T., D.V., R.V., L.E.W., H.B., E.P.B., G.D., E.F., M.F., O.H.F., P.W.F., R.A.G.,

V.G., A.H., A.T.H., C.H., A. Hofman, J.-H.J., D.L., A.L., B.A.O., C.J.O., S.P., J.S.P., M.A.P., S.S.R., P.M.R., I.R., M.B.S., B.S., A.G.U., M.W., N.J.W., H.W., T.Y.W., E.Z., J.K., M.L., I.B.B., D.I.C., B.M.P., C.M.v.D., D.M.W., E.B., W.H.L.K., R.J.F.L., T.M.F., J.I.R.

**Sample collection and phenotyping:** M.D., M.-F.H., S.R., L.L., F.K., N.G., A.S., M.G., A.S., T.A., N.A.B., Y.-D.I.C., C.Y.C., A.C., A.D., G.B.E., G.E., S.A.E., A.-E.F., O.G., M.L.G., G.H., M.K.I., M.E.J., T.J., M.K., A.T.K., J.K., I.T.L., W.-J.L., A.S.L., C.L., A.L., A.M., R. McKean-Cowdin, O. McLeod, I.N., A.P., N.W.R., I.S., J.A.S., N.T., M.T., E.T., D.M.B., G.G., E.I., C.L., J.M.N., W.H.-H.S., D.V., R.V., H.B., E.P.B., V.G., T.B.H., C.H., A.H., C.L., L.L., D.L., S.P., O.P., M.A.P., P.M.R., M.B.S., B.S., N.J.W., M.L., B.M.P., E.S.T., C.M.v.D., D.M.W., J.C.F., J.G.W., D.S.S., R.A.S.

**Genotyping:** A.Y.C., J.B., N.G., J.B.-J., M.F., J.H.Z., A.C.M., L.S., K.D.T., J.B.-J., K.H.A., J.L.A., C.B., D.W.B., Y.-D.I.C., C.Y.C., M.F., F.G., A.G., T.H., P.H., C.C.K., G.M., D.M., I.N., N.D.P., O.P., B.S., N.S., E.K.S., E.A.S., C.B., A.B., K.S., J.C.B., M.B., K.M., E.I., R.A.M., E.P.B., P.D., A.Hofman, C.L., D.L., M.A.P., A.G.U., N.J.W., D.I.C., E.S.T., C.M.v.D., D.M.W., J.I.R., R.A.S., M.O.G.

**Statistical Analysis:** J.W., A.Y.C., S.M.W., S.W., H.Y., J.B., M.D., M.-F.H., S.R., B.H., F.K., J.E.H., P.A., Y.C.L., L.J.R.-T., N.G., M.G.E., L.L., A.S.B., A.S., R.A., J.B.—J., D.F.F., XG., K.H., A.I., J.J., L.A.L., J.C.L., M.L., J.H.Z., K.M., M.A.N., M.J.P., M.S.-L., C.S., A.V.S., L.S., M.H.S., R.J.S., T.V.V., N.A., C.B., S.M.B., Y.C., J.C., F.G., W.A.G.III, S.G., Y.H., J.H., M.K.I., R.A.J., A.K., A.T.K., E.M.L., J.L., C.L., C.M.L., G.M., N.M.M., N.D.P., D.P., F.R., K.R., C.F.S., J.A.S., N.S., K.S., M.T., S.J., L.R.Y., J.B., J.B.B., G.M.P., D.I.C., D.M.W., J.D., J.I.R., R.A.S.

## Additional information

**Supplementary Information** accompanies this paper at http://www.nature.com/naturecommunications

**Competing financial interests:** J.C.F. has received consulting honoraria from PanGenX and Pfizer; T.F. consulted for Boeringer Ingelheim; J.B.M. serves as a consultant to LipoScience, and Quest Diagnostics; B.P. serves on the DSMB of a clinical trial for a device funded by the manufacturer (Zoll LifeCor) and on the Steering Committee for the Yale Open Data Access Project funded by Johnson & Johnson; D.M.W., M.G.E., L.L. and J.A. are all full time employees of GlaxoSmithKline. P.M.R. and D.I.C. have research grant support from Amgen, AstraZeneca and the NHLBI. The remaining authors declare no competing financial interests.

**Reprints and permission** information is available online at http://npg.nature.com/reprintsandpermissions/

**How to cite this article:** Wessel, J. et al. Low-frequency and rare exome chip variants associate with fasting glucose and type 2 diabetes susceptibility. *Nat. Commun.* 6:5897 doi: 10.1038/ncomms6897 (2015).

<figure><img src="image_5.png" /></figure>

This work is licensed under a Creative Commons Attribution 4.0 International License. The images or other third party material in this article are included in the article's Creative Commons license, unless indicated otherwise

in the credit line; if the material is not included under the Creative Commons license, users will need to obtain permission from the license holder to reproduce the material. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/

Jennifer Wessel<sup>1,2,*</sup>, Audrey Y. Chu<sup>3,4,*</sup>, Sara M. Willems<sup>5,6,*</sup>, Shuai Wang<sup>7,*</sup>, Hanieh Yaghootkar<sup>8</sup>, Jennifer A. Brody<sup>9,10</sup>, Marco Dauriz<sup>11,12,13</sup>, Marie-France Hivert<sup>14,15,16</sup>, Sridharan Raghavan<sup>11,12</sup>, Leonard Lipovich<sup>17,18</sup>, Bertha Hidalgo<sup>19</sup>, Keolu Fox<sup>10,20</sup>, Jennifer E. Huffman<sup>4,21</sup>, Ping An<sup>22</sup>, Yingchang Lu<sup>23,24</sup>, Laura J. Rasmussen-Torvik<sup>25</sup>, Niels Grarup<sup>26</sup>, Margaret G. Ehm<sup>27</sup>, Li Li<sup>27</sup>, Abigail S. Baldridge<sup>25</sup>, Alena Stančáková<sup>28</sup>, Ravinder Abrol<sup>29,30</sup>, Céline Besse<sup>31</sup>, Anne Boland<sup>31</sup>, Jette Bork-Jensen<sup>26</sup>, Myriam Fornage<sup>32</sup>, Daniel F. Freitag<sup>33,34</sup>, Melissa E. Garcia<sup>35</sup>, Xiuqing Guo<sup>36</sup>, Kazuo Hara<sup>23,24</sup>, Aaron Isaacs<sup>5</sup>, Johanna Jakobsdottir<sup>37</sup>, Leslie A. Lange<sup>38</sup>, Jill C. Layton<sup>39</sup>, Man Li<sup>40</sup>, Jing Hua Zhao<sup>6</sup>, Karina Meidtner<sup>41</sup>, Alanna C. Morrison<sup>42</sup>, Mike A. Nalls<sup>43</sup>, Marjolein J. Peters<sup>44,45</sup>, Maria Sabater-Lleal<sup>46</sup>, Claudia Schurmann<sup>23,24</sup>, Angela Silveira<sup>46</sup>, Albert V. Smith<sup>37,47</sup>, Lorraine Southam<sup>33,48</sup>, Marcus H. Stoiber<sup>49</sup>, Rona J. Strawbridge<sup>46</sup>, Kent D. Taylor<sup>36</sup>, Tibor V. Varga<sup>50</sup>, Kristine H. Allin<sup>26</sup>, Najaf Amin<sup>5</sup>, Jennifer L. Aponte<sup>27</sup>, Tin Aung<sup>51,52</sup>, Caterina Barbieri<sup>53</sup>, Nathan A. Bihlmeyer<sup>54,55</sup>, Michael Boehnke<sup>56</sup>, Cristina Bombieri<sup>57</sup>, Donald W. Bowden<sup>58</sup>, Sean M. Burns<sup>16</sup>, Yuning Chen<sup>7</sup>, Yii-Derl Chen<sup>36</sup>, Ching-Yu Cheng<sup>51,52,59,60</sup>, Adolfo Correa<sup>61</sup>, Jacek Czajkowski<sup>22</sup>, Abbas Dehghan<sup>62</sup>, Georg B. Ehret<sup>63,64</sup>, Gudny Eiriksdottir<sup>37</sup>, Stefan A. Escher<sup>50</sup>, Aliki-Eleni Farmaki<sup>65</sup>, Mattias Frånberg<sup>46,66</sup>, Giovanni Gambaro<sup>67</sup>, Franco Giulianini<sup>3</sup>, William A. Goddard III<sup>30</sup>, Anuj Goel<sup>68</sup>, Omri Gottesman<sup>23</sup>, Megan L. Grove<sup>42</sup>, Stefan Gustafsson<sup>69</sup>, Yang Hai<sup>36</sup>, Göran Hallmans<sup>70</sup>,
**SCARF:** We thank all participants in this study. The study was funded by the Foundation for Strategic Research, the Swedish Heart-Lung Foundation, the Swedish Research Council (8691, 12660, 20653), the European Commission (LSHM-CT-2007-037273), the Knut and Alice Wallenberg Foundation, the Torsten and Ragnar Söderberg Foundation, the Strategic Cardiovascular and Diabetes Programmes of Karolinska Institutet and the Stockholm County Council, and the Stockholm County Council (560183). Bengt Sennblad acknowledges funding from the Magnus Bergvall Foundation and the Foundation for Old Servants. Mattias Frånberg acknowledges funding from the Swedish e-Science Research Center (SeRC).

**SCES:** The Singapore Chinese Eye Study (SCES) was supported by the National Medical Research Council (NMRC), Singapore (grants 0796/2003, IRG07nov013, IRG09nov014, NMRC 1176/2008, STaR/0003/2008, CG/SERI/2010) and Biomedical Research Council (BMRC), Singapore (08/1/35/19/550 and 09/1/35/19/616).

**TEENAGE (TEENS of Attica: Genes and Environment):** This research has been co-financed by the European Union (European Social Fund—ESF) and Greek national funds through the Operational Program 'Education and Lifelong Learning' of the National Strategic Reference Framework (NSRF)—Research Funding Program: Heracleitus II. Investing in knowledge society through the European Social Fund. This work was funded by the Wellcome Trust (098051).

We thank all study participants and their families as well as all volunteers for their contribution in this study. We thank the Sample Management and Genotyping Facilities staff at the Wellcome Trust Sanger Institute for sample preparation, quality control and genotyping.

**Uppsala Longitudinal Study of Adult Men (ULSAM):** The exome chip genotyping and data analyses were supported by Uppsala University, Knut och Alice Wallenberg Foundation, European Research Council, Swedish Diabetes Foundation (grant no. 2013-024), Swedish Research Council (grant no. 2012-1397), and Swedish Heart-Lung Foundation (20120197). C.M.L. is supported by a Wellcome Trust Research Career Development Fellowship (086596/Z/08/Z).

**INGI-VB:** The Val Borbera study (INGI-VB) thanks the inhabitants of the Val Borbera for participating in the study, the local administrations and the ASL-Novi Ligure for support and Fiammetta Viganò for technical help. We also thank Professor Clara Camaschella, Professor Federico Caligaris-Cappio and the MDs of the Medicine Dept. of the San Raffaele Hospital for help with clinical data collection. The study was supported by funds from Fondazione Compagnia di San Paolo-Torino, Fondazione Cariplo-Milano, Italian Ministry of Health Progetto Finalizzato 2007 and 2012, Italian Ministry of Health Progetto CCM 2010, and PRIN 2009.

**WGHS:** The WGHS is supported by HL043851 and HL080467 from the National Heart, Lung, and Blood Institute and CA047988 from the National Cancer Institute, the Donald W. Reynolds Foundation and the Fondation Leducq, with collaborative scientific support and funding for genotyping provided by Amgen.

## Author contributions

**Writing group:** J.W., A.Y.C., S.M.W., S.W., H.Y., J.A.B., M.D., M.-F.H., S.R., K.F., L.L., B.H., R.A., J.B.B., M.S., J.C.F., J.D., J.B.M., J.I.R., R.A.S., M.O.G.

**Project, design, management and coordination:** J.D., B.M.P., D.S.S., J.B.M., J.I.R., R.A.S., M.O.G.

**Cohort PI:** R.A., A.C., Y.L., D.M.B., L.A.C., G.G., T.J., E.I., A.J.K., C.L., R.A.M., J.M.N., W.H.-H.S., D.T., D.V., R.V., L.E.W., H.B., E.P.B., G.D., E.F., M.F., O.H.F., P.W.F., R.A.G.,

V.G., A.H., A.T.H., C.H., A. Hofman, J.-H.J., D.L., A.L., B.A.O., C.J.O., S.P., J.S.P., M.A.P., S.S.R., P.M.R., I.R., M.B.S., B.S., A.G.U., M.W., N.J.W., H.W., T.Y.W., E.Z., J.K., M.L., I.B.B., D.I.C., B.M.P., C.M.v.D., D.M.W., E.B., W.H.L.K., R.J.F.L., T.M.F., J.I.R.

**Sample collection and phenotyping:** M.D., M.-F.H., S.R., L.L., F.K., N.G., A.S., M.G., A.S., T.A., N.A.B., Y.-D.I.C., C.Y.C., A.C., A.D., G.B.E., G.E., S.A.E., A.-E.F., O.G., M.L.G., G.H., M.K.I., M.E.J., T.J., M.K., A.T.K., J.K., I.T.L., W.-J.L., A.S.L., C.L., A.L., A.M., R. McKean-Cowdin, O. McLeod, I.N., A.P., N.W.R., I.S., J.A.S., N.T., M.T., E.T., D.M.B., G.G., E.I., C.L., J.M.N., W.H.-H.S., D.V., R.V., H.B., E.P.B., V.G., T.B.H., C.H., A.H., C.L., L.L., D.L., S.P., O.P., M.A.P., P.M.R., M.B.S., B.S., N.J.W., M.L., B.M.P., E.S.T., C.M.v.D., D.M.W., J.C.F., J.G.W., D.S.S., R.A.S.

**Genotyping:** A.Y.C., J.B., N.G., J.B.-J., M.F., J.H.Z., A.C.M., L.S., K.D.T., J.B.-J., K.H.A., J.L.A., C.B., D.W.B., Y.-D.I.C., C.Y.C., M.F., F.G., A.G., T.H., P.H., C.C.K., G.M., D.M., I.N., N.D.P., O.P., B.S., N.S., E.K.S., E.A.S., C.B., A.B., K.S., J.C.B., M.B., K.M., E.I., R.A.M., E.P.B., P.D., A.Hofman, C.L., D.L., M.A.P., A.G.U., N.J.W., D.I.C., E.S.T., C.M.v.D., D.M.W., J.I.R., R.A.S., M.O.G.

**Statistical Analysis:** J.W., A.Y.C., S.M.W., S.W., H.Y., J.B., M.D., M.-F.H., S.R., B.H., F.K., J.E.H., P.A., Y.C.L., L.J.R.-T., N.G., M.G.E., L.L., A.S.B., A.S., R.A., J.B.—J., D.F.F., XG., K.H., A.I., J.J., L.A.L., J.C.L., M.L., J.H.Z., K.M., M.A.N., M.J.P., M.S.-L., C.S., A.V.S., L.S., M.H.S., R.J.S., T.V.V., N.A., C.B., S.M.B., Y.C., J.C., F.G., W.A.G.III, S.G., Y.H., J.H., M.K.I., R.A.J., A.K., A.T.K., E.M.L., J.L., C.L., C.M.L., G.M., N.M.M., N.D.P., D.P., F.R., K.R., C.F.S., J.A.S., N.S., K.S., M.T., S.J., L.R.Y., J.B., J.B.B., G.M.P., D.I.C., D.M.W., J.D., J.I.R., R.A.S.

## Additional information

**Supplementary Information** accompanies this paper at http://www.nature.com/naturecommunications

**Competing financial interests:** J.C.F. has received consulting honoraria from PanGenX and Pfizer; T.F. consulted for Boeringer Ingelheim; J.B.M. serves as a consultant to LipoScience, and Quest Diagnostics; B.P. serves on the DSMB of a clinical trial for a device funded by the manufacturer (Zoll LifeCor) and on the Steering Committee for the Yale Open Data Access Project funded by Johnson & Johnson; D.M.W., M.G.E., L.L. and J.A. are all full time employees of GlaxoSmithKline. P.M.R. and D.I.C. have research grant support from Amgen, AstraZeneca and the NHLBI. The remaining authors declare no competing financial interests.

**Reprints and permission** information is available online at http://npg.nature.com/reprintsandpermissions/

**How to cite this article:** Wessel, J. et al. Low-frequency and rare exome chip variants associate with fasting glucose and type 2 diabetes susceptibility. *Nat. Commun.* 6:5897 doi: 10.1038/ncomms6897 (2015).

<figure><img src="image_10.png" /></figure>

This work is licensed under a Creative Commons Attribution 4.0 International License. The images or other third party material in this article are included in the article's Creative Commons license, unless indicated otherwise

in the credit line; if the material is not included under the Creative Commons license, users will need to obtain permission from the license holder to reproduce the material. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/

Jennifer Wessel<sup>1,2,*</sup>, Audrey Y. Chu<sup>3,4,*</sup>, Sara M. Willems<sup>5,6,*</sup>, Shuai Wang<sup>7,*</sup>, Hanieh Yaghootkar<sup>8</sup>, Jennifer A. Brody<sup>9,10</sup>, Marco Dauriz<sup>11,12,13</sup>, Marie-France Hivert<sup>14,15,16</sup>, Sridharan Raghavan<sup>11,12</sup>, Leonard Lipovich<sup>17,18</sup>, Bertha Hidalgo<sup>19</sup>, Keolu Fox<sup>10,20</sup>, Jennifer E. Huffman<sup>4,21</sup>, Ping An<sup>22</sup>, Yingchang Lu<sup>23,24</sup>, Laura J. Rasmussen-Torvik<sup>25</sup>, Niels Grarup<sup>26</sup>, Margaret G. Ehm<sup>27</sup>, Li Li<sup>27</sup>, Abigail S. Baldridge<sup>25</sup>, Alena Stančáková<sup>28</sup>, Ravinder Abrol<sup>29,30</sup>, Céline Besse<sup>31</sup>, Anne Boland<sup>31</sup>, Jette Bork-Jensen<sup>26</sup>, Myriam Fornage<sup>32</sup>, Daniel F. Freitag<sup>33,34</sup>, Melissa E. Garcia<sup>35</sup>, Xiuqing Guo<sup>36</sup>, Kazuo Hara<sup>23,24</sup>, Aaron Isaacs<sup>5</sup>, Johanna Jakobsdottir<sup>37</sup>, Leslie A. Lange<sup>38</sup>, Jill C. Layton<sup>39</sup>, Man Li<sup>40</sup>, Jing Hua Zhao<sup>6</sup>, Karina Meidtner<sup>41</sup>, Alanna C. Morrison<sup>42</sup>, Mike A. Nalls<sup>43</sup>, Marjolein J. Peters<sup>44,45</sup>, Maria Sabater-Lleal<sup>46</sup>, Claudia Schurmann<sup>23,24</sup>, Angela Silveira<sup>46</sup>, Albert V. Smith<sup>37,47</sup>, Lorraine Southam<sup>33,48</sup>, Marcus H. Stoiber<sup>49</sup>, Rona J. Strawbridge<sup>46</sup>, Kent D. Taylor<sup>36</sup>, Tibor V. Varga<sup>50</sup>, Kristine H. Allin<sup>26</sup>, Najaf Amin<sup>5</sup>, Jennifer L. Aponte<sup>27</sup>, Tin Aung<sup>51,52</sup>, Caterina Barbieri<sup>53</sup>, Nathan A. Bihlmeyer<sup>54,55</sup>, Michael Boehnke<sup>56</sup>, Cristina Bombieri<sup>57</sup>, Donald W. Bowden<sup>58</sup>, Sean M. Burns<sup>16</sup>, Yuning Chen<sup>7</sup>, Yii-Derl Chen<sup>36</sup>, Ching-Yu Cheng<sup>51,52,59,60</sup>, Adolfo Correa<sup>61</sup>, Jacek Czajkowski<sup>22</sup>, Abbas Dehghan<sup>62</sup>, Georg B. Ehret<sup>63,64</sup>, Gudny Eiriksdottir<sup>37</sup>, Stefan A. Escher<sup>50</sup>, Aliki-Eleni Farmaki<sup>65</sup>, Mattias Frånberg<sup>46,66</sup>, Giovanni Gambaro<sup>67</sup>, Franco Giulianini<sup>3</sup>, William A. Goddard III<sup>30</sup>, Anuj Goel<sup>68</sup>, Omri Gottesman<sup>23</sup>, Megan L. Grove<sup>42</sup>, Stefan Gustafsson<sup>69</sup>, Yang Hai<sup>36</sup>, Göran Hallmans<sup>70</sup>,
Jiyoung Heo<sup>71</sup>, Per Hoffmann<sup>72,73,74</sup>, Mohammad K. Ikram<sup>51,60,75</sup>, Richard A. Jensen<sup>9,10</sup>, Marit E. Jørgensen<sup>76</sup>, Torben Jørgensen<sup>77,78</sup>, Maria Karaleftheri<sup>79</sup>, Chiea C. Khor<sup>52,59,80</sup>, Andrea Kirkpatrick<sup>30</sup>, Aldi T. Kraja<sup>22</sup>, Johanna Kuusisto<sup>81</sup>, Ethan M. Lange<sup>38,82</sup>, I.T. Lee<sup>83,84</sup>, Wen-Jane Lee<sup>85</sup>, Aaron Leong<sup>11,12</sup>, Jiemin Liao<sup>51,52</sup>, Chunyu Liu<sup>4</sup>, Yongmei Liu<sup>86</sup>, Cecilia M. Lindgren<sup>87</sup>, Allan Linneberg<sup>77,88,89</sup>, Giovanni Malerba<sup>57</sup>, Vasiliki Mamakou<sup>90,91</sup>, Eirini Marouli<sup>65</sup>, Nisa M. Maruthur<sup>92,93</sup>, Angela Matchan<sup>33</sup>, Roberta McKean-Cowdin<sup>94</sup>, Olga McLeod<sup>46</sup>, Ginger A. Metcalf<sup>95</sup>, Karen L. Mohlke<sup>38</sup>, Donna M. Muzny<sup>95</sup>, Ioanna Ntalla<sup>65,96</sup>, Nicholette D. Palmer<sup>58,97</sup>, Dorota Pasko<sup>8</sup>, Andreas Peter<sup>98,99</sup>, Nigel W. Rayner<sup>33,48,100</sup>, Frida Renström<sup>50</sup>, Ken Rice<sup>9,101</sup>, Cinzia F. Sala<sup>53</sup>, Bengt Sennblad<sup>46,102</sup>, Ioannis Serafetinidis<sup>103</sup>, Jennifer A. Smith<sup>104</sup>, Nicole Soranzo<sup>33,105</sup>, Elizabeth K. Speliotes<sup>106</sup>, Eli A. Stahl<sup>107</sup>, Kathleen Stirrups<sup>33,108</sup>, Nikos Tentolouris<sup>109</sup>, Anastasia Thanopoulou<sup>110</sup>, Mina Torres<sup>94</sup>, Michela Traglia<sup>53</sup>, Emmanouil Tsafantakis<sup>111</sup>, Sundas Javad<sup>6</sup>, Lisa R. Yanek<sup>112</sup>, Eleni Zengini<sup>91,113</sup>, Diane M. Becker<sup>112</sup>, Joshua C. Bis<sup>9,10</sup>, James B. Brown<sup>49,114</sup>, L. Adrienne Cupples<sup>4,7</sup>, Torben Hansen<sup>26,115</sup>, Erik Ingelsson<sup>69,87</sup>, Andrew J. Karter<sup>116</sup>, Carlos Lorenzo<sup>117</sup>, Rasika A. Mathias<sup>112</sup>, Jill M. Norris<sup>118</sup>, Gina M. Peloso<sup>119,120</sup>, Wayne H.-H. Sheu<sup>83,84,121</sup>, Daniela Toniolo<sup>53</sup>, Dhananjay Vaidya<sup>112</sup>, Rohit Varma<sup>94</sup>, Lynne E. Wagenknecht<sup>122</sup>, Heiner Boeing<sup>123</sup>, Erwin P. Bottinger<sup>23</sup>, George Dedoussis<sup>65</sup>, Panos Deloukas<sup>108,124,125</sup>, Ele Ferrannini<sup>126</sup>, Oscar H. Franco<sup>62</sup>, Paul W. Franks<sup>50,127,128</sup>, Richard A. Gibbs<sup>95</sup>, Vilmundur Gudnason<sup>37,47</sup>, Anders Hamsten<sup>46</sup>, Tamara B. Harris<sup>35</sup>, Andrew T. Hattersley<sup>129</sup>, Caroline Hayward<sup>21</sup>, Albert Hofman<sup>62</sup>, Jan-Håkan Jansson<sup>128,130</sup>, Claudia Langenberg<sup>6</sup>, Lenore J. Launer<sup>35</sup>, Daniel Levy<sup>131,132</sup>, Ben A. Oostra<sup>5</sup>, Christopher J. O'Donnell<sup>4,12,133</sup>, Stephen O'Rahilly<sup>134</sup>, Sandosh Padmanabhan<sup>135</sup>, James S. Pankow<sup>136</sup>, Ozren Polasek<sup>137</sup>, Michael A. Province<sup>22</sup>, Stephen S. Rich<sup>138</sup>, Paul M. Ridker<sup>3,139</sup>, Igor Rudan<sup>140</sup>, Matthias B. Schulze<sup>41,99</sup>, Blair H. Smith<sup>141</sup>, André G. Uitterlinden<sup>44,62</sup>, Mark Walker<sup>142</sup>, Hugh Watkins<sup>68</sup>, Tien Y. Wong<sup>51,52,60</sup>, Eleftheria Zeggini<sup>33</sup>, The EPIC-InterAct Consortium<sup>§</sup>, Markku Laakso<sup>81</sup>, Ingrid B. Borecki<sup>22</sup>, Daniel I. Chasman<sup>3,143</sup>, Oluf Pedersen<sup>26</sup>, Bruce M. Psaty<sup>9,10,144,145,146</sup>, E. Shyong Tai<sup>59,147</sup>, Cornelia M. van Duijn<sup>5,148</sup>, Nicholas J. Wareham<sup>6</sup>, Dawn M. Waterworth<sup>149</sup>, Eric Boerwinkle<sup>42,95</sup>, W.H. Linda Kao<sup>40,93,150</sup>, Jose C. Florez<sup>12,16,119,120</sup>, Ruth J.F. Loos<sup>23,24,151</sup>, James G. Wilson<sup>152</sup>, Timothy M. Frayling<sup>8</sup>, David S. Siscovick<sup>153,154</sup>, Josée Dupuis<sup>4,7</sup>, Jerome I. Rotter<sup>36</sup>, James B. Meigs<sup>11,12</sup>, Robert A. Scott<sup>6</sup> & Mark O. Goodarzi<sup>29,155</sup>

<footnote n="1"><sup>1</sup>Department of Epidemiology, Fairbanks School of Public Health, Indianapolis, Indiana 46202, USA. <sup>2</sup>Department of Medicine, Indiana University School of Medicine, Indianapolis, Indiana 46202, USA. <sup>3</sup>Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts 02215, USA. <sup>4</sup>National Heart, Lung, and Blood Institute (NHLBI) Framingham Heart Study, Framingham, Massachusetts 01702, USA. <sup>5</sup>Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam 3000 CE, The Netherlands. <sup>6</sup>MRC Epidemiology Unit, University of Cambridge School of Clinical Medicine, Institute of Metabolic Science, Cambridge Biomedical Campus, Cambridge CB2 OSL, UK. <sup>7</sup>Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts 02118, USA. <sup>8</sup>Genetics of Complex Traits, University of Exeter Medical School, University of Exeter, Exeter EX1 2LU, UK. <sup>9</sup>Cardiovascular Health Research Unit, University of Washington, Seattle, Washington 98101, USA. <sup>10</sup>Department of Medicine, University of Washington, Seattle, Washington 98195, USA. <sup>11</sup>Massachusetts General Hospital, General Medicine Division, Boston, Massachusetts 02114, USA. <sup>12</sup>Department of Medicine, Harvard Medical School, Boston, Massachusetts 02115, USA. <sup>13</sup>Division of Endocrinology, Diabetes and Metabolism, Department of Medicine, University of Verona Medical School and Hospital Trust of Verona, Verona 37126, Italy. <sup>14</sup>Harvard Pilgrim Health Care Institute, Department of Population Medicine, Harvard Medical School, Boston, Massachusetts 02215, USA. <sup>15</sup>Division of Endocrinology and Metabolism, Department of Medicine, Université de Sherbrooke, Sherbrooke, Québec, Canada J1K 2R1. <sup>16</sup>Diabetes Unit, Department of Medicine, Massachusetts General Hospital, Boston, Massachusetts 02114, USA. <sup>17</sup>Center for Molecular Medicine and Genetics, Wayne State University, Detroit, Michigan 48201, USA. <sup>18</sup>Department of Neurology, Wayne State University School of Medicine, Detroit, Michigan 48202, USA. <sup>19</sup>Department of Epidemiology, University of Alabama at Birmingham, Birmingham, Alabama 35233, USA. <sup>20</sup>Department of Genome Sciences, University of Washington, Seattle, Washington 98195, USA. <sup>21</sup>MRC Human Genetics Unit, MRC IGMM, University of Edinburgh, Edinburgh, Scotland EH4 2XU, UK. <sup>22</sup>Division of Statistical Genomics and Department of Genetics, Washington University School of Medicine, St. Louis, Missouri 63108, USA. <sup>23</sup>The Charles Bronfman Institute for Personalized Medicine, The Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA. <sup>24</sup>The Genetics of Obesity and Related Metabolic Traits Program, The Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA. <sup>25</sup>Department of Preventive Medicine, Northwestern University Feinberg School of Medicine, Chicago, Illinois 60611, USA. <sup>26</sup>The Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen DK-2200, Denmark. <sup>27</sup>Quantitative Sciences, PCPS, GlaxoSmithKline, North Carolina 27709, USA. <sup>28</sup>Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland, Kuopio FI-70211, Finland. <sup>29</sup>Department of Medicine and Department of Biomedical Sciences, Cedars-Sinai Medical Center, Los Angeles, California 90048, USA. <sup>30</sup>Materials and Process Simulation Center, California Institute of Technology, Pasadena, California 91125, USA. <sup>31</sup>CEA, Institut de Génomique, Centre National de Génotypage, 2 Rue Gaston Crémieux, EVRY Cedex 91057, France. <sup>32</sup>Brown</footnote>
Jiyoung Heo<sup>71</sup>, Per Hoffmann<sup>72,73,74</sup>, Mohammad K. Ikram<sup>51,60,75</sup>, Richard A. Jensen<sup>9,10</sup>, Marit E. Jørgensen<sup>76</sup>, Torben Jørgensen<sup>77,78</sup>, Maria Karaleftheri<sup>79</sup>, Chiea C. Khor<sup>52,59,80</sup>, Andrea Kirkpatrick<sup>30</sup>, Aldi T. Kraja<sup>22</sup>, Johanna Kuusisto<sup>81</sup>, Ethan M. Lange<sup>38,82</sup>, I.T. Lee<sup>83,84</sup>, Wen-Jane Lee<sup>85</sup>, Aaron Leong<sup>11,12</sup>, Jiemin Liao<sup>51,52</sup>, Chunyu Liu<sup>4</sup>, Yongmei Liu<sup>86</sup>, Cecilia M. Lindgren<sup>87</sup>, Allan Linneberg<sup>77,88,89</sup>, Giovanni Malerba<sup>57</sup>, Vasiliki Mamakou<sup>90,91</sup>, Eirini Marouli<sup>65</sup>, Nisa M. Maruthur<sup>92,93</sup>, Angela Matchan<sup>33</sup>, Roberta McKean-Cowdin<sup>94</sup>, Olga McLeod<sup>46</sup>, Ginger A. Metcalf<sup>95</sup>, Karen L. Mohlke<sup>38</sup>, Donna M. Muzny<sup>95</sup>, Ioanna Ntalla<sup>65,96</sup>, Nicholette D. Palmer<sup>58,97</sup>, Dorota Pasko<sup>8</sup>, Andreas Peter<sup>98,99</sup>, Nigel W. Rayner<sup>33,48,100</sup>, Frida Renström<sup>50</sup>, Ken Rice<sup>9,101</sup>, Cinzia F. Sala<sup>53</sup>, Bengt Sennblad<sup>46,102</sup>, Ioannis Serafetinidis<sup>103</sup>, Jennifer A. Smith<sup>104</sup>, Nicole Soranzo<sup>33,105</sup>, Elizabeth K. Speliotes<sup>106</sup>, Eli A. Stahl<sup>107</sup>, Kathleen Stirrups<sup>33,108</sup>, Nikos Tentolouris<sup>109</sup>, Anastasia Thanopoulou<sup>110</sup>, Mina Torres<sup>94</sup>, Michela Traglia<sup>53</sup>, Emmanouil Tsafantakis<sup>111</sup>, Sundas Javad<sup>6</sup>, Lisa R. Yanek<sup>112</sup>, Eleni Zengini<sup>91,113</sup>, Diane M. Becker<sup>112</sup>, Joshua C. Bis<sup>9,10</sup>, James B. Brown<sup>49,114</sup>, L. Adrienne Cupples<sup>4,7</sup>, Torben Hansen<sup>26,115</sup>, Erik Ingelsson<sup>69,87</sup>, Andrew J. Karter<sup>116</sup>, Carlos Lorenzo<sup>117</sup>, Rasika A. Mathias<sup>112</sup>, Jill M. Norris<sup>118</sup>, Gina M. Peloso<sup>119,120</sup>, Wayne H.-H. Sheu<sup>83,84,121</sup>, Daniela Toniolo<sup>53</sup>, Dhananjay Vaidya<sup>112</sup>, Rohit Varma<sup>94</sup>, Lynne E. Wagenknecht<sup>122</sup>, Heiner Boeing<sup>123</sup>, Erwin P. Bottinger<sup>23</sup>, George Dedoussis<sup>65</sup>, Panos Deloukas<sup>108,124,125</sup>, Ele Ferrannini<sup>126</sup>, Oscar H. Franco<sup>62</sup>, Paul W. Franks<sup>50,127,128</sup>, Richard A. Gibbs<sup>95</sup>, Vilmundur Gudnason<sup>37,47</sup>, Anders Hamsten<sup>46</sup>, Tamara B. Harris<sup>35</sup>, Andrew T. Hattersley<sup>129</sup>, Caroline Hayward<sup>21</sup>, Albert Hofman<sup>62</sup>, Jan-Håkan Jansson<sup>128,130</sup>, Claudia Langenberg<sup>6</sup>, Lenore J. Launer<sup>35</sup>, Daniel Levy<sup>131,132</sup>, Ben A. Oostra<sup>5</sup>, Christopher J. O'Donnell<sup>4,12,133</sup>, Stephen O'Rahilly<sup>134</sup>, Sandosh Padmanabhan<sup>135</sup>, James S. Pankow<sup>136</sup>, Ozren Polasek<sup>137</sup>, Michael A. Province<sup>22</sup>, Stephen S. Rich<sup>138</sup>, Paul M. Ridker<sup>3,139</sup>, Igor Rudan<sup>140</sup>, Matthias B. Schulze<sup>41,99</sup>, Blair H. Smith<sup>141</sup>, André G. Uitterlinden<sup>44,62</sup>, Mark Walker<sup>142</sup>, Hugh Watkins<sup>68</sup>, Tien Y. Wong<sup>51,52,60</sup>, Eleftheria Zeggini<sup>33</sup>, The EPIC-InterAct Consortium<sup>§</sup>, Markku Laakso<sup>81</sup>, Ingrid B. Borecki<sup>22</sup>, Daniel I. Chasman<sup>3,143</sup>, Oluf Pedersen<sup>26</sup>, Bruce M. Psaty<sup>9,10,144,145,146</sup>, E. Shyong Tai<sup>59,147</sup>, Cornelia M. van Duijn<sup>5,148</sup>, Nicholas J. Wareham<sup>6</sup>, Dawn M. Waterworth<sup>149</sup>, Eric Boerwinkle<sup>42,95</sup>, W.H. Linda Kao<sup>40,93,150</sup>, Jose C. Florez<sup>12,16,119,120</sup>, Ruth J.F. Loos<sup>23,24,151</sup>, James G. Wilson<sup>152</sup>, Timothy M. Frayling<sup>8</sup>, David S. Siscovick<sup>153,154</sup>, Josée Dupuis<sup>4,7</sup>, Jerome I. Rotter<sup>36</sup>, James B. Meigs<sup>11,12</sup>, Robert A. Scott<sup>6</sup> & Mark O. Goodarzi<sup>29,155</sup>

<footnote n="1"><sup>1</sup>Department of Epidemiology, Fairbanks School of Public Health, Indianapolis, Indiana 46202, USA. <sup>2</sup>Department of Medicine, Indiana University School of Medicine, Indianapolis, Indiana 46202, USA. <sup>3</sup>Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts 02215, USA. <sup>4</sup>National Heart, Lung, and Blood Institute (NHLBI) Framingham Heart Study, Framingham, Massachusetts 01702, USA. <sup>5</sup>Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam 3000 CE, The Netherlands. <sup>6</sup>MRC Epidemiology Unit, University of Cambridge School of Clinical Medicine, Institute of Metabolic Science, Cambridge Biomedical Campus, Cambridge CB2 OSL, UK. <sup>7</sup>Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts 02118, USA. <sup>8</sup>Genetics of Complex Traits, University of Exeter Medical School, University of Exeter, Exeter EX1 2LU, UK. <sup>9</sup>Cardiovascular Health Research Unit, University of Washington, Seattle, Washington 98101, USA. <sup>10</sup>Department of Medicine, University of Washington, Seattle, Washington 98195, USA. <sup>11</sup>Massachusetts General Hospital, General Medicine Division, Boston, Massachusetts 02114, USA. <sup>12</sup>Department of Medicine, Harvard Medical School, Boston, Massachusetts 02115, USA. <sup>13</sup>Division of Endocrinology, Diabetes and Metabolism, Department of Medicine, University of Verona Medical School and Hospital Trust of Verona, Verona 37126, Italy. <sup>14</sup>Harvard Pilgrim Health Care Institute, Department of Population Medicine, Harvard Medical School, Boston, Massachusetts 02215, USA. <sup>15</sup>Division of Endocrinology and Metabolism, Department of Medicine, Université de Sherbrooke, Sherbrooke, Québec, Canada J1K 2R1. <sup>16</sup>Diabetes Unit, Department of Medicine, Massachusetts General Hospital, Boston, Massachusetts 02114, USA. <sup>17</sup>Center for Molecular Medicine and Genetics, Wayne State University, Detroit, Michigan 48201, USA. <sup>18</sup>Department of Neurology, Wayne State University School of Medicine, Detroit, Michigan 48202, USA. <sup>19</sup>Department of Epidemiology, University of Alabama at Birmingham, Birmingham, Alabama 35233, USA. <sup>20</sup>Department of Genome Sciences, University of Washington, Seattle, Washington 98195, USA. <sup>21</sup>MRC Human Genetics Unit, MRC IGMM, University of Edinburgh, Edinburgh, Scotland EH4 2XU, UK. <sup>22</sup>Division of Statistical Genomics and Department of Genetics, Washington University School of Medicine, St. Louis, Missouri 63108, USA. <sup>23</sup>The Charles Bronfman Institute for Personalized Medicine, The Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA. <sup>24</sup>The Genetics of Obesity and Related Metabolic Traits Program, The Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA. <sup>25</sup>Department of Preventive Medicine, Northwestern University Feinberg School of Medicine, Chicago, Illinois 60611, USA. <sup>26</sup>The Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen DK-2200, Denmark. <sup>27</sup>Quantitative Sciences, PCPS, GlaxoSmithKline, North Carolina 27709, USA. <sup>28</sup>Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland, Kuopio FI-70211, Finland. <sup>29</sup>Department of Medicine and Department of Biomedical Sciences, Cedars-Sinai Medical Center, Los Angeles, California 90048, USA. <sup>30</sup>Materials and Process Simulation Center, California Institute of Technology, Pasadena, California 91125, USA. <sup>31</sup>CEA, Institut de Génomique, Centre National de Génotypage, 2 Rue Gaston Crémieux, EVRY Cedex 91057, France. <sup>32</sup>Brown</footnote>
Foundation Institute of Molecular Medicine, University of Texas Health Science Center, Houston, Texas 77030, USA. ³³ The Wellcome Trust Sanger Institute, Hinxton CB10 1SA, UK. ³⁴ Department of Public Health and Primary Care, Strangeways Research Laboratory, University of Cambridge, Cambridge CB1 8RN, UK. ³⁵ Intramural Research Program, National Institute on Aging, Bethesda, Maryland 21224, USA. ³⁶ Institute for Translational Genomics and Population Sciences, Los Angeles Biomedical Research Institute at Harbor-UCLA Medical Center, Torrance, California 90502, USA. ³⁷ Icelandic Heart Association, Holtasmari 1, Kopavogur IS-201, Iceland. ³⁸ Department of Genetics, University of North Carolina, Chapel Hill, North Carolina 27599, USA. ³⁹ Indiana University, Fairbanks School of Public Health, Indianapolis, Indiana 46202, USA. ⁴⁰ Department of Epidemiology, Johns Hopkins University, Baltimore, Maryland 21205, USA. ⁴¹ Department of Molecular Epidemiology, German Institute of Human Nutrition Potsdam-Rehbrücke, Nuthetal DE-14558, Germany. ⁴² Human Genetics Center, School of Public Health, The University of Texas Health Science Center at Houston, Houston, Texas 77225, USA. ⁴³ Laboratory of Neurogenetics, National Institute on Aging, Bethesda, Maryland 20892, USA. ⁴⁴ Department of Internal Medicine, Erasmus University Medical Center, Rotterdam 3000 CE, The Netherlands. ⁴⁵ The Netherlands Genomics Initiative-sponsored Netherlands Consortium for Healthy Aging (NGI-NCHA), Leiden/Rotterdam 2300 RC, The Netherlands. ⁴⁶ Atherosclerosis Research Unit, Department of Medicine Solna, Karolinska Institutet, Stockholm SE-171 77, Sweden. ⁴⁷ University of Iceland, Reykjavik IS-101, Iceland. ⁴⁸ Wellcome Trust Centre for Human Genetics, Oxford OX3 7BN, UK. ⁴⁹ Department of Genome Dynamics, Lawrence Berkeley National Laboratory, Berkeley, California 94720, USA. ⁵⁰ Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Lund University, Skåne University Hospital, Malmö SE-205 02, Sweden. ⁵¹ Singapore Eye Research Institute, Singapore National Eye Centre, Singapore 168751, Singapore. ⁵² Department of Ophthalmology, National University of Singapore and National University Health System, Singapore 119228, Singapore. ⁵³ Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano 20132, Italy. ⁵⁴ Predoctoral Training Program in Human Genetics, McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Maryland 21205, USA. ⁵⁵ McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland 21205, USA. ⁵⁶ Department of Biostatistics and Center for Statistical Genetics, University of Michigan, Ann Arbor, Michigan 48109, USA. ⁵⁷ Section of Biology and Genetics, Department of Life and Reproduction Sciences, University of Verona, Verona 37100, Italy. ⁵⁸ Department of Biochemistry, Wake Forest School of Medicine, Winston-Salem, North Carolina 27157, USA. ⁵⁹ Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore 119228, Singapore. ⁶⁰ Office of Clinical Sciences, Duke-NUS Graduate Medical School, National University of Singapore, Singapore 169857, Singapore. ⁶¹ Department of Medicine, University of Mississippi Medical Center, Jackson, Mississippi 39216, USA. ⁶² Department of Epidemiology, Erasmus University Medical Center, Rotterdam 3000 CE, The Netherlands. ⁶³ McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University, Baltimore, Maryland 21205, USA. ⁶⁴ Division of Cardiology, Geneva University Hospital Geneva 1211, Switzerland. ⁶⁵ Department of Nutrition and Dietetics, School of Health Science and Education, Harokopio University, Athens 17671, Greece. ⁶⁶ Department of Numerical Analysis and Computer Science, SciLifeLab, Stockholm University, Stockholm SE-106 91, Sweden. ⁶⁷ Division of Nephrology, Department of Internal Medicine and Medical Specialties, Columbus-Gemelli University Hospital, Catholic University, Rome 00168, Italy. ⁶⁸ Department of Cardiovascular Medicine, The Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford OX3 7BN, UK. ⁶⁹ Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University, Uppsala SE-751 85, Sweden. ⁷⁰ Department of Biobank Research, Umeå University, Umeå SE-901 87, Sweden. ⁷¹ Department of Biomedical Technology, Sangmyung University, Chungnam 330-720, Korea. ⁷² Institute of Human Genetics, Department of Genomics, Life & Brain Center, University of Bonn, Bonn DE-53127, Germany. ⁷³ Human Genomics Research Group, Division of Medical Genetics, University Hospital Basel Department of Biomedicine 4031, Basel, Switzerland. ⁷⁴ Institute of Neuroscience and Medicine (INM-1) Genomic Imaging Research Center Juelich, Juelich DE-52425, Germany. ⁷⁵ Memory Aging & Cognition Centre (MACC), National University Health System, Singapore 117599, Singapore. ⁷⁶ Steno Diabetes Center, Gentofte DK-2820, Denmark. ⁷⁷ Research Centre for Prevention and Health, Glostrup University Hospital, Glostrup DK-2600, Denmark. ⁷⁸ Faculty of Medicine, University of Aalborg, Aalborg DK-9220, Denmark. ⁷⁹ Echinos Medical Centre, Echinos 67300, Greece. ⁸⁰ Division of Human Genetics, Genome Institute of Singapore, Singapore 138672, Singapore. ⁸¹ Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland and Kuopio University Hospital, Kuopio FI-70211, Finland. ⁸² Department of Biostatistics, University of North Carolina, Chapel Hill, North Carolina 27599, USA. ⁸³ Division of Endocrine and Metabolism, Department of Internal Medicine, Taichung Veterans General Hospital, Taichung 407, Taiwan. ⁸⁴ School of Medicine, National Yang-Ming University, Taipei 112, Taiwan. ⁸⁵ Department of Medical Research, Taichung Veterans General Hospital, Taichung 407, Taiwan. ⁸⁶ Department of Epidemiology & Prevention, Division of Public Health Sciences, Wake Forest University, Winston-Salem, North Carolina 27106, USA. ⁸⁷ Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford OX3 7BN, UK. ⁸⁸ Department of Clinical Experimental Research, Copenhagen University Hospital Glostrup, Glostrup DK-2600, Denmark. ⁸⁹ Department of Clinical Medicine, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen DK-2200, Denmark. ⁹⁰ National and Kapodistrian University of Athens, Faculty of Medicine, Athens 115 27, Greece. ⁹¹ Dromokaiteio Psychiatric Hospital, Athens 124 61, Greece. ⁹² Division of General Internal Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland 21205, USA. ⁹³ Welch Center for Prevention, Epidemiology, and Clinical Research, Johns Hopkins University, Baltimore, Maryland 21205, USA. ⁹⁴ Department of Preventive Medicine, Keck School of Medicine of the University of Southern California, Los Angeles 90033, USA. ⁹⁵ Human Genome Sequencing Center, Baylor College of Medicine, Houston, Texas 77030, USA. ⁹⁶ University of Leicester, Leicester LE1 7RH, UK. ⁹⁷ Center for Genomics and Personalized Medicine Research, Wake Forest School of Medicine, Winston-Salem, North Carolina 27106, USA. ⁹⁸ Department of Internal Medicine, Division of Endocrinology, Metabolism, Pathobiochemistry and Clinical Chemistry and Institute of Diabetes Research and Metabolic Diseases, University of Tübingen, Tübingen DE-72076, Germany. ⁹⁹ German Center for Diabetes Research (DZD), Neuherberg DE-85764, Germany. ¹⁰⁰ The Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford OX3 7LE, UK. ¹⁰¹ Department of Biostatistics, University of Washington, Seattle, Washington 98195, USA. ¹⁰² Science for Life Laboratory, Karolinska Institutet, Stockholm SE-171 77, Sweden. ¹⁰³ Department of Gastroenterology, Gennimatas General Hospital, Athens 11527, Greece. ¹⁰⁴ Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, Michigan 48109, USA. ¹⁰⁵ Department of Hematology, Long Road, Cambridge CB2 OXY, UK. ¹⁰⁶ Department of Internal Medicine, Division of Gastroenterology and Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, Michigan 48109, USA. ¹⁰⁷ Division of Psychiatric Genomics, The Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA. ¹⁰⁸ William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London E1 4NS, UK. ¹⁰⁹ First Department of Propaedeutic and Internal Medicine, Athens University Medical School, Laiko General Hospital, Athens 11527, Greece. ¹¹⁰ Diabetes Centre, 2nd Department of Internal Medicine, National University of Athens, Hippokration General Hospital, Athens 11527, Greece. ¹¹¹ Anogia Medical Centre, Anogia 740 51, Greece. ¹¹² The GeneSTAR Research Program, Division of General Internal Medicine, Department of Medicine, The Johns Hopkins University School of Medicine, Baltimore, Maryland 21205, USA. ¹¹³ University of Sheffield, Sheffield S10 2TN, UK. ¹¹⁴ Department of Statistics, University of California at Berkeley, Berkeley, California 94720, USA. ¹¹⁵ Faculty of Health Science, University of Copenhagen, Copenhagen 1165, Denmark. ¹¹⁶ Division of Research, Kaiser Permanente, Northern California Region, Oakland, California 94612, USA. ¹¹⁷ Department of Medicine, University of Texas Health Science Center, San Antonio, Texas 77030, USA. ¹¹⁸ Department of Epidemiology, Colorado School of Public Health, University of Colorado Denver, Aurora, Colorado 80204, USA. ¹¹⁹ Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts 02142, USA. ¹²⁰ Center for Human Genetic Research, Massachusetts General Hospital, Boston, Massachusetts 02114, USA. ¹²¹ College of Medicine, National Defense Medical Center, Taipei 114, Taiwan. ¹²² Division of Public Health Sciences, Wake Forest School of Medicine, Winston-Salem, North Carolina 27106, USA. ¹²³ Department of Epidemiology, German Institute of Human Nutrition Potsdam Rehbrücke, Nuthetal DE-14558, Germany. ¹²⁴ Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1SA, UK. ¹²⁵ Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, Jeddah 22254,
Foundation Institute of Molecular Medicine, University of Texas Health Science Center, Houston, Texas 77030, USA. ³³ The Wellcome Trust Sanger Institute, Hinxton CB10 1SA, UK. ³⁴ Department of Public Health and Primary Care, Strangeways Research Laboratory, University of Cambridge, Cambridge CB1 8RN, UK. ³⁵ Intramural Research Program, National Institute on Aging, Bethesda, Maryland 21224, USA. ³⁶ Institute for Translational Genomics and Population Sciences, Los Angeles Biomedical Research Institute at Harbor-UCLA Medical Center, Torrance, California 90502, USA. ³⁷ Icelandic Heart Association, Holtasmari 1, Kopavogur IS-201, Iceland. ³⁸ Department of Genetics, University of North Carolina, Chapel Hill, North Carolina 27599, USA. ³⁹ Indiana University, Fairbanks School of Public Health, Indianapolis, Indiana 46202, USA. ⁴⁰ Department of Epidemiology, Johns Hopkins University, Baltimore, Maryland 21205, USA. ⁴¹ Department of Molecular Epidemiology, German Institute of Human Nutrition Potsdam-Rehbrücke, Nuthetal DE-14558, Germany. ⁴² Human Genetics Center, School of Public Health, The University of Texas Health Science Center at Houston, Houston, Texas 77225, USA. ⁴³ Laboratory of Neurogenetics, National Institute on Aging, Bethesda, Maryland 20892, USA. ⁴⁴ Department of Internal Medicine, Erasmus University Medical Center, Rotterdam 3000 CE, The Netherlands. ⁴⁵ The Netherlands Genomics Initiative-sponsored Netherlands Consortium for Healthy Aging (NGI-NCHA), Leiden/Rotterdam 2300 RC, The Netherlands. ⁴⁶ Atherosclerosis Research Unit, Department of Medicine Solna, Karolinska Institutet, Stockholm SE-171 77, Sweden. ⁴⁷ University of Iceland, Reykjavik IS-101, Iceland. ⁴⁸ Wellcome Trust Centre for Human Genetics, Oxford OX3 7BN, UK. ⁴⁹ Department of Genome Dynamics, Lawrence Berkeley National Laboratory, Berkeley, California 94720, USA. ⁵⁰ Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Lund University, Skåne University Hospital, Malmö SE-205 02, Sweden. ⁵¹ Singapore Eye Research Institute, Singapore National Eye Centre, Singapore 168751, Singapore. ⁵² Department of Ophthalmology, National University of Singapore and National University Health System, Singapore 119228, Singapore. ⁵³ Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano 20132, Italy. ⁵⁴ Predoctoral Training Program in Human Genetics, McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Maryland 21205, USA. ⁵⁵ McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland 21205, USA. ⁵⁶ Department of Biostatistics and Center for Statistical Genetics, University of Michigan, Ann Arbor, Michigan 48109, USA. ⁵⁷ Section of Biology and Genetics, Department of Life and Reproduction Sciences, University of Verona, Verona 37100, Italy. ⁵⁸ Department of Biochemistry, Wake Forest School of Medicine, Winston-Salem, North Carolina 27157, USA. ⁵⁹ Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore 119228, Singapore. ⁶⁰ Office of Clinical Sciences, Duke-NUS Graduate Medical School, National University of Singapore, Singapore 169857, Singapore. ⁶¹ Department of Medicine, University of Mississippi Medical Center, Jackson, Mississippi 39216, USA. ⁶² Department of Epidemiology, Erasmus University Medical Center, Rotterdam 3000 CE, The Netherlands. ⁶³ McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University, Baltimore, Maryland 21205, USA. ⁶⁴ Division of Cardiology, Geneva University Hospital Geneva 1211, Switzerland. ⁶⁵ Department of Nutrition and Dietetics, School of Health Science and Education, Harokopio University, Athens 17671, Greece. ⁶⁶ Department of Numerical Analysis and Computer Science, SciLifeLab, Stockholm University, Stockholm SE-106 91, Sweden. ⁶⁷ Division of Nephrology, Department of Internal Medicine and Medical Specialties, Columbus-Gemelli University Hospital, Catholic University, Rome 00168, Italy. ⁶⁸ Department of Cardiovascular Medicine, The Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford OX3 7BN, UK. ⁶⁹ Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University, Uppsala SE-751 85, Sweden. ⁷⁰ Department of Biobank Research, Umeå University, Umeå SE-901 87, Sweden. ⁷¹ Department of Biomedical Technology, Sangmyung University, Chungnam 330-720, Korea. ⁷² Institute of Human Genetics, Department of Genomics, Life & Brain Center, University of Bonn, Bonn DE-53127, Germany. ⁷³ Human Genomics Research Group, Division of Medical Genetics, University Hospital Basel Department of Biomedicine 4031, Basel, Switzerland. ⁷⁴ Institute of Neuroscience and Medicine (INM-1) Genomic Imaging Research Center Juelich, Juelich DE-52425, Germany. ⁷⁵ Memory Aging & Cognition Centre (MACC), National University Health System, Singapore 117599, Singapore. ⁷⁶ Steno Diabetes Center, Gentofte DK-2820, Denmark. ⁷⁷ Research Centre for Prevention and Health, Glostrup University Hospital, Glostrup DK-2600, Denmark. ⁷⁸ Faculty of Medicine, University of Aalborg, Aalborg DK-9220, Denmark. ⁷⁹ Echinos Medical Centre, Echinos 67300, Greece. ⁸⁰ Division of Human Genetics, Genome Institute of Singapore, Singapore 138672, Singapore. ⁸¹ Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland and Kuopio University Hospital, Kuopio FI-70211, Finland. ⁸² Department of Biostatistics, University of North Carolina, Chapel Hill, North Carolina 27599, USA. ⁸³ Division of Endocrine and Metabolism, Department of Internal Medicine, Taichung Veterans General Hospital, Taichung 407, Taiwan. ⁸⁴ School of Medicine, National Yang-Ming University, Taipei 112, Taiwan. ⁸⁵ Department of Medical Research, Taichung Veterans General Hospital, Taichung 407, Taiwan. ⁸⁶ Department of Epidemiology & Prevention, Division of Public Health Sciences, Wake Forest University, Winston-Salem, North Carolina 27106, USA. ⁸⁷ Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford OX3 7BN, UK. ⁸⁸ Department of Clinical Experimental Research, Copenhagen University Hospital Glostrup, Glostrup DK-2600, Denmark. ⁸⁹ Department of Clinical Medicine, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen DK-2200, Denmark. ⁹⁰ National and Kapodistrian University of Athens, Faculty of Medicine, Athens 115 27, Greece. ⁹¹ Dromokaiteio Psychiatric Hospital, Athens 124 61, Greece. ⁹² Division of General Internal Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland 21205, USA. ⁹³ Welch Center for Prevention, Epidemiology, and Clinical Research, Johns Hopkins University, Baltimore, Maryland 21205, USA. ⁹⁴ Department of Preventive Medicine, Keck School of Medicine of the University of Southern California, Los Angeles 90033, USA. ⁹⁵ Human Genome Sequencing Center, Baylor College of Medicine, Houston, Texas 77030, USA. ⁹⁶ University of Leicester, Leicester LE1 7RH, UK. ⁹⁷ Center for Genomics and Personalized Medicine Research, Wake Forest School of Medicine, Winston-Salem, North Carolina 27106, USA. ⁹⁸ Department of Internal Medicine, Division of Endocrinology, Metabolism, Pathobiochemistry and Clinical Chemistry and Institute of Diabetes Research and Metabolic Diseases, University of Tübingen, Tübingen DE-72076, Germany. ⁹⁹ German Center for Diabetes Research (DZD), Neuherberg DE-85764, Germany. ¹⁰⁰ The Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford OX3 7LE, UK. ¹⁰¹ Department of Biostatistics, University of Washington, Seattle, Washington 98195, USA. ¹⁰² Science for Life Laboratory, Karolinska Institutet, Stockholm SE-171 77, Sweden. ¹⁰³ Department of Gastroenterology, Gennimatas General Hospital, Athens 11527, Greece. ¹⁰⁴ Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, Michigan 48109, USA. ¹⁰⁵ Department of Hematology, Long Road, Cambridge CB2 OXY, UK. ¹⁰⁶ Department of Internal Medicine, Division of Gastroenterology and Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, Michigan 48109, USA. ¹⁰⁷ Division of Psychiatric Genomics, The Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA. ¹⁰⁸ William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London E1 4NS, UK. ¹⁰⁹ First Department of Propaedeutic and Internal Medicine, Athens University Medical School, Laiko General Hospital, Athens 11527, Greece. ¹¹⁰ Diabetes Centre, 2nd Department of Internal Medicine, National University of Athens, Hippokration General Hospital, Athens 11527, Greece. ¹¹¹ Anogia Medical Centre, Anogia 740 51, Greece. ¹¹² The GeneSTAR Research Program, Division of General Internal Medicine, Department of Medicine, The Johns Hopkins University School of Medicine, Baltimore, Maryland 21205, USA. ¹¹³ University of Sheffield, Sheffield S10 2TN, UK. ¹¹⁴ Department of Statistics, University of California at Berkeley, Berkeley, California 94720, USA. ¹¹⁵ Faculty of Health Science, University of Copenhagen, Copenhagen 1165, Denmark. ¹¹⁶ Division of Research, Kaiser Permanente, Northern California Region, Oakland, California 94612, USA. ¹¹⁷ Department of Medicine, University of Texas Health Science Center, San Antonio, Texas 77030, USA. ¹¹⁸ Department of Epidemiology, Colorado School of Public Health, University of Colorado Denver, Aurora, Colorado 80204, USA. ¹¹⁹ Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts 02142, USA. ¹²⁰ Center for Human Genetic Research, Massachusetts General Hospital, Boston, Massachusetts 02114, USA. ¹²¹ College of Medicine, National Defense Medical Center, Taipei 114, Taiwan. ¹²² Division of Public Health Sciences, Wake Forest School of Medicine, Winston-Salem, North Carolina 27106, USA. ¹²³ Department of Epidemiology, German Institute of Human Nutrition Potsdam Rehbrücke, Nuthetal DE-14558, Germany. ¹²⁴ Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1SA, UK. ¹²⁵ Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, Jeddah 22254,
Saudi Arabia. ¹²⁶ CNR Institute of Clinical Physiology, Pisa 73100, Italy. ¹²⁷ Department of Nutrition, Harvard School of Public Health, Boston, Massachusetts 02115, USA. ¹²⁸ Department of Public Health & Clinical Medicine, Umeå University, Umeå SE-901 87, Sweden. ¹²⁹ Genetics of Diabetes, University of Exeter Medical School, University of Exeter, Exeter EX1 2LU, UK. ¹³⁰ Research Unit, Skellefteå SE-931 87, Sweden. ¹³¹ Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland 20892, USA. ¹³² Framingham Heart Study, Framingham, Massachusetts 01702, USA. ¹³³ Cardiology Division, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, Massachusetts 02115, USA. ¹³⁴ University of Cambridge Metabolic Research Laboratories, MRC Metabolic Diseases Unit and NIHR Cambridge Biomedical Research Centre, Wellcome Trust-MRC Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge CB2 1TN, UK. ¹³⁵ Institute of Cardiovascular and Medical Sciences, University of Glasgow, Glasgow G12 8TA, UK. ¹³⁶ Division of Epidemiology and Community Health, School of Public Health, University of Minnesota, Minneapolis, Minnesota 55455, USA. ¹³⁷ Department of Public Health, Faculty of Medicine, University of Split, Split 21000, Croatia. ¹³⁸ Center for Public Health Genomics, Department of Public Health Sciences, University of Virginia, Charlottesville, Virginia 22908, USA. ¹³⁹ Division of Cardiology, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts 02115, USA. ¹⁴⁰ Centre for Population Health Sciences, Medical School, University of Edinburgh, Edinburgh, Scotland EH8 9YL, UK. ¹⁴¹ Medical Research Institute, University of Dundee, Dundee DD1 9SY, UK. ¹⁴² Institute of Cellular Medicine, Newcastle University, Newcastle-upon-Tyne NE1 7RU, UK. ¹⁴³ Division of Genetics, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, USA. ¹⁴⁴ Department of Epidemiology, University of Washington, Seattle, Washington 98195, USA. ¹⁴⁵ Department of Health Services, University of Washington, Seattle, Washington 98195, USA. ¹⁴⁶ Group Health Research Institute, Group Health Cooperative, Seattle, Washington 98195, USA. ¹⁴⁷ Department of Medicine, Yong Loo Lin School of Medicine, National University of Singapore, Singapore 117597, Singapore. ¹⁴⁸ Center for Medical Systems Biology, Leiden 2300, The Netherlands. ¹⁴⁹ Genetics, PCPS, GlaxoSmithKline, Philadelphia, Pennsylvania 19104, USA. ¹⁵⁰ Department of Medicine. Johns Hopkins University, Baltimore, Maryland 21205, USA. ¹⁵¹ The Mindich Child Health and Development Institute, The Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA. ¹⁵² Department of Physiology and Biophysics, University of Mississippi Medical Center, Jackson, Mississippi 38677, USA. ¹⁵³ New York Academy of Medicine, New York, New York 10029, USA. ¹⁵⁴ Cardiovascular Health Research Unit, Departments of Medicine and Epidemiology, University of Washington, Seattle, Washington 98195, USA. ¹⁵⁵ Division of Endocrinology, Diabetes and Metabolism, Cedars-Sinai Medical Center, Los Angeles, California 90048, USA. * These authors contributed equally to this work. § A list of The EPIC-InterAct Consortium members is provided below.

## The EPIC-InterAct Consortium

Stephen J. Sharp⁶, Nita G. Forouhi⁶, Nicola D. Kerrison⁶, Debora ME Lucarelli⁶, Matt Sims⁶, Inês Barroso³³,¹³⁴, Mark I. McCarthy⁴⁸,¹⁰⁰,<fnref n="156" />, Larraitz Arriola¹⁵⁷,¹⁵⁸,¹⁵⁹, Beverley Balkau¹⁶⁰,¹⁶¹, Aurelio Barricarte¹⁵⁹,¹⁶², Carlos Gonzalez¹⁶³, Sara Grioni¹⁶⁴, Rudolf Kaaks¹⁶⁵, Timothy J. Key¹⁶⁶, Carmen Navarro¹⁵⁹,¹⁶⁷,¹⁶⁸, Peter M. Nilsson⁵⁰, Kim Overvad¹⁶⁹,¹⁷⁰, Domenico Palli¹⁷¹, Salvatore Panico¹⁷², J. Ramón Quirós¹⁷³, Olov Rolandsson⁷⁰, Carlotta Sacerdote¹⁷⁴,¹⁷⁵, María-José Sánchez¹⁵⁹,¹⁷⁶,¹⁷⁷, Nadia Slimani¹⁷⁸, Anne Tjonneland¹⁷⁹, Rosario Tumino¹⁸⁰,¹⁸¹, Daphne L. van der A¹⁸², Yvonne T. van der Schouw¹⁸³ & Elio Riboli¹⁸⁴

<footnote n="156">¹⁵⁶ Oxford NIHR Biomedical Research Centre, Oxford, UK. ¹⁵⁷ Public Health Division of Gipuzkoa, San Sebastian, Spain. ¹⁵⁸ Instituto BIO-Donostia, Basque Government, San Sebastian, Spain. ¹⁵⁹ CIBER Epidemiología y Salud Pública (CIBERESP), Spain. ¹⁶⁰ Inserm, CESP, U1018, Villejuif, France. ¹⁶¹ Univ Paris-Sud, UMRS 1018, Villejuif, France. ¹⁶² Navarre Public Health Institute (ISPN), Pamplona, Spain. ¹⁶³ Catalan Institute of Oncology (ICO), Barcelona, Spain. ¹⁶⁴ Epidemiology and Prevention Unit, Milan, Italy. ¹⁶⁵ German Cancer Research Centre (DKFZ), Heidelberg, Germany. ¹⁶⁶ Cancer Epidemiology Unit, Nuffield Department of Population Health, University of Oxford, Oxford, UK. ¹⁶⁷ Department of Epidemiology, Murcia Regional Health Council, Murcia, Spain. ¹⁶⁸ Unit of Preventive Medicine and Public Health, School of Medicine, University of Murcia, Murcia, Spain. ¹⁶⁹ Department of Public Health, Section for Epidemiology, Aarhus University, Aarhus, Denmark. ¹⁷⁰ Aalborg University Hospital, Aalborg, Denmark. ¹⁷¹ Cancer Research and Prevention Institute (ISPO), Florence, Italy. ¹⁷² Dipartimento di Medicina Clinica e Chirurgia, Federico II University, Naples, Italy. ¹⁷³ Public Health Directorate, Asturias, Spain. ¹⁷⁴ Unit of Cancer Epidemiology, Citta' della Salute e della Scienza Hospital-University of Turin and Center for Cancer Prevention (CPO), Torino, Italy. ¹⁷⁵ Human Genetics Foundation (HuGeF), Torino, Italy. ¹⁷⁶ Andalusian School of Public Health, Granada, Spain. ¹⁷⁷ Instituto de Investigación Biosanitaria de Granada (Granada.igs), Granada, Spain. ¹⁷⁸ International Agency for Research on Cancer, Lyon, France. ¹⁷⁹ Danish Cancer Society Research Center, Copenhagen, Denmark. ¹⁸⁰ ASP Ragusa, Italy. ¹⁸¹ Aire Onlus, Ragusa, Italy. ¹⁸² National Institute for Public Health and the Environment (RIVM), Bilthoven, The Netherlands. ¹⁸³ University Medical Center Utrecht, Utrecht, Utrecht, the Netherlands. ¹⁸⁴ School of Public Health, Imperial College London, London, UK</footnote>
Saudi Arabia. ¹²⁶ CNR Institute of Clinical Physiology, Pisa 73100, Italy. ¹²⁷ Department of Nutrition, Harvard School of Public Health, Boston, Massachusetts 02115, USA. ¹²⁸ Department of Public Health & Clinical Medicine, Umeå University, Umeå SE-901 87, Sweden. ¹²⁹ Genetics of Diabetes, University of Exeter Medical School, University of Exeter, Exeter EX1 2LU, UK. ¹³⁰ Research Unit, Skellefteå SE-931 87, Sweden. ¹³¹ Population Sciences Branch, National Heart, Lung, and Blood Institute, National Institutes of Health, Bethesda, Maryland 20892, USA. ¹³² Framingham Heart Study, Framingham, Massachusetts 01702, USA. ¹³³ Cardiology Division, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, Massachusetts 02115, USA. ¹³⁴ University of Cambridge Metabolic Research Laboratories, MRC Metabolic Diseases Unit and NIHR Cambridge Biomedical Research Centre, Wellcome Trust-MRC Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge CB2 1TN, UK. ¹³⁵ Institute of Cardiovascular and Medical Sciences, University of Glasgow, Glasgow G12 8TA, UK. ¹³⁶ Division of Epidemiology and Community Health, School of Public Health, University of Minnesota, Minneapolis, Minnesota 55455, USA. ¹³⁷ Department of Public Health, Faculty of Medicine, University of Split, Split 21000, Croatia. ¹³⁸ Center for Public Health Genomics, Department of Public Health Sciences, University of Virginia, Charlottesville, Virginia 22908, USA. ¹³⁹ Division of Cardiology, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts 02115, USA. ¹⁴⁰ Centre for Population Health Sciences, Medical School, University of Edinburgh, Edinburgh, Scotland EH8 9YL, UK. ¹⁴¹ Medical Research Institute, University of Dundee, Dundee DD1 9SY, UK. ¹⁴² Institute of Cellular Medicine, Newcastle University, Newcastle-upon-Tyne NE1 7RU, UK. ¹⁴³ Division of Genetics, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, USA. ¹⁴⁴ Department of Epidemiology, University of Washington, Seattle, Washington 98195, USA. ¹⁴⁵ Department of Health Services, University of Washington, Seattle, Washington 98195, USA. ¹⁴⁶ Group Health Research Institute, Group Health Cooperative, Seattle, Washington 98195, USA. ¹⁴⁷ Department of Medicine, Yong Loo Lin School of Medicine, National University of Singapore, Singapore 117597, Singapore. ¹⁴⁸ Center for Medical Systems Biology, Leiden 2300, The Netherlands. ¹⁴⁹ Genetics, PCPS, GlaxoSmithKline, Philadelphia, Pennsylvania 19104, USA. ¹⁵⁰ Department of Medicine, Johns Hopkins University, Baltimore, Maryland 21205, USA. ¹⁵¹ The Mindich Child Health and Development Institute, The Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA. ¹⁵² Department of Physiology and Biophysics, University of Mississippi Medical Center, Jackson, Mississippi 38677, USA. ¹⁵³ New York Academy of Medicine, New York, New York 10029, USA. ¹⁵⁴ Cardiovascular Health Research Unit, Departments of Medicine and Epidemiology, University of Washington, Seattle, Washington 98195, USA. ¹⁵⁵ Division of Endocrinology, Diabetes and Metabolism, Cedars-Sinai Medical Center, Los Angeles, California 90048, USA. * These authors contributed equally to this work. § A list of The EPIC-InterAct Consortium members is provided below.

## The EPIC-InterAct Consortium

Stephen J. Sharp⁶, Nita G. Forouhi⁶, Nicola D. Kerrison⁶, Debora ME Lucarelli⁶, Matt Sims⁶, Inês Barroso³³,¹³⁴, Mark I. McCarthy⁴⁸,¹⁰⁰,<fnref n="156" />, Larraitz Arriola¹⁵⁷,¹⁵⁸,¹⁵⁹, Beverley Balkau¹⁶⁰,¹⁶¹, Aurelio Barricarte¹⁵⁹,¹⁶², Carlos Gonzalez¹⁶³, Sara Grioni¹⁶⁴, Rudolf Kaaks¹⁶⁵, Timothy J. Key¹⁶⁶, Carmen Navarro¹⁵⁹,¹⁶⁷,¹⁶⁸, Peter M. Nilsson⁵⁰, Kim Overvad¹⁶⁹,¹⁷⁰, Domenico Palli¹⁷¹, Salvatore Panico¹⁷², J. Ramón Quirós¹⁷³, Olov Rolandsson⁷⁰, Carlotta Sacerdote¹⁷⁴,¹⁷⁵, María-José Sánchez¹⁵⁹,¹⁷⁶,¹⁷⁷, Nadia Slimani¹⁷⁸, Anne Tjonneland¹⁷⁹, Rosario Tumino¹⁸⁰,¹⁸¹, Daphne L. van der A¹⁸², Yvonne T. van der Schouw¹⁸³ & Elio Riboli¹⁸⁴

<footnote n="156">¹⁵⁶ Oxford NIHR Biomedical Research Centre, Oxford, UK. ¹⁵⁷ Public Health Division of Gipuzkoa, San Sebastian, Spain. ¹⁵⁸ Instituto BIO-Donostia, Basque Government, San Sebastian, Spain. ¹⁵⁹ CIBER Epidemiología y Salud Pública (CIBERESP), Spain. ¹⁶⁰ Inserm, CESP, U1018, Villejuif, France. ¹⁶¹ Univ Paris-Sud, UMRS 1018, Villejuif, France. ¹⁶² Navarre Public Health Institute (ISPN), Pamplona, Spain. ¹⁶³ Catalan Institute of Oncology (ICO), Barcelona, Spain. ¹⁶⁴ Epidemiology and Prevention Unit, Milan, Italy. ¹⁶⁵ German Cancer Research Centre (DKFZ), Heidelberg, Germany. ¹⁶⁶ Cancer Epidemiology Unit, Nuffield Department of Population Health, University of Oxford, Oxford, UK. ¹⁶⁷ Department of Epidemiology, Murcia Regional Health Council, Murcia, Spain. ¹⁶⁸ Unit of Preventive Medicine and Public Health, School of Medicine, University of Murcia, Murcia, Spain. ¹⁶⁹ Department of Public Health, Section for Epidemiology, Aarhus University, Aarhus, Denmark. ¹⁷⁰ Aalborg University Hospital, Aalborg, Denmark. ¹⁷¹ Cancer Research and Prevention Institute (ISPO), Florence, Italy. ¹⁷² Dipartimento di Medicina Clinica e Chirurgia, Federico II University, Naples, Italy. ¹⁷³ Public Health Directorate, Asturias, Spain. ¹⁷⁴ Unit of Cancer Epidemiology, Citta' della Salute e della Scienza Hospital-University of Turin and Center for Cancer Prevention (CPO), Torino, Italy. ¹⁷⁵ Human Genetics Foundation (HuGeF), Torino, Italy. ¹⁷⁶ Andalusian School of Public Health, Granada, Spain. ¹⁷⁷ Instituto de Investigación Biosanitaria de Granada (Granada.igs), Granada, Spain. ¹⁷⁸ International Agency for Research on Cancer, Lyon, France. ¹⁷⁹ Danish Cancer Society Research Center, Copenhagen, Denmark. ¹⁸⁰ ASP Ragusa, Italy. ¹⁸¹ Aire Onlus, Ragusa, Italy. ¹⁸² National Institute for Public Health and the Environment (RIVM), Bilthoven, The Netherlands. ¹⁸³ University Medical Center Utrecht, Utrecht, Utrecht, the Netherlands. ¹⁸⁴ School of Public Health, Imperial College London, London, UK</footnote>