The power of genetic diversity in genome-wide association studies of lipids

Sarah E. Graham, Shoa L. Clarke, Kuan-Han H. Wu, Stavroula Kanoni, Greg J. M. Zajac, Shweta Ramdas, Ida Surakka, Ioanna Ntalla, Sailaja Vedantam, Thomas W. Winkler, Adam E. Locke, Eirini Marouli, Mi Yeong Hwang, Sohee Han, Akira Narita, Ananyo Choudhury, Amy R. Bentley, Kenneth Ekoru, Anurag Verma, Bhavi Trivedi, Hilary C. Martin, Karen A. Hunt, Qin Hui, Derek Klarin, Xiang Zhu, Gudmar Thorleifsson, Anna Helgadottir, Daniel F. Gudbjartsson, Hilma Holm, Isleifur Olafsson, Masato Akiyama, Saori Sakaue, Chikashi Terao, Masahiro Kanai, Wei Zhou, Ben M. Brumpton, Humaira Rasheed, Sanni E. Ruotsalainen, Aki S. Havulinna, Yogasudha Veturi, QiPing Feng, Elisabeth A. Rosenthal, Todd Lingren, Jennifer Allen Pacheco, Sarah A. Pendergrass, Jeffrey Haessler, Franco Giulianini, Yuki Bradford, Jason E. Miller, Archie Campbell, Kuang Lin, Iona Y. Millwood, George Hindy, Asif Rasheed, Jessica D. Faul, Wei Zhao, David R. Weir, Constance Turman, Hongyan Huang, Mariaelisa Graff, Anubha Mahajan, Michael R. Brown, Weihua Zhang, Ketian Yu, Ellen M. Schmidt, Anita Pandit, Stefan Gustafsson, Xianyong Yin, Jian’an Luan, Jing-Hua Zhao, Fumihiko Matsuda, Hye-Mi Jang, Kyungheon Yoon, Carolina Medina-Gomez, Achilleas Pitsillides, Jouke Jan Hottenga, Gonneke Willemsen, Andrew R. Wood, Yingji Ji, Zishan Gao, Simon Haworth, Ruth E. Mitchell, Jin Fang Chai, Mette Aadahl, Jie Yao, Ani Manichaikul, Helen R. Warren, Julia Ramirez, Jette Bork-Jensen, Line L. Kårhus, Anuj Goel, Maria Sabater-Lleal, Raymond Noordam, Carlo Sidore, Edoardo Fiorillo, Aaron F. McDaid, Pedro Marques-Vidal, Matthias Wielscher, Stella Trompet, Naveed Sattar, Line T. Møllehave, Betina H. Thuesen, Matthias Munz, Lingyao Zeng, Jianfeng Huang, Bin Yang, Alaitz Poveda, Azra Kurbasic, Claudia Lamina, Lukas Forer, Markus Scholz, Tessel E. Galesloot, Jonathan P. Bradfield, E. Warwick Daw, Joseph M. Zmuda, Jonathan S. Mitchell, Christian Fuchsberger, Henry Christensen, Jennifer A. Brody, Mary F. Feitosa, Mary K. Wojczynski, Michael Preuss, Massimo Mangino, Paraskevi Christofidou, Niek Verweij, Jan W. Benjamins, Jorgen Engmann, Rachel L. Kember, Roderick C. Slieker, Ken Sin Lo, Nuno R. Zilhao, Phuong Le, Marcus E. Kleber, Graciela E. Delgado, Shaofeng Huo, Daisuke D. Ikeda, Hiroyuki Iha, Jian Yang, Jun Liu, Hampton L. Leonard, Jonathan Marten, Börge Schmidt, Marina Arendt, Laura J. Smyth, Marisa Cañadas-Garre, Chaolong Wang, Masahiro Nakatochi, Andrew Wong, Nina Hutri-Kähönen, Xueling Sim, Rui Xia, Alicia Huerta-Chagoya, Juan Carlos Fernandez-Lopez, Valeriya Lyssenko, Meraj Ahmed, Anne U. Jackson, Noha A. Yousri, Marguerite R. Irvin, Christopher Oldmeadow, Han-Na Kim, Seungho Ryu, Paul R. H. J. Timmers, Liubov Arbeeva, Rajkumar Dorajoo, Leslie A. Lange, Xiaoran Chai, Gauri Prasad, Laura Lorés-Motta, Marc Pauper, Jirong Long, Xiaohui Li, Elizabeth Theusch, Fumihiko Takeuchi, Cassandra N. Spracklen, Anu Loukola, Sailalitha Bollepalli, Sophie C. Warner, Ya Xing Wang, Wen B. Wei, Teresa Nutile, Daniela Ruggiero, Yun Ju Sung, Yi-Jen Hung, Shufeng Chen, Fangchao Liu, Jingyun Yang, Katherine A. Kentistou, Mathias Gorski, Marco Brumat, Karina Meidtner, Lawrence F. Bielak, Jennifer A. Smith, Prashantha Hebbar, Aliki-Eleni Farmaki, Edith Hofer, Maoxuan Lin, Chao Xue, Jifeng Zhang, Maria Pina Concas, Simona Vaccargiu, Peter J. Most, Niina Pitkänen, Brian E. Cade, Jiwon Lee, Sander W. Laan, Kumaraswamy Naidu Chitrala, Stefan Weiss, Martina E. Zimmermann, Jong Young Lee, Hyeok Sun Choi, Maria Nethander, Sandra Freitag-Wolf, Lorraine Southam, Nigel W. Rayner, Carol A. Wang, Shih-Yi Lin, Jun-Sing Wang, Christian Couture, Leo-Pekka Lyytikäinen, Kjell Nikus, Gabriel Cuellar-Partida, Henrik Vestergaard, Bertha Hildalgo, Olga Giannakopoulou, Qiuyin Cai, Morgan O. Obura, Jessica Setten, Xiaoyin Li, Karen Schwander, Natalie Terzikhan, Jae Hun Shin, Rebecca D. Jackson, Alexander P. Reiner, Lisa Warsinger Martin, Zhengming Chen, Liming Li, Heather M. Highland, Kristin L. Young, Takahisa Kawaguchi, Joachim Thiery, Joshua C. Bis, Girish N. Nadkarni, Lenore J. Launer, Huaixing Li, Mike A. Nalls, Olli T. Raitakari, Sahoko Ichihara, Sarah H. Wild, Christopher P. Nelson, Harry Campbell, Susanne Jäger, Toru Nabika, Fahd Al-Mulla, Harri Niinikoski, Peter S. Braund, Ivana Kolcic, Peter Kovacs, Tota Giardoglou, Tomohiro Katsuya, Konain Fatima Bhatti, Dominique Kleijn, Gert J. Borst, Eung Kweon Kim, Hieab H. H. Adams, M. Arfan Ikram, Xiaofeng Zhu, Folkert W. Asselbergs, Adriaan O. Kraaijeveld, Joline W. J. Beulens, Xiao-Ou Shu, Loukianos S. Rallidis, Oluf Pedersen, Torben Hansen, Paul Mitchell, Alex W. Hewitt, Mika Kähönen, Louis Pérusse, Claude Bouchard, Anke Tönjes, Yii- Der Ida Chen, Craig E. Pennell, Trevor A. Mori, Wolfgang Lieb, Andre Franke, Claes Ohlsson, Dan Mellström, Yoon Shin Cho, Hyejin Lee, Jian-Min Yuan, Woon-Puay Koh, Sang Youl Rhee, Jeong-Taek Woo, Iris M. Heid, Klaus J. Stark, Henry Völzke, Georg Homuth, Michele K. Evans, Alan B. Zonderman, Ozren Polasek, Gerard Pasterkamp, Imo E. Hoefer, Susan Redline, Katja Pahkala, Albertine J. Oldehinkel, Harold Snieder, Ginevra Biino, Reinhold Schmidt, Helena Schmidt, Y. Eugene Chen, Stefania Bandinelli, George Dedoussis, Thangavel Alphonse Thanaraj, Sharon L. R. Kardia, Norihiro Kato, Matthias B. Schulze, Giorgia Girotto, Bettina Jung, Carsten A. Böger, Peter K. Joshi, David A. Bennett, Philip L. Jager, Xiangfeng Lu, Vasiliki Mamakou, Morris Brown, Mark J. Caulfield, Patricia B. Munroe, Xiuqing Guo, Marina Ciullo, Jost B. Jonas, Nilesh J. Samani, Jaakko Kaprio, Päivi Pajukanta, Linda S. Adair, Sonny Augustin Bechayda, H. Janaka Silva, Ananda R. Wickremasinghe, Ronald M. Krauss, Jer-Yuarn Wu, Wei Zheng, Anneke I. Hollander, Dwaipayan Bharadwaj, Adolfo Correa, James G. Wilson, Lars Lind, Chew-Kiat Heng, Amanda E. Nelson, Yvonne M. Golightly, James F. Wilson, Brenda Penninx, Hyung-Lae Kim, John Attia, Rodney J. Scott, D. C. Rao, Donna K. Arnett, Steven C. Hunt, Mark Walker, Heikki A. Koistinen, Giriraj R. Chandak, Chittaranjan S. Yajnik, Josep M. Mercader, Teresa Tusié-Luna, Carlos A. Aguilar-Salinas, Clicerio Gonzalez Villalpando, Lorena Orozco, Myriam Fornage, E. Shyong Tai, Rob M. Dam, Terho Lehtimäki, Nish Chaturvedi, Mitsuhiro Yokota, Jianjun Liu, Dermot F. Reilly, Amy Jayne McKnight, Frank Kee, Karl-Heinz Jöckel, Mark I. McCarthy, Colin N. A. Palmer, Veronique Vitart, Caroline Hayward, Eleanor Simonsick, Cornelia M. Duijn, Fan Lu, Jia Qu, Haretsugu Hishigaki, Xu Lin, Winfried März, Esteban J. Parra, Miguel Cruz, Vilmundur Gudnason, Jean-Claude Tardif, Guillaume Lettre, Leen M. ’t Hart, Petra J. M. Elders, Scott M. Damrauer, Meena Kumari, Mika Kivimaki, Pim Harst, Tim D. Spector, Ruth J. F. Loos, Michael A. Province, Bruce M. Psaty, Ivan Brandslund, Peter P. Pramstaller, Kaare Christensen, Samuli Ripatti, Elisabeth Widén, Hakon Hakonarson, Struan F. A. Grant, Lambertus A. L. M. Kiemeney, Jacqueline Graaf, Markus Loeffler, Florian Kronenberg, Dongfeng Gu, Jeanette Erdmann, Heribert Schunkert, Paul W. Franks, Allan Linneberg, J. Wouter Jukema, Amit V. Khera, Minna Männikkö, Marjo-Riitta Jarvelin, Zoltan Kutalik, Francesco Cucca, Dennis O. Mook-Kanamori, Ko Willems Dijk, Hugh Watkins, David P. Strachan, Niels Grarup, Peter Sever, Neil Poulter, Jerome I. Rotter, Thomas M. Dantoft, Fredrik Karpe, Matt J. Neville, Nicholas J. Timpson, Ching-Yu Cheng, Tien-Yin Wong, Chiea Chuen Khor, Charumathi Sabanayagam, Annette Peters, Christian Gieger, Andrew T. Hattersley, Nancy L. Pedersen, Patrik K. E. Magnusson, Dorret I. Boomsma, Eco J. C. Geus, L. Adrienne Cupples, Joyce B. J. Meurs, Mohsen Ghanbari, Penny Gordon-Larsen, Wei Huang, Young Jin Kim, Yasuharu Tabara, Nicholas J. Wareham, Claudia Langenberg, Eleftheria Zeggini, Johanna Kuusisto, Markku Laakso, Erik Ingelsson, Goncalo Abecasis, John C. Chambers, Jaspal S. Kooner, Paul S. Vries, Alanna C. Morrison, Kari E. North, Martha Daviglus, Peter Kraft, Nicholas G. Martin, John B. Whitfield, Shahid Abbas, Danish Saleheen, Robin G. Walters, Michael V. Holmes, Corri Black, Blair H. Smith, Anne E. Justice, Aris Baras, Julie E. Buring, Paul M. Ridker, Daniel I. Chasman, Charles Kooperberg, Wei-Qi Wei, Gail P. Jarvik, Bahram Namjou, M. Geoffrey Hayes, Marylyn D. Ritchie, Pekka Jousilahti, Veikko Salomaa, Kristian Hveem, Bjørn Olav Åsvold, Michiaki Kubo, Yoichiro Kamatani, Yukinori Okada, Yoshinori Murakami, Unnur Thorsteinsdottir, Kari Stefansson, Yuk-Lam Ho, Julie A. Lynch, Daniel J. Rader, Philip S. Tsao, Kyong-Mi Chang, Kelly Cho, Christopher J. O’Donnell, John M. Gaziano, Peter Wilson, Charles N. Rotimi, Scott Hazelhurst, Michèle Ramsay, Richard C. Trembath, David A. Heel, Gen Tamiya, Masayuki Yamamoto, Bong-Jo Kim, Karen L. Mohlke, Timothy M. Frayling, Joel N. Hirschhorn, Sekar Kathiresan, Michael Boehnke, Pradeep Natarajan, Gina M. Peloso, Christopher D. Brown, Andrew P. Morris, Themistocles L. Assimes (), Panos Deloukas, Yan V. Sun and Cristen J. Willer ()
Additional contact information
Sarah E. Graham: University of Michigan
Shoa L. Clarke: VA Palo Alto Health Care System
Kuan-Han H. Wu: University of Michigan
Stavroula Kanoni: Queen Mary University of London
Greg J. M. Zajac: University of Michigan
Shweta Ramdas: University of Pennsylvania
Ida Surakka: University of Michigan
Ioanna Ntalla: Queen Mary University of London
Sailaja Vedantam: Boston Children’s Hospital
Thomas W. Winkler: University of Regensburg
Adam E. Locke: Washington University
Eirini Marouli: Queen Mary University of London
Mi Yeong Hwang: National Institute of Health
Sohee Han: National Institute of Health
Akira Narita: Tohoku University
Ananyo Choudhury: University of the Witwatersrand
Amy R. Bentley: National Institutes of Health
Kenneth Ekoru: National Institutes of Health
Anurag Verma: University of Pennsylvania
Bhavi Trivedi: Queen Mary University of London
Hilary C. Martin: Wellcome Sanger Institute
Karen A. Hunt: Queen Mary University of London
Qin Hui: Emory University Rollins School of Public Health
Derek Klarin: Malcolm Randall VA Medical Center
Xiang Zhu: VA Palo Alto Health Care System
Gudmar Thorleifsson: deCODE genetics/Amgen
Anna Helgadottir: deCODE genetics/Amgen
Daniel F. Gudbjartsson: deCODE genetics/Amgen
Hilma Holm: deCODE genetics/Amgen
Isleifur Olafsson: Landspitali–National University Hospital of Iceland
Masato Akiyama: RIKEN Center for Integrative Medical Sciences
Saori Sakaue: RIKEN Center for Integrative Medical Sciences
Chikashi Terao: RIKEN Center for Integrative Medical Sciences
Masahiro Kanai: Broad Institute of MIT and Harvard
Wei Zhou: University of Michigan
Ben M. Brumpton: Norwegian University of Science and Technology
Humaira Rasheed: Norwegian University of Science and Technology
Sanni E. Ruotsalainen: University of Helsinki
Aki S. Havulinna: University of Helsinki
Yogasudha Veturi: University of Pennsylvania
QiPing Feng: Division of Clinical Pharmacology, Department of Medicine, Vanderbilt University Medical Center
Elisabeth A. Rosenthal: University of Washington
Todd Lingren: Cincinnati Children’s Hospital Medical Center
Jennifer Allen Pacheco: Northwestern University
Sarah A. Pendergrass: Genentech
Jeffrey Haessler: Division of Public Health Sciences
Franco Giulianini: Brigham and Women’s Hospital
Yuki Bradford: University of Pennsylvania
Jason E. Miller: University of Pennsylvania
Archie Campbell: University of Edinburgh, Western General Hospital
Kuang Lin: University of Oxford
Iona Y. Millwood: University of Oxford
George Hindy: Qatar University College of Medicine, QU Health
Asif Rasheed: Center for Non-Communicable Diseases
Jessica D. Faul: University of Michigan
Wei Zhao: University of Michigan
David R. Weir: University of Michigan
Constance Turman: Harvard T. H. Chan School of Public Health
Hongyan Huang: Harvard T. H. Chan School of Public Health
Mariaelisa Graff: University of North Carolina
Anubha Mahajan: University of Oxford
Michael R. Brown: The University of Texas Health Science Center at Houston
Weihua Zhang: Imperial College London
Ketian Yu: University of Michigan
Ellen M. Schmidt: University of Michigan
Anita Pandit: University of Michigan
Stefan Gustafsson: Uppsala University
Xianyong Yin: University of Michigan
Jian’an Luan: University of Cambridge School of Clinical Medicine
Jing-Hua Zhao: University of Cambridge Strangeways Research Laboratory
Fumihiko Matsuda: Kyoto University Graduate School of Medicine
Hye-Mi Jang: National Institute of Health
Kyungheon Yoon: National Institute of Health
Carolina Medina-Gomez: University Medical Center Rotterdam
Achilleas Pitsillides: Boston University School of Public Health
Jouke Jan Hottenga: Vrije Universiteit Amsterdam
Gonneke Willemsen: Vrije Universiteit Amsterdam
Andrew R. Wood: University of Exeter Medical School, University of Exeter
Yingji Ji: University of Exeter Medical School, University of Exeter
Zishan Gao: Nanjing University of Chinese Medicine
Simon Haworth: MRC Integrative Epidemiology Unit at the University of Bristol
Ruth E. Mitchell: MRC Integrative Epidemiology Unit at the University of Bristol
Jin Fang Chai: National University of Singapore and National University Health System
Mette Aadahl: Bispebjerg and Frederiksberg Hospital
Jie Yao: Lundquist Institute for Biomedical Innovations (Formerly LABioMed) at Harbor-UCLA Medical Center
Ani Manichaikul: University of Virginia
Helen R. Warren: Queen Mary University of London
Julia Ramirez: Queen Mary University of London
Jette Bork-Jensen: University of Copenhagen
Line L. Kårhus: Bispebjerg and Frederiksberg Hospital
Anuj Goel: University of Oxford
Maria Sabater-Lleal: Research Institute of Hospital de la Santa Creu i Sant Pau (IIB Sant Pau)
Raymond Noordam: Leiden University Medical Center
Carlo Sidore: Italian National Council of Research (IRGB CNR)
Edoardo Fiorillo: Italian National Council of Research (IRGB CNR)
Aaron F. McDaid: University of Lausanne
Pedro Marques-Vidal: Lausanne University Hospital and University of Lausanne
Matthias Wielscher: Imperial College London
Stella Trompet: Leiden University Medical Center
Naveed Sattar: BHF Glasgow Cardiovascular Research Centre, Faculty of Medicine
Line T. Møllehave: Bispebjerg and Frederiksberg Hospital
Betina H. Thuesen: Bispebjerg and Frederiksberg Hospital
Matthias Munz: University of Lübeck, DZHK (German Research Centre for Cardiovascular Research), Partner site Hamburg/Lübeck/Kiel, University Heart Center Lübeck
Lingyao Zeng: Technische Universität München
Jianfeng Huang: State Key Laboratory of Cardiovascular Disease, Fuwai Hospital, National Center for Cardiovascular Diseases, Chinese Academy of Medical Sciences and Peking Union Medical College
Bin Yang: State Key Laboratory of Cardiovascular Disease, Fuwai Hospital, National Center for Cardiovascular Diseases, Chinese Academy of Medical Sciences and Peking Union Medical College
Alaitz Poveda: Lund University Diabetes Centre
Azra Kurbasic: Lund University Diabetes Centre
Claudia Lamina: Medical University of Innsbruck
Lukas Forer: Medical University of Innsbruck
Markus Scholz: University of Leipzig
Tessel E. Galesloot: Radboud Institute for Health Sciences
Jonathan P. Bradfield: Quantinuum Research
E. Warwick Daw: Washington University School of Medicine
Joseph M. Zmuda: University of Pittsburgh
Jonathan S. Mitchell: Affiliated Institute of the University of Lübeck
Christian Fuchsberger: Affiliated Institute of the University of Lübeck
Henry Christensen: Lillebaelt Hospital
Jennifer A. Brody: University of Washington
Mary F. Feitosa: Washington University School of Medicine
Mary K. Wojczynski: Washington University School of Medicine
Michael Preuss: Icahn School of Medicine at Mount Sinai
Massimo Mangino: King’s College London
Paraskevi Christofidou: King’s College London
Niek Verweij: University Medical Center Groningen, University of Groningen
Jan W. Benjamins: University Medical Center Groningen, University of Groningen
Jorgen Engmann: University College London
Rachel L. Kember: University of Pennsylvania
Roderick C. Slieker: Amsterdam Public Health Research Institute
Ken Sin Lo: Montreal Heart Institute
Nuno R. Zilhao: Icelandic Heart Association
Phuong Le: University of Toronto at Mississauga
Marcus E. Kleber: Heidelberg University
Graciela E. Delgado: Heidelberg University
Shaofeng Huo: University of Chinese Academy of Sciences, Chinese Academy of Sciences
Daisuke D. Ikeda: Tokushima Research Institute, Otsuka Pharmaceutical
Hiroyuki Iha: Tokushima Research Institute, Otsuka Pharmaceutical
Jian Yang: The University of Queensland
Jun Liu: University of Oxford
Hampton L. Leonard: National Institute on Aging, NIH
Jonathan Marten: University of Edinburgh, Western General Hospital
Börge Schmidt: University of Duisburg-Essen
Marina Arendt: University of Duisburg-Essen
Laura J. Smyth: Queen’s University of Belfast
Marisa Cañadas-Garre: Queen’s University of Belfast
Chaolong Wang: Tongji Medical College, Huazhong University of Science and Technology
Masahiro Nakatochi: Nagoya University Graduate School of Medicine
Andrew Wong: MRC Unit for Lifelong Health and Ageing at UCL
Nina Hutri-Kähönen: Tampere University Hospital
Xueling Sim: National University of Singapore and National University Health System
Rui Xia: University of Texas Health Science Center at Houston
Alicia Huerta-Chagoya: Instituto de Investigaciones Biomédicas, UNAM, Ciudad de Mexico
Juan Carlos Fernandez-Lopez: Instituto Nacional de Medicina Genómica, Ciudad de Mexico
Valeriya Lyssenko: University of Bergen
Meraj Ahmed: CSIR–Centre for Cellular and Molecular Biology
Anne U. Jackson: University of Michigan
Noha A. Yousri: Weill Cornell Medicine-Qatar
Marguerite R. Irvin: University of Alabama at Birmingham
Christopher Oldmeadow: Hunter Medical Research Institute
Han-Na Kim: Sungkyunkwan University School of Medicine
Seungho Ryu: Kangbuk Samsung Hospital, Sungkyunkwan University School of Medicine
Paul R. H. J. Timmers: University of Edinburgh, Western General Hospital
Liubov Arbeeva: University of North Carolina
Rajkumar Dorajoo: Genome Institute of Singapore, Agency for Science, Technology and Research
Leslie A. Lange: University of Colorado, Denver
Xiaoran Chai: Singapore Eye Research Institute, Singapore National Eye Centre
Gauri Prasad: CSIR–Institute of Genomics and Integrative Biology
Laura Lorés-Motta: Radboud University Nijmegen Medical Center
Marc Pauper: Radboud University Nijmegen Medical Center
Jirong Long: Vanderbilt University Medical Center
Xiaohui Li: Lundquist Institute for Biomedical Innovations (Formerly LABioMed) at Harbor-UCLA Medical Center
Elizabeth Theusch: University of California, San Francisco
Fumihiko Takeuchi: National Center for Global Health and Medicine
Cassandra N. Spracklen: University of North Carolina
Anu Loukola: University of Helsinki
Sailalitha Bollepalli: University of Helsinki
Sophie C. Warner: University of Leicester
Ya Xing Wang: Beijing Key Laboratory of Ophthalmology and Visual Sciences
Wen B. Wei: Beijing Tongren Hospital, Capital Medical University
Teresa Nutile: Institute of Genetics and Biophysics “Adriano Buzzati-Traverso”–CNR
Daniela Ruggiero: Institute of Genetics and Biophysics “Adriano Buzzati-Traverso”–CNR
Yun Ju Sung: Washington University
Yi-Jen Hung: Tri-Service General Hospital Songshan Branch
Shufeng Chen: State Key Laboratory of Cardiovascular Disease, Fuwai Hospital, National Center for Cardiovascular Diseases, Chinese Academy of Medical Sciences and Peking Union Medical College
Fangchao Liu: State Key Laboratory of Cardiovascular Disease, Fuwai Hospital, National Center for Cardiovascular Diseases, Chinese Academy of Medical Sciences and Peking Union Medical College
Jingyun Yang: Rush University Medical Center
Katherine A. Kentistou: University of Edinburgh
Mathias Gorski: University of Regensburg
Marco Brumat: University of Trieste
Karina Meidtner: German Institute of Human Nutrition Potsdam-Rehbruecke
Lawrence F. Bielak: School of Public Health, University of Michigan
Jennifer A. Smith: University of Michigan
Prashantha Hebbar: Dasman Diabetes Institute
Aliki-Eleni Farmaki: Harokopio University of Athens
Edith Hofer: Clinical Division of Neurogeriatrics, Medical University of Graz
Maoxuan Lin: University of North Carolina at Charlotte
Chao Xue: University of Michigan
Jifeng Zhang: University of Michigan
Maria Pina Concas: IRCCS Burlo Garofolo
Simona Vaccargiu: National Research Council of Italy, UOS of Sassari
Peter J. Most: University Medical Center Groningen, University of Groningen
Niina Pitkänen: University of Turku
Brian E. Cade: Brigham and Women’s Hospital
Jiwon Lee: Brigham and Women’s Hospital
Sander W. Laan: University Medical Center Utrecht, Utrecht University
Kumaraswamy Naidu Chitrala: National Institute on Aging Intramural Research Program, NIH Biomedical Research Center, NIA
Stefan Weiss: University of Greifswald and University Medicine Greifswald
Martina E. Zimmermann: University of Regensburg
Jong Young Lee: Oneomics
Hyeok Sun Choi: Hallym University
Maria Nethander: University of Gothenburg
Sandra Freitag-Wolf: Kiel University
Lorraine Southam: Helmholtz Zentrum München–German Research Center for Environmental Health
Nigel W. Rayner: Wellcome Sanger Institute
Carol A. Wang: University of Newcastle
Shih-Yi Lin: Taichung Veterans General Hospital
Jun-Sing Wang: Taichung Veterans General Hospital
Christian Couture: Université Laval
Leo-Pekka Lyytikäinen: Fimlab Laboratories
Kjell Nikus: Tampere University Hospital
Gabriel Cuellar-Partida: University of Queensland Diamantina Institute, Translational Research Institute
Henrik Vestergaard: University of Copenhagen
Bertha Hildalgo: University of Alabama at Birmingham
Olga Giannakopoulou: Queen Mary University of London
Qiuyin Cai: Vanderbilt University Medical Center
Morgan O. Obura: Amsterdam Public Health Research Institute
Jessica Setten: University Medical Center Utrecht, Utrecht University
Xiaoyin Li: Case Western Reserve University
Karen Schwander: Washington University School of Medicine
Natalie Terzikhan: University Medical Center Rotterdam
Jae Hun Shin: Hallym University
Rebecca D. Jackson: Ohio State University
Alexander P. Reiner: University of Washington
Lisa Warsinger Martin: George Washington University
Zhengming Chen: University of Oxford
Liming Li: Peking University Health Science Center
Heather M. Highland: University of North Carolina
Kristin L. Young: University of North Carolina
Takahisa Kawaguchi: Kyoto University Graduate School of Medicine
Joachim Thiery: University of Leipzig
Joshua C. Bis: University of Washington
Girish N. Nadkarni: Icahn School of Medicine at Mount Sinai
Lenore J. Launer: National Institute on Aging, NIH
Huaixing Li: University of Chinese Academy of Sciences, Chinese Academy of Sciences
Mike A. Nalls: National Institute on Aging, NIH
Olli T. Raitakari: University of Turku
Sahoko Ichihara: Jichi Medical University School of Medicine
Sarah H. Wild: University of Edinburgh
Christopher P. Nelson: University of Leicester
Harry Campbell: University of Edinburgh
Susanne Jäger: German Institute of Human Nutrition Potsdam-Rehbruecke
Toru Nabika: Shimane University School of Medicine
Fahd Al-Mulla: Dasman Diabetes Institute
Harri Niinikoski: Turku University Hospital and University of Turku
Peter S. Braund: University of Leicester
Ivana Kolcic: University of Split
Peter Kovacs: University of Leipzig Medical Center
Tota Giardoglou: Harokopio University, Eleftheriou Venizelou
Tomohiro Katsuya: Osaka University Graduate School of Medicine
Konain Fatima Bhatti: Queen Mary University of London
Dominique Kleijn: University Medical Center Utrecht, Utrecht University
Gert J. Borst: University Medical Center Utrecht, Utrecht University
Eung Kweon Kim: Yonsei University College of Medicine
Hieab H. H. Adams: University Medical Center Rotterdam
M. Arfan Ikram: University Medical Center Rotterdam
Xiaofeng Zhu: Case Western Reserve University
Folkert W. Asselbergs: University Medical Center Utrecht, Utrecht University
Adriaan O. Kraaijeveld: University Medical Center Utrecht, Utrecht University
Joline W. J. Beulens: Amsterdam Public Health Research Institute
Xiao-Ou Shu: Vanderbilt University Medical Center
Loukianos S. Rallidis: National and Kapodistrian University of Athens, Attikon University Hospital
Oluf Pedersen: University of Copenhagen
Torben Hansen: University of Copenhagen
Paul Mitchell: University of Sydney
Alex W. Hewitt: University of Tasmania
Mika Kähönen: Tampere University Hospital
Louis Pérusse: Université Laval
Claude Bouchard: Pennington Biomedical Research Center
Anke Tönjes: University of Leipzig Medical Center
Yii- Der Ida Chen: Lundquist Institute for Biomedical Innovations (Formerly LABioMed) at Harbor-UCLA Medical Center
Craig E. Pennell: University of Newcastle
Trevor A. Mori: University of Western Australia
Wolfgang Lieb: Kiel University
Andre Franke: Kiel University
Claes Ohlsson: University of Gothenburg
Dan Mellström: University of Gothenburg
Yoon Shin Cho: Hallym University
Hyejin Lee: Ewha Womans University School of Medicine
Jian-Min Yuan: University of Pittsburgh
Woon-Puay Koh: National University of Singapore
Sang Youl Rhee: Kyung Hee University School of Medicine
Jeong-Taek Woo: Kyung Hee University School of Medicine
Iris M. Heid: University of Regensburg
Klaus J. Stark: University of Regensburg
Henry Völzke: University Medicine Greifswald
Georg Homuth: University of Greifswald and University Medicine Greifswald
Michele K. Evans: National Institute on Aging Intramural Research Program, NIH Biomedical Research Center
Alan B. Zonderman: National Institute on Aging Intramural Research Program, NIH Biomedical Research Center
Ozren Polasek: University of Split
Gerard Pasterkamp: University Medical Center Utrecht, Utrecht University
Imo E. Hoefer: University Medical Center Utrecht, Utrecht University
Susan Redline: Brigham and Women’s Hospital
Katja Pahkala: University of Turku
Albertine J. Oldehinkel: University of Groningen, University Medical Center Groningen
Harold Snieder: University Medical Center Groningen, University of Groningen
Ginevra Biino: National Research Council of Italy
Reinhold Schmidt: Clinical Division of Neurogeriatrics, Medical University of Graz
Helena Schmidt: Medical University of Graz
Y. Eugene Chen: University of Michigan
Stefania Bandinelli: Local Health Unit Toscana Centro
George Dedoussis: Harokopio University of Athens
Thangavel Alphonse Thanaraj: Dasman Diabetes Institute
Sharon L. R. Kardia: School of Public Health, University of Michigan
Norihiro Kato: National Center for Global Health and Medicine
Matthias B. Schulze: German Institute of Human Nutrition Potsdam-Rehbruecke
Giorgia Girotto: University of Trieste
Bettina Jung: University Hospital Regensburg
Carsten A. Böger: University Hospital Regensburg
Peter K. Joshi: University of Edinburgh
David A. Bennett: Rush University Medical Center
Philip L. Jager: Columbia University Medical Center
Xiangfeng Lu: State Key Laboratory of Cardiovascular Disease, Fuwai Hospital, National Center for Cardiovascular Diseases, Chinese Academy of Medical Sciences and Peking Union Medical College
Vasiliki Mamakou: Medical School, National and Kapodistrian University Athens
Morris Brown: Queen Mary University of London
Mark J. Caulfield: Queen Mary University of London
Patricia B. Munroe: Queen Mary University of London
Xiuqing Guo: Lundquist Institute for Biomedical Innovations (Formerly LABioMed) at Harbor-UCLA Medical Center
Marina Ciullo: Institute of Genetics and Biophysics “Adriano Buzzati-Traverso”–CNR
Jost B. Jonas: Heidelberg University
Nilesh J. Samani: University of Leicester
Jaakko Kaprio: University of Helsinki
Päivi Pajukanta: University of California, Los Angeles
Linda S. Adair: University of North Carolina
Sonny Augustin Bechayda: University of San Carlos
H. Janaka Silva: University of Kelaniya
Ananda R. Wickremasinghe: Faculty of Medicine, University of Kelaniya
Ronald M. Krauss: University of California, San Francisco
Jer-Yuarn Wu: Institute of Biomedical Sciences, Academia Sinica
Wei Zheng: Vanderbilt University Medical Center
Anneke I. Hollander: Radboud University Nijmegen Medical Center
Dwaipayan Bharadwaj: CSIR–Institute of Genomics and Integrative Biology Campus
Adolfo Correa: University of Mississippi Medical Center
James G. Wilson: University of Mississippi Medical Center
Lars Lind: Uppsala University
Chew-Kiat Heng: National University of Singapore
Amanda E. Nelson: University of North Carolina
Yvonne M. Golightly: University of North Carolina
James F. Wilson: University of Edinburgh, Western General Hospital
Brenda Penninx: Vrije Universiteit Amsterdam
Hyung-Lae Kim: College of Medicine, Ewha Womans University
John Attia: Hunter Medical Research Institute
Rodney J. Scott: Hunter Medical Research Institute
D. C. Rao: Washington University School of Medicine
Donna K. Arnett: University of Kentucky, College of Public Health
Steven C. Hunt: Weill Cornell Medicine-Qatar
Mark Walker: Newcastle University
Heikki A. Koistinen: Finnish Institute for Health and Welfare
Giriraj R. Chandak: CSIR–Centre for Cellular and Molecular Biology
Chittaranjan S. Yajnik: KEM Hospital and Research Centre
Josep M. Mercader: Harvard Medical School
Teresa Tusié-Luna: Unidad de Biología Molecular y Medicina Genómica, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán
Carlos A. Aguilar-Salinas: Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán
Clicerio Gonzalez Villalpando: Instituto Nacional de Salud Publica y Centro de Estudios en Diabetes
Lorena Orozco: Instituto Nacional de Medicina Genómica
Myriam Fornage: University of Texas Health Science Center at Houston
E. Shyong Tai: National University of Singapore and National University Health System
Rob M. Dam: National University of Singapore and National University Health System
Terho Lehtimäki: Fimlab Laboratories
Nish Chaturvedi: MRC Unit for Lifelong Health and Ageing at UCL
Mitsuhiro Yokota: Kurume University School of Medicine
Jianjun Liu: Genome Institute of Singapore, Agency for Science, Technology and Research
Dermot F. Reilly: Merck Sharp & Dohme
Amy Jayne McKnight: Queen’s University of Belfast
Frank Kee: Queen’s University of Belfast
Karl-Heinz Jöckel: University of Duisburg-Essen
Mark I. McCarthy: Genentech
Colin N. A. Palmer: University of Dundee, Ninwells Hospital and Medical School
Veronique Vitart: University of Edinburgh, Western General Hospital
Caroline Hayward: University of Edinburgh, Western General Hospital
Eleanor Simonsick: Intramural Research Program, National Institute on Aging
Cornelia M. Duijn: University of Oxford
Fan Lu: Wenzhou Medical University
Jia Qu: Wenzhou Medical University
Haretsugu Hishigaki: Tokushima Research Institute, Otsuka Pharmaceutical
Xu Lin: University of Chinese Academy of Sciences, Chinese Academy of Sciences
Winfried März: Heidelberg University
Esteban J. Parra: University of Toronto at Mississauga
Miguel Cruz: Unidad de Investigacion Medica en Bioquimica, Hospital de Especialidades, Centro Medico Nacional Siglo XXI, Instituto Mexicano del Seguro Social
Vilmundur Gudnason: Icelandic Heart Association
Jean-Claude Tardif: Montreal Heart Institute
Guillaume Lettre: Montreal Heart Institute
Leen M. ’t Hart: Amsterdam Public Health Research Institute
Petra J. M. Elders: Amsterdam Public Health Research Institute
Scott M. Damrauer: University of Pennsylvania
Meena Kumari: University of Essex
Mika Kivimaki: University College London
Pim Harst: University Medical Center Groningen, University of Groningen
Tim D. Spector: King’s College London
Ruth J. F. Loos: Icahn School of Medicine at Mount Sinai
Michael A. Province: Washington University School of Medicine
Bruce M. Psaty: University of Washington
Ivan Brandslund: Lillebaelt Hospital
Peter P. Pramstaller: Affiliated Institute of the University of Lübeck
Kaare Christensen: University of Southern Denmark
Samuli Ripatti: University of Helsinki
Elisabeth Widén: University of Helsinki
Hakon Hakonarson: Center for Applied Genomics, Children’s Hospital of Philadelphia
Struan F. A. Grant: University of Pennsylvania
Lambertus A. L. M. Kiemeney: Radboud Institute for Health Sciences
Jacqueline Graaf: Radboud Institute for Health Sciences
Markus Loeffler: University of Leipzig
Florian Kronenberg: German Chronic Kidney Disease Study
Dongfeng Gu: State Key Laboratory of Cardiovascular Disease, Fuwai Hospital, National Center for Cardiovascular Diseases, Chinese Academy of Medical Sciences and Peking Union Medical College
Jeanette Erdmann: University of Lübeck, DZHK (German Research Centre for Cardiovascular Research), Partner site Hamburg/Lübeck/Kiel, and University Heart Center Lübeck
Heribert Schunkert: Deutsches Herzzentrum München, Klinik für Herz- und Kreislauferkrankungen, Technische Universität München
Paul W. Franks: Lund University Diabetes Centre
Allan Linneberg: Bispebjerg and Frederiksberg Hospital
J. Wouter Jukema: Leiden University Medical Center
Amit V. Khera: Massachusetts General Hospital
Minna Männikkö: University of Oulu
Marjo-Riitta Jarvelin: Imperial College London
Zoltan Kutalik: Swiss Institute of Bioinformatics
Francesco Cucca: Italian National Council of Research (IRGB CNR)
Dennis O. Mook-Kanamori: Leiden University Medical Center
Ko Willems Dijk: Leiden University Medical Center
Hugh Watkins: University of Oxford
David P. Strachan: St George’s University of London
Niels Grarup: University of Copenhagen
Peter Sever: Imperial College London
Neil Poulter: Imperial College London
Jerome I. Rotter: Lundquist Institute for Biomedical Innovations (Formerly LABioMed) at Harbor-UCLA Medical Center
Thomas M. Dantoft: Bispebjerg and Frederiksberg Hospital
Fredrik Karpe: University of Oxford, Churchill Hospital
Matt J. Neville: University of Oxford, Churchill Hospital
Nicholas J. Timpson: MRC Integrative Epidemiology Unit at the University of Bristol
Ching-Yu Cheng: Singapore Eye Research Institute, Singapore National Eye Centre
Tien-Yin Wong: Singapore Eye Research Institute, Singapore National Eye Centre
Chiea Chuen Khor: Genome Institute of Singapore, Agency for Science, Technology and Research
Charumathi Sabanayagam: Singapore Eye Research Institute, Singapore National Eye Centre
Annette Peters: German Research Center for Environmental Health
Christian Gieger: German Research Center for Environmental Health
Andrew T. Hattersley: University of Exeter
Nancy L. Pedersen: Karolinska Institutet
Patrik K. E. Magnusson: Karolinska Institutet
Dorret I. Boomsma: Vrije Universiteit Amsterdam
Eco J. C. Geus: Vrije Universiteit Amsterdam
L. Adrienne Cupples: Boston University School of Public Health
Joyce B. J. Meurs: University Medical Center Rotterdam
Mohsen Ghanbari: University Medical Center Rotterdam
Penny Gordon-Larsen: University of North Carolina
Wei Huang: Shanghai-MOST Key Laboratory of Health and Disease Genomics, Chinese National Human Genome Center at Shanghai
Young Jin Kim: National Institute of Health
Yasuharu Tabara: Kyoto University Graduate School of Medicine
Nicholas J. Wareham: University of Cambridge School of Clinical Medicine
Claudia Langenberg: University of Cambridge School of Clinical Medicine
Eleftheria Zeggini: Helmholtz Zentrum München–German Research Center for Environmental Health
Johanna Kuusisto: University of Eastern Finland and Kuopio University Hospital
Markku Laakso: University of Eastern Finland and Kuopio University Hospital
Erik Ingelsson: Stanford University School of Medicine
Goncalo Abecasis: University of Michigan
John C. Chambers: Imperial College London
Jaspal S. Kooner: London North West University Healthcare NHS Trust
Paul S. Vries: The University of Texas Health Science Center at Houston
Alanna C. Morrison: The University of Texas Health Science Center at Houston
Kari E. North: University of North Carolina
Martha Daviglus: University of Illinois College of Medicine
Peter Kraft: Harvard T. H. Chan School of Public Health
Nicholas G. Martin: QIMR Berghofer Medical Research Institute
John B. Whitfield: QIMR Berghofer Medical Research Institute
Shahid Abbas: Center for Non-Communicable Diseases
Danish Saleheen: Center for Non-Communicable Diseases
Robin G. Walters: University of Oxford
Michael V. Holmes: University of Oxford
Corri Black: University of Aberdeen
Blair H. Smith: Ninewells Hospital and Medical School, University of Dundee
Anne E. Justice: Geisinger Health
Aris Baras: Regeneron Pharmaceuticals
Julie E. Buring: Brigham and Women’s Hospital
Paul M. Ridker: Brigham and Women’s Hospital
Daniel I. Chasman: Brigham and Women’s Hospital
Charles Kooperberg: Division of Public Health Sciences
Wei-Qi Wei: Vanderbilt University Medical Center
Gail P. Jarvik: University of Washington Medical Center
Bahram Namjou: Cincinnati Children’s Hospital Medical Center (CCHMC)
M. Geoffrey Hayes: Northwestern University, Feinberg School of Medicine
Marylyn D. Ritchie: University of Pennsylvania
Pekka Jousilahti: Finnish Institute for Health and Welfare
Veikko Salomaa: Finnish Institute for Health and Welfare
Kristian Hveem: Norwegian University of Science and Technology
Bjørn Olav Åsvold: Norwegian University of Science and Technology
Michiaki Kubo: RIKEN Center for Integrative Medical Sciences
Yoichiro Kamatani: RIKEN Center for Integrative Medical Sciences
Yukinori Okada: RIKEN Center for Integrative Medical Sciences
Yoshinori Murakami: The University of Tokyo
Unnur Thorsteinsdottir: deCODE genetics/Amgen
Kari Stefansson: deCODE genetics/Amgen
Yuk-Lam Ho: VA Boston Healthcare System
Julie A. Lynch: VA Informatics and Computing Infrastructure, VA Salt Lake City Health Care System
Daniel J. Rader: University of Pennsylvania
Philip S. Tsao: VA Palo Alto Health Care System
Kyong-Mi Chang: University of Pennsylvania
Kelly Cho: VA Boston Healthcare System
Christopher J. O’Donnell: VA Boston Healthcare System
John M. Gaziano: VA Boston Healthcare System
Peter Wilson: Atlanta VA Medical Center
Charles N. Rotimi: National Institutes of Health
Scott Hazelhurst: University of the Witwatersrand
Michèle Ramsay: University of the Witwatersrand
Richard C. Trembath: King’s College London
David A. Heel: Queen Mary University of London
Gen Tamiya: Tohoku University
Masayuki Yamamoto: Tohoku University
Bong-Jo Kim: National Institute of Health
Karen L. Mohlke: University of North Carolina
Timothy M. Frayling: University of Exeter Medical School, University of Exeter
Joel N. Hirschhorn: Boston Children’s Hospital
Sekar Kathiresan: Broad Institute
Michael Boehnke: University of Michigan
Pradeep Natarajan: Broad Institute of Harvard and MIT
Gina M. Peloso: Boston University School of Public Health
Christopher D. Brown: University of Pennsylvania
Andrew P. Morris: The University of Manchester
Themistocles L. Assimes: VA Palo Alto Health Care System
Panos Deloukas: Queen Mary University of London
Yan V. Sun: Emory University Rollins School of Public Health
Cristen J. Willer: University of Michigan

Nature, 2021, vol. 600, issue 7890, 675-679

Abstract: Abstract Increased blood lipid levels are heritable risk factors of cardiovascular disease with varied prevalence worldwide owing to different dietary patterns and medication use1. Despite advances in prevention and treatment, in particular through reducing low-density lipoprotein cholesterol levels2, heart disease remains the leading cause of death worldwide3. Genome-wideassociation studies (GWAS) of blood lipid levels have led to important biological and clinical insights, as well as new drug targets, for cardiovascular disease. However, most previous GWAS4–23 have been conducted in European ancestry populations and may have missed genetic variants that contribute to lipid-level variation in other ancestry groups. These include differences in allele frequencies, effect sizes and linkage-disequilibrium patterns24. Here we conduct a multi-ancestry, genome-wide genetic discovery meta-analysis of lipid levels in approximately 1.65 million individuals, including 350,000 of non-European ancestries. We quantify the gain in studying non-European ancestries and provide evidence to support the expansion of recruitment of additional ancestries, even with relatively small sample sizes. We find that increasing diversity rather than studying additional individuals of European ancestry results in substantial improvements in fine-mapping functional variants and portability of polygenic prediction (evaluated in approximately 295,000 individuals from 7 ancestry groupings). Modest gains in the number of discovered loci and ancestry-specific variants were also achieved. As GWAS expand emphasis beyond the identification of genes and fundamental biology towards the use of genetic variants for preventive and precision medicine25, we anticipate that increased diversity of participants will lead to more accurate and equitable26 application of polygenic scores in clinical practice.

Date: 2021
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DOI: 10.1038/s41586-021-04064-3

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