Nat Genet. 2019 Apr;51(4):636-648. doi: 10.1038/s41588-019-0378-y. Epub 2019 Mar 29.

Multi-ancestry genome-wide gene-smoking interaction study of 387,272 individuals identifies new loci associated with serum lipids.

Bentley AR¹, Sung YJ², Brown MR³, Winkler TW⁴, Kraja AT⁵, Ntalla I⁶, Schwander K², Chasman DI^7,⁸, Lim E⁹, Deng X⁹, Guo X¹⁰, Liu J¹¹, Lu Y¹², Cheng CY^13,^14,¹⁵, Sim X¹⁶, Vojinovic D¹⁷, Huffman JE¹⁸, Musani SK¹⁹, Li C²⁰, Feitosa MF⁵, Richard MA²¹, Noordam R²², Baker J²³, Chen G²³, Aschard H^24,²⁵, Bartz TM^26,²⁷, Ding J²⁸, Dorajoo R²⁹, Manning AK^30,³¹, Rankinen T³², Smith AV^33,³⁴, Tajuddin SM³⁵, Zhao W³⁶, Graff M³⁷, Alver M³⁸, Boissel M³⁹, Chai JF¹⁶, Chen X⁴⁰, Divers J⁴¹, Evangelou E^42,⁴³, Gao C⁴⁴, Goel A^45,⁴⁶, Hagemeijer Y⁴⁷, Harris SE^48,⁴⁹, Hartwig FP^50,⁵¹, He M⁵², Horimoto ARVR⁵³, Hsu FC⁴¹, Hung YJ^54,⁵⁵, Jackson AU⁵⁶, Kasturiratne A⁵⁷, Komulainen P⁵⁸, Kühnel B^59,⁶⁰, Leander K⁶¹, Lin KH⁶², Luan J⁶³, Lyytikäinen LP^64,⁶⁵, Matoba N⁶⁶, Nolte IM⁶⁷, Pietzner M^68,⁶⁹, Prins B⁷⁰, Riaz M^71,⁷², Robino A⁷³, Said MA⁴⁷, Schupf N⁷⁴, Scott RA⁶³, Sofer T^31,⁷⁵, Stancáková A⁷⁶, Takeuchi F⁷⁷, Tayo BO⁷⁸, van der Most PJ⁶⁷, Varga TV⁷⁹, Wang TD^80,^81,⁸², Wang Y⁸³, Ware EB⁸⁴, Wen W⁸⁵, Xiang YB⁸⁶, Yanek LR⁸⁷, Zhang W^88,⁸⁹, Zhao JH⁶³, Adeyemo A²³, Afaq S⁸⁸, Amin N¹⁷, Amini M⁶⁷, Arking DE⁹⁰, Arzumanyan Z¹⁰, Aung T^13,^15,⁹¹, Ballantyne C^92,⁹³, Barr RG⁹⁴, Bielak LF³⁶, Boerwinkle E^3,⁹⁵, Bottinger EP¹², Broeckel U⁹⁶, Brown M^6,⁹⁷, Cade BE⁷⁵, Campbell A⁹⁸, Canouil M³⁹, Charumathi S^13,¹⁴, Chen YI¹⁰, Christensen K⁹⁹; COGENT-Kidney Consortium, Concas MP⁷³, Connell JM¹⁰⁰, de Las Fuentes L^2,¹⁰¹, de Silva HJ¹⁰², de Vries PS³, Doumatey A²³, Duan Q¹⁰³, Eaton CB¹⁰⁴, Eppinga RN⁴⁷, Faul JD⁸⁴, Floyd JS^27,¹⁰⁵, Forouhi NG⁶³, Forrester T¹⁰⁶, Friedlander Y¹⁰⁷, Gandin I¹⁰⁸, Gao H⁴², Ghanbari M^17,¹⁰⁹, Gharib SA¹¹⁰, Gigante B⁶¹, Giulianini F⁷, Grabe HJ¹¹¹, Gu CC², Harris TB¹¹², Heikkinen S^76,¹¹³, Heng CK^114,¹¹⁵, Hirata M¹¹⁶, Hixson JE³, Ikram MA^17,^117,¹¹⁸; EPIC-InterAct Consortium, Jia Y¹⁰, Joehanes R^119,¹²⁰, Johnson C¹²¹, Jonas JB^122,¹²³, Justice AE³⁷, Katsuya T^124,¹²⁵, Khor CC²⁹, Kilpeläinen TO^126,¹²⁷, Koh WP^16,¹²⁸, Kolcic I¹²⁹, Kooperberg C¹³⁰, Krieger JE⁵³, Kritchevsky SB¹³¹, Kubo M¹³², Kuusisto J⁷⁶, Lakka TA^58,^113,¹³³, Langefeld CD⁴¹, Langenberg C⁶³, Launer LJ¹¹², Lehne B¹³⁴, Lewis CE¹³⁵, Li Y², Liang J⁸³, Lin S⁵, Liu CT⁹, Liu J^29,¹³⁶, Liu K¹³⁷, Loh M^88,^138,^139,¹⁴⁰, Lohman KK⁴¹, Louie T¹⁴¹, Luzzi A¹⁰, Mägi R³⁸, Mahajan A⁴⁶, Manichaikul AW¹⁴², McKenzie CA¹⁴³, Meitinger T^144,^145,¹⁴⁶, Metspalu A³⁸, Milaneschi Y¹⁴⁷, Milani L³⁸, Mohlke KL¹⁰³, Momozawa Y¹⁴⁸, Morris AP^46,¹⁴⁹, Murray AD¹⁵⁰, Nalls MA^151,¹⁵², Nauck M^68,⁶⁹, Nelson CP^71,⁷², North KE³⁷, O'Connell JR^153,¹⁵⁴, Palmer ND¹⁵⁵, Papanicolau GJ¹⁵⁶, Pedersen NL⁴⁰, Peters A^60,¹⁵⁷, Peyser PA³⁶, Polasek O^129,^158,¹⁵⁹, Poulter N¹⁶⁰, Raitakari OT^161,¹⁶², Reiner AP¹³⁰, Renström F^79,¹⁶³, Rice TK², Rich SS¹⁴², Robinson JG¹⁶⁴, Rose LM⁷, Rosendaal FR¹⁶⁵, Rudan I¹⁶⁶, Schmidt CO¹⁶⁷, Schreiner PJ¹⁶⁸, Scott WR^134,¹⁶⁹, Sever P¹⁶⁹, Shi Y¹³, Sidney S¹⁷⁰, Sims M¹⁹, Smith JA^36,⁸⁴, Snieder H⁶⁷, Starr JM^48,¹⁷¹, Strauch K^172,¹⁷³, Stringham HM⁵⁶, Tan NYQ¹³, Tang H¹⁷⁴, Taylor KD¹⁰, Teo YY^16,^29,^175,^176,¹⁷⁷, Tham YC¹³, Tiemeier H^17,¹⁷⁸, Turner ST¹⁷⁹, Uitterlinden AG^17,¹⁸⁰; Understanding Society Scientific Group, van Heemst D²², Waldenberger M^59,⁶⁰, Wang H^31,⁷⁵, Wang L⁹, Wang L⁵, Wei WB¹⁸¹, Williams CA⁵, Wilson G Sr¹⁸², Wojczynski MK⁵, Yao J¹⁰, Young K³⁷, Yu C⁵², Yuan JM^183,¹⁸⁴, Zhou J²³, Zonderman AB¹⁸⁵, Becker DM⁸⁷, Boehnke M⁵⁶, Bowden DW¹⁵⁵, Chambers JC^42,^88,^89,^140,¹⁸⁶, Cooper RS⁷⁸, de Faire U⁶¹, Deary IJ^48,¹⁸⁷, Elliott P⁸⁸, Esko T^38,¹⁸⁸, Farrall M^45,⁴⁶, Franks PW^79,^189,^190,¹⁹¹, Freedman BI¹⁹², Froguel P^39,¹⁹³, Gasparini P^73,¹⁰⁸, Gieger C^60,¹⁹⁴, Horta BL⁵⁰, Juang JJ^81,⁸², Kamatani Y⁶⁶, Kammerer CM¹⁹⁵, Kato N⁷⁷, Kooner JS^88,^89,^169,¹⁸⁶, Laakso M⁷⁶, Laurie CC¹⁴¹, Lee IT^196,^197,¹⁹⁸, Lehtimäki T^64,⁶⁵; Lifelines Cohort, Magnusson PKE⁴⁰, Oldehinkel AJ¹⁹⁹, Penninx BWJH¹⁴⁷, Pereira AC⁵³, Rauramaa R⁵⁸, Redline S⁷⁵, Samani NJ^71,⁷², Scott J¹⁶⁹, Shu XO⁸⁵, van der Harst P^47,²⁰⁰, Wagenknecht LE²⁰¹, Wang JS^196,¹⁹⁸, Wang YX¹²³, Wareham NJ⁶³, Watkins H^45,⁴⁶, Weir DR⁸⁴, Wickremasinghe AR⁵⁷, Wu T⁵², Zeggini E^70,²⁰², Zheng W⁸⁵, Bouchard C³², Evans MK³⁵, Gudnason V^33,³⁴, Kardia SLR³⁶, Liu Y²⁰³, Psaty BM^27,^105,^204,²⁰⁵, Ridker PM^7,⁸, van Dam RM^16,¹³⁶, Mook-Kanamori DO^165,²⁰⁶, Fornage M^3,²¹, Province MA⁵, Kelly TN²⁰⁷, Fox ER²⁰⁸, Hayward C¹⁸, van Duijn CM^17,²⁰⁹, Tai ES^16,^128,¹³⁶, Wong TY^13,^15,⁹¹, Loos RJF^12,²¹⁰, Franceschini N³⁷, Rotter JI¹⁰, Zhu X⁸³, Bierut LJ²¹¹, Gauderman WJ²¹², Rice K¹⁴¹, Munroe PB^6,⁹⁷, Morrison AC³, Rao DC², Rotimi CN²¹³, Cupples LA^214,²¹⁵.

Author information

1: Center for Research on Genomics and Global Health, National Human Genome Research Institute, US National Institutes of Health, Bethesda, MD, USA. amy.bentley@nih.gov.
2: Division of Biostatistics, Washington University School of Medicine, St. Louis, MO, USA.
3: Human Genetics Center, Department of Epidemiology, Human Genetics, and Environmental Sciences, School of Public Health, University of Texas Health Science Center at Houston, Houston, TX, USA.
4: Department of Genetic Epidemiology, University of Regensburg, Regensburg, Germany.
5: Division of Statistical Genomics, Department of Genetics, Washington University School of Medicine, St. Louis, MO, USA.
6: Clinical Pharmacology, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, UK.
7: Division of Preventive Medicine, Brigham and Women's Hospital, Boston, MA, USA.
8: Harvard Medical School, Boston, MA, USA.
9: Department of Biostatistics, Boston University School of Public Health, Boston, MA, USA.
10: Institute for Translational Genomics and Population Sciences, Department of Pediatrics, Los Angeles Biomedical Research Institute at Harbor-UCLA Medical Center, Torrance, CA, USA.
11: Women's Health Initiative Clinical Coordinating Center, Fred Hutchinson Cancer Research Center, Seattle, WA, USA.
12: Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
13: Singapore Eye Research Institute, Singapore National Eye Centre, Singapore, Singapore.
14: Centre for Quantitative Medicine, Academic Medicine Research Institute, Ophthalmology and Visual Sciences Academic Clinical Program (Eye ACP), Duke-NUS Medical School, Singapore, Singapore.
15: Department of Ophthalmology, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore.
16: Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore.
17: Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands.
18: Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK.
19: Jackson Heart Study, Department of Medicine, University of Mississippi Medical Center, Jackson, MS, USA.
20: Epidemiology and Biostatistics, University of Georgia at Athens College of Public Health, Athens, GA, USA.
21: Brown Foundation Institute of Molecular Medicine, University of Texas Health Science Center at Houston, Houston, TX, USA.
22: Internal Medicine, Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands.
23: Center for Research on Genomics and Global Health, National Human Genome Research Institute, US National Institutes of Health, Bethesda, MD, USA.
24: Centre de Bioinformatique, Biostatistique, et Biologie Intégrative (C3BI), Institut Pasteur, Paris, France.
25: Department of Epidemiology, Harvard T. H. Chan School of Public Health, Boston, MA, USA.
26: Cardiovascular Health Research Unit, Department of Biostatistics, University of Washington, Seattle, WA, USA.
27: Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA.
28: Center on Diabetes, Obesity, and Metabolism, Gerontology and Geriatric Medicine, Wake Forest University Health Sciences, Winston-Salem, NC, USA.
29: Genome Institute of Singapore, Agency for Science, Technology and Research, Singapore, Singapore.
30: Clinical and Translational Epidemiology Unit, Massachusetts General Hospital, Boston, MA, USA.
31: Department of Medicine, Harvard Medical School, Boston, MA, USA.
32: Human Genomics Laboratory, Pennington Biomedical Research Center, Baton Rouge, LA, USA.
33: Icelandic Heart Association, Kopavogur, Iceland.
34: Faculty of Medicine, University of Iceland, Reykjavik, Iceland.
35: Health Disparities Research Section, Laboratory of Epidemiology and Population Sciences, National Institute on Aging, US National Institutes of Health, Baltimore, MD, USA.
36: Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, MI, USA.
37: Department of Epidemiology, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.
38: Estonian Genome Center, Institute of Genomics, University of Tartu, Tartu, Estonia.
39: CNRS UMR 8199, European Genomic Institute for Diabetes (EGID), Institut Pasteur de Lille, University of Lille, Lille, France.
40: Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden.
41: Department of Biostatistics and Data Science, Division of Public Health Sciences, Wake Forest School of Medicine, Winston-Salem, NC, USA.
42: Department of Epidemiology and Biostatistics, School of Public Health, Imperial College, London, UK.
43: Department of Hygiene and Epidemiology, University of Ioannina Medical School, Ioannina, Greece.
44: Molecular Genetics and Genomics Program, Wake Forest School of Medicine, Winston-Salem, NC, USA.
45: Division of Cardiovascular Medicine, Radcliffe Department of Medicine, University of Oxford, Oxford, UK.
46: Wellcome Centre for Human Genetics, University of Oxford, Oxford, UK.
47: University of Groningen, University Medical Center Groningen, Department of Cardiology, Groningen, the Netherlands.
48: Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK.
49: Medical Genetics Section, Centre for Genomic and Experimental Medicine, University of Edinburgh, Edinburgh, UK.
50: Postgraduate Programme in Epidemiology, Federal University of Pelotas, Pelotas, Brazil.
51: Medical Research Council Integrative Epidemiology Unit, University of Bristol, Bristol, UK.
52: Department of Occupational and Environmental Health and State Key Laboratory of Environmental Health for Incubating, Tongji Medical College, Huazhong University of Science and Technology, Wuhan, China.
53: Laboratory of Genetics and Molecular Cardiology, Heart Institute (InCor), University of São Paulo Medical School, São Paulo, Brazil.
54: Endocrinology and Metabolism, Tri-Service General Hospital, Taipei, Taiwan.
55: School of Medicine, National Defense Medical Center, Taipei, Taiwan.
56: Department of Biostatistics and Center for Statistical Genetics, University of Michigan, Ann Arbor, MI, USA.
57: Department of Public Health, Faculty of Medicine, University of Kelaniya, Ragama, Sri Lanka.
58: Foundation for Research in Health, Exercise, and Nutrition, Kuopio Research Institute of Exercise Medicine, Kuopio, Finland.
59: Research Unit of Molecular Epidemiology, Helmholtz Zentrum München-German Research Center for Environmental Health, Neuherberg, Germany.
60: Institute of Epidemiology, Helmholtz Zentrum München-German Research Center for Environmental Health, Neuherberg, Germany.
61: Unit of Cardiovascular Epidemiology, Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden.
62: Ophthalmology, Taichung Veterans General Hospital, Taichung, Taiwan.
63: Medical Research Council Epidemiology Unit, University of Cambridge, Cambridge, UK.
64: Department of Clinical Chemistry, Fimlab Laboratories, Tampere, Finland.
65: Department of Clinical Chemistry, Finnish Cardiovascular Research Center-Tampere, Faculty of Medicine and Technology, Tampere University, Tampere, Finland.
66: Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan.
67: University of Groningen, University Medical Center Groningen, Department of Epidemiology, Groningen, the Netherlands.
68: DZHK (German Centre for Cardiovascular Health), Partner Site Greifswald, Greifswald, Germany.
69: Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, Germany.
70: Human Genetics, Wellcome Trust Sanger Institute, Hinxton, UK.
71: Department of Cardiovascular Sciences, University of Leicester, Leicester, UK.
72: NIHR Leicester Biomedical Research Centre, Glenfield Hospital, Leicester, UK.
73: Institute for Maternal and Child Health, IRCCS 'Burlo Garofolo', Trieste, Italy.
74: Taub Institute for Research on Alzheimer's Disease and the Aging Brain, Columbia University Medical Center, New York, NY, USA.
75: Division of Sleep and Circadian Disorders, Brigham and Women's Hospital, Boston, MA, USA.
76: Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland, Kuopio, Finland.
77: Department of Gene Diagnostics and Therapeutics, Research Institute, National Center for Global Health and Medicine, Tokyo, Japan.
78: Department of Public Health Sciences, Loyola University Chicago, Maywood, IL, USA.
79: Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Lund University Diabetes Centre, Skåne University Hospital, Malmö, Sweden.
80: Cardiovascular Center, National Taiwan University Hospital, Taipei, Taiwan.
81: National Taiwan University College of Medicine, Taipei, Taiwan.
82: Division of Cardiology, Department of Internal Medicine, National Taiwan University Hospital, Taipei, Taiwan.
83: Department of Population Quantitative and Health Sciences, Case Western Reserve University, Cleveland, OH, USA.
84: Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, MI, USA.
85: Division of Epidemiology, Department of Medicine, Vanderbilt University School of Medicine, Nashville, TN, USA.
86: SKLORG and Department of Epidemiology, Shanghai Cancer Institute, Renji Hospital, Shanghai Jiaotong University School of Medicine, Shanghai, China.
87: Division of General Internal Medicine, Department of Medicine, Johns Hopkins University School of Medicine, Baltimore, MD, USA.
88: MRC-PHE Centre for Environment and Health, School of Public Health, Imperial College, London, UK.
89: Department of Cardiology, Ealing Hospital, Middlesex, UK.
90: McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, MD, USA.
91: Ophthalmology and Visual Sciences Academic Clinical Program (Eye ACP), Duke-NUS Medical School, Singapore, Singapore.
92: Section of Cardiovascular Research, Baylor College of Medicine, Houston, TX, USA.
93: Houston Methodist Debakey Heart and Vascular Center, Houston, TX, USA.
94: Departments of Medicine and Epidemiology, Columbia University Medical Center, New York, NY, USA.
95: Human Genome Sequencing Center, Baylor College of Medicine, Houston, TX, USA.
96: Section of Genomic Pediatrics, Departments of Pediatrics, Medicine, and Physiology, Medical College of Wisconsin, Milwaukee, WI, USA.
97: NIHR Barts Cardiovascular Biomedical Research Centre, Queen Mary University of London, London, UK.
98: Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK.
99: Danish Aging Research Center, Institute of Public Health, University of Southern Denmark, Odense, Denmark.
100: Ninewells Hospital and Medical School, University of Dundee, Dundee, UK.
101: Cardiovascular Division, Department of Medicine, Washington University School of Medicine, St. Louis, MO, USA.
102: Department of Medicine, Faculty of Medicine, University of Kelaniya, Ragama, Sri Lanka.
103: Department of Genetics, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.
104: Department of Family Medicine and Epidemiology, Alpert Medical School of Brown University, Providence, RI, USA.
105: Department of Epidemiology, University of Washington, Seattle, WA, USA.
106: UWI Solutions for Developing Countries, University of the West Indies, Kingston, Jamaica.
107: Braun School of Public Health, Hebrew University-Hadassah Medical Center, Jerusalem, Israel.
108: Department of Medical Sciences, University of Trieste, Trieste, Italy.
109: Department of Genetics, School of Medicine, Mashhad University of Medical Sciences, Mashhad, Iran.
110: Computational Medicine Core, Center for Lung Biology, UW Medicine Sleep Center, Department of Medicine, University of Washington, Seattle, WA, USA.
111: Department of Psychiatry and Psychotherapy, University Medicine Greifswald, Greifswald, Germany.
112: Laboratory of Epidemiology and Population Sciences, National Institute on Aging, US National Institutes of Health, Bethesda, MD, USA.
113: Institute of Biomedicine, School of Medicine, University of Eastern Finland, Kuopio, Finland.
114: Department of Paediatrics, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore.
115: Khoo Teck Puat-National University Children's Medical Institute, National University Health System, Singapore, Singapore.
116: Laboratory of Genome Technology, Human Genome Center, Institute of Medical Science, University of Tokyo, Minato-ku, Japan.
117: Department of Radiology and Nuclear Medicine, Erasmus University Medical Center, Rotterdam, the Netherlands.
118: Department of Neurology, Erasmus University Medical Center, Rotterdam, the Netherlands.
119: Hebrew SeniorLife, Beth Israel Deaconess Medical Center, Harvard Medical School, Boston, MA, USA.
120: Framingham Heart Study, National Heart, Lung, and Blood Institute, US National Institutes of Health, Bethesda, MD, USA.
121: Collaborative Health Studies Coordinating Center, University of Washington, Seattle, WA, USA.
122: Department of Ophthalmology, Medical Faculty Mannheim, University of Heidelberg, Mannheim, Germany.
123: Beijing Institute of Ophthalmology, Beijing Ophthalmology and Visual Science Key Laboratory, Beijing Tongren Eye Center, Capital Medical University, Beijing, China.
124: Department of Clinical Gene Therapy, Osaka University Graduate School of Medicine, Suita, Japan.
125: Department of Geriatric and General Medicine, Osaka University Graduate School of Medicine, Suita, Japan.
126: Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark.
127: Department of Environmental Medicine and Public Health, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
128: Duke-NUS Medical School, Singapore, Singapore.
129: Department of Public Health, Department of Medicine, University of Split, Split, Croatia.
130: Fred Hutchinson Cancer Research Center, University of Washington School of Public Health, Seattle, WA, USA.
131: Sticht Center for Healthy Aging and Alzheimer's Prevention, Department of Internal Medicine, Wake Forest School of Medicine, Winston-Salem, NC, USA.
132: RIKEN Center for Integrative Medical Sciences, Yokohama, Japan.
133: Department of Clinical Physiology and Nuclear Medicine, Kuopio University Hospital, Kuopio, Finland.
134: Institute of Clinical Sciences, Department of Molecular Sciences, Imperial College, London, UK.
135: Department of Medicine, University of Alabama at Birmingham, Birmingham, AL, USA.
136: Department of Medicine, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore.
137: Epidemiology, Department of Preventive Medicine, Northwestern University Feinberg School of Medicine, Chicago, IL, USA.
138: Translational Laboratory in Genetic Medicine, Agency for Science, Technology, and Research, Singapore, Singapore.
139: Department of Biochemistry, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore.
140: Lee Kong Chian School of Medicine, Nanyang Technological University, Singapore, Singapore.
141: Department of Biostatistics, University of Washington, Seattle, WA, USA.
142: Center for Public Health Genomics, University of Virginia, Charlottesville, VA, USA.
143: Tropical Metabolism Research Unit, Caribbean Institute for Health Research, University of the West Indies, Mona, Jamaica.
144: Institute of Human Genetics, Helmholtz Zentrum München-German Research Center for Environmental Health, Neuherberg, Germany.
145: Institute of Human Genetics, Technische Universität München, Munich, Germany.
146: Technische Universität München, Munich, Germany.
147: Department of Psychiatry, Amsterdam Neuroscience and Amsterdam Public Health Research Institute, Amsterdam University Medical Center, Vrije Universiteit, Amsterdam, the Netherlands.
148: Laboratory for Genotyping Development, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan.
149: Department of Biostatistics, University of Liverpool, Liverpool, UK.
150: Institute of Medical Sciences, Aberdeen Biomedical Imaging Centre, University of Aberdeen, Aberdeen, UK.
151: Data Tecnica International, Glen Echo, MD, USA.
152: Laboratory of Neurogenetics, National Institute on Aging, Bethesda, MD, USA.
153: Division of Endocrinology, Diabetes, and Nutrition, University of Maryland School of Medicine, Baltimore, MD, USA.
154: Program for Personalized and Genomic Medicine, University of Maryland School of Medicine, Baltimore, MD, USA.
155: Biochemistry, Wake Forest School of Medicine, Winston-Salem, NC, USA.
156: Epidemiology Branch, Division of Cardiovascular Sciences, National Heart, Lung, and Blood Institute, US National Institutes of Health, Bethesda, MD, USA.
157: Epidemiology, Faculty of Medicine, Institute for Medical Information Processing, Biometry, and Epidemiology, Ludwig Maximilian University, Munich, Germany.
158: Psychiatric Hospital 'Sveti Ivan', Zagreb, Croatia.
159: Gen-info, Ltd, Zagreb, Croatia.
160: School of Public Health, Imperial College, London, UK.
161: Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland.
162: Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland.
163: Department of Biobank Research, Umeå University, Umeå, Sweden.
164: Department of Epidemiology and Medicine, University of Iowa, Iowa City, IA, USA.
165: Clinical Epidemiology, Leiden University Medical Center, Leiden, the Netherlands.
166: Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Edinburgh, UK.
167: Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.
168: Division of Epidemiology and Community Health, School of Public Health, University of Minnesota, Minneapolis, MN, USA.
169: National Heart and Lung Institute, Imperial College, London, UK.
170: Division of Research, Kaiser Permanente Northern California, Oakland, CA, USA.
171: Alzheimer Scotland Dementia Research Centre, University of Edinburgh, Edinburgh, UK.
172: Genetic Epidemiology, Faculty of Medicine, Institute for Medical Information Processing, Biometry, and Epidemiology, Ludwig Maximilian University, Munich, Germany.
173: Institute of Genetic Epidemiology, Helmholtz Zentrum München-German Research Center for Environmental Health, Neuherberg, Germany.
174: Department of Genetics, Stanford University, Stanford, CA, USA.
175: Department of Statistics and Applied Probability, National University of Singapore, Singapore, Singapore.
176: Life Sciences Institute, National University of Singapore, Singapore, Singapore.
177: NUS Graduate School for Integrative Science and Engineering, National University of Singapore, Singapore, Singapore.
178: Department of Social and Behavioral Sciences, Harvard T. H. Chan School of Public Health, Boston, MA, USA.
179: Division of Nephrology and Hypertension, Mayo Clinic, Rochester, MN, USA.
180: Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, the Netherlands.
181: Beijing Tongren Eye Center, Beijing Tongren Hospital, Capital Medical University, Beijing, China.
182: Jackson Heart Study, School of Public Health, Jackson State University, Jackson, MS, USA.
183: Department of Epidemiology, Graduate School of Public Health, University of Pittsburgh, Pittsburgh, PA, USA.
184: Division of Cancer Control and Population Sciences, UPMC Hillman Cancer, University of Pittsburgh, Pittsburgh, PA, USA.
185: Behavioral Epidemiology Section, Laboratory of Epidemiology and Population Sciences, National Institute on Aging, US National Institutes of Health, Baltimore, MD, USA.
186: Imperial College Healthcare NHS Trust, London, UK.
187: Psychology, University of Edinburgh, Edinburgh, UK.
188: Broad Institute of MIT and Harvard, Boston, MA, USA.
189: Department of Nutrition, Harvard T. H. Chan School of Public Health, Harvard University, Boston, MA, USA.
190: Department of Public Health and Clinical Medicine, Umeå University, Umeå, Sweden.
191: OCDEM, Radcliffe Department of Medicine, University of Oxford, Oxford, UK.
192: Nephrology, Internal Medicine, Wake Forest School of Medicine, Winston-Salem, NC, USA.
193: Department of Genomics of Common Disease, Imperial College, London, UK.
194: German Center for Diabetes Research (DZD), Neuherberg, Germany.
195: Department of Human Genetics, Graduate School of Public Health, University of Pittsburgh, Pittsburgh, PA, USA.
196: Endocrinology and Metabolism, Internal Medicine, Taichung Veterans General Hospital, Taichung, Taiwan.
197: School of Medicine, Chung Shan Medical University, Taichung, Taiwan.
198: School of Medicine, National Yang-Ming University, Taipei, Taiwan.
199: University of Groningen, University Medical Center Groningen, Department of Psychiatry, Groningen, the Netherlands.
200: University of Groningen, University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands.
201: Public Health Sciences, Wake Forest School of Medicine, Winston-Salem, NC, USA.
202: Institute of Translational Genomics, Helmholtz Zentrum München-German Research Center for Environmental Health, Neuherberg, Germany.
203: Public Health Sciences, Epidemiology and Prevention, Wake Forest University Health Sciences, Winston-Salem, NC, USA.
204: Department of Health Services, University of Washington, Seattle, WA, USA.
205: Kaiser Permanente Washington Health Research Institute, Seattle, WA, USA.
206: Public Health and Primary Care, Leiden University Medical Center, Leiden, the Netherlands.
207: Epidemiology, Tulane University School of Public Health and Tropical Medicine, New Orleans, LA, USA.
208: Cardiology, Medicine, University of Mississippi Medical Center, Jackson, MS, USA.
209: Nuffield Department of Population Health, University of Oxford, Oxford, UK.
210: Mindich Child Health Development Institute, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
211: Psychiatry, Washington University School of Medicine, St. Louis, MO, USA.
212: Biostatistics, Department of Preventive Medicine, University of Southern California, Los Angeles, CA, USA.
213: Center for Research on Genomics and Global Health, National Human Genome Research Institute, US National Institutes of Health, Bethesda, MD, USA. rotimic@mail.nih.gov.
214: Department of Biostatistics, Boston University School of Public Health, Boston, MA, USA. adrienne@bu.edu.
215: Framingham Heart Study, National Heart, Lung, and Blood Institute, US National Institutes of Health, Bethesda, MD, USA. adrienne@bu.edu.

Abstract

The concentrations of high- and low-density-lipoprotein cholesterol and triglycerides are influenced by smoking, but it is unknown whether genetic associations with lipids may be modified by smoking. We conducted a multi-ancestry genome-wide gene-smoking interaction study in 133,805 individuals with follow-up in an additional 253,467 individuals. Combined meta-analyses identified 13 new loci associated with lipids, some of which were detected only because association differed by smoking status. Additionally, we demonstrate the importance of including diverse populations, particularly in studies of interactions with lifestyle factors, where genomic and lifestyle differences by ancestry may contribute to novel findings.