Whole-genome sequencing and deep imputation identifies non-coding variants near Engrailed-1 with large effects on bone mineral density and fracture

Hou-Feng Zheng 1,2 *, Vincenzo Forgetta *, Yi-Hsiang Hsu *, Karol Estrada *, Alberto Rosello-Diez *, Paul J Leo *, Chitra L Dahia *, Kyung Hyun Park-Min*, Jonathan H Tobias *, Charles Kooperberg *, Aaron Kleinman, Unnur Styrkarsdottir, Ching-Ti Liu , Charlotta Uggla , Daniel S Evans , Carrie M Nielson , Klaudia Walter , Ulrika Pettersson-Kymmer , Shane McCarthy , Joel Eriksson , Tony Kwan , Mila Jhamai, Katerina Trajanoska, Yasin Memari , Josine Min , Jie Huang , Petr Danecek , Beth Wilmot , Rui Li , Wen-Chi Chou , Lauren E Mokry , Alireza Moayyeri , Melina Claussnitzer , Chia-Ho Cheng, Warren Cheung , Carolina Medina-Gómez , Bing Ge , Shu-Huang Chen , Kwangbom Choi , Ling Oei , James Fraser , Robert Kraaij , Matthew A Hibbs , Celia L Gregson , Denis Paquette , Albert Hofman , Carl Wibom , Gregory J Tranah, Mhairi Marshall , Brooke B Gardiner , Paul Auer, Li Hsu, Sue Ring, Joyce Y Tung, Gudmar Thorleifsson, Anke W Enneman , Natasja M van Schoor, Lisette C.P.G.M. de Groot, Nathalie van der Velde, Beatrice Melin , John P Kemp , Claus Christiansen, Adrian Sayers , Yanhua Zhou , Sophie Calderari , Jeroen van Rooij , Chris Carlson , Ulrike Peters , Soizik Berlivet , Josée Dostie , Andre G Uitterlinden , Stephen R. Williams, Charles Farber, Daniel Grinberg, Andrea Z LaCroix, Jeff Haessler, Daniel I Chasman, Franco Giulianini, Lynda M Rose, Paul M Ridker, John A Eisman, Tuan V Nguyen, Jacqueline R Center, Xavier Nogues, Natalia Garcia-Giralt, Lenore L Launer, Vilmunder Gudnason, Dan Mellström, Liesbeth Vandenput, Magnus K Karlsson, Östen Ljunggren, Olle Svensson, Göran Hallmans, François Rousseau, Sylvie Giroux, Johanne Bussière, Pascal P Arp, Fjorda Koromani, Richard L Prince, Joshua R Lewis, Bente L Langdahl, A Pernille Hermann, JensErik B Jensen, Stephen Kaptoge, Kay-Tee Khaw, Jonathan Reeve, Melissa M Formosa, Angela Xuereb-Anastasi, Kristina Åkesson, Fiona E McGuigan, Gaurav Garg, Jose M Olmos, Maria T Zarrabeitia, Jose A Riancho, Stuart H Ralston, Nerea Alonso, Xi Jiang, David Goltzman , Tomi Pastinen , Elin Grundberg , Dominique Gauguier , Eric S Orwoll , David Karasik , George Davey-Smith , AOGC Consortium, Albert V Smith, Kristin Siggeirsdottir, Tamara B Harris, M Carola Zillikens, Joyce BJ van Meurs, Unnur Thorsteinsdottir, Matthew T Maurano , Nicholas J Timpson , Nicole Soranzo , Richard Durbin , Scott G Wilson , Evangelia E Ntzani, Matthew A Brown , Kari Stefansson David A Hinds, Tim Spector , L Adrienne Cupples , Claes Ohlsson , Celia Greenwood , Rebecca D Jackson †, David W Rowe†, Cynthia A Loomis †, David M Evans †, Cheryl L Ackert-Bicknell †, Alexandra L Joyner †, Emma L Duncan †, Douglas P Kiel †, Fernando Rivadeneira †, J Brent Richards † for the GEFOS and UK10K Consortia.