Whole-genome sequencing reveals host factors underlying critical COVID-19
Authors: Kousathanas Athanasios , Pairo-Castineira Erola , Rawlik Konrad , Stuckey Alex , Odhams Christopher A. , Walker Susan , Russell Clark D. , Malinauskas Tomas , Wu Yang , Millar Jonathan , Shen Xia , Elliott Katherine S. , Griffiths Fiona , Oosthuyzen Wilna , Morrice Kirstie , Keating Sean , Wang Bo , Rhodes Daniel , Klaric Lucija , Zechner Marie , Parkinson Nick , Siddiq Afshan , Goddard Peter , Donovan Sally , Maslove David , Nichol Alistair , Semple Malcolm G. , Zainy Tala , Maleady-Crowe Fiona , Todd Linda , Salehi Shahla , Knight Julian , Elgar Greg , Chan Georgia , Arumugam Prabhu , Patch Christine , Rendon Augusto , Bentley David , Kingsley Clare , Kosmicki Jack A. , Horowitz Julie E. , Baras Aris , Abecasis Goncalo R. , Ferreira Manuel A. R. , Justice Anne , Mirshahi Tooraj , Oetjens Matthew , Rader Daniel J. , Ritchie Marylyn D. , Verma Anurag , Fowler Tom A. , Shankar-Hari Manu , Summers Charlotte , Hinds Charles , Horby Peter , Ling Lowell , McAuley Danny , Montgomery Hugh , Openshaw Peter J. M. , Elliott Paul , Walsh Timothy , Tenesa Albert ,   GenOMICC investigators,   GenOMICC co-investigators, Baillie J. Kenneth , Begg Colin , Clohisey Hendry Sara , Hinds Charles , Horby Peter , Knight Julian , Ling Lowell , Maslove David , McAuley Danny , Millar Johnny , Montgomery Hugh , Nichol Alistair , Openshaw Peter J. M. , Pereira Alexandre C. , Ponting Chris P. , Rowan Kathy , Semple Malcolm G. , Shankar-Hari Manu , Summers Charlotte , Walsh Timothy ,   Management, laboratory and data team, Aravindan Latha , Armstrong Ruth , Biggs Heather , Boz Ceilia , Brown Adam , Clark Richard , Coutts Audrey , Coyle Judy , Cullum Louise , Das Sukamal , Day Nicky , Donnelly Lorna , Duncan Esther , Fawkes Angie , Finernan Paul , Fourman Max Head , Furlong Anita , Furniss James , Gallagher Bernadette , Gilchrist Tammy , Golightly Ailsa , Griffiths Fiona , Hafezi Katarzyna , Hamilton Debbie , Hendry Ross , Law Andy , Law Dawn , Law Rachel , Law Sarah , Lidstone-Scott Rebecca , Macgillivray Louise , Maclean Alan , Mal Hanning , McCafferty Sarah , Mcmaster Ellie , Meikle Jen , Moore Shona C. , Morrice Kirstie , Murphy Lee , Murphy Sheena , Hellen Mybaya , Oosthuyzen Wilna , Zheng Chenqing , Chen Jiantao , Parkinson Nick , Paterson Trevor , Schon Katherine , Stenhouse Andrew , Das Mihaela , Swets Maaike , Szoor-McElhinney Helen , Taneski Filip , Turtle Lance , Wackett Tony , Ward Mairi , Weaver Jane , Wrobel Nicola , Zechner Marie ,   Guy’s and St Thomas’ Hospital team, Arbane Gill , Bociek Aneta , Campos Sara , Grau Neus , Jones Tim Owen , Lim Rosario , Marotti Martina , Ostermann Marlies , Shankar-Hari Manu , Whitton Christopher ,   Barts Health NHS Trust team, Alldis Zoe , Astin-Chamberlain Raine , Bibi Fatima , Biddle Jack , Blow Sarah , Bolton Matthew , Borra Catherine , Bowles Ruth , Burton Maudrian , Choudhury Yasmin , Collier David , Cox Amber , Easthope Amy , Ebano Patrizia , Fotiadis Stavros , Gurasashvili Jana , Halls Rosslyn , Hartridge Pippa , Kallon Delordson , Kassam Jamila , Lancoma-Malcolm Ivone , Matharu Maninderpal , May Peter , Mitchelmore Oliver , Newman Tabitha , Patel Mital , Pheby Jane , Pinzuti Irene , Prime Zoe , Prysyazhna Oleksandra , Shiel Julian , Taylor Melanie , Tierney Carey , Wood Suzanne , Zak Anne , Zongo Olivier ,   James Cook University Hospital team, Bonner Stephen , Hugill Keith , Jones Jessica , Liggett Steven , Headlam Evie ,   Royal Stoke University Hospital team, Bandla Nageswar , Gellamucho Minnie , Davies Michelle , Thompson Christopher ,   North Middlesex University Hospital NHS Trust team, Abdelrazik Marwa , Bakthavatsalam Dhanalakshmi , Elhassan Munzir , Ganesan Arunkumar , Haldeos Anne , Moreno-Cuesta Jeronimo , Purohit Dharam , Vincent Rachel , Xavier Kugan , Kumar Rohit , Frater Alasdair , Saleem Malik , Carter David , Jenkins Samuel , Lamond Zoe , Wall Alanna ,   The Royal Liverpool University Hospital team, Fernandez-Roman Jaime , Hamilton David O. , Johnson Emily , Johnston Brian , Martinez Maria Lopez , Mulla Suleman , Shaw David , Waite Alicia A. C. , Waugh Victoria , Welters Ingeborg D. , Williams Karen ,   King’s College Hospital team, Cavazza Anna , Cockrell Maeve , Corcoran Eleanor , Depante Maria , Finney Clare , Jerome Ellen , McPhail Mark , Nayak Monalisa , Noble Harriet , O’Reilly Kevin , Pappa Evita , Saha Rohit , Saha Sian , Smith John , Knighton Abigail ,   Charing Cross Hospital team, Antcliffe David , Banach Dorota , Brett Stephen , Coghlan Phoebe , Fernandez Ziortza , Gordon Anthony , Rojo Roceld , Arias Sonia Sousa , Templeton Maie ,   Nottingham University Hospital team, Meredith Megan , Morris Lucy , Ryan Lucy , Clark Amy , Sampson Julia , Peters Cecilia , Dent Martin , Langley Margaret , Ashraf Saima , Wei Shuying , Andrew Angela ,   John Radcliffe Hospital team, Bashyal Archana , Davidson Neil , Hutton Paula , McKechnie Stuart , Wilson Jean ,   Kingston Hospital team, Baptista David , Crowe Rebecca , Fernandes Rita , Herdman-Grant Rosaleen , Joseph Anna , O’Connor Denise , Allen Meryem , Loveridge Adam , McKenley India , Morino Eriko , Naranjo Andres , Simms Richard , Sollesta Kathryn , Swain Andrew , Venkatesh Harish , Khera Jacyntha , Fox Jonathan ,   Royal Infirmary of Edinburgh team, Andrew Gillian , Baillie J. Kenneth , Barclay Lucy , Callaghan Marie , Campbell Rachael , Clark Sarah , Hope Dave , Marshall Lucy , McCulloch Corrienne , Briton Kate , Singleton Jo , Birch Sophie ,   Queen Alexandra Hospital team, Brimfield Lutece , Daly Zoe , Pogson David , Rose Steve , Nown Angela ,   Morriston Hospital team, Battle Ceri , Brinkworth Elaine , Harford Rachel , Murphy Carl , Newey Luke , Rees Tabitha , Williams Marie , Arnold Sophie ,   Addenbrooke’s Hospital team, Polgarova Petra , Stroud Katerina , Summers Charlotte , Meaney Eoghan , Jones Megan , Ng Anthony , Agrawal Shruti , Pathan Nazima , White Deborah , Daubney Esther , Elston Kay ,   BHRUT (Barking Havering)—Queen’s Hospital and King George Hospital team, Grauslyte Lina , Hussain Musarat , Phull Mandeep , Pogreban Tatiana , Rosaroso Lace , Salciute Erika , Franke George , Wong Joanna , George Aparna ,   Royal Sussex County Hospital team, de Gordoa Laura Ortiz-Ruiz , Peasgood Emily , Phillips Claire ,   Queen Elizabeth Hospital team, Bates Michelle , Dasgin Jo , Gill Jaspret , Nilsson Annette , Scriven James , Collins Amy , Khaliq Waqas , Gude Estefania Treus ,   St George’s Hospital team, Delgado Carlos Castro , Dawson Deborah , Ding Lijun , Durrant Georgia , Ezeobu Obiageri , Farnell-Ward Sarah , Harrison Abiola , Kanu Rebecca , Leaver Susannah , Maccacari Elena , Manna Soumendu , Saluzzio Romina Pepermans , Queiroz Joana , Samakomva Tinashe , Sicat Christine , Texeira Joana , Da Gloria Edna Fernandes , Lisboa Ana , Rawlins John , Mathew Jisha , Kinch Ashley , Hurt William James , Shah Nirav , Clark Victoria , Thanasi Maria , Yun Nikki , Patel Kamal ,   Stepping Hill Hospital team, Bennett Sara , Goodwin Emma , Jackson Matthew , Kent Alissa , Tibke Clare , Woodyatt Wiesia , Zaki Ahmed ,   Countess of Chester Hospital team, Abraheem Azmerelda , Bamford Peter , Cawley Kathryn , Dunmore Charlie , Faulkner Maria , Girach Rumanah , Jeffrey Helen , Jones Rhianna , London Emily , Nagra Imrun , Nasir Farah , Sainsbury Hannah , Smedley Clare ,   Royal Blackburn Teaching Hospital team, Patel Tahera , Smith Matthew , Chukkambotla Srikanth , Kazi Aayesha , Hartley Janice , Dykes Joseph , Hijazi Muhammad , Keith Sarah , Khan Meherunnisa , Ryan-Smith Janet , Springle Philippa , Thomas Jacqueline , Truman Nick , Saad Samuel , Coleman Dabheoc , Fine Christopher , Matt Roseanna , Gay Bethan , Dalziel Jack , Ali Syamlan , Goodchild Drew , Harling Rhiannan , Bhatterjee Ravi , Goddard Wendy , Davison Chloe , Duberly Stephen , Hargreaves Jeanette , Bolton Rachel ,   The Tunbridge Wells Hospital and Maidstone Hospital team, Davey Miriam , Golden David , Seaman Rebecca ,   Royal Gwent Hospital team, Cherian Shiney , Cutler Sean , Heron Anne Emma , Roynon-Reed Anna , Szakmany Tamas , Williams Gemma , Richards Owen , Cheema Yusuf ,   Pinderfields General Hospital team, Brooke Hollie , Buckley Sarah , Suarez Jose Cebrian , Charlesworth Ruth , Hansson Karen , Norris John , Poole Alice , Rose Alastair , Sandhu Rajdeep , Sloan Brendan , Smithson Elizabeth , Thirumaran Muthu , Wagstaff Veronica , Metcalfe Alexandra ,   Royal Berkshire NHS Foundation Trust team, Brunton Mark , Caterson Jess , Coles Holly , Frise Matthew , Rai Sabi Gurung , Jacques Nicola , Keating Liza , Tilney Emma , Bartley Shauna , Bhuie Parminder ,   Broomfield Hospital team, Gibson Sian , Lyle Amanda , McNeela Fiona , Radhakrishnan Jayachandran , Hughes Alistair ,   Northumbria Healthcare NHS Foundation Trust team, Yates Bryan , Reynolds Jessica , Campbell Helen , Thompsom Maria , Dodds Steve , Duffy Stacey ,   Whiston Hospital team, Greer Sandra , Shuker Karen , Tridente Ascanio ,   Croydon University Hospital team, Khade Reena , Sundar Ashok , Tsinaslanidis George ,   York Hospital team, Birkinshaw Isobel , Carter Joseph , Howard Kate , Ingham Joanne , Joy Rosie , Pearson Harriet , Roche Samantha , Scott Zoe ,   Heartlands Hospital team, Bancroft Hollie , Bellamy Mary , Carmody Margaret , Daglish Jacqueline , Moore Faye , Rhodes Joanne , Sangombe Mirriam , Kadiri Salma , Scriven James ,   Ashford and St Peter’s Hospital team, Croft Maria , White Ian , Frost Victoria , Aquino Maia ,   Barnet Hospital team, Jha Rajeev , Krishnamurthy Vinodh , Lim Lai , Lim Li ,   East Surrey Hospital team, Combes Edward , Joefield Teishel , Monnery Sonja , Beech Valerie , Trotman Sallyanne ,   Ninewells Hospital team, Almaden-Boyle Christine , Austin Pauline , Cabrelli Louise , Cole Stephen , Casey Matt , Chapman Susan , Whyte Clare ,   Worthing Hospital team, Baird Yolanda , Butler Aaron , Chadbourn Indra , Folkes Linda , Fox Heather , Gardner Amy , Gomez Raquel , Hobden Gillian , Hodgson Luke , King Kirsten , Margarson Michael , Martindale Tim , Meadows Emma , Raynard Dana , Thirlwall Yvette , Helm David , Margalef Jordi ,   Southampton General Hospital team, Criste Kristine , Cusack Rebecca , Golder Kim , Golding Hannah , Jones Oliver , Leggett Samantha , Male Michelle , Marani Martyna , Prager Kirsty , Williams Toran , Roberts Belinda , Salmon Karen ,   The Alexandra Hospital team, Anderson Peter , Archer Katie , Austin Karen , Davis Caroline , Durie Alison , Kelsall Olivia , Thrush Jessica , Vigurs Charlie , Wild Laura , Wood Hannah-Louise , Tranter Helen , Harrison Alison , Cowley Nicholas , McAlindon Michael , Burtenshaw Andrew , Digby Stephen , Low Emma , Morgan Aled , Cother Naiara , Rankin Tobias , Clayton Sarah , McCurdy Alex ,   Sandwell General Hospital and City Hospital team, Ahmed Cecilia , Baines Balvinder , Clamp Sarah , Colley Julie , Haq Risna , Hayes Anne , Hulme Jonathan , Hussain Samia , Joseph Sibet , Kumar Rita , Maqsood Zahira , Purewal Manjit ,   Blackpool Victoria Hospital team, Benham Leonie , Bradshaw Zena , Brown Joanna , Caswell Melanie , Cupitt Jason , Melling Sarah , Preston Stephen , Slawson Nicola , Stoddard Emma , Warden Scott ,   Royal Glamorgan Hospital team, Deacon Bethan , Lynch Ceri , Pothecary Carla , Roche Lisa , Howe Gwenllian Sera , Singh Jayaprakash , Turner Keri , Ellis Hannah , Stroud Natalie ,   The Royal Oldham Hospital team, Hunt Jodie , Dearden Joy , Dobson Emma , Drummond Andy , Mulcahy Michelle , Munt Sheila , O’Connor Grainne , Philbin Jennifer , Rishton Chloe , Tully Redmond , Winnard Sarah ,   Glasgow Royal Infirmary team, Cathcart Susanne , Duffy Katharine , Puxty Alex , Puxty Kathryn , Turner Lynne , Ireland Jane , Semple Gary ,   St James’s University Hospital and Leeds General Infirmary team, Long Kate , Whiteley Simon , Wilby Elizabeth , Ogg Bethan ,   University Hospital North Durham team, Cowton Amanda , Kay Andrea , Kent Melanie , Potts Kathryn , Wilkinson Ami , Campbell Suzanne , Brown Ellen ,   Fairfield General Hospital team, Melville Julie , Naisbitt Jay , Joseph Rosane , Lazo Maria , Walton Olivia , Neal Alan ,   Wythenshawe Hospital team, Alexander Peter , Allen Schvearn , Bradley-Potts Joanne , Brantwood Craig , Egan Jasmine , Felton Timothy , Padden Grace , Ward Luke , Moss Stuart , Glasgow Susannah ,   Royal Alexandra Hospital team, Abel Lynn , Brett Michael , Digby Brian , Gemmell Lisa , Hornsby James , MacGoey Patrick , O’Neil Pauline , Price Richard , Rodden Natalie , Rooney Kevin , Sundaram Radha , Thomson Nicola ,   Good Hope Hospital team, Hopkins Bridget , Scriven James , Thrasyvoulou Laura , Willis Heather ,   Tameside General Hospital team, Clark Martyn , Coulding Martina , Jude Edward , McCormick Jacqueline , Mercer Oliver , Potla Darsh , Rehman Hafiz , Savill Heather , Turner Victoria ,   Royal Derby Hospital team, Downes Charlotte , Holding Kathleen , Riches Katie , Hilton Mary , Hayman Mel , Subramanian Deepak , Daniel Priya ,   Medway Maritime Hospital team, Adanini Oluronke , Bhatia Nikhil , Msiska Maines , Collins Rebecca ,   Royal Victoria Infirmary team, Clement Ian , Patel Bijal , Gulati A. , Hays Carole , Webster K. , Hudson Anne , Webster Andrea , Stephenson Elaine , McCormack Louise , Slater Victoria , Nixon Rachel , Hanson Helen , Fearby Maggie , Kelly Sinead , Bridgett Victoria , Robinson Philip ,   Poole Hospital team, Camsooksai Julie , Humphrey Charlotte , Jenkins Sarah , Reschreiter Henrik , Wadams Beverley , Death Yasmin ,   Bedford Hospital team, Bastion Victoria , Clarke Daphene , David Beena , Kent Harriet , Lorusso Rachel , Lubimbi Gamu , Murdoch Sophie , Penacerrada Melchizedek , Thomas Alastair , Valentine Jennifer , Vochin Ana , Wulandari Retno , Djeugam Brice ,   Queens Hospital Burton team, Bell Gillian , English Katy , Katary Amro , Wilcox Louise ,   North Manchester General Hospital team, Bruce Michelle , Connolly Karen , Duncan Tracy , Michael Helen T. , Lindergard Gabriella , Hey Samuel , Fox Claire , Alfonso Jordan , Durrans Laura Jayne , Guerin Jacinta , Blackledge Bethan , Harris Jade , Hruska Martin , Eltayeb Ayaa , Lamb Thomas , Hodgkiss Tracey , Cooper Lisa , Rothwell Joanne ,   Aberdeen Royal Infirmary team, Allan Angela , Anderson Felicity , Kaye Callum , Liew Jade , Medhora Jasmine , Scott Teresa , Trumper Erin , Botello Adriana ,   Derriford Hospital team, Lankester Liana , Nikitas Nikitas , Wells Colin , Stowe Bethan , Spencer Kayleigh ,   Manchester Royal Infirmary team, Brandwood Craig , Smith Lara , Clark Richard , Birchall Katie , Kolakaluri Laurel , Baines Deborah , Sukumaran Anila ,   Salford Royal Hospital team, Apetri Elena , Basikolo Cathrine , Blackledge Bethan , Catlow Laura , Charles Bethan , Dark Paul , Doonan Reece , Harris Jade , Harvey Alice , Horner Daniel , Knowles Karen , Lee Stephanie , Lomas Diane , Lyons Chloe , Marsden Tracy , McLaughlan Danielle , McMorrow Liam , Pendlebury Jessica , Perez Jane , Poulaka Maria , Proudfoot Nicola , Slaughter Melanie , Slevin Kathryn , Taylor Melanie , Thomas Vicky , Walker Danielle , Michael Angiy , Collis Matthew ,   William Harvey Hospital team, Cosier Tracey , Millen Gemma , Richardson Neil , Schumacher Natasha , Weston Heather , Rand James ,   Queen Elizabeth University Hospital team, Baxter Nicola , Henderson Steven , Kennedy-Hay Sophie , McParland Christopher , Rooney Laura , Sim Malcolm , McCreath Gordan ,   Bradford Royal Infirmary team, Akeroyd Louise , Bano Shereen , Bromley Matt , Gurr Lucy , Lawton Tom , Morgan James , Sellick Kirsten , Warren Deborah , Wilkinson Brian , McGowan Janet , Ledgard Camilla , Stacey Amelia , Pye Kate , Bellwood Ruth , Bentley Michael ,   Bristol Royal Infirmary team, Bewley Jeremy , Garland Zoe , Grimmer Lisa , Gumbrill Bethany , Johnson Rebekah , Sweet Katie , Webster Denise , Efford Georgia ,   Norfolk and Norwich University Hospital (NNUH) team, Convery Karen , Fottrell-Gould Deirdre , Hudig Lisa , Keshet-Price Jocelyn , Randell Georgina , Stammers Katie ,   Queen Elizabeth Hospital Gateshead team, Bokhari Maria , Linnett Vanessa , Lucas Rachael , McCormick Wendy , Ritzema Jenny , Sanderson Amanda , Wild Helen ,   Sunderland Royal Hospital team, Rostron Anthony , Roy Alistair , Woods Lindsey , Cornell Sarah , Wakinshaw Fiona , Rogerson Kimberley , Jarmain Jordan ,   Aintree University Hospital team, Parker Robert , Reddy Amie , Turner-Bone Ian , Wilding Laura , Harding Peter ,   Hull Royal Infirmary team, Abernathy Caroline , Foster Louise , Gratrix Andrew , Martinson Vicky , Parkinson Priyai , Stones Elizabeth , Carbral-Ortega Llucia ,   University College Hospital team, Bercades Georgia , Brealey David , Hass Ingrid , MacCallum Niall , Martir Gladys , Raith Eamon , Reyes Anna , Smyth Deborah ,   Royal Devon and Exeter Hospital team, Zitter Letizia , Benyon Sarah , Marriott Suzie , Park Linda , Keenan Samantha , Gordon Elizabeth , Quinn Helen , Baines Kizzy ,   The Royal Papworth Hospital team, Cagova Lenka , Fofano Adama , Garner Lucie , Holcombe Helen , Mepham Sue , Mitchell Alice Michael , Mwaura Lucy , Praman Krithivasan , Vuylsteke Alain , Zamikula Julie ,   Ipswich Hospital team, Purewal Bally , Rivers Vanessa , Bell Stephanie ,   Southmead Hospital team, Blakemore Hayley , Borislavova Borislava , Faulkner Beverley , Gendall Emma , Goff Elizabeth , Hayes Kati , Thomas Matt , Worner Ruth , Smith Kerry , Stephens Deanna ,   Milton Keynes University Hospital team, Mew Louise , Mwaura Esther , Stewart Richard , Williams Felicity , Wren Lynn , Sutherland Sara-Beth ,   Royal Hampshire County Hospital team, Bevan Emily , Martin Jane , Trodd Dawn , Watson Geoff , Wrey Brown Caroline ,   Great Ormond St Hospital and UCL Great Ormond St Institute of Child Health NIHR Biomedical Research Centre team, Akinkugbe Olugbenga , Bamford Alasdair , Beech Emily , Belfield Holly , Bell Michael , Davies Charlene , Jones Gareth A. L. , McHugh Tara , Meghari Hamza , O’Neill Lauran , Peters Mark J. , Ray Samiran , Tomas Ana Luisa ,   Stoke Mandeville Hospital team, Burn Iona , Hambrook Geraldine , Manso Katarina , Penn Ruth , Shanmugasundaram Pradeep , Tebbutt Julie , Thornton Danielle ,   University Hospital of Wales team, Cole Jade , Davies Michelle , Davies Rhys , Duffin Donna , Hill Helen , Player Ben , Thomas Emma , Williams Angharad ,   Basingstoke and North Hampshire Hospital team, Griffin Denise , Muchenje Nycola , Mupudzi Mcdonald , Partridge Richard , Conyngham Jo-Anna , Thomas Rachel , Wright Mary , Corral Maria Alvarez ,   Arrowe Park Hospital team, Jacob Reni , Jones Cathy , Denmade Craig ,   Chesterfield Royal Hospital Foundation Trust team, Beavis Sarah , Dale Katie , Gascoyne Rachel , Hawes Joanne , Pritchard Kelly , Stevenson Lesley , Whileman Amanda ,   Musgrove Park Hospital team, Doble Patricia , Hutter Joanne , Pawley Corinne , Shovelton Charmaine , Vaida Marius ,   Peterborough City Hospital team, Butcher Deborah , O’Sullivan Susie , Butterworth-Cowin Nicola ,   Royal Hallamshire Hospital and Northern General Hospital team, Ahmad Norfaizan , Barker Joann , Bauchmuller Kris , Bird Sarah , Cawthron Kay , Harrington Kate , Jackson Yvonne , Kibutu Faith , Lenagh Becky , Masuko Shamiso , Mills Gary H. , Raithatha Ajay , Wiles Matthew , Willson Jayne , Newell Helen , Lye Alison , Nwafor Lorenza , Jarman Claire , Rowland-Jones Sarah , Foote David , Cole Joby , Thompson Roger , Watson James , Hesseldon Lisa , Macharia Irene , Chetam Luke , Smith Jacqui , Ford Amber , Anderson Samantha , Birchall Kathryn , Housley Kay , Walker Sara , Milner Leanne , Hanratty Helena , Trower Helen , Phillips Patrick , Oxspring Simon , Donne Ben ,   Dumfries and Galloway Royal Infirmary team, Jardine Catherine , Williams Dewi , Hay Alasdair ,   Royal Bolton Hospital team, Flanagan Rebecca , Hughes Gareth , Latham Scott , McKenna Emma , Anderson Jennifer , Hull Robert , Rhead Kat ,   Lister Hospital team, Cruz Carina , Pattison Natalie ,   Craigavon Area Hospital team, Charnock Rob , McFarland Denise , Cosgrove Denise ,   Southport and Formby District General Hospital team, Ahmed Ashar , Morris Anna , Jakkula Srinivas , Nune Arvind ,   Calderdale Royal Hospital team, Ali Asifa , Brady Megan , Dale Sam , Dance Annalisa , Gledhill Lisa , Greig Jill , Hanson Kathryn , Holdroyd Kelly , Home Marie , Kelly Diane , Kitson Ross , Matapure Lear , Melia Deborah , Mellor Samantha , Nortcliffe Tonicha , Pinnell Jez , Robinson Matthew , Shaw Lisa , Shaw Ryan , Thomis Lesley , Wilson Alison , Wood Tracy , Bayo Lee-Ann , Merwaha Ekta , Ishaq Tahira , Hanley Sarah ,   Prince Charles Hospital team, Deacon Bethan , Hibbert Meg , Pothecary Carla , Tetla Dariusz , Woodford Christopher , Durga Latha , Kennard-Holden Gareth ,   Royal Bournemouth Hospital team, Branney Debbie , Frankham Jordan , Pitts Sally , White Nigel ,   Royal Preston Hospital team, Laha Shondipon , Verlander Mark , Williams Alexandra ,   Whittington Hospital team, Altabaibeh Abdelhakim , Alvaro Ana , Gilbert Kayleigh , Ma Louise , Mostoles Loreta , Parmar Chetan , Simpson Kathryn , Jetha Champa , Booker Lauren , Pratley Anezka ,   Princess Royal Hospital team, Adams Colene , Agasou Anita , Arden Tracie , Bowes Amy , Boyle Pauline , Beekes Mandy , Button Heather , Capps Nigel , Carnahan Mandy , Carter Anne , Childs Danielle , Donaldson Denise , Hard Kelly , Hurford Fran , Hussain Yasmin , Javaid Ayesha , Jones James , Jose Sanal , Leigh Michael , Martin Terry , Millward Helen , Motherwell Nichola , Rikunenko Rachel , Stickley Jo , Summers Julie , Ting Louise , Tivenan Helen , Tonks Louise , Wilcox Rebecca , Skinner Denise , Gaylard Jane , Mullan Dee , Newman Julie ,   Macclesfield District General Hospital team, Holland Maureen , Keenan Natalie , Lyons Marc , Wassall Helen , Marsh Chris , Mahenthran Mervin , Carter Emma , Kong Thomas ,   Royal Surrey County Hospital team, Blackman Helen , Creagh-Brown Ben , Donlon Sinead , Michalak-Glinska Natalia , Mtuwa Sheila , Pristopan Veronika , Salberg Armorel , Smith Eleanor , Stone Sarah , Piercy Charles , Verula Jerik , Burda Dorota , Montaser Rugia , Harden Lesley , Mayangao Irving , Marriott Cheryl , Bradley Paul , Harris Celia ,   Hereford County Hospital team, Anderson Susan , Andrews Eleanor , Birch Janine , Collins Emma , Hammerton Kate , O’Leary Ryan ,   University Hospital of North Tees team, Clark Michele , Purvis Sarah ,   Lincoln County Hospital team, Barber Russell , Hewitt Claire , Hilldrith Annette , Jackson-Lawrence Karen , Shepardson Sarah , Wills Maryanne , Butler Susan , Tavares Silvia , Cunningham Amy , Hindale Julia , Arif Sarwat ,   Royal Cornwall Hospital team, Bean Sarah , Burt Karen , Spivey Michael ,   Royal United Hospital team, Demetriou Carrie , Eckbad Charlotte , Hierons Sarah , Howie Lucy , Mitchard Sarah , Ramos Lidia , Serrano-Ruiz Alfredo , White Katie , Kelly Fiona ,   Royal Brompton Hospital team, Cristiano Daniele , Dormand Natalie , Farzad Zohreh , Gummadi Mahitha , Liyanage Kamal , Patel Brijesh , Salmi Sara , Sloane Geraldine , Thwaites Vicky , Varghese Mathew , Zborowski Anelise C. ,   University Hospital Crosshouse team, Allan John , Geary Tim , Houston Gordon , Meikle Alistair , O’Brien Peter ,   Basildon Hospital team, Forsey Miranda , Kaliappan Agilan , Nicholson Anne , Riches Joanne , Vertue Mark ,   Glan Clwyd Hospital team, Allan Elizabeth , Darlington Kate , Davies Ffyon , Easton Jack , Kumar Sumit , Lean Richard , Menzies Daniel , Pugh Richard , Qiu Xinyi , Davies Llinos , Williams Hannah , Scanlon Jeremy , Davies Gwyneth , Mackay Callum , Lewis Joanne , Rees Stephanie ,   West Middlesex Hospital team, Oblak Metod , Popescu Monica , Thankachen Mini ,   Royal Lancaster Infirmary team, Higham Andrew , Simpson Kerry , Craig Jayne ,   Western General Hospital team, Baruah Rosie , Morris Sheila , Ferguson Susie , Shepherd Amy ,   Chelsea and Westminster NHS Foundation Trust team, Moore Luke Stephen Prockter , Vizcaychipi Marcela Paola , de Almeida Martins Laura Gomes , Carungcong Jaime ,   The Queen Elizabeth Hospital team, Ali Inthakab Ali Mohamed , Beaumont Karen , Blunt Mark , Coton Zoe , Curgenven Hollie , Elsaadany Mohamed , Fernandes Kay , Mohamed Ally Sameena , Rangarajan Harini , Sarathy Varun , Selvanayagam Sivarupan , Vedage Dave , White Matthew ,   King’s Mill Hospital team, Gill Mandy , Paul Paul , Ratnam Valli , Shelton Sarah , Wynter Inez ,   Watford General Hospital team, Carmody Siobhain , Page Valerie Joan ,   University Hospital Wishaw team, Beith Claire Marie , Black Karen , Clements Suzanne , Morrison Alan , Strachan Dominic , Taylor Margaret , Clarkson Michelle , D’Sylva Stuart , Norman Kathryn ,   Forth Valley Royal Hospital team, Auld Fiona , Donnachie Joanne , Edmond Ian , Prentice Lynn , Runciman Nikole , Salutous Dario , Symon Lesley , Todd Anne , Turner Patricia , Short Abigail , Sweeney Laura , Murdoch Euan , Senaratne Dhaneesha ,   George Eliot Hospital NHS Trust team, Hill Michaela , Kannan Thogulava , Wild Laura ,   Barnsley Hospital team, Crawley Rikki , Crew Abigail , Cunningham Mishell , Daniels Allison , Harrison Laura , Hope Susan , Inweregbu Ken , Jones Sian , Lancaster Nicola , Matthews Jamie , Nicholson Alice , Wray Gemma ,   The Great Western Hospital team, Langton Helen , Prout Rachel , Watters Malcolm , Novis Catherine ,   Harefield Hospital team, Barron Anthony , Collins Ciara , Kaul Sundeep , Passmore Heather , Prendergast Claire , Reed Anna , Rogers Paula , Shokkar Rajvinder , Woodruff Meriel , Middleton Hayley , Polgar Oliver , Nolan Claire , Thwaites Vicky , Mahay Kanta ,   Rotherham General Hospital team, Collier Dawn , Hormis Anil , Maynard Victoria , Graham Cheryl , Walker Rachel ,   Ysbyty Gwynedd team, Knights Ellen , Price Alicia , Thomas Alice , Thorpe Chris ,   Diana Princess of Wales Hospital team, Behan Teresa , Burnett Caroline , Hatton Jonathan , Heeney Elaine , Mitra Atideb , Newton Maria , Pollard Rachel , Stead Rachael ,   Russell’s Hall Hospital team, Amin Vishal , Anastasescu Elena , Anumakonda Vikram , Karthik Komala , Kausar Rizwana , Reid Karen , Smith Jacqueline , Imeson-Wood Janet ,   St Mary’s Hospital team, Brown Alison , Crickmore Vikki , Debreceni Gabor , Wilkins Joy , Nicol Liz ,   University Hospital Lewisham team, Khaliq Waqas , Reece-Anthony Rosie , Birt Mark ,   Colchester General Hospital team, Ghosh Alison , Williams Emma ,   Queen Elizabeth the Queen Mother Hospital team, Allen Louise , Beranova Eva , Crisp Nikki , Deery Joanne , Hazelton Tracy , Knight Alicia , Price Carly , Tilbey Sorrell , Turki Salah , Turney Sharon ,   Royal Albert Edward Infirmary team, Cooper Joshua , Finch Cheryl , Liderth Sarah , Quinn Alison , Waddington Natalia ,   Victoria Hospital team, Coventry Tina , Fowler Susan , MacMahon Michael , McGregor Amanda ,   Eastbourne District General Hospital team, Cowley Anne , Highgate Judith ,   Cumberland Infirmary team, Brown Alison , Gregory Jane , O’Connell Susan , Smith Tim , Barberis Luigi ,   New Cross Hospital team, Gopal Shameer , Harris Nichola , Lake Victoria , Metherell Stella , Radford Elizabeth ,   The Princess Alexandra Hospital team, Daniel Amelia , Finn Joanne , Saha Rajnish , White Nikki , Easthope Amy ,   Salisbury District Hospital team, Donnison Phil , Trim Fiona , Eapen Beena ,   Dorset County Hospital team, Birch Jenny , Bough Laura , Goodsell Josie , Tutton Rebecca , Williams Patricia , Williams Sarah , Winter-Goodwin Barbara ,   University College Dublin team, Nichol Ailstair , Brickell Kathy , Smyth Michelle , Murphy Lorna ,   Glangwili General Hospital team, Coetzee Samantha , Gales Alistair , Otahal Igor , Raj Meena , Sell Craig ,   Gloucestershire Royal Hospital team, Hilltout Paula , Evitts Jayne , Tyler Amanda , Waldron Joanne ,   Yeovil Hospital team, Beesley Kate , Board Sarah , Kubisz-Pudelko Agnieszka , Lewis Alison , Perry Jess , Pippard Lucy , Wood Di , Buckley Clare ,   Leicester Royal Infirmary team, Barry Peter , Flint Neil , Rekha Patel , Hales Dawn ,   Royal Manchester Children’s Hospital team, Bunni Lara , Jennings Claire , Latif Monica , Marshall Rebecca , Subramanian Gayathri ,   Royal Victoria Hospital team, McGuigan Peter J. , Wasson Christopher , Finn Stephanie , Green Jackie , Collins Erin , King Bernadette ,   Wrexham Maelor Hospital team, Campbell Andy , Smuts Sara , Duffield Joseph , Smith Oliver , Mallon Lewis , Watkins Claire ,   Walsall Manor Hospital team, Botfield Liam , Butler Joanna , Dexter Catherine , Fletcher Jo , Garg Atul , Kuravi Aditya , Ranga Poonam , Virgilio Emma ,   Darent Valley Hospital team, Belagodu Zakaula , Fuller Bridget , Gherman Anca , Olufuwa Olumide , Paramsothy Remi , Stuart Carmel , Oakley Naomi , Kamundi Charlotte , Tyl David , Collins Katy , Silva Pedro , Taylor June , King Laura , Coates Charlotte , Crowley Maria , Wakefield Phillipa , Beadle Jane , Johnson Laura , Sargeant Janet , Anderson Madeleine ,   Warrington General Hospital team, Brady Ailbhe , Chan Rebekah , Little Jeff , McIvor Shane , Prady Helena , Whittle Helen , Mathew Bijoy ,   Warwick Hospital team, Attwood Ben , Parsons Penny ,   University Hospitals Coventry and Warwickshire NHS Trust team, Ward Geraldine , Bremmer Pamela ,   University Hospital Monklands team, Joe West , Tracy Baird , Jim Ruddy ,   Princess of Wales Hospital team, Davies Ellie , Roche Lisa , Sathe Sonia ,   Northwick Park Hospital team, Dennis Catherine , McGregor Alastair , Parris Victoria , Srikaran Sinduya , Sukha Anisha ,   Raigmore Hospital team, Campbell Rachael , Clarke Noreen , Whiteside Jonathan , Mascarenhas Mairi , Donaldson Avril , Matheson Joanna , Barrett Fiona , O’Hara Marianne , Okeefe Laura , Bradley Clare ,   Royal Free Hospital team, Eastgate-Jackson Christine , Filipe Helder , Martin Daniel , Maharajh Amitaa , Garcia Sara Mingo , Pakou Glykeria , De Neef Mark ,   Scunthorpe General Hospital team, Dent Kathy , Horsley Elizabeth , Akhtar Muhammad Nauman , Pearson Sandra , Potoczna Dorota , Spencer Sue ,   West Cumberland Hospital team, Clapham Melanie , Harper Rosemary , Poultney Una , Rice Polly , Smith Tim , Mutch Rachel , Barberis Luigi ,   Airedale General Hospital team, Armstrong Lisa , Bates Hayley , Dooks Emma , Farquhar Fiona , Hairsine Brigid , McParland Chantal , Packham Sophie ,   Birmingham Children’s Hospital team, Bi Rehana , Scholefield Barney , Ashton Lydia ,   Liverpool Heart and Chest Hospital team, George Linsha , Twiss Sophie , Wright David ,   Pilgrim Hospital team, Chablani Manish , Kirkby Amy , Netherton Kimberley ,   Prince Philip Hospital team, Davies Kim , O’Brien Linda , Omar Zohra , Otahal Igor , Perkins Emma , Lewis Tracy , Sutherland Isobel ,   Furness General Hospital team, Burns Karen , Higham Andrew ,   Scarborough General Hospital team, Chandler Ben , Elliott Kerry , Mallinson Janine , Turnbull Alison ,   Southend University Hospital team, Gondo Prisca , Hadebe Bernard , Kayani Abdul , Masunda Bridgett ,   Alder Hey Children’s Hospital team, Anderson Taya , Hawcutt Dan , O’Malley Laura , Rad Laura , Rogers Naomi , Saunderson Paula , Allison Kathryn Sian , Afolabi Deborah , Whitbread Jennifer , Jones Dawn , Dore Rachael ,   Torbay Hospital team, Halkes Matthew , Mercer Pauline , Thornton Lorraine ,   Borders General Hospital team, Dawson Joy , Garrioch Sweyn , Tolson Melanie , Aldridge Jonathan ,   Kent and Canterbury Hospital team, Kapoor Ritoo , Loader David , Castle Karen ,   West Suffolk Hospital team, Humphreys Sally , Tampsett Ruth ,   James Paget University Hospital NHS Trust team, Mackintosh Katherine , Ayers Amanda , Harrison Wendy , North Julie ,   The Christie NHS Foundation Trust team, Allibone Suzanne , Genetu Roman , Kasipandian Vidya , Patel Amit , Mac Ainhi , Murphy Anthony , Mahjoob Parisa , Nazari Roonak , Worsley Lucy , Fagan Andrew ,   The Royal Marsden Hospital team, Bemand Thomas , Black Ethel , Dela Rosa Arnold , Howle Ryan , Jhanji Shaman , Baikady Ravishankar Rao , Tatham Kate Colette , Thomas Benjamin ,   University Hospital Hairmyres team, Bell Dina , Boyle Rosalind , Douglas Katie , Glass Lynn , Lee Emma , Lennon Liz , Rattray Austin ,   Withybush General Hospital team, Taylor Abigail , Hughes Rachel Anne , Thomas Helen , Rees Alun , Duskova Michaela , Phipps Janet , Brooks Suzanne , Edwards Michelle ,   Ealing Hospital team, Parris Victoria , Quaid Sheena , Watson Ekaterina ,   North Devon District Hospital team, Brayne Adam , Fisher Emma , Hunt Jane , Jackson Peter , Kaye Duncan , Love Nicholas , Parkin Juliet , Tuckey Victoria , van Koutrik Lynne , Carter Sasha , Andrew Benedict , Findlay Louise , Adams Katie ,   St John’s Hospital Livingston team, Service Jen , Williams Alison , Cheyne Claire , Saunderson Anne , Moultrie Sam , Odam Miranda ,   Northampton General Hospital NHS Trust team, Hall Kathryn , Mapfunde Isheunesu , Willis Charlotte , Lyon Alex ,   Harrogate and District NHS Foundation Trust team, Sri-Chandana Chunda , Scherewode Joslan , Stephenson Lorraine , Marsh Sarah ,   National Hospital for Neurology and Neurosurgery team, Brealey David , Hardy John , Houlden Henry , Moncur Eleanor , Raith Eamon , Tariq Ambreen , Tucci Arianna ,   Bronglais General Hospital team, Hobrok Maria , Loosley Ronda , McGuinness Heather , Tench Helen , Wolf-Roberts Rebecca ,   Golden Jubilee National Hospital team, Irvine Val , Shelley Benjamin ,   Homerton University Hospital Foundation NHS Trust team, Easthope Amy , Gorman Claire , Gupta Abhinav , Timlick Elizabeth , Brady Rebecca ,   Royal Hospital for Children team, Milligan Barry ,   Sheffield Children’s Hospital team, Bellini Arianna , Bryant Jade , Mayer Anton , Pickard Amy , Roe Nicholas , Sowter Jason , Howlett Alex ,   The Royal Alexandra Children’s Hospital team, Fidler Katy , Tagliavini Emma , Donnelly Kevin ,   23andMe investigators, Shelton Janie F. , Shastri Anjali J. , Ye Chelsea , Weldon Catherine H. , Filshtein-Sonmez Teresa , Coker Daniella , Symons Antony , Esparza-Gordillo Jorge , Aslibekyan Stella , Auton Adam ,   COVID-19 Human Genetics Initiative, Pathak Gita A. , Karjalainen Juha , Stevens Christine , Andrews Shea J. , Kanai Masahiro , Cordioli Mattia , Polimanti Renato , Pirinen Matti , Harerimana Nadia , Veerapen Kumar , Wolford Brooke , Nguyen Huy , Solomonson Matthew , Liao Rachel G. , Chwialkowska Karolina , Trankiem Amy , Balaconis Mary K. , Hayward Caroline , Richmond Anne , Campbell Archie , Morris Marcela , Fawns-Ritchie Chloe , Glessner Joseph T. , Shaw Douglas M. , Chang Xiao , Polikowski Hannah , Petty Lauren E. , Chen Hung-Hsin , Wanying Zhu , Hakonarson Hakon , Porteous David J. , Below Jennifer , North Kari , McCormick Joseph B. , Timmers Paul R. H. J. , Wilson James F. , Tenesa Albert , D’Mellow Kenton , Kerr Shona M. , Niemi Mari E. K. , Nkambul Lindokuhle , von Hohenstaufen Kathrin Aprile , Sobh Ali , Eltoukhy Madonna M. , Yassen Amr M. , Hegazy Mohamed A. F. , Okasha Kamal , Eid Mohammed A. , Moahmed Hanteera S. , Shahin Doaa , El-Sherbiny Yasser M. , Elhadidy Tamer A. , Abd Elghafar Mohamed S. , El-Jawhari Jehan J. , Mohamed Attia A. S. , Elnagdy Marwa H. , Samir Amr , Abdel-Aziz Mahmoud , Khafaga Walid T. , El-Lawaty Walaa M. , Torky Mohamed S. , El-shanshory Mohamed R. , Batini Chiara , Lee Paul H. , Shrine Nick , Williams Alexander T. , Tobin Martin D. , Guyatt Anna L. , John Catherine , Packer Richard J. , Ali Altaf , Free Robert C. , Wang Xueyang , Wain Louise V. , Hollox Edward J. , Venn Laura D. , Bee Catherine E. , Adams Emma L. , Niavarani Ahmadreza , Sharififard Bahareh , Aliannejad Rasoul , Amirsavadkouhi Ali , Naderpour Zeinab , Tadi Hengameh Ansari , Aleagha Afshar Etemadi , Ahmadi Saeideh , Moghaddam Seyed Behrooz Mohseni , Adamsara Alireza , Saeedi Morteza , Abdollahi Hamed , Hosseini Abdolmajid , Chariyavilaskul Pajaree , Chamnanphon Monpat , Suttichet Thitima B. , Shotelersuk Vorasuk , Pongpanich Monnat , Phokaew Chureerat , Chetruengchai Wanna , Jantarabenjakul Watsamon , Putchareon Opass , Torvorapanit Pattama , Puthanakit Thanyawee , Suchartlikitwong Pintip , Hirankarn Nattiya , Nilaratanakul Voraphoj , Sodsai Pimpayao , Brumpton Ben M. , Hveem Kristian , Willer Cristen , Zhou Wei , Rogne Tormod , Solligard Erik , Åsvold Bjørn Olav , Abedalthagafi Malak , Alaamery Manal , Alqahtani Saleh , Barakeh Duna , Al Harthi Fawz , Alsolm Ebtehal , Safieh Leen Abu , Alowayn Albandary M. , Alqubaishi Fatimah , Al Mutairi Amal , Mangul Serghei , Alshareef Abdulraheem , Sawaji Mona , Almutairi Mansour , Aljawini Nora , Albesher Nour , Arabi Yaseen M. , Mahmoud Ebrahim S. , Khattab Amin K. , Halawani Roaa T. , Alahmadey Ziab Z. , Albakri Jehad K. , Felemban Walaa A. , Suliman Bandar A. , Hasanato Rana , Al-Awdah Laila , Alghamdi Jahad , AlZahrani Deema , AlJohani Sameera , Al-Afghani Hani , Alrashed May , AlDhawi Nouf , AlBardis Hadeel , Alkwai Sarah , Alswailm Moneera , Almalki Faisal , Albeladi Maha , Almohammed Iman , Barhoush Eman , Albader Anoud , Massadeh Salam , AlMalik Abdulaziz , Alotaibi Sara , Alghamdi Bader , Jung Junghyun , Fawzy Mohammad S. , Lee Yunsung , Magnus Per , Trogstad Lill-Iren S. , Helgeland Øyvind , Harris Jennifer R. , Mangino Massimo , Spector Tim D. , Duncan Emma , Smieszek Sandra P. , Przychodzen Bartlomiej P. , Polymeropoulos Christos , Polymeropoulos Vasilios , Polymeropoulos Mihael H. , Fernandez-Cadenas Israel , Perez-Tur Jordi , Llucià-Carol Laia , Cullell Natalia , Muiño Elena , Cárcel-Márquez Jara , DeDiego Marta L. , Iglesias Lara Lloret , Planas Anna M. , Soriano Alex , Rico Veronica , Agüero Daiana , Bedini Josep L. , Lozano Francisco , Domingo Carlos , Robles Veronica , Ruiz-Jaén Francisca , Márquez Leonardo , Gomez Juan , Coto Eliecer , Albaiceta Guillermo M. , García-Clemente Marta , Dalmau David , Arranz Maria J. , Dietl Beatriz , Serra-Llovich Alex , Soler Pere , Colobrán Roger , Martín-Nalda Andrea , Martínez Alba Parra , Bernardo David , Rojo Silvia , Fiz-López Aida , Arribas Elisa , de la Cal-Sabater Paloma , Segura Tomás , González-Villa Esther , Serrano-Heras Gemma , Martí-Fàbregas Joan , Jiménez-Xarrié Elena , de Felipe Mimbrera Alicia , Masjuan Jaime , García-Madrona Sebastian , Domínguez-Mayoral Anna , Villalonga Joan Montaner , Menéndez-Valladares Paloma , Chasman Daniel I. , Buring Julie E. , Ridker Paul M. , Franco Giulianini , Sesso Howard D. , Manson JoAnn E. , Glessner Joseph R. , Hakonarson Hakon , Medina-Gomez Carolina , Uitterlinden Andre G. , Ikram M. Arfan , Kristiansson Kati , Koskelainen Sami , Perola Markus , Donner Kati , Kivinen Katja , Palotie Aarno , Ripatti Samuli , Ruotsalainen Sanni , Kaunisto Mari , Nakanishi Tomoko , Butler-Laporte Guillaume , Forgetta Vincenzo , Morrison David R. , Ghosh Biswarup , Laurent Laetitia , Belisle Alexandre , Henry Danielle , Abdullah Tala , Adeleye Olumide , Mamlouk Noor , Kimchi Nofar , Afrasiabi Zaman , Rezk Nardin , Vulesevic Branka , Bouab Meriem , Guzman Charlotte , Petitjean Louis , Tselios Chris , Xue Xiaoqing , Schurr Erwin , Afilalo Jonathan , Afilalo Marc , Oliveira Maureen , Brenner Bluma , Lepage Pierre , Ragoussis Jiannis , Auld Daniel , Brassard Nathalie , Durand Madeleine , Chassé Michaël , Kaufmann Daniel E. , Lathrop G. Mark , Mooser Vincent , Richards J. Brent , Li Rui , Adra Darin , Rahmouni Souad , Georges Michel , Moutschen Michel , Misset Benoit , Darcis Gilles , Guiot Julien , Guntz Julien , Azarzar Samira , Gofflot Stéphanie , Beguin Yves , Claassen Sabine , Malaise Olivier , Huynen Pascale , Meuris Christelle , Thys Marie , Jacques Jessica , Léonard Philippe , Frippiat Frederic , Giot Jean-Baptiste , Sauvage Anne-Sophie , von Frenckell Christian , Belhaj Yasmine , Lambermont Bernard , Pigazzini Sara , Nkambule Lindokuhle , Daya Michelle , Shortt Jonathan , Rafaels Nicholas , Wicks Stephen J. , Crooks Kristy , Barnes Kathleen C. , Gignoux Christopher R. , Chavan Sameer , Laisk Triin , Läll Kristi , Lepamets Maarja , Mägi Reedik , Esko Tõnu , Reimann Ene , Milani Lili , Alavere Helene , Metsalu Kristjan , Puusepp Mairo , Metspalu Andres , Naaber Paul , Laane Edward , Pesukova Jaana , Peterson Pärt , Kisand Kai , Tabri Jekaterina , Allos Raili , Hensen Kati , Starkopf Joel , Ringmets Inge , Tamm Anu , Kallaste Anne , Bochud Pierre-Yves , Rivolta Carlo , Bibert Stéphanie , Quinodoz Mathieu , Kamdar Dhryata , Boillat Noémie , Nussle Semira Gonseth , Albrich Werner , Suh Noémie , Neofytos Dionysios , Erard Véronique , Voide Cathy , de Cid Rafael , Galván-Femenía Iván , Blay Natalia , Carreras Anna , Cortés Beatriz , Farré Xavier , Sumoy Lauro , Moreno Victor , Mercader Josep Maria , Guindo-Martinez Marta , Torrents David , Kogevinas Manolis , Garcia-Aymerich Judith , Castaño-Vinyals Gemma , Dobaño Carlota , Renieri Alessandra , Mari Francesca , Fallerini Chiara , Daga Sergio , Benetti Elisa , Baldassarri Margherita , Fava Francesca , Frullanti Elisa , Valentino Floriana , Doddato Gabriella , Giliberti Annarita , Tita Rossella , Amitrano Sara , Bruttini Mirella , Croci Susanna , Meloni Ilaria , Mencarelli Maria Antonietta , Rizzo Caterina Lo , Pinto Anna Maria , Beligni Giada , Tommasi Andrea , Di Sarno Laura , Palmieri Maria , Carriero Miriam Lucia , Alaverdian Diana , Busani Stefano , Bruno Raffaele , Vecchia Marco , Belli Mary Ann , Picchiotti Nicola , Sanarico Maurizio , Gori Marco , Furini Simone , Mantovani Stefania , Ludovisi Serena , Mondelli Mario Umberto , Castelli Francesco , Quiros-Roldan Eugenia , Antoni Melania Degli , Zanella Isabella , Vaghi Massimo , Rusconi Stefano , Siano Matteo , Montagnani Francesca , Emiliozzi Arianna , Fabbiani Massimiliano , Rossetti Barbara , Bargagli Elena , Bergantini Laura , D’Alessandro Miriana , Cameli Paolo , Bennett David , Anedda Federico , Marcantonio Simona , Scolletta Sabino , Franchi Federico , Mazzei Maria Antonietta , Guerrini Susanna , Conticini Edoardo , Cantarini Luca , Frediani Bruno , Tacconi Danilo , Spertilli Chiara , Feri Marco , Donati Alice , Scala Raffaele , Guidelli Luca , Spargi Genni , Corridi Marta , Nencioni Cesira , Croci Leonardo , Bandini Maria , Caldarelli Gian Piero , Piacentini Paolo , Desanctis Elena , Cappelli Silvia , Canaccini Anna , Verzuri Agnese , Anemoli Valentina , Ognibene Agostino , Pancrazzi Alessandro , Lorubbio Maria , D’Arminio Monforte Antonella , Miraglia Federica Gaia , Girardis Massimo , Venturelli Sophie , Cossarizza Andrea , Antinori Andrea , Vergori Alessandra , Gabrieli Arianna , Riva Agostino , Francisci Daniela , Schiaroli Elisabetta , Paciosi Francesco , Scotton Pier Giorgio , Andretta Francesca , Panese Sandro , Scaggiante Renzo , Gatti Francesca , Parisi Saverio Giuseppe , Baratti Stefano , Della Monica Matteo , Piscopo Carmelo , Capasso Mario , Russo Roberta , Andolfo Immacolata , Iolascon Achille , Fiorentino Giuseppe , Carella Massimo , Castori Marco , Merla Giuseppe , Squeo Gabriella Maria , Aucella Filippo , Raggi Pamela , Marciano Carmen , Perna Rita , Bassetti Matteo , Di Biagio Antonio , Sanguinetti Maurizio , Masucci Luca , Valente Serafina , Mandalà Marco , Giorli Alessia , Salerni Lorenzo , Zucchi Patrizia , Parravicini Pierpaolo , Menatti Elisabetta , Trotta Tullio , Giannattasio Ferdinando , Coiro Gabriella , Lena Fabio , Coviello Domenico A. , Mussini Cristina , Martinelli Enrico , Mancarella Sandro , Tavecchia Luisa , Crotti Lia , Gabbi Chiara , Rizzi Marco , Maggiolo Franco , Ripamonti Diego , Bachetti Tiziana , La Rovere Maria Teresa , Sarzi-Braga Simona , Bussotti Maurizio , Ceri Stefano , Pinoli Pietro , Raimondi Francesco , Biscarini Filippo , Stella Alessandra , Zguro Kristina , Capitani Katia , Suardi Claudia , Dei Simona , Parati Gianfranco , Ravaglia Sabrina , Artuso Rosangela , Bottà Giordano , Di Domenico Paolo , Rancan Ilaria , Perrella Antonio , Bianchi Francesco , Romani Davide , Bergomi Paola , Catena Emanuele , Colombo Riccardo , Tanfoni Marco , Vincenti Antonella , Ferri Claudio , Grassi Davide , Pessina Gloria , Tumbarello Mario , Di Pietro Massimo , Sabrina Ravaglia , Luchi Sauro , Barbieri Chiara , Acquilini Donatella , Andreucci Elena , Segala Francesco Vladimiro , Tiseo Giusy , Falcone Marco , Lista Mirjam , Poscente Monica , De Vivo Oreste , Petrocelli Paola , Guarnaccia Alessandra , Baroni Silvia , Smith Albert V. , Boughton Andrew P. , Li Kevin W. , LeFaive Jonathon , Annis Aubrey , Justice Anne E. , Mirshahi Tooraj , Chittoor Geetha , Josyula Navya Shilpa , Kosmicki Jack A. , Ferreira Manuel A. R. , Leader Joseph B. , Carey Dave J. , Gass Matthew C. , Horowitz Julie E. , Cantor Michael N. , Yadav Ashish , Baras Aris , Abecasis Goncalo R. , van Heel David A. , Hunt Karen A. , Mason Dan , Huang Qin Qin , Finer Sarah , Trivedi Bhavi , Griffiths Christopher J. , Martin Hilary C. , Wright John , Trembath Richard C. , Soranzo Nicole , Zhao Jing Hua , Butterworth Adam S. , Danesh John , Di Angelantonio Emanuele , Franke Lude , Boezen Marike , Deelen Patrick , Claringbould Annique , Lopera Esteban , Warmerdam Robert , Vonk Judith M. , van Blokland Irene , Lanting Pauline , Ori Anil P. S. , Zöllner Sebastian , Wang Jiongming , Beck Andrew , Peloso Gina , Ho Yuk-Lam , Sun Yan V. , Huffman Jennifer E. , O’Donnell Christopher J. , Cho Kelly , Tsao Phil , Gaziano J. Michael , Nivard Michel , de Geus Eco , Bartels Meike , Jan Hottenga Jouke , Weiss Scott T. , Karlson Elizabeth W. , Smoller Jordan W. , Green Robert C. , Feng Yen-Chen Anne , Mercader Josep , Murphy Shawn N. , Meigs James B. , Woolley Ann E. , Perez Emma F. , Rader Daniel , Verma Anurag , Ritchie Marylyn D. , Li Binglan , Verma Shefali S. , Lucas Anastasia , Bradford Yuki , Zeberg Hugo , Frithiof Robert , Hultström Michael , Lipcsey Miklos , Nkambul Lindo , Tardif Nicolas , Rooyackers Olav , Grip Jonathan , Maricic Tomislav , Karczewski Konrad J. , Atkinson Elizabeth G. , Tsuo Kristin , Baya Nikolas , Turley Patrick , Gupta Rahul , Callier Shawneequa , Walters Raymond K. , Palmer Duncan S. , Sarma Gopal , Cheng Nathan , Lu Wenhan , Bryant Sam , Churchhouse Claire , Cusick Caroline , Goldstein Jacqueline I. , King Daniel , Seed Cotton , Finucane Hilary , Martin Alicia R. , Satterstrom F. Kyle , Wilson Daniel J. , Armstrong Jacob , Rudkin Justine K. , Band Gavin , Earle Sarah G. , Lin Shang-Kuan , Arning Nicolas , Crook Derrick W. , Wyllie David H. , O’Connell Anne Marie , Spencer Chris C. A. , Koelling Nils , Caulfield Mark J. , Scott Richard H. , Fowler Tom , Moutsianas Loukas , Kousathanas Athanasios , Pasko Dorota , Walker Susan , Rendon Augusto , Stuckey Alex , Odhams Christopher A. , Rhodes Daniel , Chan Georgia , Arumugam Prabhu , Ball Catherine A. , Hong Eurie L. , Rand Kristin , Girshick Ahna , Guturu Harendra , Baltzell Asher Haug , Roberts Genevieve , Park Danny , Coignet Marie , McCurdy Shannon , Knight Spencer , Partha Raghavendran , Rhead Brooke , Zhang Miao , Berkowitz Nathan , Gaddis Michael , Noto Keith , Ruiz Luong , Pavlovic Milos , Sloofman Laura G. , Charney Alexander W. , Beckmann Noam D. , Schadt Eric E. , Jordan Daniel M. , Thompson Ryan C. , Gettler Kyle , Abul-Husn Noura S. , Ascolillo Steven , Buxbaum Joseph D. , Chaudhary Kumardeep , Cho Judy H. , Itan Yuval , Kenny Eimear E. , Belbin Gillian M. , Sealfon Stuart C. , Sebra Robert P. , Salib Irene , Collins Brett L. , Levy Tess , Britvan Bari , Keller Katherine , Tang Lara , Peruggia Michael , Hiester Liam L. , Niblo Kristi , Aksentijevich Alexandra , Labkowsky Alexander , Karp Avrohom , Zlatopolsky Menachem , Preuss Michael , Loos Ruth J. F. , Nadkarni Girish N. , Do Ron , Hoggart Clive , Choi Sam , Underwood Slayton J. , O’Reilly Paul , Huckins Laura M. , Zyndorf Marissa , Daly Mark J. , Neale Benjamin M. , Ganna Andrea , Fawkes Angie , Murphy Lee , Rowan Kathy , Ponting Chris P. , Vitart Veronique , Wilson James F. , Yang Jian , Bretherick Andrew D. , Scott Richard H. , Hendry Sara Clohisey , Moutsianas Loukas , Law Andy , Caulfield Mark J. , Baillie J. Kenneth 
Published: March 2022

Critical COVID-19 is caused by immune-mediated inflammatory lung injury. Host genetic variation influences the development of illness requiring critical care1 or hospitalization2–4 after infection with SARS-CoV-2. The GenOMICC (Genetics of Mortality in Critical Care) study enables the comparison of genomes from individuals who are critically ill with those of population controls to find underlying disease mechanisms. Here we use whole-genome sequencing in 7,491 critically ill individuals compared with 48,400 controls to discover and replicate 23 independent variants that significantly predispose to critical COVID-19. We identify 16 new independent associations, including variants within genes that are involved in interferon signalling (IL10RB and PLSCR1), leucocyte differentiation (BCL11A) and blood-type antigen secretor status (FUT2). Using transcriptome-wide association and colocalization to infer the effect of gene expression on disease severity, we find evidence that implicates multiple genes—including reduced expression of a membrane flippase (ATP11A), and increased expression of a mucin (MUC1)—in critical disease. Mendelian randomization provides evidence in support of causal roles for myeloid cell adhesion molecules (SELE, ICAM5 and CD209) and the coagulation factor F8, all of which are potentially druggable targets. Our results are broadly consistent with a multi-component model of COVID-19 pathophysiology, in which at least two distinct mechanisms can predispose to life-threatening disease: failure to control viral replication; or an enhanced tendency towards pulmonary inflammation and intravascular coagulation. We show that comparison between cases of critical illness and population controls is highly efficient for the detection of therapeutically relevant mechanisms of disease.