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Cited by 225Genetic Associations and EpidemiologyCOVID-19 Clinical Research StudiesSARS-CoV-2 and COVID-19 Research

GWAS and meta-analysis identifies 49 genetic variants underlying critical COVID-19

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Pairo-Castineira, Erola, Rawlik, Konrad, Bretherick, Andrew D., Qi, Ting, Wu, Yang, Nassiri, Isar, McConkey, Glenn A., Zechner, Marie +2814 more·2023·Nature
DOI 10.1038/s41586-023-06034-3PMC PMC10208981
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Abstract

Critical illness in COVID-19 is an extreme and clinically homogeneous disease phenotype that we have previously shown 1 to be highly efficient for discovery of genetic associations 2 . Despite the advanced stage of illness at presentation, we have shown that host genetics in patients who are critically ill with COVID-19 can identify immunomodulatory therapies with strong beneficial effects in this group 3 . Here we analyse 24,202 cases of COVID-19 with critical illness comprising a combination of microarray genotype and whole-genome sequencing data from cases of critical illness in the international GenOMICC (11,440 cases) study, combined with other studies recruiting hospitalized patients with a strong focus on severe and critical disease: ISARIC4C (676 cases) and the SCOURGE consortium (5,934 cases). To put these results in the context of existing work, we conduct a meta-analysis of the new GenOMICC genome-wide association study (GWAS) results with previously published data. We find 49 genome-wide significant associations, of which 16 have not been reported previously. To investigate the therapeutic implications of these findings, we infer the structural consequences of protein-coding variants, and combine our GWAS results with gene expression data using a monocyte transcriptome-wide association study (TWAS) model, as well as gene and protein expression using Mendelian randomization. We identify potentially druggable targets in multiple systems, including inflammatory signalling ( JAK1 ), monocyte–macrophage activation and endothelial permeability ( PDE4A ), immunometabolism ( SLC2A5 and AK5 ), and host factors required for viral entry and replication ( TMPRSS2 and RAB2A ).

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  • rs67579710's A allele is associated with lower CSF and higher plasma abundance of THBS3 as well as decreased risk of multiple phenotypes related to COVID-19 severity.

    “Last, rs67579710’s A allele is associated with lower CSF and higher plasma abundance of thrombospondin 3 (THBS3) as well as decreased risk of multiple phenotypes related to severe acute respiratory syndrome coronavirus 2 (COVID-19) severity (fig. S4E) (30).”

    From Low overlap of plasma and CSF protein quantitative trait loci affects protein discovery for neurological disease (2026) · cites this paper as [30]

    Table 1 (Main; page unavailable) reports an association between a variant near THBS3 and critical COVID-19, supporting the broader link between this region and disease severity. It lists rs41264915, not rs67579710, and does not establish decreased risk for rs67579710’s A allele. Searches for rs67579710 and THBS3 protein abundance also found no passage documenting the claimed CSF or plasma associations. (LLM, conf=0.76)

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Pairo-Castineira, E., Rawlik, K., Bretherick, A. D., Qi, T., Wu, Y., Nassiri, I., McConkey, G. A., Zechner, M., Klaric, L., Griffiths, F., Oosthuyzen, W., Kousathanas, A., Richmond, A., Millar, J., Russell, C. D., Malinauskas, T., Thwaites, R., Morrice, K., Keating, S., Maslove, D., Nichol, A., Semple, M. G., Knight, J., Shankar-Hari, M., Summers, C., Hinds, C., Horby, P., Ling, L., McAuley, D., Montgomery, H., Openshaw, P. J. M., Begg, C., Walsh, T., Tenesa, A., Flores, C., Riancho, J. A., Rojas-Martinez, A., Lapunzina, P., GenOMICC Investigators, Co-Investigators, Clohisey, S., Millar, J., Shankar-Hari, M., Management, L. A. D. T., Aitkin, E., Aravindan, L., Armstrong, R., Baillie, J. K., Biggs, H., Boz, C., Brown, A., Chikowore, P., Clark, R., Coutts, A., Coyle, J., Cullum, L., Das, S., Day, N., Donnelly, L., Duncan, E., Finernan, P., Fourman, M. H., Furlong, A., Furniss, J., Gallagher, B., Gilchrist, T., Golightly, A., Griffiths, F., Hafezi, K., Hamilton, D., Hendry, R., Kearns, N., Law, D., Law, R., Law, S., Lidstone-Scott, R., Lauder, C., Macgillivray, L., Maclean, A., Mal, H., McCafferty, S., McMaster, E., Meikle, J., Moore, S. C., Murphy, S., Mybaya, H., Odam, M., Oosthuyzen, W., Zheng, C., Chen, J., Parkinson, N., Paterson, T., Tucker, P., Schon, K., Stenhouse, A., Das, M., Swets, M., Szoor-McElhinney, H., Taneski, F., Turtle, L., Wackett, T., Ward, M., Weaver, J., Wrobel, N., Zechner, M., Guys and St Thomas’ Hospital, L., Pan, J., Grau, N., Jones, T. O., Lim, R., Marotti, M., Whitton, C., Bociek, A., Campos, S., Arbane, G., Shankar-Hari, M., Ostermann, M., Cha, M., DAmato, F., Kosifidou, E., Lorah, S., Morera, K., James Cook University Hospital, M., Bircham, S., Brady, L., Hugill, K., Henning, J., Bonner, S., Headlam, E., Jones, J., List, A., Morley, J., Welford, A., Kamangu, B., Ratnakumar, A., Shoremekun, A., Barts Health NHS Trust, L., Alldis, Z., Astin-Chamberlain, R., Bibi, F., Biddle, J., Blow, S., Bolton, M., Borra, C., Bowles, R., Burton, M., Choudhury, Y., Cox, A., Easthope, A., Ebano, P., Fotiadis, S., Gurasashvili, J., Halls, R., Hartridge, P., Kallon, D., Kassam, J., Lancoma-Malcolm, I., Matharu, M., May, P., Mitchelmore, O., Newman, T., Patel, M., Pheby, J., Pinzuti, I., Prime, Z., Prysyazhna, O., Shiel, J., Taylor, M., Tierney, C., Zongo, O., Wood, S., Zak, A., Collier, D., Royal Stoke University Hospital, S., Mundy, M., Thompson, C., Pritchard, L., Gellamucho, M., Cartlidge, D., Bandla, N., Bailey, L., Davies, M., Delaney, J., Scott, L., North Middlesex University Hospital NHS trust, L., Abdelrazik, M., Alasdair, F., Carter, D., Elhassan, M., Ganesan, A., Jenkins, S., Lamond, Z., Purohit, D., Rohit, K., Saleem, M., Wall, A., Xavier, K., Bakthavatsalam, D., Gehad, K., Gnanapragasam, P., Jain, K., Jain, S., Malik, A., Pappachan, N., Moreno-Cuesta, J., Haldeos, A., Vincent, R., Oziegb, M., King’s College Hospital, L., Cavazza, A., Cockrell, M., Corcoran, E., Depante, M., Finney, C., Jerome, E., Knighton, A., Nayak, M., Pappa, E., Saha, R., Saha, S., Dodd, A., O’Reilly, K., McPhail, M., Clarey, E., Noble, H., Smith, J., Charing Cross Hospital, S. M. H. A. H. H., Coghlan, P., Brett, S., Gordon, A., Templeton, M., Antcliffe, D., Banach, D., Darnell, S., Fernandez, Z., Jepson, E., Mohammed, A., Rojo, R., Arias, S. S., Gurung, A. T., Wong, J., The Royal Liverpool University Hospital, L., Fernandez-Roman, J., Hamilton, D. O., Johnson, E., Johnston, B., Martinez, M. L., Mulla, S., Waite, A. A. C., Williams, K., Waugh, V., Welters, I., Emblem, J., Norris, M., Shaw, D., John Radcliffe Hospital, O., Bashyal, A., Beer, S., Hutton, P., McKechnie, S., Davidson, N., Mathew, S., Readion, G., Ryu, J., Wilson, J., Addenbrooke’s Hospital, C., Agrawal, S., Elston, K., Jones, M., Meaney, E., Polgarova, P., Elbehery, M., Summers, C., Daubney, E., Ng, A., Marshall, J., Pathan, N., Stroud, K., White, D., Nottingham University Hospital, N., Andrew, A., Ashraf, S., Clark, A., Dent, M., Langley, M., Peters, C., Ryan, L., Sampson, J., Wei, S., Baddeley, A., Meredith, M., Morris, L., Gibbons, A., McLoughlin, L., St George’s Hospital, L., Delgado, C. C., Clark, V., Dawson, D., Ding, L., Durrant, G., Ezeobu, O., Harrison, A., Hurt, W. J., Kanu, R., Kinch, A., Leaver, S., Lisboa, A., Mathew, J., Patel, K., Saluzzio, R. P., Rawlins, J., Samakomva, T., Shah, N., Sicat, C., Texeira, J., De Queiroz, J. G., Da Gloria, E. F., Maccacari, E., Yun, N., Manna, S., Farnell-Ward, S., Maizcordoba, M., Thanasi, M., Ali, H. H., BHRUT (Barking Havering)—Queens Hospital and King George Hospital, E., Hastings, J., Grauslyte, L., Hussain, M., Ruge, B., King, S., Pogreban, T., Rosaroso, L., Smith, H., Phull, M.-K., Adams, N., Franke, G., George, A., Salciute, E., Wong, J., Dunne, K., Flower, L., Sharland, E., Sra, S., Royal Infirmary of Edinburgh, E., Andrew, G., Callaghan, M., Barclay, L., Marshall, L., Baillie, K., Amamio, M., Birch, S., Briton, K., Clark, S., Doverman, K., Hope, D., Mcculloch, C., Simpson, S., Singleton, J., Kingston Hospital, S., Fernandez, R., Allen, M., Baptista, D., Crowe, R., Fox, J., Khera, J., Loveridge, A., McKenley, I., Morino, E., Naranjo, A., O’Connor, D., Simms, R., Sollesta, K., Swain, A., Venkatesh, H., Herdman-Grant, R., Joseph, A., Queen Alexandra Hospital, P., Nown, A., Rose, S., Pogson, D., Boxall, H., Brimfield, L., Claridge, H., Daly, Z., George, S., Gribbin, A., Royal Gwent Hospital, N., Cheema, Y., Cutler, S., Richards, O., Roynon-Reed, A., Cherian, S., Heron, A. E., Williams, G., Szakmany, T., Waters, A., Collins, K., Dunhill, J., Jones, F., Morris, R., Ship, L., Cardwell, A., Royal Blackburn Teaching Hospital, B., Ali, S., Bhatterjee, R., Bolton, R., Chukkambotla, S., Coleman, D., Dalziel, J., Dykes, J., Fine, C., Gay, B., Goddard, W., Goodchild, D., Harling, R., Hijazi, M., Keith, S., Khan, M., Matt, R., Ryan-Smith, J., Saad, S., Springle, P., Thomas, J., Truman, N., Kazi, A., Smith, M., Collier, H., Davison, C., Duberley, S., Hargreaves, J., Hartley, J., Patel, T., Smith, E., Stepping Hill Hospital, S., Kent, A., Goodwin, E., Zaki, A., Tibke, C., Hopkins, S., Gerrard, H., Jackson, M., Bennett, S., Marsh, L., Mills, R., Northumbria Healthcare NHS Foundation Trust, N. S., Bell, J., Campbell, H., Dawson, A., Dodds, S., Duffy, S., Gallagher, L., McCafferty, G., Short, S., Smith, T., Thomas, K., Walker, C., Reynolds, J., Yates, B., McKie, H., Panteli, M., Thompson, M., Waddell, G., Countess of Chester Hospital, C., De Beger, S., Abraheem, A., Dunmore, C., Girach, R., Jones, R., London, E., Nagra, I., Nasir, F., Sainsbury, H., Smedley, C., Brearey, S., Burchett, C., Cawley, K., Faulkner, M., Jeffrey, H., Bamford, P., Shaikh, F., Slack, L., Davies, A., Pinderfields General Hospital, W., Brooke, H., Suarez, J. C., Charlesworth, R., Hansson, K., Norris, J., Poole, A., Sandhu, R., Smithson, E., Thirumaran, M., Wagstaff, V., Buckley, S., Sloan, B., Rose, A., Major, A., Metcalfe, A., Ninewells Hospital, D., Almaden-Boyle, C., Austin, P., Chapman, S., Eros, A., Cabrelli, L., Cole, S., Whyte, C., Casey, M., Croydon University Hospital, C., Bafitis, V., Tsinaslanidis, G., George, C., Khade, R., Black, C., Ashok, S. R., Morriston Hospital, S., Farley, S., Brinkworth, E., Harford, R., Murphy, C., Williams, M., Newey, L., Toghill, H., Lewis, S., Rees, T., Battle, C., Baker, M., Travers, J., Chesters, K., Queen Elizabeth University Hospital, G., Baxter, N., Arnott, A., McCreath, G., McParland, C., Rooney, L., Sim, M., Henderson, S., Abel, L., Dalton, C., Kennedy-Hay, S., O’Donohoe, L., O’Hare, M., Orlikowska, I., Parker, N., Broomfield Hospital, C., McNeela, F., Lyle, A., Hughes, A., Radhakrishnan, J., Gibson, S., Heartlands Hospital, B., Bancroft, H., Bellamy, M., Daglish, J., Kadiri, S., Moore, F., Rhodes, J., Sangombe, M., Peterkin, Z., Scriven, J., Carmody, M., Royal Sussex County Hospital, B., Cottle, J., Peasgood, E., de Gordoa, L. 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S., Singh, J., Stroud, N., Roche, L., Lynch, C., Deacon, B., Pothecary, C., Smeaton, J., Agravante, K., Barnet Hospital, L., Krishnamurthy, V., Diaba, C., John, L., Lim, L., Jha, R., Wythenshawe Hospital, M., Egan, J., Felton, T., Glasgow, S., Padden, G., Choudhr, O., Bradley-Potts, J., Moss, S., Lingeswaran, S., Alexander, P., Brandwood, C., Fiouni, S., Ward, L., Allen, S., Shaw, J., Smith, C., Medway Maritime Hospital, G., Adanini, O., Collins, R., Msiska, M., Ofori, L., Bhatia, N., Dolan, H., Royal Berkshire NHS Foundation Trust, B., Brunton, M., Caterson, J., Coles, H., Keating, L., Tilney, E., Jacques, N., Frise, M., Armistead, J., Bartley, S., Bhuie, P., Rai, S., Tomkova, G., Whiston Hospital, P., Greer, S., Shuker, K., Tridente, A., The Royal Oldham Hospital, M., Dobson, E., Hunt, J., Tully, R., Dearden, J., Drummond, A., Kamath, P., Bullock, E., Mulcahy, M., Munt, S., O’Connor, G., Philbin, J., Rishton, C., Scott, C., Winnard, S., Chesterfield Royal Hospital Foundation Trust, C., Hasni, N., Gascoyne, R., Hawes, J., Pritchard, K., Stevenson, L., Whileman, A., Beavis, S., Bishop, L., Cart, C., Dale, K., Kelly-Baxter, M., Mendelski, A., Moakes, E., Smith, R., Woodward, J., Wright, S., Aberdeen Royal Infirmary, A., Allan, A., Botello, A., Liew, J., Medhora, J., Trumper, E., Savage, F., Scott, T., Place, M., Kaye, C., Royal Devon and Exeter Hospital, E., Benyon, S., Marriott, S., Park, L., Quinn, H., Skyes, D., Zitter, L., Baines, K., Gordon, E., Keenan, S., Pitt, A., Glasgow Royal Infirmary, G., Duffy, K., Ireland, J., Semple, G., Turner, L., Cathcart, S., Rimmer, D., Puxty, A., Puxty, K., Hurst, A., Miller, J., Speirs, S., Walker, L., Blackpool Victoria Hospital, B., Bradshaw, Z., Brown, J., Melling, S., Preston, S., Slawson, N., Warden, S., Beasley, A., Stoddard, E., Benham, L., Cupitt, J., Caswell, M., Elawamy, L., Wignall, A., Southampton General Hospital, S., Roberts, B., Golding, H., Leggett, S., Male, M., Marani, M., Prager, K., Williams, T., Golder, K., Jones, O., Cusack, R., Bolger, C., Burnish, R., Carter, M., Jackson, S., Salmon, K., Biss, J., Ashford and St Peter’s Hospital, S., Aquino, M., Croft, M., Frost, V., White, I., Govender, K., Derriford Hospital, P., Webb, N., Stapleton, L., Wells, C., Nikitas, N., Sanchez-Rodriguez, A., Spencer, K., Stowe, B., East Surrey Hospital, R., Izzard, Y., Poole, M., Monnery, S., Trotman, S., Beech, V., Combes, E., Joefield, T., Poole Hospital, P., Covernton, P., Savage, S., Woodward, E., Camsooksai, J., Reschreiter, H., Barclay, C., DeAth, Y., Dube, J., Humphrey, C., Jenkins, S., Langridge, E., Milne, R., Wadams, B., Woolcock, M., Royal Alexandra Hospital, P., Brett, M., Digby, B., Gemmell, L., Hornsby, J., MacGoey, P., O’Neil, P., Price, R., Sundaram, R., Abel, L., Rodden, N., Thomson, N., Rooney, K., Currie, S., Parker, N., Walker, L., Henderson, P., St James’s University Hospital and Leeds General Infirmary, L., Ogg, B., Whiteley, S., Wilby, L., Long, K., Matthew, S., Salada, S., Trott, S., Watts, S., Friar, Z., Speight, A., Bedford Hospital, B., Bastion, V., Chandna, H., Djeugam, B., Haseeb, M., Kent, H., Lubimbi, G., Murdoch, S., Thomas, A., David, B., Lorusso, R., Vochin, A., Penacerrada, M., Wulandari, R., Southport and Formby District General Hospital, O., Heath, C., Jakkula, S., Morris, A., Ahmed, A., Nune, A., Buttriss, C., Whitaker, E., The Tunbridge Wells Hospital and Maidstone Hospital, K., Davey, M., Golden, D., Acklery, A., Fernandes, F., Seaman, B., Earl, V., Queen Elizabeth Hospital, W., Collins, A., Khaliq, W., Adam, R., Treus, E., North Manchester General Hospital, M., Holland, S., Alfonso, J., Blackledge, B., Bruce, M., Durrans, L. J., Eltayeb, A., Harris, J., Hey, S., Hruska, M., Lamb, T., Rothwell, J., Fitzgerald, A., Lindergard, G., T-Michael, H., Duncan, T., Baxter-Dore, S., Cooper, L., Fox, C., Guerin, J., Hodgkiss, T., Connolly, K., Royal Victoria Infirmary, N. U. 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K., Mwaura, E., Sutherland, S.-B., Stewart, R., Mew, L., Wren, L., Good Hope Hospital, B., Thrasyvoulou, L., Willis, H., Scriven, J., Hopkins, B., Lenton, D., Roberts, A., Queen Elizabeth Hospital Gateshead, G., Bokhari, M., Lucas, R., McCormick, W., Ritzema, J., Linnett, V., Sanderson, A., Wild, H., Royal Bolton Hospital, B., Flanagan, R., Hull, R., Rhead, K., McKenna, E., Hughes, G., Anderson, J., Jones, K., Latham, S., Riley, H., Tameside General Hospital, A. U. L., Coulding, M., Clark, M., McCormick, J., Mercer, O., Potla, D., Rehman, H., Savill, H., Turner, V., Jude, E., Kilroy, S., Salford Royal Hospital, M., Apetri, E., Basikolo, C., Blackledge, B., Catlow, L., Collis, M., Doonan, R., Harris, J., Harvey, A., Knowles, K., Lee, S., Lomas, D., Lyons, C., McMorrow, L., Michael, A., Pendlebury, J., Perez, J., Poulaka, M., Proudfoot, N., Slevin, K., Thomas, V., Walker, D., Dark, P., Charles, B., McLaughlan, D., Slaughter, M., Horner, D., Cawley, K., Marsden, T., Great Ormond St Hospital and UCL Great Ormond St Institute of Child Health NIHR Biomedical Research Centre, L., Andrews, J., Beech, E., Akinkugbe, O., Bamford, A., Belfield, H., Jones, G. A. L., McHugh, T., Meghari, H., Ray, S., Tomas, A. L., O’Neill, L., Peters, M., Bell, M., Benkenstein, S., Chisholm, C., Davies, C., Kupiec, K., Payne, C., Southmead Hospital, B., Halls, J., Blakemore, H., Goff, E., Hayes, K., Smith, K., Stephens, D., Worner, R., Borislavova, B., Faulkner, B., Thomas, M., Cookson, R., Gendall, E., Larman, G., Pope, R., Smalira, A., William Harvey Hospital, A., Priestley, V., Cosier, T., Millen, G., Rand, J., Schumacher, N., Sandhar, R., Weston, H., Richardson, N., Cooper, L., Arrowe Park Hospital, W., Jones, C., Huang, Y.-W. J., Jacob, R., Denmade, C., McIntyre, L., Royal Hampshire County Hospital, H., Trodd, D., Martin, J., Watson, G., Bevan, E., Wreybrown, C., Bradford Royal Infirmary, B., Bano, S., Bellwood, R., Bentley, M., Bromley, M., Gurr, L., Ledgard, C., McGowan, J., Pye, K., Sellick, K., Stacey, A., Warren, D., Wilkinson, B., Akeroyd, L., Shafique, H., Morgan, J., Shorter, S., Swinger, R., Waters, E., Lawton, T., Glan Clwyd Hospital, B., Allan, E., Darlington, K., Davies, F., Davies, L., Easton, J., Kumar, S., Lean, R., Mackay, C., Pugh, R., Qiu, X., Rees, S., Scanlon, J., Lewis, J., Menzies, D., Bolger, A., Davies, G., Davies, J., Garrod, E., Jones, H., Manley, R., Williams, H., Royal Bournemouth Hospital, B., Frankham, J., Pitts, S., White, N., Branney, D., Tiller, H., Bristol Royal Infirmary, B., Efford, G., Garland, Z., Grimmer, L., Gumbrill, B., Johnson, R., Sweet, K., Bewley, J., Coleman, C., Corcoran, K., Morano, E. M. H., Shiel, R., Webster, D., Bonnici, J., Daniel, E., Dell, A., University Hospital North Durham, D., Kent, M., Wilkinson, A., Brown, E., Kay, A., Campbell, S., Cowton, A., Birt, M., Greenaway, V., Potts, K., Hutton, C., Shepperson, A., Basildon Hospital, B., Forsey, M., Nicholson, A., Vertue, M., Riches, J., Kaliappan, A., Nicholson, A., University College Hospital, L., MacCallum, N., Raith, E., Bercades, G., Hass, I., Brealey, D., Martir, G., Reyes, A., Smyth, D., Zapatamartinez, M., Whittington Hospital, L., Alvaro, A., Jetha, C., Ma, L., Booker, L., Mostoles, L., Pratley, A., Altabaibeh, A., Parmar, C., Gilbert, K., Western General Hospital, E., Ferguson, S., Shepherd, A., Morris, S., Singleton, J., Baruah, R., Amamio, M., Birch, S., Briton, K., Clark, S., Doverman, K., Marshall, L., Simpson, S., Ipswich Hospital, I., Lloyd, G., Bell, S., Rivers, V., Purewal, B., Hereford County Hospital, H., Hammerton, K., Anderson, S., Birch, J., Collins, E., Oleary, R., Sunderland Royal Hospital, S., Cornell, S., Jarmain, J., Rogerson, K., Wakinshaw, F., Woods, L., Rostron, A., Elcioglu, Z., Roy, A., Queens Hospital Burton, B.-O.-T., Bell, G., Dickson, H., Wilcox, L., Katary, A., English, K., Musgrove Park Hospital, T., Hutter, J., Pawley, C., Doble, P., Shovelton, C., Vaida, M., Purnell, R., Thomas, A., The Royal Papworth Hospital, C., Cagova, L., Fofano, A., Holcombe, H., Mitchell, A. M., Mwaura, L., Raman, K. P., Garnr, L., Mepham, S., Paques, K., Vuylsteke, A., Mackie, J., Pearn, C., Zamikula, J., University Hospital Lewisham, L., Birt, M., Gude, E. T., Nyirenda, M., Capozzi, L., Reece-Anthony, R., Khaliq, W., Noor, H., Nilo, A. C., The Princess Alexandra Hospital, H., Grove, M., Daniel, A., Easthope, A., Finn, J., White, N., Saha, R., Badal, B., Ixer, K., University Hospital of Wales, C., Duffin, D., Player, B., Hill, H., Cole, J., Brooks, J., Davies, M., Davies, R., Hunt, L., Thomas, E., Williams, A., West Middlesex Hospital, I., Oblak, M., Thankachen, M., Irisari, J., Sayan, A., Popescu, M., Royal Albert Edward Infirmary, W., Finch, C., Jamieson, A., Quinn, A., Cooper, J., Liderth, S., Waddington, N., Stoke Mandeville Hospital, B., Burn, I., Manso, K., Penn, R., Tebbutt, J., Thornton, D., Winchester, J., Hambrook, G., Shanmugasundaram, P., Royal Lancaster Infirmary, L., Craig, J., Simpson, K., Higham, A., Sibbett, L., Basingstoke and North Hampshire Hospital, B., Paine, S., Reed, A., Conyngham, J.-A., Mupudzi, M., Thomas, R., Wright, M., Griffin, D., Partridge, R., Corral, M. A., Muchenje, N., Sitonik, M., Brown, C. W., Worthing Hospital, W., Butler, A., Folkes, L., Fox, H., Gardner, A., Helm, D., Hobden, G., King, K., Margalef, J., Margarson, M., Martindale, T., Meadows, E., Raynard, D., Thirlwall, Y., Baird, Y., Gomez, R., Martin, D., Hodgson, L., Corin, C., Sidall, E., Szabo, D., Floyd, S., The Alexandra Hospital, R. A. W. R. H., Davies, H., Austin, K., Kelsall, O., Wood, H., Anderson, P., Archer, K., Burtenshaw, A., Clayton, S., Cother, N., Cowley, N., Davis, C., Digby, S., Durie, A., Harrison, A., Low, E., McAlindon, M., McCurdy, A., Morgan, A., Rankin, T., Thrush, J., Tranter, H., Vigurs, C., Wild, L., Royal Cornwall Hospital, T., Cornell, T., Ralph, K., Bean, S., Burt, K., Spivey, M., Richards, C., Tedstone, R., Watford General Hospital, W., Carmody, S., Zhao, X., Page, V., Guanco, M. L., Hoxha, E., Zorloni, C., Macclesfield District General Hospital, M., Dean, C., Jones, E., Carter, E., Dunn, J., Kong, T., Mahenthran, M., Marsh, C., Holland, M., Keenan, N., Mahmoud, M., Lyons, M., Bradley-Potts, J., Wassall, H., Young, M., Royal Surrey County Hospital, G., Bradley, P., Burda, D., Donlon, S., Harden, L., Harris, C., Mayangao, I., Montaser, R., Mtuwa, S., Piercy, C., Smith, E., Stone, S., Verula, J., Blackman, H., Marriott, C., Michalak, N., Creagh-Brown, B., Salberg, A., Boyer, N., Pristopan, V., Rotherham General Hospital, R., Maynard, V., Walker, R., Hormis, A., Collier, D., Graham, C., Maynard, V., McCormick, J., Warrington, J., Craigavon Area Hospital, C. A., Cosgrove, D., McFarland, D., Ratcliffe, J., Charnock, R., King’s Mill Hospital, N., Wynter, I., Gill, M., Kirk, J., Paul, P., Ratnam, V., Shelton, S., Dumfries and Galloway Royal Infirmary, D., Jardine, C., Hay, A., Williams, D., Prince Charles Hospital, M. T., Deacon, B., Durga, L., Hibbert, M., Kennard-Holden, G., Woodford, C., Pothecary, C., Roche, L., Tetla, D., Agravante, K., Smeaton, J., Ysbyty Gwynedd, B., Price, A., Thomas, A., Thorpe, C., Knights, E., Ward, D., Royal Preston Hospital, P., Laha, S., Verlander, M., Williams, A., The Great Western Hospital, S., Prout, R., Langton, H., Watters, M., Hunt, C., Novis, C., Lincoln County Hospital, L., Arif, S., Cunningham, A., Hewitt, C., Hindale, J., Jackson-Lawrence, K., Shepardson, S., Wills, M., Butler, S., Tavares, S., Barber, R., Hilldrith, A., Hubbard, K., University Hospital of North Tees, S. O. T., Egginton, D., Clark, M., Purvis, S., Sinclair, S., Collins, V., Glangwili General Hospital, C., Landeg, B., Sell, C., Coetzee, S., Gales, A., Otahal, I., Icke, B., Raj, M., Williams, C., Williams, J., Hill, L., Southend University Hospital, W.-O.-S., Kayani, A., Masunda, B., Gondo, P., Atayeva, N., Lister Hospital, S., Cruz, C., Pattison, N., Diana Princess of Wales Hospital, G., Burnett, C., Hatton, J., Heeney, E., Newton, M., Al-Moasseb, H., Behan, T., Player, J., Stead, R., Mitra, A., Nauyokas, K., West Suffolk Hospital, B. S. E., Humphreys, S., Cockerill, H., Tampsett, R., Victoria Hospital, K., Postovalova, E., Coventry, T., McGregor, A., Fowler, S., Macmahon, M., Cochrane, P., Pirie, S., Calderdale Royal Hospital, H., Hanley, S., Ali, A., Brady, M., Dale, S., Dance, A., Gledhill, L., Greig, J., Hanson, K., Holdroyd, K., Home, M., Ishaq, T., Kelly, D., Matapure, L., Melia, D., Mellor, S., Merwaha, E., Nortcliffe, T., Shaw, L., Shaw, R., Wood, T., Bayo, L.-A., Usher, M., Wilson, A., Kitson, R., Pinnell, J., Robinson, M., Boltwood, K., Dorset County Hospital, D., Birch, J., Bough, L., Tutton, R., Winter-Goodwin, B., Goodsell, J., Taylor, K., Williams, P., Williams, S., Cave, A., Rees, J., Russell’s Hall Hospital, D., Imeson-Wood, J., Smith, J., Amin, V., Karthik, K., Kausar, R., Anastasescu, E., Reid, K., Anumakonda, V., Stoddart, E., Royal United Hospital, B., Demetriou, C., Eckbad, C., Howie, L., Mitchard, S., Ramos, L., White, K., Hierons, S., Kelly, F., Serrano-Ruiz, A., Evans, G., St Mary’s Hospital, N., Nicol, L., Wilkins, J., Hulacka, K., Debreceni, G., Brown, A., Crickmore, V., George Eliot Hospital NHS Trust, N., Hill, K., Kannan, T., Yeovil Hospital, Y., Dagutao, Z., Beesley, K., Lewis, A., Perry, J., Antony, S., Board, S., Buckley, C., Pippard, L., Tanate, A., Wood, D., Kubisz-Pudelko, A., Gouda, A., Forth Valley Royal Hospital, F., Auld, F., Donnachie, J., Murdoch, E., Prentice, L., Runciman, N., Senaratne, D., Short, A., Sweeney, L., Symon, L., Todd, A., Turner, P., McCann, E., Salutous, D., Edmond, I., Whitelaw, L., Frimley Park Hospital, S., Venkatesh, H., Bland, Y., Kajtor, I., Kavanagh, L., Singler, K., Linfield-Brown, G., Chelsea & Westminster NHS Foundation Trust, L., Moore, L. S. P., Vizcaychipi, M., Martins, L., Moore, L., Bull, R., Carungcong, J., Queen Elizabeth the Queen Mother Hospital, M., Allen, L., Beranova, E., Knight, A., Price, C., Tilbey, S., Turney, S., Hazelton, T., Tutt, G., Arora, M., Turki, S., Sinfield, E., Deery, J., Ramos, H., Royal Brompton Hospital, L., Cristiano, D., Dormand, N., Farzad, Z., Gummadi, M., Salmi, S., Sloane, G., Varghese, M., Thwaites, V., Patel, B., Kamal, L., Zborowski, A. C., Darent Valley Hospital, D., Coe, R., Anderson, M., Beadle, J., Coates, C., Collins, K., Crowley, M., Johnson, L., King, L., Paramsothy, R., Sargeant, J., Silva, P., Stuart, C., Taylor, J., Tyl, D., Wakefield, P., Kamundi, C., Olufuwa, O., Belagodu, Z., Gherman, A., Oakley, N., University Hospital Crosshouse, K., Allan, J., Geary, T., Meikle, A., O’Brien, P., Wood, S., Clark, A., Houston, G., University Hospital Wishaw, W., Black, K., Clarkson, M., D’Sylva, S., Morrison, A., Norman, K., Taylor, M., Clements, S., Cohrane, C., Gonzalez, N., Strachan, D., Beith, C., Moar, K., University College Dublin, S. V. U. H., Murphy, L., Smythe, M., Nichol, A., Brickell, K., The Queen Elizabeth Hospital, K. L., Ali, I. A. M., Beaumont, K., Elsaadany, M., Fernandes, K., Ally, S. M., Rangarajan, H., Sarathy, V., Selvanayagam, S., Vedage, D., White, M., Coton, Z., Joshy, A., Blunt, M., Curgenven, H., Walsall Manor Hospital, W., Botfield, L., Dexter, C., Kuravi, A., Butler, J., Chadwick, R., Ranga, P., Richardson, L., Virgilio, E., Anwer, M., Garg, A., Botfield, D., Marriott, X., Princess Royal Hospital, B., Stewart, K., Mullan, D., Phillips, C., Gaylard, J., Nowak, J., Skinner, D., Barnsley Hospital, B., Jones, S., Crawley, R., Crew, A., Cunningham, M., Daniels, A., Harrison, L., Hope, S., Lancaster, N., Matthews, J., Wray, G., Nicholson, A., Inweregbu, K., Cutts, S., Miller, K., Warrington General Hospital, W., Brady, A., Chan, R., McIvor, S., Prady, H., Mathew, B., Little, J., Furniss, T., Royal Victoria Hospital, B., Wright, C., King, B., Wasson, C., O’Neill, A., Turley, C., McGuigan, P., Collins, E., Finn, S., Green, J., McAuley, J., Nair, A., Quinn, C., Tauro, S., Ward, K., McGinlay, M., Reddy, K., Royal Hallamshire Hospital and Northern General Hospital, S., Ahmad, N., Anderson, S., Barker, J., Bauchmuller, K., Birchall, K., Bird, S., Cawthron, K., Chetam, L., Cole, J., Donne, B., Foote, D., Ford, A., Hanratty, H., Harrington, K., Hesseldon, L., Housley, K., Jackson, Y., Jarman, C., Kibutu, F., Lenagh, B., Macharia, I., Masuko, S., Milner, L., Newell, H., Nwafor, L., Oxspring, S., Phillips, P., Raithatha, A., Rowland-Jones, S., Smith, J., Thompson, R., Trower, H., Walker, S., Watson, J., Wiles, M., Lye, A., Willson, J., Mills, G., Harris, S., Hartill, E., Harefield Hospital, L., Barron, A., Collins, C., Kaul, S., Nolan, C., Polgar, O., Prendergast, C., Rogers, P., Shokkar, R., Woodruff, M., Mahay, K., Thwaites, V., Reed, A., Meyrick, H., Passmore, H., Farwell, J., Cumberland Infirmary, C., Brown, A., O’Connell, S., Gregory, J., Barberis, L., Harper, R., Smith, T., Armstrong, D., Eastbourne District General Hospital, E. S., Bowey, A., Cowley, A., Corner, A., Highgate, J., Rutherfurd, C., Taylor, J.-A., Goodwin, S., Rutherford, C., Salisbury District Hospital, S., Eapen, B., Trim, F., Donnison, P., Airedale General Hospital, K., Armstrong, L., Bates, H., Dooks, E., Farquhar, F., Kitching, A., McParland, C., Packham, S., Hairsine, B., Leicester Royal Infirmary, L., Patil, A., Andreou, P., Hales, D., Mathews, M., Patel, R., Barry, P., Flint, N., Hailstone, J., Ghuman, N., Leonard, B., Lees, R., Peterborough City Hospital, P., Butcher, D., Leng, K., Butterworth-Cowin, N., O’Sullivan, S., Colchester General Hospital, C., Ghosh, A., Williams, E., Princess Royal Hospital, T. A. R. S. H., Adams, C., Agasou, A., Arden, T., Beekes, M., Bowes, A., Boyle, P., Button, H., Carnahan, M., Carter, A., Childs, D., Gaylard, J., Hurford, F., Hussain, Y., Javaid, A., Jones, J., Leigh, M., Martin, T., Millward, H., Motherwell, N., Mullan, D., Newman, J., Rikunenko, R., Stickley, J., Summers, J., Ting, L., Tivenan, H., Donaldson, D., Capps, N., Cale, E., Jose, S., Osbourne, W., Pajak, S., Rankin, J., Tonks, L., University Hospital Monklands, A., Baird, T., Harkins, M., Ruddy, J., West, J., Wrexham Maelor Hospital, W., Duffield, J., Mallon, L., Smith, O., Smuts, S., Campbell, A., Davies, C., Davies, S., Hughes, R., Jobes, L., Whitehead, V., Watkins, C., Royal Hospital for Children, G., Bowman, F., Milligan, B., McPherson, L., New Cross Hospital, W., Metherell, S., Harris, N., Lake, V., Radford, E., Smallwood, A., Gopal, S., Vassell, K., University Hospital Hairmyres, E. K., Bell, D., Boyle, R., Douglas, K., Glass, L., Lennon, L., Rattray, A., Beith, C., Lee, E., Warwick Hospital, W., Jones, D., Parsons, P., Attwood, B., Jefferson, P., Ranganathan, M., Atwal, I., Campbell, B., Day, A., Stagg, C., Sandwell General Hospital and City Hospital, B., Haynes, E., Ahmed, C., Clamp, S., Colley, J., Haq, R., Hayes, A., Joseph, S., Maqsood, Z., Hussain, S., Hulme, J., Domingos, P., Kumar, R., Purewal, M., Taylor, B., Royal Manchester Children’s Hospital, M., Bunni, L., Latif, M., Jennings, C., Jose, S., Marshall, R., Metryka, A., Subramanian, G., Gloucestershire Royal Hospital, G., Burgoyne, A., O’Connell, S., Tyler, A., Waldron, J., Hilltout, P., Evitts, J., University Hospitals Coventry & Warwickshire NHS Trust, C., Ward, G., Bremmer, P., Hawkins, C., Jackman, S., Ogorek, M., Torbay Hospital, T., Ashby, K., Thornton, L., Mercer, P., Halkes, M., Revill, A., Pilgrim Hospital, L., Saint, B., Fletcher, J., Netherton, K., Chablani, M., Kirkby, A., Roper, A., Szymiczek, K., Prince Philip Hospital, L., Sutherland, I., O’Brien, L., Otahal, I., Connell, J., Davies, K., Lewis, T., Omar, Z., Perkins, E., Princess of Wales Hospital, L., Roche, L., Sathe, S., Davies, E., Northampton General Hospital NHS Trust, N., Lyon, A., Mapfunde, I., Willis, C., Hitchcock, R., Hall, K., King, C., The Christie NHS Foundation Trust, M., Fagan, A., Nazari, R., Worsley, L., Allibone, S., Kasipandian, V., Patel, A., Cutting, P., Genetu, R., Mac, A., Murphy, A., Ward, S., Butt, F., James Paget University Hospital NHS Trust, G. Y., Ayers, A., Harrison, W., Mackintosh, K., North, J., Birmingham Children’s Hospital, B., Ashton, L., Bi, R., Owen, S., Winmill, H., Scholefield, B., Withybush General Hospital, P., Blowing, H., Williams, E., Duskova, M., Edwards, M., Rees, A., Thomas, H., Hughes, R., Otahal, I., Brooks, J., Phipps, J., Brooks, S., Northwick Park Hospital, L., Dennis, C., Parris, V., Srikaran, S., Sukha, A., McGregor, A., Tiongson, G., North Devon District Hospital, B., Adams, K., Andrew, B., Brayne, A., Carter, S., Findlay, L., Fisher, E., Jackson, P., Kaye, D., Parkin, J., Tuckey, V., Hunt, J., Love, N., van Koutrick, L., Hanson, A., Scunthorpe General Hospital, S., Dent, K., Horsley, E., Pearson, S., Spencer, S., Hutchinson, D., Player, J., Potoczna, D., Akhtar, M. N., Cottam, L.-J., Nauyokas, K., Sanders, J., Royal Free Hospital, L., Garcia, S. M., Pakou, G., Diaba, C., Filipe, H., John, L., Maharajh, A., de Neef, M., Martin, D., Eastgate, C., Teoh, P. C., Raigmore Hospital, I., Barrett, F., Bradley, C., Donaldson, A., Mascarenhas, M., O’Hara, M., Okeefe, L., Clarke, N., Whiteside, J., Campbell, R., Matheson, J., McDonald, D., Patience, D., West Cumberland Hospital, W., Rice, P., Smith, T., Clapham, M., Mutch, R., Barberis, L., Harper, R., Craig, H., Poultney, U., Furness General Hospital, B.-I.-F., Burns, K., Higham, A., Liverpool Heart and Chest Hospital, L., Twiss, S., Barton, J., George, L., Harrop, C., Mathew, S., Wright, D. J., Scarborough General Hospital, S., Harrison, R., Toohie, J., Chandler, B., Turnbull, A., Mallinson, J., Elliott, K., Bronglais General Hospital, A., Wolf-Roberts, R., Tench, H., Otahal, I., Hobrok, M., Loosley, R., McGuinness, H., Sims, T., Alder Hey Children’s Hospital, L., Afolabi, D., Allison, K. S., Anderson, T., Dore, R., Jones, D., Rogers, N., Saunderson, P., Whitbread, J., O’Malley, L., Rad, L., Hawcutt, D., Borders General Hospital, M., Aldridge, J., Tolson, M., Garrioch, S., Leighton Hospital, C., Tomlinson, J., Grosdenier, M., Kent & Canterbury Hospital, C., Loader, D., Kapoor, R., Hector, G., Harrogate and District NHS Foundation Trust, H., Scherewode, J., Sri-Chandana, C., Stephenson, L., Marsh, S., The Royal Marsden Hospital, L., Dela Rosa, A., Jhanji, S., Bemand, T., Howle, R., Baikady, R. R., Thomas, B., Black, E., Tatham, K., Ealing Hospital, S., Gurram, S., Watson, E., Parris, V., Quaid, S., McGregor, A., St John’s Hospital Livingston, L., Saunderson, A., O’Brien, R., Moultrie, S., Service, J., Cheyne, C., Odam, M., Wiliams, A., Wexham Park Hospital, S., Barnes, N., Csabi, P., Da Rocha, J., Glynou, L., Sheffield Children’s Hospital, S., Huffenberger, A., Bryant, J., Pickard, A., Roe, N., Bellini, A., Mayer, A., Burrow, A., Colley, N., Evans, J., Howlett, A., Khalifeh, Z., Homerton University Hospital Foundation NHS Trust, L. U., Pryce, J., Gorman, C., Easthope, A., Brady, R., Timlick, E., Antoine, P., Gupta, A., National Hospital for Neurology and Neurosurgery, L., Hardy, J., Houlden, H., Moncur, E., Tucci, A., Raith, E., Tariq, A., Brealey, D., The Royal Alexandra Children’s Hospital, B., Tagliavini, E., Ramsay, B., Fidler, K., Donnelly, K., Hollis, R., Golden Jubilee National Hospital, C., Barr, J., Boyd, E., Irvine, V., Shelley, B., Buckley, J., Hamilton, C., Valdeavella, K., SCOURGE Consortium, Abellan, J., Acosta-Isaac, R., Aguado, J. M., Aguilar, C., Aguilera-Albesa, S., Sabbagh, A. A., Alba, J., Albu, S., Alcalá-Gallardo, K. A. M., Alcoba-Florez, J., Batres, S. A., Algarin-Lara, H. R., Almadana, V., Almeida, J., Almoguera, B., Alonso, M. R., Alvarez, N., Álvarez-Benítez, Y., Álvarez-Navia, F., Walther, R. A.-S., Andreu-Bernabeu, Á., Antonijoan, M. R., Arana-Arri, E., Aranda, C., Arango, C., Araque, C., Araujo, N. K., Araujo, I. M. T., Arcanjo, A. C., Arnaiz, A., Fernández, F. A., Arranz, M. J., Lopez, J. R. A., Artiga, M.-J., Avello-Malaver, Y., Ayuso, C., Baldión-Elorza, A. M., Martín, B. B., Baptista-Rosas, R. C., Barranco-Díaz, A., Barreda-Sánchez, M., Barrera-Penagos, V., Belhassen-Garcia, M., Bernal, E., Bernal-Bello, D., Bezerra, J. F., Bezerra, M. A. C., Blanca-López, N., Blancas, R., Boix-Palop, L., Borobia, A., Bravo, E., Brion, M., Brochado-Kith, Ó., Brugada, R., Bustos, M., Cabello, A., Caceres-Agra, J. J., Calbo, E., Calderón, E. J., Camacho, S., Campos, M. C., Cañadas, Y., Carbonell, C., Cardona-Huerta, S., Carioca, A. A. F., Carpintero, M. S., Segura, C. C., Carratto, T. M. T., Carrillo-Avila, J. A., Carvalho, M. C. C., Casasnovas, C., Castano, L., Castaño, C. F., Castelao, J. E., Candalija, A. C., Castillo, M. A., Ceballos, F. C., Chaux, J. G., Chaves-Santiago, W. G., Chiquillo-Gómez, S., Cid-Lopez, M. A., Cienfuegos-Jimenez, O., Conde-Vicente, R., Cordero-Lorenzana, M. L., Corella, D., Corrales, A., Cortes-Sanchez, J. L., Corton, M., Costa, T. X., Cruz, R., Cruz, M. S., Cuesta, L., Cunha, G. C. R., da Silva, G. V., Dalmau, D., Dantas-Komatsu, R. C. S., Darnaude, M. T., de Andrés, R., de Araújo, J. N. G., de Juan, C., De la Cruz Troca, J., de la Horra, C., de la Hoz, A. B., De Martino-Rodríguez, A., de Sousa Alves Neri, J. L., del Campo-Pérez, V., Delgado-Cuesta, J., Diaz-Caneja, C. M., Díaz-Pérez, A., de Bustamante, A. D., Dietl, B., Diz-de Almeida, S., Alves, M. D. M., Domínguez-Garrido, E., dos Santos, K. A., Duarte, A. M., Echave-Sustaeta, J., Eiros, R., Enciso-Olivera, C. O., Escudero, G., España, P. P., Sanabria, G. M. E., Fariñas, M. C., Fernandes, M. R., Fernández, R., Fernandez-Caballero, L., Fernández-Cruz, A., Fernandez-Nestosa, M. J., Fernández-Robelo, U., Fernández-Rodríguez, A., Fernández-Sampedro, M., Fernández-Sánchez, R., Fernández-Villa, T., Ferrero, S. F., Martínez, Y. F., Capitán, C. F., Flores-Pérez, P., Friaza, V., Fuenmayor-Hernández, L., Núñez, M. F., Fumadó, V., Gadea, I., Gagliardi, L., Gago-Domínguez, M., Gallego, N., Galoppo, C., García, I., García, M., García, L., Garcia-Cerrada, C., García-de-Vicuña, A., Garcia-García, J., García-García, I., García-Ibarbia, C., García-Montero, A. C., García-Soidán, A., García-Vázquez, E., Torrejón, M. C. G., Garza-Frias, E., Gentile, A., Gil-Fournier, B., Gómez-Arrue, J., Gómez-Duque, M., Carrera, L. G., García, M. G., Sacristán, Á. G., González-Neira, A., González-Peñas, J., Gonzalez-Sagrado, M., Álvarez, B. G., de Quirós, F. G. B., Benito, H. G., Gorgojo-Galindo, O., Górgolas, M., Guaragna, F., Guegel, G. P., Guillen-Guio, B., Guillen-Navarro, E., Guisado-Vasco, P., Gutiérrez-Bautista, J. F., Gutierrez-Castañeda, L. D., Heili-Frades, S., Hernandez, E., Hernandez-Ortega, L. D., Hernández-Pérez, G., Hernández-Vaquero, R., Moro, C. H., Herraez, B., Herranz, M. T., Herrera, M., Herrero, M. J., Herrero-Gonzalez, A., Horcajada, J. P., Imaz-Ayo, N., Intxausti-Urrutibeaskoa, M., Íñiguez, M., Jacomo, R. H., Jara, R., Jazmin, P. M., Jiménez, Á., Jiménez, P., Jiménez-Alfaro, I., Jimenez-Sousa, M. A., Jordan, I., Laguna-Goya, R., Laorden, D., Lasa-Lazaro, M., Lattig, M. C., Lauriente, A., Borja, A. L., Llanos, L., López-Bernús, A., Lopez-Garcia, E., Lopez-Rodriguez, R., López-Ruz, M. A., Granados, E. L., Lorente, L., Lozano, J. E., Lozano-Espinosa, M., Luchessi, A. D., Mahillo, I., Mancebo, E., Mar, C., Calvo, C. M., Marcos, M., Marcos-Delgado, A., Candon, A. M., Aguilar, P. M., Martín, M. M., Martín, M. D., Martín, V., Martin-Fernandez, M., Martín-López, C., Martín-Oterino, J.-Á., Martin-Pedraza, L., Martín-Vicente, M., Martinez, A., Martínez, R., Martínez, J. J., Martínez, S., Martínez-Aquino, E., Martínez-González, Ó., Martinez-Lopez, I., Martinez-Nieto, O., Martinez-Paz, P., Martinez-Perez, A., Martínez-Ramas, A., Martinez-Resendez, M. F., Robles, V. M., Marzal, L., Mazzeu, J. F., Medeiros, J. F. P., Medeiros, K. A., Medrano, F. J., Meijome, X. M., Mejuto-Montero, N., Méndez-Echevarria, A., Charris, H. M., Macías, E. M., Mercadillo, F., Mercado-Sesma, A. R., Minguez, P., Molina, A. J. J., Molina-Roldán, E., Montoya, J. J., Moraes, V. M. S., Moreira-Escriche, P., Morelos-Arnedo, X., Moreno-Docón, A., Moreno-Escalante, J., Cuerda, V. M., Fernández, A. M., Morilla, R., García, P. M., Neira, P., Nevado, J., Nieto-Gañán, I., Nunes, J. F. R., Nuñez-Torres, R., Obrador-Hevia, A., Ocejo-Vinyals, J. G., Olivar, V., Oliveira, S. F., Ondo, L., Orfao, A., Ortega, L., Ortega-Paino, E., Ortiz-Flores, F., Ortiz-Lopez, R., Oteo, J. A., Pachajoa, H., Pacheco, M., Pacheco-Miranda, F. J., Conejo, I. P., Panadero-Fajardo, S., Parellada, M., Pariente-Rodríguez, R., Paz-Artal, E., Peces-Barba, G., Pedromingo Kus, M. S., Perales, C., Perez, P., Pérez, C., Perez-de-Nanclares, G., Pérez-García, F., Pérez-Matute, P., Pérez-Serra, A., Pérez-Tomás, M. E., Perucho, T., Pichardo, L. A., Pinho, S. M. T., Pinsach-Abuin, M., Pinzón, L. A., Pita, G., Pla-Junca, F., Planas-Serra, L., Pompa-Mera, E. N., Porras-Hurtado, G. L., Pujol, A., Chávez, M. E. Q., Quijada, M. A., Quintela, I., Ramirez-Montaño, D., León, S. R., Sedes, P. R., Recalde, D., Recio-Fernández, E., Resino, S., Ribeiro, A. P., Rivadeneira-Chamorro, C. S., Roa-Agudelo, D., Pardo, M. R., Rodriguez, M. J., Rodriguez-Artalejo, F., Rodríguez-Ferrer, M., Rodriguez-Gallego, C., Rodriguez-Garcia, J. A., Rodriguez-Hernandez, M. A., Rodriguez-Nicolas, A., Rodriguez-Palmero, A., Rodríguez-Ruiz, E., Rodriguez-Urrego, P. A., Maya, B. R., Novoa, G. E. R., Rojo, F., Romero-Coronado, A., García, F. R., Rosa, L. S., Rosales-Castillo, A., Rubio, C., Olivera, M. R., Ruiz, M., Ruiz-Cabello, F., Ruiz-Casares, E., Ruiz-Cubillan, J. J., Ruiz-Hornillos, J., Ryan, P., Salamanca, H. D., Salazar-García, L., Origlia, G. G. S., Sánchez, P.-L., Sánchez-Pablo, C., Sánchez-Pernaute, O., Sánchez López, A. J., Prados, M. C. S., Real, J. S., Redondo, J. S., Sancho-Sainz, C., Sangil, A., Santos, A., Santos, N. P. C., Schlüter, A., Segovia, S., Serra-Llovich, A., Puras, F. S., Porras, M. S., Sicolo, M. A., Silbiger, V. N., Silva, N. S., Silva, F. T. C., Fuentes, C. S., Solé-Violán, J., Soria, J. M., Sorlí, J. V., Sousa, R. R., Souto, J. C., Souza, K. S. C., Souza, V. S., Sprockel, J. J., Suárez-Rama, J. J., Suarez-Zamora, D. A., Taboada-Fraga, X., Tamayo, E., Tamayo-Velasco, A., Taracido-Fernandez, J. C., Tavares, N. A. C., Tellería, C., Castaño, J. A. T., Teper, A., Torres-Macho, J., Torres-Tobar, L., Torres Gutiérrez, R. P., Troya, J., Urioste, M., Valencia-Ramos, J., Valido, A., Gallo, J. P. V., Varón, B., Vasconcelos, R. H. T., Vega, T., Velasco-Quirce, S., Vélez-Santamaría, V., Víctor, V., Estévez, J. V., Vieitez-Santiago, M., Vilches, C., Villalobos, L., Villar, F., Villar-Garcia, J., Villaverde, C., Villoslada-Blanco, P., Virseda-Berdices, A., Yáñez, Z., Zapatero-Gaviria, A., Zarate, R., Zazo, S., de Heredia, M. L., Mendes, I., Moreno, R., Sande, E., Lapunzina, P., Carracedo, A., ISARICC Investigators, Co-Investigators, Alex, B., Andrikopoulos, P., Bach, B., Barclay, W. S., Bogaert, D., Chand, M., Chechi, K., Cooke, G. S., da Silva Filipe, A., de Silva, T., Docherty, A. B., dos Santos Correia, G., Dumas, M.-E., Dunning, J., Fletcher, T., Green, C. A., Greenhalf, W., Griffin, J., Gupta, R. K., Harrison, E. M., Ho, A. Y. W., Holden, K., Horby, P. W., Ijaz, S., Khoo, S., Klenerman, P., Law, A., Lewis, M., Liggi, S., Lim, W. S., Maslen, L., Mentzer, A. J., Merson, L., Meynert, A. M., Moore, S. C., Noursadeghi, M., Olanipekun, M., Osagie, A., Palmarini, M., Palmieri, C., Paxton, W. A., Pollakis, G., Price, N., Rambaut, A., Robertson, D. L., Russell, C. D., Sancho-Shimizu, V., Sands, C., Scott, J. T., Sigfrid, L., Solomon, T., Sriskandan, S., Stuart, D., Swann, O. V., Takats, Z., Takis, P., Tedder, R. S., Thompson, A. A. R., Thomson, E. C., Thwaites, R. S., Turtle, L. C. W., Zambon, M., Carson, G., Data Analysis Team, Drake, T. M., Fairfield, C. J., Knight, S. R., Mclean, K. A., Murphy, D., Norman, L., Pius, R., Shaw, C. A., Data Architecture Team, Connor, M., Dalton, J., Gamble, C., Girvan, M., Halpin, S., Harrison, J., Jackson, C., Marsh, L., Roberts, S., Saviciute, E., Knight, S., Lahnsteiner, E., Leeming, G., Norris, L., Scott-Brown, J., Tait, S., Wham, M., Data Analysis and Management Team, Lee, J., Plotkin, D., Keating, S., Project Administration Team, Donegan, C., Spencer, R. G., Project Management Team, Donohue, C., Hardwick, H., The 23andMe COVID-19 Team, Shelton, J. F., Shastri, A. J., Ye, C., Weldon, C. H., Filshtein-Sonmez, T., Coker, D., Symons, A., Esparza-Gordillo, J., Aslibekyan, S., Auton, A., Yang, J., Ponting, C. P., Wilson, J. F., Vitart, V., Abedalthagafi, M., Luchessi, A. D., Parra, E. J., Cruz, R., Carracedo, A., Fawkes, A., Murphy, L., Rowan, K., Pereira, A. C., Law, A., Fairfax, B., Hendry, S. C., & Baillie, J. K.
Affiliations
  • Centre for Inflammation Research (GB)
  • Western General Hospital (GB)
  • Westlake University (CN)
  • The University of Queensland (AU)
  • Centre for Human Genetics (GB)
  • University of Leeds (GB)
  • Genomics England (GB)
  • Roslin Institute (GB)
  • Queen's University (CA)
  • University of Liverpool (GB)
  • Queen Mary University of London (GB)
  • Chinese University of Hong Kong (HK)
  • University College London (GB)
  • Imperial College Healthcare NHS Trust (GB)
  • Royal Hospital for Children (GB)
  • Edinburgh Royal Infirmary (GB)
  • Instituto de Salud Carlos III (ES)
  • Universidad de Cantabria (ES)
  • Hammersmith Hospital (GB)
  • St Thomas' Hospital (GB)
  • James Cook University Hospital (GB)
  • Barts Health NHS Trust (GB)
  • Royal Stoke University Hospital (GB)
  • North Middlesex University Hospital NHS Trust (GB)
  • Poole Hospital (GB)
  • King's College Hospital (GB)
  • Princess Alexandra Hospital (GB)
  • Russells Hall Hospital (GB)
  • Royal Liverpool University Hospital (GB)
  • John Radcliffe Hospital (GB)
  • Addenbrooke's Hospital (GB)
  • University Hospital Crosshouse (GB)
  • St George's Hospital (GB)
  • King George Hospital (GB)
  • Royal Berkshire NHS Foundation Trust (GB)
  • Instituto de Investigación Marqués de Valdecilla (ES)
  • Kingston Hospital (GB)
  • Queen Alexandra Hospital (GB)
  • Royal Gwent Hospital (GB)
  • Royal Blackburn Teaching Hospital (GB)
  • Stepping Hill Hospital (GB)
  • Royal Surrey County Hospital (GB)
  • Northumbria Healthcare NHS Foundation Trust (GB)
  • West Cumberland Hospital (GB)
  • Countess of Chester Hospital (GB)
  • Pinderfields Hospital (GB)
  • Ninewells Hospital (GB)
  • Croydon University Hospital (GB)
  • Morriston Hospital (GB)
  • Queen Elizabeth University Hospital (GB)
  • Airedale General Hospital (GB)
  • Broomfield Hospital (GB)
  • Heartlands Hospital (GB)
  • Royal Sussex County Hospital (GB)
  • York Hospital (GB)
  • Queen Elizabeth Hospital Birmingham (GB)
  • Royal Glamorgan Hospital (GB)
  • Barnet Hospital (GB)
  • Wythenshawe Hospital (GB)
  • Medway Maritime Hospital (GB)
  • Royal Oldham Hospital (GB)
  • North Devon District Hospital (GB)
  • Chesterfield Royal Hospital NHS Foundation Trust (GB)
  • Aberdeen Royal Infirmary (GB)
  • Royal Devon and Exeter Hospital (GB)
  • Glasgow Royal Infirmary (GB)
  • Blackpool Victoria Hospital (GB)
  • Southampton General Hospital (GB)
  • Derriford Hospital (GB)
  • East Surrey Hospital (GB)
  • Royal Alexandra Hospital (GB)
  • Leeds General Infirmary (GB)
  • Bedford Hospital (GB)
  • Musgrove Park Hospital (GB)
  • Ormskirk District General Hospital (GB)
  • Maidstone and Tunbridge Wells NHS Trust (GB)
  • North Manchester General Hospital (GB)
  • Royal Victoria Infirmary (GB)
  • William Harvey Hospital (GB)
  • Hull Royal Infirmary (GB)
  • Manchester Royal Infirmary (GB)
  • Royal Derby Hospital (GB)
  • Fairfield General Hospital (GB)
  • Norfolk and Norwich University Hospital (GB)
  • Good Hope Hospital (GB)
  • Milton Keynes Hospital (GB)
  • Bolton NHS Foundation Trust (GB)
  • Tameside Hospital (GB)
  • University Hospital of North Tees (GB)
  • Salford Royal Hospital (GB)
  • NIHR Great Ormond Street Hospital Biomedical Research Centre (GB)
  • Southmead Hospital (GB)
  • Wrexham Maelor Hospital (GB)
  • Arrowe Park Hospital (GB)
  • Bradford Royal Infirmary (GB)
  • Glan Clwyd Hospital (GB)
  • Royal Bournemouth Hospital (GB)
  • Bristol Royal Infirmary (GB)
  • University Hospital of North Durham (GB)
  • University College Hospital (GB)
  • Whittington Hospital (GB)
  • Sunderland Royal Hospital (GB)
  • Royal Hallamshire Hospital (GB)
  • Queen's Hospital (GB)
  • Royal Papworth Hospital (GB)
  • University Hospital of Wales (GB)
  • West Middlesex University Hospital (GB)
  • Withybush General Hospital (GB)
  • Stoke Mandeville Hospital (GB)
  • Royal Lancaster Infirmary (GB)
  • Basingstoke and North Hampshire Hospital (GB)
  • Worthing Hospital (GB)
  • Alexandra Hospital (GB)
  • Royal Cornwall Hospital (GB)
  • Watford General Hospital (GB)
  • Macclesfield District General Hospital (GB)
  • Rotherham General Hospital (GB)
  • Dumfries and Galloway Royal Infirmary (GB)
  • Craigavon Area Hospital (GB)
  • Prince Charles Hospital (GB)
  • Ysbyty Gwynedd Hospital NHS Trust (GB)
  • Royal Preston Hospital (GB)
  • Great Western Hospital (GB)
  • Lincoln County Hospital (GB)
  • Lister Hospital (GB)
  • Diana Princess of Wales Hospital (GB)
  • Southend Hospital (GB)
  • Calderdale Royal Hospital (GB)
  • Ealing Hospital (GB)
  • Royal United Hospital (GB)
  • St Mary's Hospital (GB)
  • Forth Valley Royal Hospital (GB)
  • Frimley Park Hospital (GB)
  • Chelsea and Westminster Hospital NHS Foundation Trust (GB)
  • Royal Brompton Hospital (GB)
  • Darent Valley Hospital (GB)
  • Wishaw General Hospital (GB)
  • University College Dublin (IE)
  • Walsall Manor Hospital (GB)
  • Princess Royal Hospital (GB)
  • Warrington Hospital (GB)
  • Royal Victoria Hospital (GB)
  • Cumberland Infirmary (GB)
  • Harefield Hospital (GB)
  • Salisbury District Hospital (GB)
  • Conquest Hospital (GB)
  • Leicester Royal Infirmary (GB)
  • Peterborough City Hospital (GB)
  • Colchester Hospital (GB)
  • Royal Shrewsbury Hospital (GB)
  • Monklands Hospital (GB)
  • New Cross Hospital (GB)
  • Hairmyres Hospital (GB)
  • Warwick Hospital (GB)
  • Sandwell General Hospital (GB)
  • Gloucestershire Royal Hospital (GB)
  • University Hospitals Coventry and Warwickshire NHS Trust (GB)
  • Torbay Hospital (GB)
  • Pilgrim Hospital (GB)
  • Prince Philip Hospital (GB)
  • Princess of Wales Hospital (GB)
  • Northampton General Hospital NHS Trust (GB)
  • James Paget University Hospital (GB)
  • Birmingham Children's Hospital (GB)
  • Northwick Park Hospital (GB)
  • Scunthorpe General Hospital (GB)
  • The Royal Free Hospital (GB)
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  • Hospital de Sant Pau (ES)
  • Universitat Autònoma de Barcelona (ES)
  • Hospital Universitario La Paz (ES)
  • Universidad de Salamanca (ES)
  • Universidad Francisco de Vitoria (ES)
  • Universidad Complutense de Madrid (ES)
  • BioCruces Health research Institute (ES)
  • Hospital Universitario Infantil San José (CO)
  • Universidade de Brasília (BR)
  • Universidad Simón Bolívar (CO)
  • Hospital Universitario Fundación Jiménez Díaz (ES)
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ORCID
Venue
Nature
Publisher
Springer Science and Business Media LLC
Published
2023-05-17
Volume
617
Issue
7962
Pages
764-768
ISSN
  • 0028-0836 (ISSN-L)
  • 1476-4687
DOI
10.1038/s41586-023-06034-3
PMC
PMC10208981
OpenAlex
W4376959222
Cited by
225
References
4
Citations by year
2026: 76 · 2025: 58 · 2024: 66 · 2023: 25
FWCI
8.30
Type
journal article
Language
en
Update notices
correction (2023-07-11)
Open access
Licence
Locations
  • Nature (publishedVersion, cc-by)
  • PubMed Central (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • Discovery Research Portal (University of Dundee) (publishedVersion, cc-by)
  • Hispana (publishedVersion)
  • Research Explorer (The University of Manchester) (publishedVersion, cc-by)
  • Edinburgh Research Explorer (University of Edinburgh) (publishedVersion, cc-by)
  • White Rose Research Online (University of Leeds, The University of Sheffield, University of York) (acceptedVersion)
  • Research Portal (Queen's University Belfast) (acceptedVersion, cc-by)
  • Brunel University Research Archive (BURA) (Brunel University London) (submittedVersion, cc-by)
  • Institutional Repository of the University of Granada (University of Granada) (submittedVersion, cc-by)
  • DIGITAL.CSIC (Spanish National Research Council (CSIC)) (submittedVersion, cc-by)
  • UCL Discovery (University College London) (submittedVersion, cc-by)
  • idUS (Universidad de Sevilla) (submittedVersion, cc-by)
  • University of Birmingham Research Portal (University of Birmingham) (submittedVersion, cc-by)
  • Queen Mary Research Online (Queen Mary University of London) (submittedVersion, cc-by)
  • UCrea (University of Cantabria) (submittedVersion, cc-by)
  • RUNA. Repositorio Institucional del Sergas (Servicio Gallego de Salud (SERGAS)) (submittedVersion, cc-by)
  • Spiral (Imperial College London) (submittedVersion, cc-by)
  • Apollo (University of Cambridge) (submittedVersion, cc-by)
  • Digital Commons-TMC (Texas Medical Center) (submittedVersion)
  • RePEc: Research Papers in Economics (submittedVersion)
  • Oxford University Research Archive (ORA) (University of Oxford) (submittedVersion)
  • Repositori digital de la UPF (Universitat Pompeu Fabra) (submittedVersion, cc-by)
  • Apollo (University of Cambridge) (submittedVersion)
Funders
  • Wellcome Trust (MR/R026408/1, 201488/Z/16/Z, 201488, 204979/Z/16/Z, 204979, 223164, 223164/Z/21/Z)
  • LifeArc
  • UK Research and Innovation (MC_PC_1905, MC_PC_20004, MC_PC_19025, BBS/E/D/20002172, MC_PC_20029, MR/R026408/1, MRNO2995X/1)
  • National Institute for Health and Care Research (NIHR201385, NF-SI-0513-10150, COVID-19)
  • British Heart Foundation
  • Department of Health and Social Care (COVID-19)
  • Diabetes UK
  • Research Councils UK (MR/R026408/1, BBS/E/D/10002070, BBS/E/D/20002172, BBS/E/D/30002275)
  • European Commission (34,35, COVID-19)
  • Intensive Care Society
  • Public Health England
  • Conselho Nacional de Desenvolvimento Científico e Tecnológico (309173/2019-1)
  • Northwest Regional Development Agency
  • Medical Research Council (MR/V001329/1, MR/R026408/1, MC_UU_00007/10, MC_PC_20029, MC_PC_19025, MC_UU_00007/15, COVID-19, MRNO2995X/1)
  • Biotechnology and Biological Sciences Research Council (BBS/E/RL/230002A, BBS/E/D/20002172, /30002275, BBS/E/D/30002275, BBS/E/D/10002070)
  • Instituto de Salud Carlos III (COV20/00622, PI20/00876)
  • European Regional Development Fund (COV20_00622, CGIEU0000219140, PI20/00876)
  • Health Data Research UK
Topics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetic Associations and Epidemiology
  • Health Sciences › Medicine › Infectious Diseases › COVID-19 Clinical Research Studies
  • Health Sciences › Medicine › Infectious Diseases › SARS-CoV-2 and COVID-19 Research
Keywords
critical COVID-19, host genetics, genetic variants, Mendelian randomization, druggable targets, JAK1, PDE4A, SLC2A5, TMPRSS2, RAB2A, immunometabolism, monocyte/macrophage activation, endothelial permeability, inflammatory signalling, scourge
MeSH terms
Whole Genome Sequencing, COVID-19, Adenylate Kinase, Genotype, Humans, Monocytes, Phenotype, Serine Endopeptidases, Genetic Variation, Critical Illness, Genetic Predisposition to Disease, rab GTP-Binding Proteins, rab2 GTP-Binding Protein, Glucose Transporter Type 5, Janus Kinase 1, Cyclic Nucleotide Phosphodiesterases, Type 4, Genome-Wide Association Study, Transcriptome, Genotyping Techniques

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