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Nature. 2023; 617(7962): 764–768.
Published online 2023 May 17. doi: 10.1038/s41586-023-06034-3
PMCID: PMC10208981
PMID: 37198478

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

Associated Data

Supplementary Materials
Data Availability Statement

Abstract

Critical illness in COVID-19 is an extreme and clinically homogeneous disease phenotype that we have previously shown1 to be highly efficient for discovery of genetic associations2. 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 group3. 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).

Subject terms: Genome-wide association studies, Genetics research, SARS-CoV-2, Viral infection

An analysis of 24,202 critical cases of COVID-19 identifies potentially druggable targets in 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).

Main

The design of the GenOMICC study and the rationale for focusing on critical illness has been previously described1,2. In brief, patients with confirmed COVID-19 requiring continuous cardiorespiratory monitoring or organ support (a generalizable definition for critical illness) were recruited in 2020–2022. We first performed ancestry-specific GWAS analyses according to the methods that we described previously1,2. Using the results of these GWAS analyses, previously reported results obtained using GenOMICC participants with whole-genome sequencing data2 and data from GenOMICC Brazil, we performed trans-ancestry and -platform meta-analyses within the GenOMICC study for a critically ill COVID-19 phenotype and a hospitalized COVID-19 phenotype (Extended Data Fig. Fig.1).1). The results of these GenOMICC-only meta-analyses are presented for both critically ill and hospitalized phenotypes (Table (Table11 and Extended Data Fig. Fig.2).2). To put these results into the context of existing knowledge, we performed comprehensive meta-analyses, drawing on further GWAS results, including data shared by the SCOURGE consortium and published data from the COVID-19 Human Genetics Initiative (HGIv6, 2021)4. The characteristics of the contributing studies are summarized in Supplementary Tables 13 and 14 for the critically ill and hospitalized phenotypes, with further details on each study provided in the Supplementary Information. We used a mathematical subtraction approach, as done in our previous work2, to remove signals of previous GenOMICC releases from HGIv6, yielding an independent dataset.

Table 1

Genome-wide significant associations with critical COVID-19, listing independent lead variants

Chr:pos(b38)rsidEAORORCIPPcondNearest geneCitation
1:9067157rs2478868A0.920.9–0.951.5 × 10−10SLC2A5GenOMICCnew
1:64948270rs12046291A1.11.07–1.135.1 × 10−11JAK1GenOMICCnew
1:77501822rs71658797A1.11.09–1.182.8 × 10−9AK5GenOMICCnew
1:155066988rs114301457T2.41.81–3.181.5 × 10−9EFNA4GenOMICC2
1:155175305rs7528026A1.31.25–1.441.5 × 10−15TRIM46GenOMICC2
1:155197995rs41264915A1.21.17–1.267.6 × 10−24THBS3HGI18
1:155278322rs11264349A0.940.92–0.977.3 × 10−53.9 × 10−13HCN3GenOMICCnew
2:60480453rs1123573A1.11.09–1.151 × 10−14BCL11AGenOMICC2
3:45796521rs2271616T1.21.14–1.231.1 × 10−16SLC6A20HGI18
3:45818159rs17713054*A21.96–2.137.7 × 10−254LZTFL1SCGG19
3:45873093rs35482426CTT0.530.5–0.576.1 × 10−91LZTFL1SCGG19
3:101790631rs11706494A1.11.05–1.119.4 × 10−9NXPE3GenOMICCnew
3:146522652rs343314T1.21.09–1.214.6 × 10−8PLSCR1GenOMICC2
4:25446871rs7664615A1.11.07–1.141.5 × 10−8ANAPC4GenOMICCnew
4:105673359rs72670002A1.11.09–1.214.4 × 10−8ARHGEF38GenOMICCnew
4:167824478rs1073165A1.11.05–1.111.1 × 10−9GenOMICCnew
5:132422622rs2269821A1.11.08–1.163 × 10−10IRF1-AS1GenOMICC2
6:31153455rs111837807T0.80.77–0.848.6 × 10−26CCHCR1GenOMICC1
6:31571991rs2071590A1.11.06–1.113.1 × 10−10LTAGenOMICCnew
6:32702687rs2858305T0.930.9–0.952.1 × 10−9HLA-DQA1GenOMICC2
6:41522644rs41435745C1.41.31–1.511.5 × 10−20FOXP4HGI18
7:75623396rs1179620T0.920.9–0.952.3 × 10−9HIP1GenOMICCnew
7:100032719rs2897075T1.11.06–1.118.9 × 10−11ZKSCAN1GenOMICCnew
8:60532539rs13276831T1.11.05–1.11.7 × 10−8RAB2AGenOMICCnew
9:21206606rs28368148C0.590.49–0.75.3 × 10−9IFNA10GenOMICC2
9:33425186rs60840586G1.11.07–1.149.7 × 10−9AQP3SCOURGE20
9:133271182rs879055593T1.11.1–1.161 × 10−16ABOSCGG19
10:79946568rs721917A0.930.9–0.957.6 × 10−9SFTPDHGI4
11:1219991rs35705950T0.860.82–0.893.8 × 10−14MUC5BHGI4
11:34482745rs61882275A0.880.86–0.911 × 10−22ELF5GenOMICC2
12:112919637rs2660A1.11.08–1.132.8 × 10−15OAS1GenOMICC1
12:132481571rs11614702A1.11.08–1.132.1 × 10−16FBRSL1GenOMICC2
13:112881427rs12585036T1.21.12–1.189.6 × 10−22ATP11AGenOMICC2
16:89196249rs117169628A1.21.12–1.22.6 × 10−16SLC22A31GenOMICC2
17:40003082rs12941811T0.930.91–0.951.1 × 10−9PSMD3GenOMICCnew
17:46085231rs8080583A0.890.86–0.911.8 × 10−16KANSL121
17:49863303rs77534576T1.31.24–1.438.7 × 10−16TAC421
19:4717660rs12610495A0.80.77–0.829.1 × 10−51DPP9GenOMICC1
19:10352442rs34536443C1.51.39–1.612.2 × 10−28TYK2GenOMICC1
19:10414696rs142770866A1.21.19–1.39.4 × 10−21PDE4AGenOMICCnew
19:48702915rs516246T0.90.88–0.931.4 × 10−15FUT2GenOMICC2
19:50374423rs35463555A1.11.07–1.131.9 × 10−13NR1H2HGI4
20:6489447rs2326788A0.930.9–0.951.5 × 10−8CASC20GenOMICCnew
21:33229937rs188401375C0.740.66–0.843.1 × 10−64.7 × 10−9IFNAR2GenOMICC1
21:33237639rs9636867A0.830.81–0.855.2 × 10−48IFNAR2GenOMICC1
21:33287378rs8178521T1.11.1–1.176.2 × 10−15IL10RBGenOMICC2
21:33980963rs76608815T1.21.14–1.237.4 × 10−17ATP5POGenOMICC2
21:41479527rs915823A1.11.06–1.132.1 × 10−9TMPRSS2GenOMICCnew
X:15523993rs35697037A11.03–1.066.8 × 10−9ACE2HGI4

Chr:pos(b38), chromosome and position on human genome build 38; rsid, lead variant rsid; EA, effect allele; OR, odds ratio; ORCI, odds ratio confidence interval; P, P value against null hypothesis of odds ratio of one; Pcond, P value in conditional analysis in variants with P > 5 × 10−8; nearest gene, the nearest or most plausible nearby gene; citation, the first report of genome-wide significant association. ‘GenOMICCnew’ indicates findings presented in this Article. Additional details are provided in Supplementary Table 15 and full results of conditional analysis are provided in Supplementary Table 16. Supplementary Table 17 contains details of lead variants from the analysis of the hospitalized phenotype.

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Pipeline of meta-analysis and post-GWAS analyses.

Red border indicates that the data is only available for the hospitalized phenotype, while a black border indicates that the analysis was performed for the critical illness phenotype.

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Miami plots.

Meta-analysis results are shown for a) critical and b) hospitalized phenotypes. In each plot results obtained using all cohorts are shown at the top and using GenOMICC data only at the bottom. Independent lead variants in the analyses of all cohorts are annotated with associated genes. Genome-wide significant associations that have not been previously reported are indicated in bold.

As no replication cohorts exist for these meta-analyses, we used the heterogeneity across studies to assess the reliability of individual findings (Supplementary Table 15). Owing to the unusually extreme phenotype in the GenOMICC study, some heterogeneity is expected for the strongest associations when compared with studies with more permissive inclusion criteria. Importantly, significant heterogeneity was not detected for any of the findings that we report here (Supplementary Table 15). Comparing effect estimates between studies using a regression approach that takes into account estimation errors (Methods), we detected systematic differences in effect sizes between studies (Extended Data Fig. Fig.3).3). For example, effects for the HGI critical illness phenotype (which was designed to parallel the GenOMICC inclusion criteria) are smaller than those obtained using prospective recruitment in GenOMICC by a factor of 0.68. As the effect sizes in GenOMICC are consistently larger than other studies, and GenOMICC contributes a disproportionately large signal to meta-analyses of both critical and hospitalized phenotypes (Extended Data Fig. Fig.4),4), between-study heterogeneity is likely to reflect the careful case ascertainment and extreme phenotype in GenOMICC compared with other studies.

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Comparison of effect size estimates.

GenOMICC is compared with the critical and hospitalized phenotype definitions in the SCOURGE, 23andMe, and HGI analyses. The black line indicates the best linear fit, given by the equation in each plot, obtained using Orthogonal Distance Regression to account for estimation errors in both sets of effects in the comparison.

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Study weightings for (a) critical and (b) hospitalized COVID-19.

Mean +/− standard deviation of weights assigned to each data source in meta-analyses for all significant SNPs.

We found 49 common genetic associations with critical COVID-19 meeting our criteria for genome-wide significance in the absence of heterogeneity (Extended Data Fig. Fig.22 and Table Table1).1). Findings from previous reports were consistently replicated (Extended Data Table Table2).2). Conditional analysis revealed two additional lead variants (Table (Table1)1) and statistical fine-mapping provided credible sets of putative causal variants for a majority of lead variants (Supplementary Figs. 2744 and Supplementary Table 5). Gene-level analyses found 196 significantly associated genes at a Bonferroni-corrected threshold (Supplementary Table 10). There were no genome-wide significant differences in the effects between sexes in a sex-stratified meta-analysis using a subset of cohorts (Supplementary Fig. 1).

Extended Data Table 2

Replication table

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Each report of genome-wide significant associations with severe COVID-19 is shown, with associations that were first reported by the GenOMICC consortium are highlighted in blue.

Therapeutic implications

Our analysis is limited to common variants that are detectable on genotyping arrays and imputation panels. Although most lead variants are not directly causal, in some cases, they highlight molecular mechanisms that alter clinical outcomes in COVID-19, and may have direct therapeutic relevance. To investigate the disease mechanisms, we first quantified the effect of inferred gene expression on critical illness in three relevant tissue/cell types. Many of the genes that we have found to be implicated in critical COVID-19 (refs. 1,2) are highly expressed in the monocyte–macrophage system, which has poor coverage in existing expression quantitative trait loci (eQTL) datasets. For this reason, we constructed a new TWAS model in primary monocytes obtained from 176 individuals (Methods). We found significant associations after Bonferroni correction between critical COVID-19 and predicted gene expression in lung (33), blood (21), monocyte (37) and all-tissue (107) meta-analysis (Supplementary Table 2 and Supplementary Table 11). We extended these findings using generalized summary-level data Mendelian randomization (GSMR) for RNA expression (Fig. (Fig.2,2, Extended Data Table Table1,1, Supplementary Figs. 1118 and Supplementary Table 4).

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GSMR effect sizes.

a,b, The predicted effect of change in protein concentration (a) and gene expression (b) on the risk of critical COVID-19 is shown for proteins and genes significantly linked to critical COVID-19 by GSMR (false-discovery rate (FDR) < 0.01). The bars show 95% confidence intervals.

Extended Data Table 1

pQTL GSMR results table

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Proteins significantly linked to COVID-19 severity (false discovery rate (FDR) <0.05). Exposure: protein name; b: effect-size estimate of the protein on COVID-19 severity from GSMR; se: standard-error of b; p: p-value of the GSMR result; N: Number of independent SNPs included in the analysis. : indicates proteins with GSMR evidence previously presented in Kousathanas et al.2.

In parallel, we assessed the effect of genetically determined variation in circulating protein levels on the critical illness phenotype using GSMR5. We identified 15 unique proteins linked to critical illness, as summarized in Extended Data Table Table11 (Supplementary Table 3). Of the significant results, we found causal evidence implicating five new proteins in comparison to our previous GSMR analysis2: QSOX2, CREB3L4, myeloperoxidase (MPO), ADAMTS13 and mannose-binding lectin-2 (MBL2) (Supplementary Fig. 10). These include well-studied biomarkers and potential drug targets in sepsis—the innate immune pattern recognition receptor MBL2 and the neutrophil effector enzyme MPO. ADAMTS13 modulates von Willebrand-factor-mediated platelet thrombus formation and may have a role in the hypercoagulable state in critical COVID-19 (Extended Data Fig. Fig.55).

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Cartoon showing postulated roles for genes and mediators implicated in the pathogenesis of critical COVID-19 by GenOMICC GWAS, TWAS and Mendelian randomization.

Postulated roles for genetic variants are shown in a highly simplified format to illustrate potential roles in pathogenesis, with the shaded background indicating the hypothetical impact of the host immune response over time17. Host immune processes are divided into those that are thought to play a role in controlling viral replication early in disease (orange section, showing “adaptive” response), and those implicated in driving hypoxaemic respiratory failure later in disease (green section, showing “maladaptive” response). Bold type gene names indicate a higher level of confidence in both the gene identification and the biological role.

Three genes containing non-synonymous protein-coding changes associated with severe disease were also found to have significant effects from differential gene expression: SLC22A31 (ref. 2) (Fig. (Fig.1),1), SFTPD4 (Fig. (Fig.1)1) and TKY2 (ref. 1) (Extended Data Fig. Fig.6).6). Further biological and clinical research will be required to dissect the genetic evidence at these loci. In the example of TYK2, there is now a therapeutic test of the genetic predictions. Our previous report of association between higher expression and critical illness1 led directly to the inclusion of a new drug, baricitinib, in a large clinical trial; the result demonstrated a clear therapeutic benefit3. This therapeutic signal is consistent across multiple trials, providing the first proof-of-concept for drug target identification using genetics in critical illness and infectious disease.

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Functional genomics analyses for SLC22A31 and SFTPD.

a, Effect-size plot for the effect of multiple variants on SLC22A31 expression (eQTLgen, x axis) against increasing susceptibility to critical COVID-19 (βxy = 0.11; Pxy = 1.3 × 10−9). The colour shows linkage disequilibrium (LD) with the missense variant rs117169628. b, Three cartoon views of an AlphaFold22 model of putative solute carrier family 22 member 31 (SLC22A31; UniProtKB: A6NKX4). The side chains of Pro474 and interacting amino acids are shown as connected spheres. A putative channel for small-molecule transport across the cell membrane is indicated by a dashed circle. Pro474 is predicted to be located in the transmembrane helix and point towards a putative transport pathway of a small molecule. The risk variant, P474L (Ala at rs117169628) would be expected to introduce more flexibility to the transmembrane helix and might therefore affect the transport properties of SLC22A31. Pro474 is predicted to be in a tightly packed environment, and may therefore affect the folding of SLC22A31. c, Effect-size plot for effect of multiple variants on SFTPD expression (eQTLgen, x axis) against increasing susceptibility to critical COVID-19 (βxy = 0.16; Pxy = 9.7 × 10−6). Colour shows linkage disequilibrium with the missense variant rs721917. d, Three cartoon views of an AlphaFold22 model of pulmonary surfactant-associated protein D (SFTPD; UniProtKB: P35247). The side chain of the variant Met31 is shown as connected spheres. Met31 is predicted to be located in the secondary-structure-lacking region of SFTPD. In the diagram on the right, oxygen and nitrogen atoms are coloured red and blue respectively, and the sulfur atom is coloured yellow.

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Functional genomics analyses for TYK2.

(a) Effect size plot for effect of multiple variants on TYK2 expression (eQTLgen, x-axis) against increasing susceptibility to critical COVID-19 (βxy = 0.53; Pxy=1.2×1023). Colour shows linkage disequilibrium (LD) with the missense variant rs34536443. (b) Crystal structure of TYK2 kinase domain (Protein Data Bank ID 4GVJ39) in two views that differ by a 45° rotation around a horizontal axis. The side chain of P1104 is shown as connected spheres with a nitrogen atom coloured in blue. Carbon, oxygen, nitrogen and phosphorus atoms of ATP are shown as magenta, red, blue and orange connected spheres, respectively. The N-terminal region of the kinase domain is not shown in the second view for clarity. The right-most panel shows a close view of P1104 and neighbouring residues with their side chains shown as sticks. Numbering of residues corresponds to UniProtKB entry P29597. P1104 is in the catalytic kinase domain and proximal to the ATP-binding site; TYK2 P1104A is catalytically impaired40.

To assess the immediate therapeutic use of our results for repurposing of existing compounds, we considered the drug therapies under consideration by the UK COVID-19 Therapeutic Advisory Panel (UK-CTAP), a national independent review group supported by an expert due-diligence panel6. Consistent evidence from gene-level GWAS (Supplementary Table 6 and Supplementary Table 10) and post-GWAS analyses was identified for several licensed compounds (Supplementary Table 12). For example, we found an association in another gene encoding a protein that is inhibited by baricitinib and other JAK inhibitors—the intracellular signalling kinase, JAK1, which is stimulated by numerous cytokines including type I interferons and IL-6. Mendelian randomization analysis of RNA expression revealed a significant positive association between the expression of the gene encoding a canonical inflammatory cytokine, tumour necrosis factor (TNF), and severe disease (Fig. (Fig.2).2). This suggests that inhibition of TNF signalling may be an effective therapy in severe COVID-19.

Our additional expression data in monocytes reveal a marked tissue-specific effect on expression of PDE4A. This phosphodiesterase regulates the production of multiple inflammatory cytokines by myeloid cells. In contrast to the negative correlations seen in the lungs and blood, we show that a genetic tendency for higher expression of PDE4A in monocytes is associated with critical COVID-19 (Supplementary Table 11). Inhibition of PDE4A by several existing drugs is under investigation in multiple inflammatory diseases7, reduces pulmonary endothelial permeability8 and appears to be safe in small clinical trials in patients with COVID-19.

The postulated biological role of genes associated with critical COVID-19 in GWAS, TWAS and GSMR results is shown in Extended Data Fig. Fig.5,5, which highlights the preponderance of genes with expression or functions in the mononuclear phagocyte system. This includes SLC2A5, encoding the GLUT5 fructose transporter, which is strongly inducible in primary macrophages in response to inflammatory stimulation9, and XCR1, a dendritic cell receptor with a critical role in cytotoxic T cell-mediated antiviral immunity10. NPNT, a significant meta-TWAS association in the genome-wide significant region on chromosome 4 (chr4:105673359; Supplementary Table 11), encodes a pulmonary basement membrane protein that may have a protective role in acute lung injury11.

Host–pathogen interaction

Our results also demonstrate the capacity of host genetics to reveal core mechanisms of disease. Multiple genes implicated in viral entry are associated with severe disease. In addition to ACE2, we detect a genome-wide significant association in TMPRSS2, a key host protease that facilitates viral entry that we have previously studied as a candidate gene12. This effect may be viral-lineage specific13. A strong GWAS association is seen in RAB2A (Table (Table1),1), with TWAS evidence suggesting that more expression of this gene is associated with worse disease (Supplementary Table 11). RAB2A is highly ranked in our previous meta-analysis by information content14 study of host genes implicated in SARS-CoV-2 interaction using in vitro and clinical data15, and is consistent with CRISPR screen data showing that RAB2A is required for viral replication16.

Although our focus on critical illness enhances discovery power (Extended Data Fig. Fig.4),4), it has the disadvantage of combining genetic signals for multiple stages in disease progression, including viral exposure, infection and replication, and development of inflammatory lung disease. From these data alone we cannot identify when in disease progression the causal effect is mediated, although clinical evidence helps to make some predictions17 (Extended Data Fig. Fig.5).5). As most cases included were recruited before vaccinations and treatments became available (Extended Data Fig. Fig.7),7), at present, our study does not have sufficient statistical power to dissect the genetic effects of treatments or vaccination. These effects may include the masking of true associations, or the detection of genetic effects mediated by vaccine or drug response, rather than COVID-19 susceptibility. However, the absence of divergent genetic effects between studies (Supplementary Figs. 25) or consistent changes in effect allele frequency among cases over time (Supplementary Figs. 4548) suggests that treatment and vaccination have not substantially affected the association between the specific variants that we report and the risk of critical illness.

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Steroid treatment and vaccination status.

Data are shown for a subset of GenOMICC cases who were also recruited to the ISARIC4C study in the UK.

As we performed a meta-analysis of multiple studies that may have slightly different definitions of the phenotype, effect sizes differ between studies (Supplementary Figs. 25). This, together with ancestry-specific effects1, may explain the heterogeneity in strong GWAS signals, such as the LZTFL1 signal in Table Table1.1. Different studies also have sets of variants that are not completely overlapping, so P values between variants in high linkage disequilibrium are more different than expected. Although most of the studies contain individuals from multiple ancestries, a large majority of the individuals are of European ancestry. In future research, there is a scientific and moral imperative to include the full diversity of human populations.

Together, these results deepen our understanding of the pathogenesis of critical COVID-19 and highlight new biological mechanisms of disease, several of which have immediate potential for therapeutic targeting.

Methods

Hospitalization meta-analysis

The hospitalized phenotype includes patients who were hospitalized with a laboratory-confirmed SARS-Cov2 infection. In this analysis we included GenOMICC, GenOMICC Brazil, GenOMICC Saudi Arabia, ISARIC4C, HGIv6 B2 phenotype with subtraction of GenOMICC data, SCOURGE hospitalized versus population and mild cases, and 23andMe broad respiratory phenotype. A summary description of each analysis is given above, a table with the included studies can be found in Supplementary Table 14 and an extended description can be found in Supplementary Table 1.

Critical illness meta-analysis

The critically ill COVID-19 group included patients who were hospitalized owing to symptoms associated with laboratory-confirmed SARS-CoV-2 infection and who required respiratory support or whose cause of death was associated with COVID-19. In the critical illness analysis, we included GenOMICC, patients with critical illness from ISARIC4C, HGIv6 phenotype A2 with subtraction of GenOMICC data, SCOURGE severity grades 3 and 4 versus population controls, and 23andMe respiratory support phenotype. A summary description of each analysis can be found above, a table with the included studies can be found in Supplementary Table 13 and an extended description can be found in Supplementary Table 1.

Meta-analyses

All meta-analyses across studies were performed using a fixed-effect inverse-variance weighting method and control for population stratification in the METAL software23. Allele frequency was calculated as the average frequency across studies with the METAL option AVERAGEFREQ. P values for heterogeneity in effect sizes between studies were calculated using a Cochran’s Q-test implemented in METAL. For variants in the same position with different REF and ALT alleles across studies, the GenoMICC variant in the European population was selected and the rest were removed. Finally, variants with switched ALT and REF alleles between HGIv6 and GenOMICC were also removed on the basis of differences in allele frequency of the alternative allele. Variants were annotated to the closest genes using dbsnp v.b151 GRCh38p7 and bionrRt R package (v.2.46.3)24. As each single-nucleotide polymorphism (SNP) of the meta-analysis can be present in different subsets of cohorts, there may be large differences in P values in SNPs with a high level of linkage disequilibrium, which may have an effect on downstream analyses. For this reason, variants that were not present in one of the three biggest studies—GenOMICC European ancestry, HGIv6 or SCOURGE—were filtered out from post-GWAS analysis.

Conditional analysis

We performed a step-wise conditional analysis to find independent signals. As European-specific data are not available in some cohorts but European ancestry is largely predominant (87.2% of cases with critical illness), we performed the conditional analysis using a European reference panel and the meta-analysis results of the whole cohort. To perform the conditional analysis, we used the GCTA (v.1.9.3) --cojo-slct function25. The parameters for the function were P = 5 × 10−8, a distance of 10,000 kb and a co-linear threshold of 0.9 (ref. 26), and the reference population for the conditional analysis was individuals of European ancestry with whole-genome sequence available in the GenOMICC study and whole genomes from the 100,000 Genomics England project2.

Credible set fine-mapping

We performed fine-mapping using the SuSiE model27 to construct credible sets for the independent signals identified using conditional analysis. As for conditional analysis, we used a European reference panel and the meta-analysis results of the whole critical illness cohort. We performed analyses in 1 Mb windows centred on the lead variants identified through conditional analysis. In cases in which windows for multiple variants overlapped, they were joined into a single window. For each window, we fitted the SuSiE summary statistics model setting the expected number of independent signals to the number of identified though conditional analysis. Models for three windows did not converge in 500 iterations and have been excluded. As a reference, we used the publically available linkage disequilibrium information for non-Finish Europeans from the GNOMAD 2.1.1 release. Full data for all variants included in credible sets are included in Supplementary Table 5.

Gene-level analysis

We performed an analysis summarizing the genetic associations at the gene level using the mBAT-combo method28. We used the COVID ‘all critical cohorts’ meta-analysis (GenOMICC, HGIv6 phenotype A2, SCOURGE and 23andMe) summary statistics. As this is a trans-ethnic meta-analysis, we used a mixed ancestry linkage disequilibrium reference panel, consisting of 3,202 1000 Genomes phase 3 samples. We considered a list of protein-coding genes with unique ensemble gene ID based on the release from GENCODE (v.40) for hg38, which can be found on the mBAT-combo website (https://yanglab.westlake.edu.cn/software/gcta/#mBAT-combo). A gene region was taken to span 50 kb upstream to 50 kb downstream of the gene’s untranslated regions.

Sex-stratified meta-analysis

To test for differences in genetic effects, we performed sex-stratified GWAS of the COVID-19 critical illness phenotype in the European ancestry GenOMICC WGS and genotyped cohorts and SCOURGE. We then performed a meta-analysis for each sex following the same methods as for the main analysis. We tested for differences in effects between the meta-analyses of the two sexes following previously described methods29.

Mendelian randomization

GSMR5 was performed. We used the COVID ‘all critical cohorts’ meta-analysis (GenOMICC, HGIv6 phenotype A2, SCOURGE and 23andMe) as the outcome, protein expression quantitative-trait loci (pQTLs) from ref. 30 and RNA expression quantitative-trait loci (eQTLs) from eQTLgen31 (2019-12-23 data release) as exposures, and 10,000 individuals of European ancestry randomly sampled from the UK Biobank as the linkage disequilibrium reference cohort (50,000 for linkage disequilibrium to missense variant plots). GSMR was performed for all exposures for which we were able to identify two or more suitable SNPs. SNPs were chosen to meet the following criteria: (1) SNP to exposure association P < 5 × 10−8; (2) linkage disequilibrium clumping lead SNPs only (±1 Mb, r2 < 0.05); (3) SNP not removed by HEIDI-outlier filtering (for the removal of SNPs with evidence of horizontal pleiotropy) at the default threshold value of 0.01. eQTLGen effect sizes and standard errors were estimated as described in supplementary note 2 of ref. 32. We considered as significant those exposure–outcome pairs with FDR < 0.05.

TWAS analysis

To perform TWAS analysis in GTExv8 tissues33, we used the MetaXcan framework and the GTExv8 eQTL and sQTL MASHR-M models available for download online (http://predictdb.org/) and the ‘all critical cohorts’ meta-analysis. We first calculated individual TWAS for whole blood and lungs using the S-PrediXcan function34,35. We next performed a metaTWAS including data from all tissues to increase the statistical power using s-MultiXcan36. We applied Bonferroni correction to the results to choose significant genes and introns for each analysis.

Monocyte gene expression

To detect eQTLs, untreated primary monocytes were prepared from 174 healthy individuals of Northern European (British) ancestry recruited through the Oxford Biobank. Poly(A) RNA was paired-end 100 bp sequenced in the Oxford Genome Centre using the Illumina HiSeq-4000 machines (median = 47,735,438 reads per sample). Reads were aligned to CRGh38/hg38 using HISAT2 with the default parameters. High mapping quality reads were selected on the basis of MAPQ score using bamtools. Duplicate reads were marked and removed using picard (v.1.105). Samtools was used to pass through the mapped reads and calculate statistics. Read count information was generated using HTSeq and normalized using DESeq2. Sample contamination and swaps were detected by comparing the imputed SNP-array genotypes with genotypes called from RNA-seq using verifyBamID. Genotyping was performed with Illumina HumanOmniExpress with coverage of 733,202 separate markers. Genotypes were pre-phased with SHAPEIT2, and missing genotypes were imputed with PBWT. Poly(A) RNA was paired-end sequenced at the Oxford Genome Centre using the Illumina HiSeq-4000 machines. vcftools (v0.1.12b) was applied on genetic variation data in the form of variant call format (VCF) files to filter out indels and SNPs with a minor allele frequency of less than 0.04.

TWAS analysis for monocyte data was performed using genotyping and monocyte RNA-sequencing data from 174 individuals. Using a region of 500 kb around each gene, we calculated gene expression models using the Fusion R package37. For each gene, three models were calculated adding as covariates the two first principal components calculated from the genotype: blup, elastic networks and lasso. The model with a better r2 between predicted and measured expression in a fivefold cross-validation was chosen. Then SNP genetic heritability was calculated for the 500 kb region for each gene and those genes with a nominal significant SNP heritability estimate (P ≤ 0.01) were chosen for the TWAS analysis. Summary statistics for the ‘all critical cohorts’ meta-analysis and the best model for each gene were then used to perform the TWAS.

Colocalization

Significant genes in the TWAS and metaTWAS were selected for a colocalization analysis using the coloc R package. The lead SNPs and a region of 200 Mb around the gene were used to colocalize with significant genes in the TWAS with eQTL summary statistics data on the region from GTExv8 lung, GTExv8 whole blood, eQTLgen or monocyte eqtl. As in our previous analysis2, we first performed a sensitivity analysis of the posterior probability of colocalization (PPH4) on the prior probability of colocalization (P12), going from P12 = 10−8 to P12 = 10−4, with the default threshold being P12 = 10−5. eQTL signal and GWAS signals were deemed to colocalize if these two criteria were met: (1) at P12 = 5 × 10−5 the probability of colocalization PPH4 > 0.5; and (2) at P12 = 10−5 the probability of independent signal (PPH3) was not the main hypothesis (PPH3 < 0.5). These criteria were chosen to allow eQTLs with weaker P values, owing to lack of power in GTEx v.8, to be colocalized with the signal when the main hypothesis using small priors was that there was not any signal in the eQTL data.

Effect comparison

We compared the estimates of effect sizes between the individual GWASs used in the meta-analysis, for all variants that were genome-wide significant in at least one of the individual GWASs. To this end, we regressed the effects obtained using critical illness and hospitalization in the SCOURGE and 23andMe cohorts, as well as the HGI meta-analyses on the effect estimates obtained using the GenOMICC cohort. To account for estimation errors present in both the dependent and independent variables of the regression we used orthogonal distance regression38.

Weight of studies

To calculate the weight of GenOMICC, we downloaded the leave-one-out data of HGIv7. As the meta-analysis is performed using a variance-weighted method, we can recover the variance for each SNP as v=1s.e.2, for the meta-analysis of all of the cohorts and for each one of the leave-one-out analysis. The total weight is wtot=1v and the weight leaving out a specific study is wloo=1vloo. The weight of a cohort is then wtotwloo. We calculated the weight for each the significant SNPs in our analysis for each study and normalized it using the total weight. Finally, we calculated the mean and s.d. from the significant SNPs for each cohort.

Forest plots

To compare effects between cohorts, we first performed a trans-ancestry meta-analysis for GenOMICC and 23andMe using METAL23. Then, we used the metagen and forest functions of the meta R package to produce forest plots for critical illness and hospitalization separately.

Reporting summary

Further information on research design is available in the Nature Portfolio Reporting Summary linked to this article.

Online content

Any methods, additional references, Nature Portfolio reporting summaries, source data, extended data, supplementary information, acknowledgements, peer review information; details of author contributions and competing interests; and statements of data and code availability are available at 10.1038/s41586-023-06034-3.

Supplementary information

Supplementary Information(12M, pdf)

Supplementary Sections 1–13, including Supplementary Figs. 1–54 and Supplementary Tables 7–17.

Peer Review File(1.9M, pdf)

Supplementary Table 1(16K, xlsx)

Description of the cohorts used in critical and hospitalized meta-analyses. Cohorts are divided by ancestry and genotyping method (whole-genome sequencing or microarray genotyping). In cohorts in which data are available, the median age with s.d. in parentheses, percentage of female cases, number of female and male cases and controls is shown. NA, data are not available for the cohort. Country of origin indicates the country in which individuals in the cohort were recruited; GenotypingPlatform indicates the array or WGS platform used for genotyping; reference indicates the reference of the publication (if the data have already been published). ‘In GenOMICC v2’ is an indication of whether the dataset was included in a previous GenOMICC paper1.

Supplementary Table 2(37K, xlsx)

Full results for colocalization and TWAS analyses in lungs, blood, monocytes and across multiple tissue types (metaTWAS). Colocalization results are reported between significant TWAS genes and eQTLs in GTExv8 in lungs, blood and monocytes, and in eqtlGEN. rsid indicates chromosome, position, reference and alternative alleles; gene.tested is the significant TWAS gene, and ensembl.id is the Ensembl ID corresponding to the significant gene. PP.H3.1e-5 is the posterior probability of independent signals with a prior for colocalization of 5 × 10−5, PP.H4.1e-5 is the posterior probability of colocalization with a prior of 5 × 10−5, PP.H3.5e-5 is the posterior probability of independent signals with a prior for colocalization of 5 × 10−5 and PP.H4.5e-5 is the probability of colocalization with a prior of 5 × 10−5. Colocalization was considered to be significant when PP.H3 was lower than 0.5 and PP.H4 was the highest posterior probability. Significant genes after Bonferroni correction in a TWAS meta-analysis of lungs, blood, monocytes and all tissues in GTExv8 and the ‘all critical cohorts’ GWAS. Ensembl ID, gene name, P value of the meta-analysis, number of SNPs used in to model gene expression in all tissues, mean z score for all tissues and its s.d. are shown.

Supplementary Table 3(32K, txt)

The full results from GSMR analysis for protein level. Exposure indicates the protein name used as exposure for the analysis, bxy is the effect size, se is the standard error, p is the P value of the analysis, nsnp is the number of SNPs used as instruments and multi_snp_based_heidi_outlier is the P value of the Heidi test.

Supplementary Table 4(406K, txt)

Full results from GSMR analysis for RNA-seq data from eQTLGEN. Exposure indicates the gene name used as exposure for the analysis, bxy is the effect size, se is the standard error, p is the P value of the analysis, nsnp is the number of SNPs used as instruments and multi_snp_based_heidi_outlier is the P value of the Heidi test.

Supplementary Table 5(104K, xlsx)

Table of variants in credible sets with 95% probability of containing the causal SNP. Credible set ID is the credible set index to which the variant belongs; posterior is the posterior probability of causality for the variant. Variant indicates chromosome, position, reference and alternative alleles; beta is the effect, beta.se is the error and P is the P value.

Supplementary Table 6(1.1M, xlsx)

The full results of the gene-level analysis performed using the mBAT-combo method. Gene indicates ensembl ID, gene_name indicates name of the gene, gene_p is the P value of the gene-level test, nsps is the number of SNPs used for the test, lead_snp is the chromosome, position, reference and alternative alleles for the SNPs with lowest P value in the region, and lead_snp_p indicates the P value of this lead SNP.

Acknowledgements

We thank the patients and their loved ones who volunteered to contribute to this study at one of the most difficult times in their lives, and the research staff in every intensive care unit who recruited patients at personal risk during the most extreme conditions ever witnessed in most hospitals. GenOMICC was funded by Sepsis Research (the Fiona Elizabeth Agnew Trust), the Intensive Care Society, a Wellcome Trust Senior Research Fellowship (to J.K.B., 223164/Z/21/Z), the Department of Health and Social Care (DHSC), Illumina, LifeArc, the Medical Research Council, UKRI, a BBSRC Institute Program Support Grant to the Roslin Institute (BBS/E/D/20002172, BBS/E/D/10002070 and BBS/E/D/30002275) and UKRI grants MC_PC_20004, MC_PC_19025, MC_PC_1905 and MRNO2995X/1. A.D.B. acknowledges funding from the Wellcome PhD training fellowship for clinicians (204979/Z/16/Z), the Edinburgh Clinical Academic Track (ECAT) programme. This research is supported in part by the Data and Connectivity National Core Study, led by Health Data Research UK in partnership with the Office for National Statistics and funded by UK Research and Innovation (grant MC_PC_20029). Laboratory work was funded by a Wellcome Intermediate Clinical Fellowship to B.F. (201488/Z/16/Z). We acknowledge the staff at NHS Digital, Public Health England and the Intensive Care National Audit and Research Centre who provided clinical data on the participants; and the National Institute for Healthcare Research Clinical Research Network (NIHR CRN) and the Chief Scientist’s Office (Scotland), who facilitate recruitment into research studies in NHS hospitals, and to the global ISARIC and InFACT consortia. GenOMICC genotype controls were obtained using UK Biobank Resource under project 788 funded by Roslin Institute Strategic Programme Grants from the BBSRC (BBS/E/D/10002070 and BBS/E/D/30002275) and Health Data Research UK (HDR-9004 and HDR-9003). UK Biobank data were used in the GSMR analyses presented here under project 66982. The UK Biobank was established by the Wellcome Trust medical charity, Medical Research Council, Department of Health, Scottish Government and the Northwest Regional Development Agency. It has also had funding from the Welsh Assembly Government, British Heart Foundation and Diabetes UK. The work of L.K. was supported by an RCUK Innovation Fellowship from the National Productivity Investment Fund (MR/R026408/1). J.Y. is supported by the Westlake Education Foundation. SCOURGE is funded by the Instituto de Salud Carlos III (COV20_00622 to A.C., PI20/00876 to C.F.), European Union (ERDF) ‘A way of making Europe’, Fundación Amancio Ortega, Banco de Santander (to A.C.), Cabildo Insular de Tenerife (CGIEU0000219140 ‘Apuestas científicas del ITER para colaborar en la lucha contra la COVID-19’ to C.F.) and Fundación Canaria Instituto de Investigación Sanitaria de Canarias (PIFIISC20/57 to C.F.). We also acknowledge the contribution of the Centro National de Genotipado (CEGEN) and Centro de Supercomputación de Galicia (CESGA) for funding this project by providing supercomputing infrastructures. A.D.L. is a recipient of fellowships from the National Council for Scientific and Technological Development (CNPq)-Brazil (309173/2019-1 and 201527/2020-0). We thank the members of the Banco Nacional de ADN and the GRA@CE cohort group; and the research participants and employees of 23andMe for making this work possible. A full list of contributors who have provided data that were collated in the HGI project, including previous iterations, is available online (https://www.covid19hg.org/acknowledgements).

Extended data figures and tables

Author contributions

E.P.-C., K. Rawlik, K.M., S.K., C.P.P., J.F.W., V.V., M.A., A.D.L., E.J.P., R.C., A.C., A.F., L.M., K. Rowan, A.C.P., A.L., S.C.H. and J.K.B. contributed to design. E.P.-C., K. Rawlik, A.D.B., T.Q., Y.W., I.N., G.A.M., M.Z., L.K., A.K., A.R., T.M., J.Y., A.L., B.F., S.C.H. and J.K.B. contributed to data analysis. E.P.-C., K. Rawlik, I.N., A.K., A.R., J.M., C.D.R., A.L., B.F. and S.C.H. contributed to bioinformatics. E.P.-C., K. Rawlik, I.N., G.A.M., M.Z., A.K., J.M., C.D.R., R.T., D. McAuley, A.N., M.G.S., B.F., S.C.H. and J.K.B. contributed to writing and reviewing the manuscript. I.N., F.G., W.O., K.M., S.K., D. Maslove, A.N., M.G.S., J.K., M.S.-H., C.S., C.H., P.H., L.L., D. McAuley, H.M., P.J.M.O., C.B., T.W., A.T., C.F., J.A.R., A.R.-M., P.L., C.P.P., A.F., L.M., K. Rowan, A.L., B.F. and S.C.H. contributed to oversight. F.G. and W.O. contributed to project management. F.G., W.O. and J.K.B. contributed to ethics and governance. K.M., A.F. and L.M. contributed to sample handling and sequencing. C.P.P., K. Rowan, S.C.H. and J.K.B. contributed to conception. C.P.P., J.F.W., V.V., M.A., A.D.L., E.J.P., R.C., A.C., K. Rowan and A.C.P. contributed to reviewing the manuscript. K. Rowan and A.L. contributed to clinical data management. J.K.B. contributed to scientific leadership.

Peer review

Peer review information

Nature thanks Jacques Fellay and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Peer reviewer reports are available.

Data availability

Downloadable summary data are available through the GenOMICC data site (https://genomicc.org/data). Summary statistics are available, but without the 23andMe summary statistics, except for the 10,000 most significant hits, for which full summary statistics are available. The full GWAS summary statistics for the 23andMe discovery dataset will be made available through 23andMe to qualified researchers under an agreement with 23andMe that protects the privacy of the 23andMe participants. For further information and to apply for access to the data, see the 23andMe website (https://research.23andMe.com/dataset-access/). All individual-level genotype and whole-genome sequencing data (for both academic and commercial uses) can be accessed through the UKRI/HDR UK Outbreak Data Analysis Platform (https://odap.ac.uk). A restricted dataset for a subset of GenOMICC participants is also available through the Genomics England data service. Monocyte RNA-seq data are available under the title ‘Monocyte gene expression data’ within the Oxford University Research Archives (10.5287/ora-ko7q2nq66). Sequencing data will be made freely available to organizations and researchers to conduct research in accordance with the UK Policy Framework for Health and Social Care Research through a data access agreement. Sequencing data have been deposited at the European Genome–Phenome Archive (EGA), which is hosted by the EBI and the CRG, under accession number EGAS00001007111.

Code availability

Code to calculate the imputation of P values on the basis of SNPs in linkage disequilibrium is available at GitHub (https://github.com/baillielab/GenOMICC_GWAS).

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

These authors contributed equally: Erola Pairo-Castineira, Konrad Rawlik

A list of authors and their affiliations appears online

Change history

7/11/2023

A Correction to this paper has been published: 10.1038/s41586-023-06383-z

Contributor Information

GenOMICC Investigators

Sara Clohisey,3 Johnny Millar,3 and Manu Shankar-Hari18, for the Co-Investigators
Manuela Mundy,56 Christopher Thompson,56 Lisa Pritchard,56 Minnie Gellamucho,56 David Cartlidge,56 Nageswar Bandla,56 Lucy Bailey,56 Michelle Davies,56 Jane Delaney,56 and Leanne Scott56, for the Royal Stoke University Hospital, Staffordshire, UK
Phoebe Coghlan,59 Stephen Brett,59 Anthony Gordon,59 Maie Templeton,59 David Antcliffe,59 Dorota Banach,59 Sarah Darnell,59 Ziortza Fernandez,59 Eleanor Jepson,59 Amal Mohammed,59 Roceld Rojo,59 Sonia Sousa Arias,59 Anita Tamang Gurung,59 and Jenny Wong59, for the Charing Cross Hospital, St Mary’s Hospital and Hammersmith Hospital, London, UK
Archana Bashyal,61 Sally Beer,61 Paula Hutton,61 Stuart McKechnie,61 Neil Davidson,61 Soya Mathew,61 Grace Readion,61 Jung Ryu,61 and Jean Wilson61, for the John Radcliffe Hospital, Oxford, UK
Janice Hastings,65 Lina Grauslyte,65 Musarat Hussain,65 Bobby Ruge,65 Sam King,65 Tatiana Pogreban,65 Lace Rosaroso,65 Helen Smith,65 Mandeep-Kaur Phull,65 Nikkita Adams,65 George Franke,65 Aparna George,65 Erika Salciute,65 Joanna Wong,65 Karen Dunne,65 Luke Flower,65 Emma Sharland,65 and Sukhmani Sra65, for the BHRUT (Barking Havering)—Queens Hospital and King George Hospital, Essex, UK
Angela Nown,68 Steve Rose,68 David Pogson,68 Helen Boxall,68 Lutece Brimfield,68 Helen Claridge,68 Zoe Daly,68 Shenu George,68 and Andrew Gribbin68, for the Queen Alexandra Hospital, Portsmouth, UK
Alissa Kent,71 Emma Goodwin,71 Ahmed Zaki,71 Clare Tibke,71 Susan Hopkins,71 Hywel Gerrard,71 Matthew Jackson,71 Sara Bennett,71 Liane Marsh,71 and Rebecca Mills71, for the Stepping Hill Hospital, Stockport, UK
Vasileios Bafitis,76 George Tsinaslanidis,76 Cassandra George,76 Reena Khade,76 Christopher Black,76 and Sundar Raj Ashok76, for the Croydon University Hospital, Croydon, UK
Fiona McNeela,79 Amanda Lyle,79 Alistair Hughes,79 Jayachandran Radhakrishnan,79 and Sian Gibson79, for the Broomfield Hospital, Chelmsford, UK
Juliet Cottle,81 Emily Peasgood,81 Laura Ortiz-Ruiz de Gordoa,81 Claire Phillips,81 and Denise Skinner81, for the Royal Sussex County Hospital, Brighton, UK
Jo Dasgin,83 Jaspret Gill,83 Annette Nilsson,83 Amy Bamford,83 Diana Hull,83 James Scriven,83 Nafeesah Ahmadhaider,83 Michelle Bates,83 and Christopher McGhee83, for the Queen Elizabeth Hospital, Birmingham, UK
Vinodh Krishnamurthy,85 Cynthia Diaba,85 Lincy John,85 Lai Lim,85 and Rajeev Jha85, for the Barnet Hospital, London, UK
Oluronke Adanini,87 Rebecca Collins,87 Maines Msiska,87 Linda Ofori,87 Nikhil Bhatia,87 and Hayley Dolan87, for the Medway Maritime Hospital, Gillingham, UK
Mark Brunton,88 Jess Caterson,88 Holly Coles,88 Liza Keating,88 Emma Tilney,88 Nicola Jacques,88 Matthew Frise,88 Jennifer Armistead,88 Shauna Bartley,88 Parminder Bhuie,88 Sabi Rai,88 and Gabriela Tomkova88, for the Royal Berkshire NHS Foundation Trust, Berkshire, UK
Sandra Greer,89 Karen Shuker,89 and Ascanio Tridente89, for the Whiston Hospital, Prescot, UK
Angela Allan,92 Adriana Botello,92 Jade Liew,92 Jasmine Medhora,92 Erin Trumper,92 Felicity Savage,92 Teresa Scott,92 Marc Place,92 and Callum Kaye92, for the Aberdeen Royal Infirmary, Aberdeen, UK
Sarah Benyon,93 Suzie Marriott,93 Linda Park,93 Helen Quinn,93 Daisy Skyes,93 Lily Zitter,93 Kizzy Baines,93 Elizabeth Gordon,93 Samantha Keenan,93 and Andrew Pitt93, for the Royal Devon and Exeter Hospital, Exeter, UK
Maia Aquino,97 Maria Croft,97 Victoria Frost,97 Ian White,97 and Keshnie Govender97, for the Ashford and St Peter’s Hospital, Surrey, UK
Natasha Webb,98 Liana Stapleton,98 Colin Wells,98 Nikitas Nikitas,98 Ana Sanchez-Rodriguez,98 Kayleigh Spencer,98 and Bethan Stowe98, for the Derriford Hospital, Plymouth, UK
Yvonne Izzard,99 Michelle Poole,99 Sonja Monnery,99 Sallyanne Trotman,99 Valerie Beech,99 Edward Combes,99 and Teishel Joefield99, for the East Surrey Hospital, Redhill, UK
Bethan Ogg,102 Simon Whiteley,102 Liz Wilby,102 Kate Long,102 Shailamma Matthew,102 Sheila Salada,102 Susan Trott,102 Sarah Watts,102 Zoe Friar,102 and Abigail Speight102, for the St James’s University Hospital and Leeds General Infirmary, Leeds, UK
Charlotte Heath,104 Srinivas Jakkula,104 Anna Morris,104 Ashar Ahmed,104 Arvind Nune,104 Claire Buttriss,104 and Emma Whitaker104, for the Southport and Formby District General Hospital, Ormskirk, UK
Miriam Davey,105 David Golden,105 Amy Acklery,105 Fabio Fernandes,105 Bec Seaman,105 and Victoria Earl105, for the The Tunbridge Wells Hospital and Maidstone Hospital, Kent, UK
Amy Collins,106 Waqas Khaliq,106 Rachel Adam,106 and Estefania Treus106, for the Queen Elizabeth Hospital, Woolwich, London, UK
Charlotte Downes,111 Melanie Hayman,111 Katie Riches,111 Priya Daniel,111 Deepak Subramanian,111 Kathleen Holding,111 Mary Hilton,111 Carly McDonald,111 and Georgina Richardson111, for the Royal Derby Hospital, Derby, UK
Georgia Halladay,112 Peter Harding,112 Amie Reddy,112 Ian Turner-Bone,112 Laura Wilding,112 Robert Parker,112 Michaela Lloyd,112 Leanne Smith,112 and Charlie Kelly112, for the Aintree University Hospital, Liverpool, UK
Maria Lazo,113 Alan Neal,113 Olivia Walton,113 Julie Melville,113 Jay Naisbitt,113 Emily Bullock,113 and Rosane Joseph113, for the Fairfield General Hospital, Bury, UK
Sara Callam,114 Lisa Hudig,114 Jocelyn Keshet-Price,114 Katie Stammers,114 Karen Convery,114 Georgina Randell,114 and Deirdre Fottrell-Gould114, for the Norfolk and Norwich University hospital (NNUH), Norwich, UK
Esther Mwaura,115 Sara-Beth Sutherland,115 Richard Stewart,115 Louise Mew,115 and Lynn Wren115, for the Milton Keynes University Hospital, Milton Keynes, UK
Laura Thrasyvoulou,116 Heather Willis,116 James Scriven,116 Bridget Hopkins,116 Daniel Lenton,116 and Abigail Roberts116, for the Good Hope Hospital, Birmingham, UK
Maria Bokhari,117 Rachael Lucas,117 Wendy McCormick,117 Jenny Ritzema,117 Vanessa Linnett,117 Amanda Sanderson,117 and Helen Wild117, for the Queen Elizabeth Hospital Gateshead, Gateshead, UK
Rebecca Flanagan,118 Robert Hull,118 Kat Rhead,118 Emma McKenna,118 Gareth Hughes,118 Jennifer Anderson,118 Kelly Jones,118 Scott Latham,118 and Heather Riley118, for the Royal Bolton Hospital, Bolton, UK
Martina Coulding,119 Martyn Clark,119 Jacqueline McCormick,119 Oliver Mercer,119 Darsh Potla,119 Hafiz Rehman,119 Heather Savill,119 Victoria Turner,119 Edward Jude,119 and Susan Kilroy119, for the Tameside General Hospital, Ashton Under Lyne, UK
Joyann Andrews,121 Emily Beech,121 Olugbenga Akinkugbe,121 Alasdair Bamford,121 Holly Belfield,121 Gareth A. L. Jones,121 Tara McHugh,121 Hamza Meghari,121 Samiran Ray,121 Ana Luisa Tomas,121 Lauran O’Neill,121 Mark Peters,121 Michael Bell,121 Sarah Benkenstein,121 Catherine Chisholm,121 Charlene Davies,121 Klaudia Kupiec,121 and Caroline Payne121, for the Great Ormond St Hospital and UCL Great Ormond St Institute of Child Health NIHR Biomedical Research Centre, London, UK
Victoria Priestley,123 Tracey Cosier,123 Gemma Millen,123 James Rand,123 Natasha Schumacher,123 Roxana Sandhar,123 Heather Weston,123 Neil Richardson,123 and Lucy Cooper123, for the William Harvey Hospital, Ashford, UK
Cathy Jones,124 Ya-Wen Jessica Huang,124 Reni Jacob,124 Craig Denmade,124 and Lewis McIntyre124, for the Arrowe Park Hospital, Wirral, UK
Dawn Trodd,125 Jane Martin,125 Geoff Watson,125 Emily Bevan,125 and Caroline Wreybrown125, for the Royal Hampshire County Hospital, Hampshire, UK
Jordan Frankham,128 Sally Pitts,128 Nigel White,128 Debbie Branney,128 and Heather Tiller128, for the Royal Bournemouth Hospital, Bournemouth, UK
Melanie Kent,130,131 Ami Wilkinson,130,131 Ellen Brown,130,131 Andrea Kay,130,131 Suzanne Campbell,130,131 Amanda Cowton,130,131 Mark Birt,130,131 Vicki Greenaway,130,131 Kathryn Potts,130,131 Clare Hutton,130,131 and Andrew Shepperson130,131, for the University Hospital North Durham, Darlington, UK and Darlington Memorial Hospital, Darlington, UK
Miranda Forsey,132 Alice Nicholson,132 Mark Vertue,132 Joanne Riches,132 Agilan Kaliappan,132 and Anne Nicholson132, for the Basildon Hospital, Basildon, UK
Niall MacCallum,133 Eamon Raith,133 Georgia Bercades,133 Ingrid Hass,133 David Brealey,133 Gladys Martir,133 Anna Reyes,133 Deborah Smyth,133 and Maria Zapatamartinez133, for the University College Hospital, London, UK
Ana Alvaro,134 Champa Jetha,134 Louise Ma,134 Lauren Booker,134 Loreta Mostoles,134 Anezka Pratley,134 Abdelhakim Altabaibeh,134 Chetan Parmar,134 and Kayleigh Gilbert134, for the Whittington Hospital, London, UK
Susie Ferguson,135 Amy Shepherd,135 Sheila Morris,135 Jo Singleton,135 Rosie Baruah,135 Maria Amamio,135 Sophie Birch,135 Kate Briton,135 Sarah Clark,135 Katherine Doverman,135 Lucy Marshall,135 and Scott Simpson135, for the Western General Hospital, Edinburgh, UK
Georgina Lloyd,136 Stephanie Bell,136 Vanessa Rivers,136 and Bally Purewal136, for the Ipswich Hospital, Ipswich, UK
Kate Hammerton,137 Susan Anderson,137 Janine Birch,137 Emma Collins,137 and Ryan Oleary137, for the Hereford County Hospital, Hereford, UK
Sarah Cornell,138 Jordan Jarmain,138 Kimberley Rogerson,138 Fiona Wakinshaw,138 Lindsey Woods,138 Anthony Rostron,138 Zeynep Elcioglu,138 and Alistair Roy138, for the Sunderland Royal Hospital, Sunderland, UK
Gillian Bell,139 Holly Dickson,139 Louise Wilcox,139 Amro Katary,139 and Katy English139, for the Queens Hospital Burton, Burton-On-Trent, UK
Joanne Hutter,140 Corinne Pawley,140 Patricia Doble,140 Charmaine Shovelton,140 Marius Vaida,140 Rebecca Purnell,140 and Ashly Thomas140, for the Musgrove Park Hospital, Taunton, UK
Mark Birt,142 Estefania Treus Gude,142 Maggie Nyirenda,142 Lisa Capozzi,142 Rosie Reece-Anthony,142 Waqas Khaliq,142 Hazma Noor,142 and Alfa Cresia Nilo142, for the University Hospital Lewisham, London, UK
Michelle Grove,143 Amelia Daniel,143 Amy Easthope,143 Joanne Finn,143 Nikki White,143 Rajnish Saha,143 Bibi Badal,143 and Karen Ixer143, for the The Princess Alexandra Hospital, Harlow, UK
Donna Duffin,144 Ben Player,144 Helen Hill,144 Jade Cole,144 Jenny Brooks,144 Michelle Davies,144 Rhys Davies,144 Lauren Hunt,144 Emma Thomas,144 and Angharad Williams144, for the University Hospital of Wales, Cardiff, UK
Metod Oblak,145 Mini Thankachen,145 Jamie Irisari,145 Amrinder Sayan,145 and Monica Popescu145, for the West Middlesex Hospital, Isleworth, UK
Cheryl Finch,146 Andrew Jamieson,146 Alison Quinn,146 Joshua Cooper,146 Sarah Liderth,146 and Natalia Waddington146, for the Royal Albert Edward Infirmary, Wigan, UK
Iona Burn,147 Katarina Manso,147 Ruth Penn,147 Julie Tebbutt,147 Danielle Thornton,147 James Winchester,147 Geraldine Hambrook,147 and Pradeep Shanmugasundaram147, for the Stoke Mandeville Hospital, Buckinghamshire, UK
Jayne Craig,148 Kerry Simpson,148 Andrew Higham,148 and Louise Sibbett148, for the Royal Lancaster Infirmary, Lancaster, UK
Sheila Paine,149 Annabel Reed,149 Jo-Anna Conyngham,149 McDonald Mupudzi,149 Rachel Thomas,149 Mary Wright,149 Denise Griffin,149 Richard Partridge,149 Maria Alvarez Corral,149 Nycola Muchenje,149 Mildred Sitonik,149 and Caroline Wrey Brown149, for the Basingstoke and North Hampshire Hospital, Basingstoke, UK
Aaron Butler,150,151 Linda Folkes,150,151 Heather Fox,150,151 Amy Gardner,150,151 David Helm,150,151 Gillian Hobden,150,151 Kirsten King,150,151 Jordi Margalef,150,151 Michael Margarson,150,151 Tim Martindale,150,151 Emma Meadows,150,151 Dana Raynard,150,151 Yvette Thirlwall,150,151 Yolanda Baird,150,151 Raquel Gomez,150,151 Darren Martin,150,151 Luke Hodgson,150,151 Clinton Corin,150,151 Erikka Sidall,150,151 Densie Szabo,150,151 and Sharon Floyd150,151, for the Worthing Hospital, Worthing, UK and St Richard’s Hospital, Chichester, UK
Hannah Davies,152 Karen Austin,152 Olivia Kelsall,152 Hannah Wood,152 Peter Anderson,152 Katie Archer,152 Andrew Burtenshaw,152 Sarah Clayton,152 Naiara Cother,152 Nicholas Cowley,152 Caroline Davis,152 Stephen Digby,152 Alison Durie,152 Alison Harrison,152 Emma Low,152 Michael McAlindon,152 Alex McCurdy,152 Aled Morgan,152 Tobias Rankin,152 Jessica Thrush,152 Helen Tranter,152 Charlie Vigurs,152 and Laura Wild152, for the The Alexandra Hospital, Redditch and Worcester Royal Hospital, Worcester, UK
Thomas Cornell,153 Kate Ralph,153 Sarah Bean,153 Karen Burt,153 Michael Spivey,153 Carol Richards,153 and Rachel Tedstone153, for the Royal Cornwall Hospital, Truro, UK
Siobhain Carmody,154 Xiaobei Zhao,154 Valerie Page,154 Mark Louie Guanco,154 Elvira Hoxha,154 and Camilla Zorloni154, for the Watford General Hospital, Watford, UK
Charlotte Dean,155 Emma Jones,155 Emma Carter,155 Joshua Dunn,155 Thomas Kong,155 Mervin Mahenthran,155 Chris Marsh,155 Maureen Holland,155 Natalie Keenan,155 Mohamed Mahmoud,155 Marc Lyons,155 Joanne Bradley-Potts,155 Helen Wassall,155 and Meghan Young155, for the Macclesfield District General Hospital, Macclesfield, UK
Victoria Maynard,157 Rachel Walker,157 Anil Hormis,157 Dawn Collier,157 Cheryl Graham,157 Vicky Maynard,157 Jake McCormick,157 and Jake Warrington157, for the Rotherham General Hospital, Rotherham, UK
Denise Cosgrove,158 Denise McFarland,158 Judith Ratcliffe,158 and Rob Charnock158, for the Craigavon Area Hospital, County Armagh, Northern Ireland
Inez Wynter,159 Mandy Gill,159 Jill Kirk,159 Paul Paul,159 Valli Ratnam,159 and Sarah Shelton159, for the King’s Mill Hospital, Nottingham, UK
Catherine Jardine,160 Alasdair Hay,160 and Dewi Williams160, for the Dumfries and Galloway Royal Infirmary, Dumfries, UK
Bethan Deacon,161 Latha Durga,161 Meg Hibbert,161 Gareth Kennard-Holden,161 Chrsitopher Woodford,161 Carla Pothecary,161 Lisa Roche,161 Dariusz Tetla,161 Kevin Agravante,161 and Justyna Smeaton161, for the Prince Charles Hospital, Merthyr Tydfil, UK
Alicia Price,162 Alice Thomas,162 Chris Thorpe,162 Ellen Knights,162 and Donna Ward162, for the Ysbyty Gwynedd, Bangor, UK
Shondipon Laha,163 Mark Verlander,163 and Alexandra Williams163, for the Royal Preston Hospital, Preston, UK
Rachel Prout,164 Helen Langton,164 Malcolm Watters,164 Charlotte Hunt,164 and Catherine Novis164, for the The Great Western Hospital, Swindon, UK
Dawn Egginton,166 Michele Clark,166 Sarah Purvis,166 Simon Sinclair,166 and Vicky Collins166, for the University Hospital of North Tees, Stockton on Tees, UK
Bethan Landeg,167 Craig Sell,167 Samantha Coetzee,167 Alistair Gales,167 Igor Otahal,167 Becky Icke,167 Meena Raj,167 Caroline Williams,167 Jill Williams,167 and Lucy Hill167, for the Glangwili General Hospital, Camarthen, UK
Abdul Kayani,168 Bridgett Masunda,168 Prisca Gondo,168 and Nigara Atayeva168, for the Southend University Hospital, Westcliff-on-Sea, UK
Carina Cruz169 and Natalie Pattison169, for the Lister Hospital, Stevenage, UK
Caroline Burnett,170 Jonathan Hatton,170 Elaine Heeney,170 Maria Newton,170 Hassan Al-Moasseb,170 Teresa Behan,170 Jasmine Player,170 Rachael Stead,170 Atideb Mitra,170 and Kirsty Nauyokas170, for the Diana Princess of Wales Hospital, Grimsby, UK
Sally Humphreys,171 Helen Cockerill,171 and Ruth Tampsett171, for the West Suffolk Hospital, Bury St Edmunds, UK
Evgeniya Postovalova,172 Tina Coventry,172 Amanda McGregor,172 Susan Fowler,172 Mike Macmahon,172 Patricia Cochrane,172 and Sandra Pirie172, for the Victoria Hospital, Kirkcaldy, UK
Sarah Hanley,173,174 Asifa Ali,173,174 Megan Brady,173,174 Sam Dale,173,174 Annalisa Dance,173,174 Lisa Gledhill,173,174 Jill Greig,173,174 Kathryn Hanson,173,174 Kelly Holdroyd,173,174 Marie Home,173,174 Tahira Ishaq,173,174 Diane Kelly,173,174 Lear Matapure,173,174 Deborah Melia,173,174 Samantha Mellor,173,174 Ekta Merwaha,173,174 Tonicha Nortcliffe,173,174 Lisa Shaw,173,174 Ryan Shaw,173,174 Tracy Wood,173,174 Lee-Ann Bayo,173,174 Miranda Usher,173,174 Alison Wilson,173,174 Ross Kitson,173,174 Jez Pinnell,173,174 Matthew Robinson,173,174 and Kaitlin Boltwood173,174, for the Calderdale Royal Hospital, Halifax, UK and Huddersfield Royal Infirmary, Huddersfield, UK
Jenny Birch,175 Laura Bough,175 Rebecca Tutton,175 Barbara Winter-Goodwin,175 Josie Goodsell,175 Kate Taylor,175 Patricia Williams,175 Sarah Williams,175 Ashleigh Cave,175 and James Rees175, for the Dorset County Hospital, Dorchester, UK
Janet Imeson-Wood,176 Jacqueline Smith,176 Vishal Amin,176 Komala Karthik,176 Rizwana Kausar,176 Elena Anastasescu,176 Karen Reid,176 Vikram Anumakonda,176 and Ella Stoddart176, for the Russell’s Hall Hospital, Dudley, UK
Carrie Demetriou,177 Charlotte Eckbad,177 Lucy Howie,177 Sarah Mitchard,177 Lidia Ramos,177 Katie White,177 Sarah Hierons,177 Fiona Kelly,177 Alfredo Serrano-Ruiz,177 and Gabrielle Evans177, for the Royal United Hospital, Bath, UK
Liz Nicol,178 Joy Wilkins,178 Kim Hulacka,178 Gabor Debreceni,178 Alison Brown,178 and Vikki Crickmore178, for the St Mary’s Hospital, Newport, UK
Kay Hill179 and Thogulava Kannan179, for the George Eliot Hospital NHS Trust, Nuneaton, UK
Harish Venkatesh,182 Yvonne Bland,182 Istvan Kajtor,182 Lisa Kavanagh,182 Karen Singler,182 and George Linfield-Brown182, for the Frimley Park Hospital, Surrey, UK
Luke Stephen Prockter Moore,183 Marcela Vizcaychipi,183 Laura Martins,183 Luke Moore,183 Rhian Bull,183 and Jaime Carungcong183, for the Chelsea & Westminster NHS Foundation Trust, London, UK
Louise Allen,184 Eva Beranova,184 Alicia Knight,184 Carly Price,184 Sorrell Tilbey,184 Sharon Turney,184 Tracy Hazelton,184 Gabriella Tutt,184 Mansi Arora,184 Salah Turki,184 Emily Sinfield,184 Joanne Deery,184 and Hazel Ramos184, for the Queen Elizabeth the Queen Mother Hospital, Margate, UK
John Allan,187 Tim Geary,187 Alistair Meikle,187 Peter O’Brien,187 Stephen Wood,187 Andrew Clark,187 and Gordon Houston187, for the University Hospital Crosshouse, Kilmarnock, UK
Lorna Murphy,189 Michelle Smythe,189 Alistair Nichol,189 and Kathy Brickell189, for the University College Dublin, St Vincent’s University Hospital, Dublin, Ireland
Liam Botfield,191 Catherine Dexter,191 Aditya Kuravi,191 Joanne Butler,191 Robert Chadwick,191 Poonam Ranga,191 Lisa Richardson,191 Emma Virgilio,191 Maddiha Anwer,191 Atul Garg,191 Donna Botfield,191 and Xana Marriott191, for the Walsall Manor Hospital, Walsall, UK
Keely Stewart,192 Dee Mullan,192 Claire Phillips,192 Jane Gaylard,192 Justyna Nowak,192 and Denise Skinner192, for the Princess Royal Hospital, Brighton, UK
Ailbhe Brady,194 Rebekah Chan,194 Shane McIvor,194 Helena Prady,194 Bijoy Mathew,194 Jeff Little,194 and Tim Furniss194, for the Warrington General Hospital, Warrington, UK
Alison Brown,198 Susan O’Connell,198 Jane Gregory,198 Luigi Barberis,198 Rosemary Harper,198 Tim Smith,198 and Diane Armstrong198, for the Cumberland Infirmary, Carlisle, UK
Angie Bowey,199,200 Anne Cowley,199,200 Andrew Corner,199,200 Judith Highgate,199,200 Claire Rutherfurd,199,200 Jo-Anne Taylor,199,200 Sarah Goodwin,199,200 and Claire Rutherford199,200, for the Eastbourne District General Hospital, East Sussex, UK and Conquest Hospital, East Sussex, UK
Beena Eapen,201 Fiona Trim,201 and Phil Donnison201, for the Salisbury District Hospital, Salisbury, UK
Lisa Armstrong,202 Hayley Bates,202 Emma Dooks,202 Fiona Farquhar,202 Amy Kitching,202 Chantal McParland,202 Sophie Packham,202 and Brigid Hairsine202, for the Airedale General Hospital, Keighley, UK
Anand Patil,203 Premetie Andreou,203 Dawn Hales,203 Megha Mathews,203 Rekha Patel,203 Peter Barry,203 Neil Flint,203 Jessica Hailstone,203 Navneet Ghuman,203 Bethany Leonard,203 and Rachel Lees203, for the Leicester Royal Infirmary, Leicester, UK
Deborah Butcher,204,205 Katy Leng,204,205 Nicola Butterworth-Cowin,204,205 and Susie O’Sullivan204,205, for the Peterborough City Hospital, Peterborough, UK and Hinchingbrooke Hospital, Huntingdon, UK
Alison Ghosh206 and Emma Williams206, for the Colchester General Hospital, Colchester, UK
Tracy Baird,208 Margaret Harkins,208 Jim Ruddy,208 and Joe West208, for the University Hospital Monklands, Airdrie, UK
Joseph Duffield,209 Lewis Mallon,209 Oliver Smith,209 Sara Smuts,209 Andy Campbell,209 Cate Davies,209 Sarah Davies,209 Rachel Hughes,209 Lisa Jobes,209 Victoria Whitehead,209 and Clare Watkins209, for the Wrexham Maelor Hospital, Wrexham, Wales
Fiona Bowman,27 Barry Milligan,27 and Liane McPherson27, for the Royal Hospital for Children, Glasgow, UK
Stella Metherell,210 Nichola Harris,210 Victoria Lake,210 Elizabeth Radford,210 Andy Smallwood,210 Shameer Gopal,210 and Katherine Vassell210, for the New Cross Hospital, Wolverhampton, UK
Dina Bell,211 Rosalind Boyle,211 Katie Douglas,211 Lynn Glass,211 Liz Lennon,211 Austin Rattray,211 Claire Beith,211 and Emma Lee211, for the University Hospital Hairmyres, East Kilbride, UK
Danielle Jones,212 Penny Parsons,212 Ben Attwood,212 Paul Jefferson,212 Mohan Ranganathan,212 Inderjit Atwal,212 Bridget Campbell,212 Angela Day,212 and Camilla Stagg212, for the Warwick Hospital, Warwick, UK
Emma Haynes,213 Cecilia Ahmed,213 Sarah Clamp,213 Julie Colley,213 Risna Haq,213 Anne Hayes,213 Sibet Joseph,213 Zahira Maqsood,213 Samia Hussain,213 Jonathan Hulme,213 Patience Domingos,213 Rita Kumar,213 Manjit Purewal,213 and Becky Taylor213, for the Sandwell General Hospital and City Hospital, Birmingham, UK
Lara Bunni,214 Monica Latif,214 Claire Jennings,214 Shilu Jose,214 Rebecca Marshall,214 Aleksandra Metryka,214 and Gayathri Subramanian214, for the Royal Manchester Children’s Hospital, Manchester, UK
Adam Burgoyne,215 Susan O’Connell,215 Amanda Tyler,215 Joanne Waldron,215 Paula Hilltout,215 and Jayne Evitts215, for the Gloucestershire Royal Hospital, Gloucester, UK
Geraldine Ward,216 Pamela Bremmer,216 Carl Hawkins,216 Sophie Jackman,216 and Michal Ogorek216, for the University Hospitals Coventry & Warwickshire NHS Trust, Coventry, UK
Kylie Ashby,217 Lorraine Thornton,217 Pauline Mercer,217 Matthew Halkes,217 and Adam Revill217, for the Torbay Hospital, Torquay, UK
Bryony Saint,218 Jo Fletcher,218 Kimberley Netherton,218 Manish Chablani,218 Amy Kirkby,218 Amanda Roper,218 and Kinga Szymiczek218, for the Pilgrim Hospital, Lincoln, UK
Isobel Sutherland,219 Linda O’Brien,219 Igor Otahal,219 Joanne Connell,219 Kim Davies,219 Tracy Lewis,219 Zohra Omar,219 and Emma Perkins219, for the Prince Philip Hospital, Lianelli, UK
Lisa Roche,220 Sonia Sathe,220 and Ellie Davies220, for the Princess of Wales Hospital, Llantrisant, UK
Alex Lyon,221 Isheunesu Mapfunde,221 Charlotte Willis,221 Rachael Hitchcock,221 Kathryn Hall,221 and Christopher King221, for the Northampton General Hospital NHS Trust, Northampton, UK
Andrew Fagan,222 Roonak Nazari,222 Lucy Worsley,222 Suzanne Allibone,222 Vidya Kasipandian,222 Amit Patel,222 Parisa Cutting,222 Roman Genetu,222 Ainhi Mac,222 Anthony Murphy,222 Sinead Ward,222 and Fatima Butt222, for the The Christie NHS Foundation Trust, Manchester, UK
Amanda Ayers,223 Wendy Harrison,223 Katherine Mackintosh,223 and Julie North223, for the James Paget University Hospital NHS Trust, Great Yarmouth, UK
Lydia Ashton,224 Rehana Bi,224 Samantha Owen,224 Helen Winmill,224 and Barney Scholefield224, for the Birmingham Children’s Hospital, Birmingham, UK
Hannah Blowing,225 Erin Williams,225 Michaela Duskova,225 Michelle Edwards,225 Alun Rees,225 Helen Thomas,225 Rachel Hughes,225 Igor Otahal,225 Jolene Brooks,225 Janet Phipps,225 and Suzanne Brooks225, for the Withybush General Hospital, Pembrokeshire, Wales
Catherine Dennis,226 Vicki Parris,226 Sinduya Srikaran,226 Anisha Sukha,226 Alistair McGregor,226 and Gerlynn Tiongson226, for the Northwick Park Hospital, London, UK
Katie Adams,227 Benedict Andrew,227 Adam Brayne,227 Sasha Carter,227 Louise Findlay,227 Emma Fisher,227 Peter Jackson,227 Duncan Kaye,227 Juliet Parkin,227 Victoria Tuckey,227 Jane Hunt,227 Nicholas Love,227 Lynne van Koutrick,227 and Ashley Hanson227, for the North Devon District Hospital, Barnstaple, UK
Kathy Dent,228 Elizabeth Horsley,228 Sandra Pearson,228 Sue Spencer,228 Dorothy Hutchinson,228 Jasmine Player,228 Dorota Potoczna,228 Muhammad Nauman Akhtar,228 Lisa-Jayne Cottam,228 Kirsty Nauyokas,228 and Jack Sanders228, for the Scunthorpe General Hospital, Scunthorpe, UK
Sara Mingo Garcia,229 Glykeria Pakou,229 Cynthia Diaba,229 Helder Filipe,229 Lincy John,229 Amitaa Maharajh,229 Mark de Neef,229 Daniel Martin,229 Christine Eastgate,229 and Poh Choo Teoh229, for the Royal Free Hospital, London, UK
Polly Rice,231 Tim Smith,231 Melanie Clapham,231 Rachel Mutch,231 Luigi Barberis,231 Rosemary Harper,231 Hannah Craig,231 and Una Poultney231, for the West Cumberland Hospital, Whitehaven, UK
Karen Burns232 and Andrew Higham232, for the Furness General Hospital, Barrow-in-Furness, UK
Sophie Twiss,233 Janet Barton,233 Linsha George,233 Clare Harrop,233 Sherly Mathew,233 and David Justin Wright233, for the Liverpool Heart and Chest Hospital, Liverpool, UK
Rachel Harrison,234 Jordan Toohie,234 Ben Chandler,234 Alison Turnbull,234 Janine Mallinson,234 and Kerry Elliott234, for the Scarborough General Hospital, Scarborough, UK
Rebecca Wolf-Roberts,235 Helen Tench,235 Igor Otahal,235 Maria Hobrok,235 Ronda Loosley,235 Heather McGuinness,235 and Tanya Sims235, for the Bronglais General Hospital, Aberystwyth, UK
Deborah Afolabi,236 Kathryn Sian Allison,236 Taya Anderson,236 Rachael Dore,236 Dawn Jones,236 Naomi Rogers,236 Paula Saunderson,236 Jennifer Whitbread,236 Laura O’Malley,236 Laura Rad,236 and Daniel Hawcutt236, for the Alder Hey Children’s Hospital, Liverpool, UK
Jonathan Aldridge,237 Melanie Tolson,237 and Sweyn Garrioch237, for the Borders General Hospital, Melrose, UK
Joanne Tomlinson238 and Michael Grosdenier238, for the Leighton Hospital, Cheshire, UK
David Loader,239 Ritoo Kapoor,239 and Gemma Hector239, for the Kent & Canterbury Hospital, Canterbury, UK
Joslan Scherewode,240 Chunda Sri-Chandana,240 Lorraine Stephenson,240 and Sarah Marsh240, for the Harrogate and District NHS Foundation Trust, Harrogate, UK
Arnold Dela Rosa,241 Shaman Jhanji,241 Thomas Bemand,241 Ryan Howle,241 Ravishankar Rao Baikady,241 Benjamin Thomas,241 Ethel Black,241 and Kate Tatham241, for the The Royal Marsden Hospital, London, UK
Sambasivarao Gurram,242 Ekaterina Watson,242 Vicki Parris,242 Sheena Quaid,242 and Alistair McGregor242, for the Ealing Hospital, Southall, UK
Anne Saunderson,243 Rachel O’Brien,243 Sam Moultrie,243 Jen Service,243 Clare Cheyne,243 Miranda Odam,243 and Alison Wiliams243, for the St John’s Hospital Livingston, Livingston, UK
Nicky Barnes,244 Peter Csabi,244 Joana Da Rocha,244 and Louika Glynou244, for the Wexham Park Hospital, Slough, UK
Amy Huffenberger,245 Jade Bryant,245 Amy Pickard,245 Nicholas Roe,245 Arianna Bellini,245 Anton Mayer,245 Amy Burrow,245 Natalie Colley,245 Jayne Evans,245 Alex Howlett,245 and Zeinab Khalifeh245, for the Sheffield Children’s Hospital, Sheffield, UK
Jerldine Pryce,246 Claire Gorman,246 Amy Easthope,246 Rebecca Brady,246 Elizabeth Timlick,246 Pierre Antoine,246 and Abhinhav Gupta246, for the Homerton University Hospital Foundation NHS Trust, London UK
John Hardy,247 Henry Houlden,247 Eleanor Moncur,247 Arianna Tucci,247 Eamon Raith,247 Ambreen Tariq,247 and David Brealey247, for the National Hospital for Neurology and Neurosurgery, London, UK
Emma Tagliavini,248 Becky Ramsay,248 Katy Fidler,248 Kevin Donnelly,248 and Rebecca Hollis248, for the The Royal Alexandra Children’s Hospital, Brighton, UK
:
Sara Clohisey,3 Johnny Millar,3 and Manu Shankar-Hari18, for the Co-Investigators
Manuela Mundy,56 Christopher Thompson,56 Lisa Pritchard,56 Minnie Gellamucho,56 David Cartlidge,56 Nageswar Bandla,56 Lucy Bailey,56 Michelle Davies,56 Jane Delaney,56 and Leanne Scott56, for the Royal Stoke University Hospital, Staffordshire, UK
Phoebe Coghlan,59 Stephen Brett,59 Anthony Gordon,59 Maie Templeton,59 David Antcliffe,59 Dorota Banach,59 Sarah Darnell,59 Ziortza Fernandez,59 Eleanor Jepson,59 Amal Mohammed,59 Roceld Rojo,59 Sonia Sousa Arias,59 Anita Tamang Gurung,59 and Jenny Wong59, for the Charing Cross Hospital, St Mary’s Hospital and Hammersmith Hospital, London, UK
Archana Bashyal,61 Sally Beer,61 Paula Hutton,61 Stuart McKechnie,61 Neil Davidson,61 Soya Mathew,61 Grace Readion,61 Jung Ryu,61 and Jean Wilson61, for the John Radcliffe Hospital, Oxford, UK
Janice Hastings,65 Lina Grauslyte,65 Musarat Hussain,65 Bobby Ruge,65 Sam King,65 Tatiana Pogreban,65 Lace Rosaroso,65 Helen Smith,65 Mandeep-Kaur Phull,65 Nikkita Adams,65 George Franke,65 Aparna George,65 Erika Salciute,65 Joanna Wong,65 Karen Dunne,65 Luke Flower,65 Emma Sharland,65 and Sukhmani Sra65, for the BHRUT (Barking Havering)—Queens Hospital and King George Hospital, Essex, UK
Angela Nown,68 Steve Rose,68 David Pogson,68 Helen Boxall,68 Lutece Brimfield,68 Helen Claridge,68 Zoe Daly,68 Shenu George,68 and Andrew Gribbin68, for the Queen Alexandra Hospital, Portsmouth, UK
Alissa Kent,71 Emma Goodwin,71 Ahmed Zaki,71 Clare Tibke,71 Susan Hopkins,71 Hywel Gerrard,71 Matthew Jackson,71 Sara Bennett,71 Liane Marsh,71 and Rebecca Mills71, for the Stepping Hill Hospital, Stockport, UK
Vasileios Bafitis,76 George Tsinaslanidis,76 Cassandra George,76 Reena Khade,76 Christopher Black,76 and Sundar Raj Ashok76, for the Croydon University Hospital, Croydon, UK
Fiona McNeela,79 Amanda Lyle,79 Alistair Hughes,79 Jayachandran Radhakrishnan,79 and Sian Gibson79, for the Broomfield Hospital, Chelmsford, UK
Juliet Cottle,81 Emily Peasgood,81 Laura Ortiz-Ruiz de Gordoa,81 Claire Phillips,81 and Denise Skinner81, for the Royal Sussex County Hospital, Brighton, UK
Jo Dasgin,83 Jaspret Gill,83 Annette Nilsson,83 Amy Bamford,83 Diana Hull,83 James Scriven,83 Nafeesah Ahmadhaider,83 Michelle Bates,83 and Christopher McGhee83, for the Queen Elizabeth Hospital, Birmingham, UK
Vinodh Krishnamurthy,85 Cynthia Diaba,85 Lincy John,85 Lai Lim,85 and Rajeev Jha85, for the Barnet Hospital, London, UK
Oluronke Adanini,87 Rebecca Collins,87 Maines Msiska,87 Linda Ofori,87 Nikhil Bhatia,87 and Hayley Dolan87, for the Medway Maritime Hospital, Gillingham, UK
Mark Brunton,88 Jess Caterson,88 Holly Coles,88 Liza Keating,88 Emma Tilney,88 Nicola Jacques,88 Matthew Frise,88 Jennifer Armistead,88 Shauna Bartley,88 Parminder Bhuie,88 Sabi Rai,88 and Gabriela Tomkova88, for the Royal Berkshire NHS Foundation Trust, Berkshire, UK
Sandra Greer,89 Karen Shuker,89 and Ascanio Tridente89, for the Whiston Hospital, Prescot, UK
Angela Allan,92 Adriana Botello,92 Jade Liew,92 Jasmine Medhora,92 Erin Trumper,92 Felicity Savage,92 Teresa Scott,92 Marc Place,92 and Callum Kaye92, for the Aberdeen Royal Infirmary, Aberdeen, UK
Sarah Benyon,93 Suzie Marriott,93 Linda Park,93 Helen Quinn,93 Daisy Skyes,93 Lily Zitter,93 Kizzy Baines,93 Elizabeth Gordon,93 Samantha Keenan,93 and Andrew Pitt93, for the Royal Devon and Exeter Hospital, Exeter, UK
Maia Aquino,97 Maria Croft,97 Victoria Frost,97 Ian White,97 and Keshnie Govender97, for the Ashford and St Peter’s Hospital, Surrey, UK
Natasha Webb,98 Liana Stapleton,98 Colin Wells,98 Nikitas Nikitas,98 Ana Sanchez-Rodriguez,98 Kayleigh Spencer,98 and Bethan Stowe98, for the Derriford Hospital, Plymouth, UK
Yvonne Izzard,99 Michelle Poole,99 Sonja Monnery,99 Sallyanne Trotman,99 Valerie Beech,99 Edward Combes,99 and Teishel Joefield99, for the East Surrey Hospital, Redhill, UK
Bethan Ogg,102 Simon Whiteley,102 Liz Wilby,102 Kate Long,102 Shailamma Matthew,102 Sheila Salada,102 Susan Trott,102 Sarah Watts,102 Zoe Friar,102 and Abigail Speight102, for the St James’s University Hospital and Leeds General Infirmary, Leeds, UK
Charlotte Heath,104 Srinivas Jakkula,104 Anna Morris,104 Ashar Ahmed,104 Arvind Nune,104 Claire Buttriss,104 and Emma Whitaker104, for the Southport and Formby District General Hospital, Ormskirk, UK
Miriam Davey,105 David Golden,105 Amy Acklery,105 Fabio Fernandes,105 Bec Seaman,105 and Victoria Earl105, for the The Tunbridge Wells Hospital and Maidstone Hospital, Kent, UK
Amy Collins,106 Waqas Khaliq,106 Rachel Adam,106 and Estefania Treus106, for the Queen Elizabeth Hospital, Woolwich, London, UK
Charlotte Downes,111 Melanie Hayman,111 Katie Riches,111 Priya Daniel,111 Deepak Subramanian,111 Kathleen Holding,111 Mary Hilton,111 Carly McDonald,111 and Georgina Richardson111, for the Royal Derby Hospital, Derby, UK
Georgia Halladay,112 Peter Harding,112 Amie Reddy,112 Ian Turner-Bone,112 Laura Wilding,112 Robert Parker,112 Michaela Lloyd,112 Leanne Smith,112 and Charlie Kelly112, for the Aintree University Hospital, Liverpool, UK
Maria Lazo,113 Alan Neal,113 Olivia Walton,113 Julie Melville,113 Jay Naisbitt,113 Emily Bullock,113 and Rosane Joseph113, for the Fairfield General Hospital, Bury, UK
Sara Callam,114 Lisa Hudig,114 Jocelyn Keshet-Price,114 Katie Stammers,114 Karen Convery,114 Georgina Randell,114 and Deirdre Fottrell-Gould114, for the Norfolk and Norwich University hospital (NNUH), Norwich, UK
Esther Mwaura,115 Sara-Beth Sutherland,115 Richard Stewart,115 Louise Mew,115 and Lynn Wren115, for the Milton Keynes University Hospital, Milton Keynes, UK
Laura Thrasyvoulou,116 Heather Willis,116 James Scriven,116 Bridget Hopkins,116 Daniel Lenton,116 and Abigail Roberts116, for the Good Hope Hospital, Birmingham, UK
Maria Bokhari,117 Rachael Lucas,117 Wendy McCormick,117 Jenny Ritzema,117 Vanessa Linnett,117 Amanda Sanderson,117 and Helen Wild117, for the Queen Elizabeth Hospital Gateshead, Gateshead, UK
Rebecca Flanagan,118 Robert Hull,118 Kat Rhead,118 Emma McKenna,118 Gareth Hughes,118 Jennifer Anderson,118 Kelly Jones,118 Scott Latham,118 and Heather Riley118, for the Royal Bolton Hospital, Bolton, UK
Martina Coulding,119 Martyn Clark,119 Jacqueline McCormick,119 Oliver Mercer,119 Darsh Potla,119 Hafiz Rehman,119 Heather Savill,119 Victoria Turner,119 Edward Jude,119 and Susan Kilroy119, for the Tameside General Hospital, Ashton Under Lyne, UK
Joyann Andrews,121 Emily Beech,121 Olugbenga Akinkugbe,121 Alasdair Bamford,121 Holly Belfield,121 Gareth A. L. Jones,121 Tara McHugh,121 Hamza Meghari,121 Samiran Ray,121 Ana Luisa Tomas,121 Lauran O’Neill,121 Mark Peters,121 Michael Bell,121 Sarah Benkenstein,121 Catherine Chisholm,121 Charlene Davies,121 Klaudia Kupiec,121 and Caroline Payne121, for the Great Ormond St Hospital and UCL Great Ormond St Institute of Child Health NIHR Biomedical Research Centre, London, UK
Victoria Priestley,123 Tracey Cosier,123 Gemma Millen,123 James Rand,123 Natasha Schumacher,123 Roxana Sandhar,123 Heather Weston,123 Neil Richardson,123 and Lucy Cooper123, for the William Harvey Hospital, Ashford, UK
Cathy Jones,124 Ya-Wen Jessica Huang,124 Reni Jacob,124 Craig Denmade,124 and Lewis McIntyre124, for the Arrowe Park Hospital, Wirral, UK
Dawn Trodd,125 Jane Martin,125 Geoff Watson,125 Emily Bevan,125 and Caroline Wreybrown125, for the Royal Hampshire County Hospital, Hampshire, UK
Jordan Frankham,128 Sally Pitts,128 Nigel White,128 Debbie Branney,128 and Heather Tiller128, for the Royal Bournemouth Hospital, Bournemouth, UK
Melanie Kent,130,131 Ami Wilkinson,130,131 Ellen Brown,130,131 Andrea Kay,130,131 Suzanne Campbell,130,131 Amanda Cowton,130,131 Mark Birt,130,131 Vicki Greenaway,130,131 Kathryn Potts,130,131 Clare Hutton,130,131 and Andrew Shepperson130,131, for the University Hospital North Durham, Darlington, UK and Darlington Memorial Hospital, Darlington, UK
Miranda Forsey,132 Alice Nicholson,132 Mark Vertue,132 Joanne Riches,132 Agilan Kaliappan,132 and Anne Nicholson132, for the Basildon Hospital, Basildon, UK
Niall MacCallum,133 Eamon Raith,133 Georgia Bercades,133 Ingrid Hass,133 David Brealey,133 Gladys Martir,133 Anna Reyes,133 Deborah Smyth,133 and Maria Zapatamartinez133, for the University College Hospital, London, UK
Ana Alvaro,134 Champa Jetha,134 Louise Ma,134 Lauren Booker,134 Loreta Mostoles,134 Anezka Pratley,134 Abdelhakim Altabaibeh,134 Chetan Parmar,134 and Kayleigh Gilbert134, for the Whittington Hospital, London, UK
Susie Ferguson,135 Amy Shepherd,135 Sheila Morris,135 Jo Singleton,135 Rosie Baruah,135 Maria Amamio,135 Sophie Birch,135 Kate Briton,135 Sarah Clark,135 Katherine Doverman,135 Lucy Marshall,135 and Scott Simpson135, for the Western General Hospital, Edinburgh, UK
Georgina Lloyd,136 Stephanie Bell,136 Vanessa Rivers,136 and Bally Purewal136, for the Ipswich Hospital, Ipswich, UK
Kate Hammerton,137 Susan Anderson,137 Janine Birch,137 Emma Collins,137 and Ryan Oleary137, for the Hereford County Hospital, Hereford, UK
Sarah Cornell,138 Jordan Jarmain,138 Kimberley Rogerson,138 Fiona Wakinshaw,138 Lindsey Woods,138 Anthony Rostron,138 Zeynep Elcioglu,138 and Alistair Roy138, for the Sunderland Royal Hospital, Sunderland, UK
Gillian Bell,139 Holly Dickson,139 Louise Wilcox,139 Amro Katary,139 and Katy English139, for the Queens Hospital Burton, Burton-On-Trent, UK
Joanne Hutter,140 Corinne Pawley,140 Patricia Doble,140 Charmaine Shovelton,140 Marius Vaida,140 Rebecca Purnell,140 and Ashly Thomas140, for the Musgrove Park Hospital, Taunton, UK
Mark Birt,142 Estefania Treus Gude,142 Maggie Nyirenda,142 Lisa Capozzi,142 Rosie Reece-Anthony,142 Waqas Khaliq,142 Hazma Noor,142 and Alfa Cresia Nilo142, for the University Hospital Lewisham, London, UK
Michelle Grove,143 Amelia Daniel,143 Amy Easthope,143 Joanne Finn,143 Nikki White,143 Rajnish Saha,143 Bibi Badal,143 and Karen Ixer143, for the The Princess Alexandra Hospital, Harlow, UK
Donna Duffin,144 Ben Player,144 Helen Hill,144 Jade Cole,144 Jenny Brooks,144 Michelle Davies,144 Rhys Davies,144 Lauren Hunt,144 Emma Thomas,144 and Angharad Williams144, for the University Hospital of Wales, Cardiff, UK
Metod Oblak,145 Mini Thankachen,145 Jamie Irisari,145 Amrinder Sayan,145 and Monica Popescu145, for the West Middlesex Hospital, Isleworth, UK
Cheryl Finch,146 Andrew Jamieson,146 Alison Quinn,146 Joshua Cooper,146 Sarah Liderth,146 and Natalia Waddington146, for the Royal Albert Edward Infirmary, Wigan, UK
Iona Burn,147 Katarina Manso,147 Ruth Penn,147 Julie Tebbutt,147 Danielle Thornton,147 James Winchester,147 Geraldine Hambrook,147 and Pradeep Shanmugasundaram147, for the Stoke Mandeville Hospital, Buckinghamshire, UK
Jayne Craig,148 Kerry Simpson,148 Andrew Higham,148 and Louise Sibbett148, for the Royal Lancaster Infirmary, Lancaster, UK
Sheila Paine,149 Annabel Reed,149 Jo-Anna Conyngham,149 McDonald Mupudzi,149 Rachel Thomas,149 Mary Wright,149 Denise Griffin,149 Richard Partridge,149 Maria Alvarez Corral,149 Nycola Muchenje,149 Mildred Sitonik,149 and Caroline Wrey Brown149, for the Basingstoke and North Hampshire Hospital, Basingstoke, UK
Aaron Butler,150,151 Linda Folkes,150,151 Heather Fox,150,151 Amy Gardner,150,151 David Helm,150,151 Gillian Hobden,150,151 Kirsten King,150,151 Jordi Margalef,150,151 Michael Margarson,150,151 Tim Martindale,150,151 Emma Meadows,150,151 Dana Raynard,150,151 Yvette Thirlwall,150,151 Yolanda Baird,150,151 Raquel Gomez,150,151 Darren Martin,150,151 Luke Hodgson,150,151 Clinton Corin,150,151 Erikka Sidall,150,151 Densie Szabo,150,151 and Sharon Floyd150,151, for the Worthing Hospital, Worthing, UK and St Richard’s Hospital, Chichester, UK
Hannah Davies,152 Karen Austin,152 Olivia Kelsall,152 Hannah Wood,152 Peter Anderson,152 Katie Archer,152 Andrew Burtenshaw,152 Sarah Clayton,152 Naiara Cother,152 Nicholas Cowley,152 Caroline Davis,152 Stephen Digby,152 Alison Durie,152 Alison Harrison,152 Emma Low,152 Michael McAlindon,152 Alex McCurdy,152 Aled Morgan,152 Tobias Rankin,152 Jessica Thrush,152 Helen Tranter,152 Charlie Vigurs,152 and Laura Wild152, for the The Alexandra Hospital, Redditch and Worcester Royal Hospital, Worcester, UK
Thomas Cornell,153 Kate Ralph,153 Sarah Bean,153 Karen Burt,153 Michael Spivey,153 Carol Richards,153 and Rachel Tedstone153, for the Royal Cornwall Hospital, Truro, UK
Siobhain Carmody,154 Xiaobei Zhao,154 Valerie Page,154 Mark Louie Guanco,154 Elvira Hoxha,154 and Camilla Zorloni154, for the Watford General Hospital, Watford, UK
Charlotte Dean,155 Emma Jones,155 Emma Carter,155 Joshua Dunn,155 Thomas Kong,155 Mervin Mahenthran,155 Chris Marsh,155 Maureen Holland,155 Natalie Keenan,155 Mohamed Mahmoud,155 Marc Lyons,155 Joanne Bradley-Potts,155 Helen Wassall,155 and Meghan Young155, for the Macclesfield District General Hospital, Macclesfield, UK
Victoria Maynard,157 Rachel Walker,157 Anil Hormis,157 Dawn Collier,157 Cheryl Graham,157 Vicky Maynard,157 Jake McCormick,157 and Jake Warrington157, for the Rotherham General Hospital, Rotherham, UK
Denise Cosgrove,158 Denise McFarland,158 Judith Ratcliffe,158 and Rob Charnock158, for the Craigavon Area Hospital, County Armagh, Northern Ireland
Inez Wynter,159 Mandy Gill,159 Jill Kirk,159 Paul Paul,159 Valli Ratnam,159 and Sarah Shelton159, for the King’s Mill Hospital, Nottingham, UK
Catherine Jardine,160 Alasdair Hay,160 and Dewi Williams160, for the Dumfries and Galloway Royal Infirmary, Dumfries, UK
Bethan Deacon,161 Latha Durga,161 Meg Hibbert,161 Gareth Kennard-Holden,161 Chrsitopher Woodford,161 Carla Pothecary,161 Lisa Roche,161 Dariusz Tetla,161 Kevin Agravante,161 and Justyna Smeaton161, for the Prince Charles Hospital, Merthyr Tydfil, UK
Alicia Price,162 Alice Thomas,162 Chris Thorpe,162 Ellen Knights,162 and Donna Ward162, for the Ysbyty Gwynedd, Bangor, UK
Shondipon Laha,163 Mark Verlander,163 and Alexandra Williams163, for the Royal Preston Hospital, Preston, UK
Rachel Prout,164 Helen Langton,164 Malcolm Watters,164 Charlotte Hunt,164 and Catherine Novis164, for the The Great Western Hospital, Swindon, UK
Dawn Egginton,166 Michele Clark,166 Sarah Purvis,166 Simon Sinclair,166 and Vicky Collins166, for the University Hospital of North Tees, Stockton on Tees, UK
Bethan Landeg,167 Craig Sell,167 Samantha Coetzee,167 Alistair Gales,167 Igor Otahal,167 Becky Icke,167 Meena Raj,167 Caroline Williams,167 Jill Williams,167 and Lucy Hill167, for the Glangwili General Hospital, Camarthen, UK
Abdul Kayani,168 Bridgett Masunda,168 Prisca Gondo,168 and Nigara Atayeva168, for the Southend University Hospital, Westcliff-on-Sea, UK
Carina Cruz169 and Natalie Pattison169, for the Lister Hospital, Stevenage, UK
Caroline Burnett,170 Jonathan Hatton,170 Elaine Heeney,170 Maria Newton,170 Hassan Al-Moasseb,170 Teresa Behan,170 Jasmine Player,170 Rachael Stead,170 Atideb Mitra,170 and Kirsty Nauyokas170, for the Diana Princess of Wales Hospital, Grimsby, UK
Sally Humphreys,171 Helen Cockerill,171 and Ruth Tampsett171, for the West Suffolk Hospital, Bury St Edmunds, UK
Evgeniya Postovalova,172 Tina Coventry,172 Amanda McGregor,172 Susan Fowler,172 Mike Macmahon,172 Patricia Cochrane,172 and Sandra Pirie172, for the Victoria Hospital, Kirkcaldy, UK
Sarah Hanley,173,174 Asifa Ali,173,174 Megan Brady,173,174 Sam Dale,173,174 Annalisa Dance,173,174 Lisa Gledhill,173,174 Jill Greig,173,174 Kathryn Hanson,173,174 Kelly Holdroyd,173,174 Marie Home,173,174 Tahira Ishaq,173,174 Diane Kelly,173,174 Lear Matapure,173,174 Deborah Melia,173,174 Samantha Mellor,173,174 Ekta Merwaha,173,174 Tonicha Nortcliffe,173,174 Lisa Shaw,173,174 Ryan Shaw,173,174 Tracy Wood,173,174 Lee-Ann Bayo,173,174 Miranda Usher,173,174 Alison Wilson,173,174 Ross Kitson,173,174 Jez Pinnell,173,174 Matthew Robinson,173,174 and Kaitlin Boltwood173,174, for the Calderdale Royal Hospital, Halifax, UK and Huddersfield Royal Infirmary, Huddersfield, UK
Jenny Birch,175 Laura Bough,175 Rebecca Tutton,175 Barbara Winter-Goodwin,175 Josie Goodsell,175 Kate Taylor,175 Patricia Williams,175 Sarah Williams,175 Ashleigh Cave,175 and James Rees175, for the Dorset County Hospital, Dorchester, UK
Janet Imeson-Wood,176 Jacqueline Smith,176 Vishal Amin,176 Komala Karthik,176 Rizwana Kausar,176 Elena Anastasescu,176 Karen Reid,176 Vikram Anumakonda,176 and Ella Stoddart176, for the Russell’s Hall Hospital, Dudley, UK
Carrie Demetriou,177 Charlotte Eckbad,177 Lucy Howie,177 Sarah Mitchard,177 Lidia Ramos,177 Katie White,177 Sarah Hierons,177 Fiona Kelly,177 Alfredo Serrano-Ruiz,177 and Gabrielle Evans177, for the Royal United Hospital, Bath, UK
Liz Nicol,178 Joy Wilkins,178 Kim Hulacka,178 Gabor Debreceni,178 Alison Brown,178 and Vikki Crickmore178, for the St Mary’s Hospital, Newport, UK
Kay Hill179 and Thogulava Kannan179, for the George Eliot Hospital NHS Trust, Nuneaton, UK
Harish Venkatesh,182 Yvonne Bland,182 Istvan Kajtor,182 Lisa Kavanagh,182 Karen Singler,182 and George Linfield-Brown182, for the Frimley Park Hospital, Surrey, UK
Luke Stephen Prockter Moore,183 Marcela Vizcaychipi,183 Laura Martins,183 Luke Moore,183 Rhian Bull,183 and Jaime Carungcong183, for the Chelsea & Westminster NHS Foundation Trust, London, UK
Louise Allen,184 Eva Beranova,184 Alicia Knight,184 Carly Price,184 Sorrell Tilbey,184 Sharon Turney,184 Tracy Hazelton,184 Gabriella Tutt,184 Mansi Arora,184 Salah Turki,184 Emily Sinfield,184 Joanne Deery,184 and Hazel Ramos184, for the Queen Elizabeth the Queen Mother Hospital, Margate, UK
John Allan,187 Tim Geary,187 Alistair Meikle,187 Peter O’Brien,187 Stephen Wood,187 Andrew Clark,187 and Gordon Houston187, for the University Hospital Crosshouse, Kilmarnock, UK
Lorna Murphy,189 Michelle Smythe,189 Alistair Nichol,189 and Kathy Brickell189, for the University College Dublin, St Vincent’s University Hospital, Dublin, Ireland
Liam Botfield,191 Catherine Dexter,191 Aditya Kuravi,191 Joanne Butler,191 Robert Chadwick,191 Poonam Ranga,191 Lisa Richardson,191 Emma Virgilio,191 Maddiha Anwer,191 Atul Garg,191 Donna Botfield,191 and Xana Marriott191, for the Walsall Manor Hospital, Walsall, UK
Keely Stewart,192 Dee Mullan,192 Claire Phillips,192 Jane Gaylard,192 Justyna Nowak,192 and Denise Skinner192, for the Princess Royal Hospital, Brighton, UK
Ailbhe Brady,194 Rebekah Chan,194 Shane McIvor,194 Helena Prady,194 Bijoy Mathew,194 Jeff Little,194 and Tim Furniss194, for the Warrington General Hospital, Warrington, UK
Alison Brown,198 Susan O’Connell,198 Jane Gregory,198 Luigi Barberis,198 Rosemary Harper,198 Tim Smith,198 and Diane Armstrong198, for the Cumberland Infirmary, Carlisle, UK
Angie Bowey,199,200 Anne Cowley,199,200 Andrew Corner,199,200 Judith Highgate,199,200 Claire Rutherfurd,199,200 Jo-Anne Taylor,199,200 Sarah Goodwin,199,200 and Claire Rutherford199,200, for the Eastbourne District General Hospital, East Sussex, UK and Conquest Hospital, East Sussex, UK
Beena Eapen,201 Fiona Trim,201 and Phil Donnison201, for the Salisbury District Hospital, Salisbury, UK
Lisa Armstrong,202 Hayley Bates,202 Emma Dooks,202 Fiona Farquhar,202 Amy Kitching,202 Chantal McParland,202 Sophie Packham,202 and Brigid Hairsine202, for the Airedale General Hospital, Keighley, UK
Anand Patil,203 Premetie Andreou,203 Dawn Hales,203 Megha Mathews,203 Rekha Patel,203 Peter Barry,203 Neil Flint,203 Jessica Hailstone,203 Navneet Ghuman,203 Bethany Leonard,203 and Rachel Lees203, for the Leicester Royal Infirmary, Leicester, UK
Deborah Butcher,204,205 Katy Leng,204,205 Nicola Butterworth-Cowin,204,205 and Susie O’Sullivan204,205, for the Peterborough City Hospital, Peterborough, UK and Hinchingbrooke Hospital, Huntingdon, UK
Alison Ghosh206 and Emma Williams206, for the Colchester General Hospital, Colchester, UK
Tracy Baird,208 Margaret Harkins,208 Jim Ruddy,208 and Joe West208, for the University Hospital Monklands, Airdrie, UK
Joseph Duffield,209 Lewis Mallon,209 Oliver Smith,209 Sara Smuts,209 Andy Campbell,209 Cate Davies,209 Sarah Davies,209 Rachel Hughes,209 Lisa Jobes,209 Victoria Whitehead,209 and Clare Watkins209, for the Wrexham Maelor Hospital, Wrexham, Wales
Fiona Bowman,27 Barry Milligan,27 and Liane McPherson27, for the Royal Hospital for Children, Glasgow, UK
Stella Metherell,210 Nichola Harris,210 Victoria Lake,210 Elizabeth Radford,210 Andy Smallwood,210 Shameer Gopal,210 and Katherine Vassell210, for the New Cross Hospital, Wolverhampton, UK
Dina Bell,211 Rosalind Boyle,211 Katie Douglas,211 Lynn Glass,211 Liz Lennon,211 Austin Rattray,211 Claire Beith,211 and Emma Lee211, for the University Hospital Hairmyres, East Kilbride, UK
Danielle Jones,212 Penny Parsons,212 Ben Attwood,212 Paul Jefferson,212 Mohan Ranganathan,212 Inderjit Atwal,212 Bridget Campbell,212 Angela Day,212 and Camilla Stagg212, for the Warwick Hospital, Warwick, UK
Emma Haynes,213 Cecilia Ahmed,213 Sarah Clamp,213 Julie Colley,213 Risna Haq,213 Anne Hayes,213 Sibet Joseph,213 Zahira Maqsood,213 Samia Hussain,213 Jonathan Hulme,213 Patience Domingos,213 Rita Kumar,213 Manjit Purewal,213 and Becky Taylor213, for the Sandwell General Hospital and City Hospital, Birmingham, UK
Lara Bunni,214 Monica Latif,214 Claire Jennings,214 Shilu Jose,214 Rebecca Marshall,214 Aleksandra Metryka,214 and Gayathri Subramanian214, for the Royal Manchester Children’s Hospital, Manchester, UK
Adam Burgoyne,215 Susan O’Connell,215 Amanda Tyler,215 Joanne Waldron,215 Paula Hilltout,215 and Jayne Evitts215, for the Gloucestershire Royal Hospital, Gloucester, UK
Geraldine Ward,216 Pamela Bremmer,216 Carl Hawkins,216 Sophie Jackman,216 and Michal Ogorek216, for the University Hospitals Coventry & Warwickshire NHS Trust, Coventry, UK
Kylie Ashby,217 Lorraine Thornton,217 Pauline Mercer,217 Matthew Halkes,217 and Adam Revill217, for the Torbay Hospital, Torquay, UK
Bryony Saint,218 Jo Fletcher,218 Kimberley Netherton,218 Manish Chablani,218 Amy Kirkby,218 Amanda Roper,218 and Kinga Szymiczek218, for the Pilgrim Hospital, Lincoln, UK
Isobel Sutherland,219 Linda O’Brien,219 Igor Otahal,219 Joanne Connell,219 Kim Davies,219 Tracy Lewis,219 Zohra Omar,219 and Emma Perkins219, for the Prince Philip Hospital, Lianelli, UK
Lisa Roche,220 Sonia Sathe,220 and Ellie Davies220, for the Princess of Wales Hospital, Llantrisant, UK
Alex Lyon,221 Isheunesu Mapfunde,221 Charlotte Willis,221 Rachael Hitchcock,221 Kathryn Hall,221 and Christopher King221, for the Northampton General Hospital NHS Trust, Northampton, UK
Andrew Fagan,222 Roonak Nazari,222 Lucy Worsley,222 Suzanne Allibone,222 Vidya Kasipandian,222 Amit Patel,222 Parisa Cutting,222 Roman Genetu,222 Ainhi Mac,222 Anthony Murphy,222 Sinead Ward,222 and Fatima Butt222, for the The Christie NHS Foundation Trust, Manchester, UK
Amanda Ayers,223 Wendy Harrison,223 Katherine Mackintosh,223 and Julie North223, for the James Paget University Hospital NHS Trust, Great Yarmouth, UK
Lydia Ashton,224 Rehana Bi,224 Samantha Owen,224 Helen Winmill,224 and Barney Scholefield224, for the Birmingham Children’s Hospital, Birmingham, UK
Hannah Blowing,225 Erin Williams,225 Michaela Duskova,225 Michelle Edwards,225 Alun Rees,225 Helen Thomas,225 Rachel Hughes,225 Igor Otahal,225 Jolene Brooks,225 Janet Phipps,225 and Suzanne Brooks225, for the Withybush General Hospital, Pembrokeshire, Wales
Catherine Dennis,226 Vicki Parris,226 Sinduya Srikaran,226 Anisha Sukha,226 Alistair McGregor,226 and Gerlynn Tiongson226, for the Northwick Park Hospital, London, UK
Katie Adams,227 Benedict Andrew,227 Adam Brayne,227 Sasha Carter,227 Louise Findlay,227 Emma Fisher,227 Peter Jackson,227 Duncan Kaye,227 Juliet Parkin,227 Victoria Tuckey,227 Jane Hunt,227 Nicholas Love,227 Lynne van Koutrick,227 and Ashley Hanson227, for the North Devon District Hospital, Barnstaple, UK
Kathy Dent,228 Elizabeth Horsley,228 Sandra Pearson,228 Sue Spencer,228 Dorothy Hutchinson,228 Jasmine Player,228 Dorota Potoczna,228 Muhammad Nauman Akhtar,228 Lisa-Jayne Cottam,228 Kirsty Nauyokas,228 and Jack Sanders228, for the Scunthorpe General Hospital, Scunthorpe, UK
Sara Mingo Garcia,229 Glykeria Pakou,229 Cynthia Diaba,229 Helder Filipe,229 Lincy John,229 Amitaa Maharajh,229 Mark de Neef,229 Daniel Martin,229 Christine Eastgate,229 and Poh Choo Teoh229, for the Royal Free Hospital, London, UK
Polly Rice,231 Tim Smith,231 Melanie Clapham,231 Rachel Mutch,231 Luigi Barberis,231 Rosemary Harper,231 Hannah Craig,231 and Una Poultney231, for the West Cumberland Hospital, Whitehaven, UK
Karen Burns232 and Andrew Higham232, for the Furness General Hospital, Barrow-in-Furness, UK
Sophie Twiss,233 Janet Barton,233 Linsha George,233 Clare Harrop,233 Sherly Mathew,233 and David Justin Wright233, for the Liverpool Heart and Chest Hospital, Liverpool, UK
Rachel Harrison,234 Jordan Toohie,234 Ben Chandler,234 Alison Turnbull,234 Janine Mallinson,234 and Kerry Elliott234, for the Scarborough General Hospital, Scarborough, UK
Rebecca Wolf-Roberts,235 Helen Tench,235 Igor Otahal,235 Maria Hobrok,235 Ronda Loosley,235 Heather McGuinness,235 and Tanya Sims235, for the Bronglais General Hospital, Aberystwyth, UK
Deborah Afolabi,236 Kathryn Sian Allison,236 Taya Anderson,236 Rachael Dore,236 Dawn Jones,236 Naomi Rogers,236 Paula Saunderson,236 Jennifer Whitbread,236 Laura O’Malley,236 Laura Rad,236 and Daniel Hawcutt236, for the Alder Hey Children’s Hospital, Liverpool, UK
Jonathan Aldridge,237 Melanie Tolson,237 and Sweyn Garrioch237, for the Borders General Hospital, Melrose, UK
Joanne Tomlinson238 and Michael Grosdenier238, for the Leighton Hospital, Cheshire, UK
David Loader,239 Ritoo Kapoor,239 and Gemma Hector239, for the Kent & Canterbury Hospital, Canterbury, UK
Joslan Scherewode,240 Chunda Sri-Chandana,240 Lorraine Stephenson,240 and Sarah Marsh240, for the Harrogate and District NHS Foundation Trust, Harrogate, UK
Arnold Dela Rosa,241 Shaman Jhanji,241 Thomas Bemand,241 Ryan Howle,241 Ravishankar Rao Baikady,241 Benjamin Thomas,241 Ethel Black,241 and Kate Tatham241, for the The Royal Marsden Hospital, London, UK
Sambasivarao Gurram,242 Ekaterina Watson,242 Vicki Parris,242 Sheena Quaid,242 and Alistair McGregor242, for the Ealing Hospital, Southall, UK
Anne Saunderson,243 Rachel O’Brien,243 Sam Moultrie,243 Jen Service,243 Clare Cheyne,243 Miranda Odam,243 and Alison Wiliams243, for the St John’s Hospital Livingston, Livingston, UK
Nicky Barnes,244 Peter Csabi,244 Joana Da Rocha,244 and Louika Glynou244, for the Wexham Park Hospital, Slough, UK
Amy Huffenberger,245 Jade Bryant,245 Amy Pickard,245 Nicholas Roe,245 Arianna Bellini,245 Anton Mayer,245 Amy Burrow,245 Natalie Colley,245 Jayne Evans,245 Alex Howlett,245 and Zeinab Khalifeh245, for the Sheffield Children’s Hospital, Sheffield, UK
Jerldine Pryce,246 Claire Gorman,246 Amy Easthope,246 Rebecca Brady,246 Elizabeth Timlick,246 Pierre Antoine,246 and Abhinhav Gupta246, for the Homerton University Hospital Foundation NHS Trust, London UK
John Hardy,247 Henry Houlden,247 Eleanor Moncur,247 Arianna Tucci,247 Eamon Raith,247 Ambreen Tariq,247 and David Brealey247, for the National Hospital for Neurology and Neurosurgery, London, UK
Emma Tagliavini,248 Becky Ramsay,248 Katy Fidler,248 Kevin Donnelly,248 and Rebecca Hollis248, for the The Royal Alexandra Children’s Hospital, Brighton, UK
Jocelyn Barr,249 Elizabeth Boyd,249 Val Irvine,249 Ben Shelley,249 Julie Buckley,249 Charlene Hamilton,249 and Kathryn Valdeavella249, for the Golden Jubilee National Hospital, Clydebank, UK

SCOURGE Consortium:

Javier Abellan,250,251 René Acosta-Isaac,252 Jose María Aguado,253,254,255,256 Carlos Aguilar,257 Sergio Aguilera-Albesa,258,259 Abdolah Ahmadi Sabbagh,260 Jorge Alba,261 Sergiu Albu,262,263,264 Karla A. M. Alcalá-Gallardo,265 Julia Alcoba-Florez,266 Sergio Alcolea Batres,267 Holmes Rafael Algarin-Lara,268,269 Virginia Almadana,270 Julia Almeida,271,272,273,274 Berta Almoguera,37,275 María R. Alonso,276 Nuria Alvarez,276 Yady Álvarez-Benítez,268,269 Felipe Álvarez-Navia,277,278 Rodolfo Alvarez-Sala Walther,267 Álvaro Andreu-Bernabeu,255,279 Maria Rosa Antonijoan,280 Eunate Arana-Arri,281,282 Carlos Aranda,283,284 Celso Arango,255,279,285 Carolina Araque,286,287 Nathalia K. Araujo,288 Izabel M. T. Araujo,289 Ana C. Arcanjo,290,291,292 Ana Arnaiz,33,34,35 Francisco Arnalich Fernández,293 María J. Arranz,294 José Ramon Arribas Lopez,293 Maria-Jesus Artiga,295 Yubelly Avello-Malaver,296 Carmen Ayuso,37,275 Ana Margarita Baldión-Elorza,296 Belén Ballina Martín,260 Raúl C. Baptista-Rosas,297,298,299 Andrea Barranco-Díaz,269 María Barreda-Sánchez,300,301 Viviana Barrera-Penagos,296 Moncef Belhassen-Garcia,278,302 Enrique Bernal,300 David Bernal-Bello,303 Joao F. Bezerra,304 Marcos A. C. Bezerra,305 Natalia Blanca-López,306 Rafael Blancas,307 Lucía Boix-Palop,308 Alberto Borobia,309 Elsa Bravo,310 María Brion,311,312 Óscar Brochado-Kith,256,313 Ramón Brugada,312,314,315,316 Matilde Bustos,317 Alfonso Cabello,318 Juan J. Caceres-Agra,319 Esther Calbo,320 Enrique J. Calderón,321,322,323 Shirley Camacho,324 Marcela C. Campos,290 Yolanda Cañadas,284 Cristina Carbonell,277,278 Servando Cardona-Huerta,36 Antonio Augusto F. Carioca,325 Maria Sanchez Carpintero,283,284 Carlos Carpio Segura,267 Thássia M. T. Carratto,326 José Antonio Carrillo-Avila,327 Maria C. C. Carvalho,328 Carlos Casasnovas,37,329,330 Luis Castano,37,281,331,332,333 Carlos F. Castaño,283,284 Jose E. Castelao,334 Aranzazu Castellano Candalija,335 María A. Castillo,324 Francisco C. Ceballos,313 Jessica G. Chaux,287 Walter G. Chaves-Santiago,287,336 Sylena Chiquillo-Gómez,268,269 Marco A. Cid-Lopez,265 Oscar Cienfuegos-Jimenez,36 Rosa Conde-Vicente,337 M. Lourdes Cordero-Lorenzana,338 Dolores Corella,339,340 Almudena Corrales,30,31 Jose L. Cortes-Sanchez,36,341 Marta Corton,37,275 Tatiana X. Costa,342 Raquel Cruz,37,44,45,343 Marina S. Cruz,288 Luisa Cuesta,344 Gabriela C. R. Cunha,345 Gabriela V. da Silva,289 David Dalmau,320,346 Raquel C. S. Dantas-Komatsu,288 M. Teresa Darnaude,347 Raimundo de Andrés,348 Jéssica N. G. de Araújo,349 Carmen de Juan,350 Juan De la Cruz Troca,322,351,352 Carmen de la Horra,323 Ana B. de la Hoz,281 Alba De Martino-Rodríguez,353,354 Julianna Lys de Sousa Alves Neri,355 Victor del Campo-Pérez,356 Juan Delgado-Cuesta,357 Covadonga M. Diaz-Caneja,255,279,285 Anderson Díaz-Pérez,269 Aranzazu Diaz de Bustamante,347 Beatriz Dietl,320 Silvia Diz-de Almeida,37,44 Manoella do Monte Alves,358,359 Elena Domínguez-Garrido,360 Katiusse A. dos Santos,328 Alice M. Duarte,289 Jose Echave-Sustaeta,361 Rocío Eiros,362 César O. Enciso-Olivera,286,287 Gabriela Escudero,363 Pedro Pablo España,364 Gladys Mercedes Estigarribia Sanabria,365 María Carmen Fariñas,33,34,35 Marianne R. Fernandes,366,367 Ramón Fernández,33,368 Lidia Fernandez-Caballero,37,275 Ana Fernández-Cruz,369 María J. Fernandez-Nestosa,370 Uxía Fernández-Robelo,371 Amanda Fernández-Rodríguez,256,313 Marta Fernández-Sampedro,33,34,35 Ruth Fernández-Sánchez,37,275 Tania Fernández-Villa,372 Silvia Fernández Ferrero,260 Yolanda Fernández Martínez,260 Carmen Fernéndez Capitán,335 Patricia Flores-Pérez,373 Vicente Friaza,322,323 Lácides Fuenmayor-Hernández,269 Marta Fuertes Núñez,260 Victoria Fumadó,374 Ignacio Gadea,375 Lidia Gagliardi,283,284 Manuela Gago-Domínguez,45,46 Natalia Gallego,38 Cristina Galoppo,376 Inés García,37,275 Mercedes García,283,284 Leticia García,283,284 Carlos Garcia-Cerrada,37,250,251 Aitor García-de-Vicuña,281,331 Josefina Garcia-García,300 Irene García-García,309 Carmen García-Ibarbia,33,34,35 Andrés C. García-Montero,377 Ana García-Soidán,378 Elisa García-Vázquez,300 María Carmen García Torrejón,251,379 Emiliano Garza-Frias,36 Angela Gentile,376 Belén Gil-Fournier,380 Javier Gómez-Arrue,353,354 Mario Gómez-Duque,287,336 Luis Gómez Carrera,267 María Gómez García,343 Ángela Gómez Sacristán,381 Anna González-Neira,276 Javier González-Peñas,255,279,285 Manuel Gonzalez-Sagrado,337 Beatriz González Álvarez,353,354 Fernan Gonzalez Bernaldo de Quirós,382 Hugo Gonzalo Benito,383 Oscar Gorgojo-Galindo,384 Miguel Górgolas,318 Florencia Guaragna,376 Genilson P. Guegel,385 Beatriz Guillen-Guio,30 Encarna Guillen-Navarro,300,386,387,388 Pablo Guisado-Vasco,361 Juan F. Gutiérrez-Bautista,389 Luz D. Gutierrez-Castañeda,287,390 Sarah Heili-Frades,391 Estefania Hernandez,392 Luis D. Hernandez-Ortega,299,393 Guillermo Hernández-Pérez,277 Rebeca Hernández-Vaquero,394 Cristina Hernández Moro,260 Belen Herraez,276 M. Teresa Herranz,300 María Herrera,283,284 María José Herrero,395,396 Antonio Herrero-Gonzalez,397 Juan P. Horcajada,256,263,398,399,400 Natale Imaz-Ayo,281 Maider Intxausti-Urrutibeaskoa,401 María Íñiguez,402 Rafael H. Jacomo,403 Rubén Jara,300 Perez Maria Jazmin,376 Ángel Jiménez,283,284 Pilar Jiménez,389 Ignacio Jiménez-Alfaro,404 María A. Jimenez-Sousa,256,313 Iolanda Jordan,322,405,406 Rocío Laguna-Goya,407,408 Daniel Laorden,267 María Lasa-Lazaro,407,408 María Claudia Lattig,324,409 Ailen Lauriente,376 Anabel Liger Borja,410 Lucía Llanos,411 Amparo López-Bernús,277,278 Esther Lopez-Garcia,322,351,352,412 Rosario Lopez-Rodriguez,37,275 Miguel A. López-Ruz,413,414,415 Eduardo López Granados,37,416,417 Leonardo Lorente,418 José E. Lozano,419 María Lozano-Espinosa,410 Andre D. Luchessi,420 Ignacio Mahillo,31,421,422 Esther Mancebo,407,408 Carmen Mar,364 Cristina Marcelo Calvo,335 Miguel Marcos,277,278 Alba Marcos-Delgado,423 Alicia Marín Candon,309 Pablo Mariscal Aguilar,267 María M. Martín,424 María Dolores Martín,425 Vicente Martín,322,423 Marta Martin-Fernandez,426 Caridad Martín-López,410 José-Ángel Martín-Oterino,277,278 Laura Martin-Pedraza,306 María Martín-Vicente,313 Amalia Martinez,427 Ricardo Martínez,392 Juan José Martínez,37,330 Silvia Martínez,33,35 Eleno Martínez-Aquino,428 Óscar Martínez-González,307 Iciar Martinez-Lopez,429,430 Oscar Martinez-Nieto,296,409 Pedro Martinez-Paz,383 Angel Martinez-Perez,431 Andrea Martínez-Ramas,37,275 Michel F. Martinez-Resendez,36 Violeta Martínez Robles,260 Laura Marzal,37,275 Juliana F. Mazzeu,432,433,434 Jeane F. P. Medeiros,288 Kelliane A. Medeiros,435,436 Francisco J. Medrano,321,322,323 Xose M. Meijome,437,438 Natalia Mejuto-Montero,439 Ana Méndez-Echevarria,293 Humberto Mendoza Charris,269,310 Eleuterio Merayo Macías,440 Fátima Mercadillo,441 Arieh R. Mercado-Sesma,299,393 Pablo Minguez,37,275 Antonio J. J. Molina,322,423 Elena Molina-Roldán,442 Juan José Montoya,392 Vitor M. S. Moraes,326 Patricia Moreira-Escriche,350 Xenia Morelos-Arnedo,269,310 Antonio Moreno-Docón,300 Junior Moreno-Escalante,269 Victor Moreno Cuerda,250,251 Alberto Moreno Fernández,335 Rubén Morilla,323,443 Patricia Muñoz García,31,255,444 Pablo Neira,376 Julian Nevado,37,38,445 Israel Nieto-Gañán,378 Joana F. R. Nunes,290 Rocio Nuñez-Torres,276 Antònia Obrador-Hevia,446,447 J. Gonzalo Ocejo-Vinyals,33,35 Virginia Olivar,376 Silviene F. Oliveira,432,433,434,448,449 Lorena Ondo,37,275 Alberto Orfao,271,272,273,274 Luis Ortega,450 Eva Ortega-Paino,295 Fernando Ortiz-Flores,33,35 Rocio Ortiz-Lopez,36,451 José A. Oteo,261,402 Harry Pachajoa,452,453 Manuel Pacheco,392 Fredy Javier Pacheco-Miranda,269 Irene Padilla Conejo,260 Sonia Panadero-Fajardo,327 Mara Parellada,255,279,285 Roberto Pariente-Rodríguez,378 Estela Paz-Artal,407,408,454 Germán Peces-Barba,31,455 Miguel S. Pedromingo Kus,456 Celia Perales,375 Patricia Perez,457 César Pérez,458 Gustavo Perez-de-Nanclares,281,331 Felipe Pérez-García,459,460 Patricia Pérez-Matute,402 Alexandra Pérez-Serra,312,314 M. Elena Pérez-Tomás,300 Teresa Perucho,461 Lisbeth A. Pichardo,260 Susana M. T. Pinho,435,462,463 Mel·lina Pinsach-Abuin,312,314 Luz Adriana Pinzón,287,336 Guillermo Pita,276 Francesc Pla-Junca,37,464 Laura Planas-Serra,37,330 Ericka N. Pompa-Mera,465 Gloria L. Porras-Hurtado,392 Aurora Pujol,37,330,466 María Eugenia Quevedo Chávez,268,269 Maria Angeles Quijada,280,467 Inés Quintela,343 Diana Ramirez-Montaño,468 Soraya Ramiro León,380 Pedro Rascado Sedes,394 Delia Recalde,353,354 Emma Recio-Fernández,402 Salvador Resino,256,313 Adriana P. Ribeiro,435,436,463 Carlos S. Rivadeneira-Chamorro,287 Diana Roa-Agudelo,296 Montserrat Robelo Pardo,394 Marilyn Johanna Rodriguez,287 Fernando Rodriguez-Artalejo,322,351,352,412 Marena Rodríguez-Ferrer,269 Carlos Rodriguez-Gallego,32,469 José A. Rodriguez-Garcia,260 María A. Rodriguez-Hernandez,317 Antonio Rodriguez-Nicolas,389 Agustí Rodriguez-Palmero,330,470 Emilio Rodríguez-Ruiz,45,394 Paula A. Rodriguez-Urrego,296 Belén Rodríguez Maya,250 German Ezequiel Rodriguez Novoa,376 Federico Rojo,274,471 Andrea Romero-Coronado,269 Filomeno Rondón García,260 Lidia S. Rosa,472 Antonio Rosales-Castillo,473 Cladelis Rubio,474,475 María Rubio Olivera,283,284 Montserrat Ruiz,37,330 Francisco Ruiz-Cabello,389,414,476 Eva Ruiz-Casares,461 Juan J. Ruiz-Cubillan,33,35 Javier Ruiz-Hornillos,284,477,478 Pablo Ryan,479,480,481 Hector D. Salamanca,286,287 Lorena Salazar-García,324 Giorgina Gabriela Salgueiro Origlia,335 Pedro-Luis Sánchez,278,362 Clara Sánchez-Pablo,362 Olga Sánchez-Pernaute,482 Antonio J. Sánchez López,483 María Concepción Sánchez Prados,267 Javier Sánchez Real,260 Jorge Sánchez Redondo,250,484 Cristina Sancho-Sainz,401 Anna Sangil,308 Arnoldo Santos,458 Ney P. C. Santos,366 Agatha Schlüter,37,330 Sonia Segovia,464,485,486 Alex Serra-Llovich,346 Fernando Sevil Puras,257 Marta Sevilla Porras,37,38 Miguel A. Sicolo,487,488 Vivian N. Silbiger,420 Nayara S. Silva,349 Fabiola T. C. Silva,290 Cristina Silván Fuentes,37 Jordi Solé-Violán,31,489,490 José Manuel Soria,431 Jose V. Sorlí,339,340 Renata R. Sousa,432 Juan Carlos Souto,252 Karla S. C. Souza,328 Vanessa S. Souza,345 John J. Sprockel,287,336 José Javier Suárez-Rama,343 David A. Suarez-Zamora,296 Xiana Taboada-Fraga,439 Eduardo Tamayo,384,491 Alvaro Tamayo-Velasco,492 Juan Carlos Taracido-Fernandez,397 Nathali A. C. Tavares,493 Carlos Tellería,353,354 Jair Antonio Tenorio Castaño,37,38,445 Alejandro Teper,376 Juan Torres-Macho,494 Lilian Torres-Tobar,287 Ronald P. Torres Gutiérrez,456 Jesús Troya,479 Miguel Urioste,441 Juan Valencia-Ramos,495 Agustín Valido,270,496 Juan Pablo Vargas Gallo,497,498 Belén Varón,499 Romero H. T. Vasconcelos,493 Tomas Vega,500 Santiago Velasco-Quirce,501 Valentina Vélez-Santamaría,329,330 Virginia Víctor,283,284 Julia Vidán Estévez,260 Miriam Vieitez-Santiago,33,35 Carlos Vilches,502 Lavinia Villalobos,260 Felipe Villar,455 Judit Villar-Garcia,503,504,505 Cristina Villaverde,37,275 Pablo Villoslada-Blanco,402 Ana Virseda-Berdices,313 Zuleima Yáñez,269 Antonio Zapatero-Gaviria,506 Ruth Zarate,507 Sandra Zazo,471 Miguel López de Heredia,37 Ingrid Mendes,37 Rocío Moreno,37 Esther Sande,37,44,45 Pablo Lapunzina,37,38,445 and Angel Carracedo37,44,45,46,343

ISARICC Investigators

James Lee,560 Daniel Plotkin,560 and Seán Keating11, for the Data Analysis and Management Team
Cara Donegan561 and Rebecca G. Spencer561, for the Project Administration Team
:
James Lee,560 Daniel Plotkin,560 and Seán Keating11, for the Data Analysis and Management Team
Cara Donegan561 and Rebecca G. Spencer561, for the Project Administration Team
Chloe Donohue556 and Hayley Hardwick16, for the Project Management Team

The 23andMe COVID-19 Team:

Extended data

is available for this paper at 10.1038/s41586-023-06034-3.

Supplementary information

The online version contains supplementary material available at 10.1038/s41586-023-06034-3.

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