Nature | Vol 621 | 7 September 2023 | E7 Matters arising A second update on mapping the human genetic architecture of COVID-19 Investigating the role of host genetic factors in COVID-19 severity and susceptibility can inform our understanding of the underlying bio- logical mechanisms that influence adverse outcomes and drug devel- opment1,2. Here we present a second updated genome-wide association study (GWAS) on COVID-19 severity and infection susceptibility to SARS-CoV-2 from the COVID-19 Host Genetic Initiative (data release 7). We performed a meta-analysis of up to 219,692 cases and over 3 million controls, identifying 51 distinct genome-wide significant loci— adding 28 loci from the previous data release2. The increased number of candidate genes at the identified loci helped to map three major biological pathways that are involved in susceptibility and severity: viral entry, airway defence in mucus and type I interferon. We conducted a meta-analysis for 3 phenotypes across 82 studies from 35 countries, including 36 studies of individuals with non- European ancestry (Fig. 1, Supplementary Figs. 1 and 2 and Supplemen- tary Table 1): critical illness (respiratory support or death; 21,194 cases), hospitalization (49,033 cases) and SARS-CoV-2 infection (219,692 cases). Most of the studies were collected before the widespread introduction of COVID-19 vaccination. We found 30, 40 and 21 loci that are associated with critical illness, hospitalization and infection due to SARS-CoV-2, respectively, for a total of 51 distinct genome-wide significant loci across all three phenotypes (P < 5 × 10−8; Fig. 2, Supplementary Fig. 3 and Supplementary Table 2), adding 28 genome-wide significant loci to the 23 previously identified by the COVID-19 Host Genomics Initia- tive (HGI; data release 6)1,2. We observed a median increase of 2.9-fold in statistical power across lead variants owing to a median increase of 1.6-fold in effective sample sizes from the previous release (Supplemen- tary Table 3). After correcting for the number of phenotypes examined, 46 loci remained significant (P < 1.67 × 10−8). Of the 28 additional loci, 6 loci were originally reported by the GenOMICC study3, which also contributed to the current meta-analysis, and 9 other loci were identi- fied by the new GenOMICC meta-analysis4 during the preparation of this paper. We found nine more loci that reached genome-wide sig- nificance, but we excluded them as they were probably false positives, as determined using a leave-most-significant-biobank-out analysis (Supplementary Table 4 and Supplementary Note). Comparing the effect sizes and statistical significance between the previous2 and cur- rent analysis indicated that all of the previously identified loci were replicated and showed an increase in statistical significance (Supple- mentary Fig. 4). Using our previously developed two-class Bayesian model for classifying loci as being more likely involved in infection susceptibility or severity2, we determined that 36 loci are substantially more likely (higher than 99% posterior probability) to affect disease severity (hospitalization) and 9 loci are substantially more likely to influence susceptibility to SARS-CoV-2 infection, while the remaining 6 loci could not be classified (Supplementary Fig. 5, Supplementary Table 5 and Supplementary Note). We observed that the 1q22 locus (lead variant: rs12752585:G>A) showed significant effect-size heterogeneity across ancestries (Phet < 9.80 × 10 −4 = 0.05/51), whereas the previously reported heterogenous locus (FOXP4) remained at the same level of sig- nificance as before2, despite an increase in sample size (Phet = 2.01 × 10 −3; Supplementary Fig. 6 and Supplementary Table 6). We found signifi- cant observed-scale single-nucleotide polymorphism heritabilities of all the three phenotypes (1.2–8.2%, P < 0.0001). We also estimated liability-scale heritabilities for a range of population prevalences (Sup- plementary Fig. 7, Supplementary Table 7 and Supplementary Note). To better understand the biological mechanisms underlying COVID-19 susceptibility and severity, we further characterized candidate causal genes by mapping them onto biological pathways and performing a phenome-wide association analysis (Extended Data Fig. 1, Supplemen- tary Fig. 8 and Supplementary Tables 2, 8 and 9). In total, 15 out of 51 loci could be linked to three major pathways involved in susceptibility and severity defined by expert-driven classification (Supplementary Note): (1) viral entry; (2) entry defence in airway mucus; and (3) type I interferon response. Moreover, the phenome-wide association analysis identified nine loci involved in the upkeep of healthy lung tissue. First, five loci include candidate causal genes involved in the viral entry pathway (Extended Data Fig. 1a), such as previously reported SLC6A20 (3p21.31), ABO (9q34.2), SFTPD (10q22.3) and ACE2 (Xp22.2), as well as TMPRSS2 (21q22.3), which was also identified in the data release 7. We found that the lead variant rs9305744:G>A, an intronic variant of TMPRSS2, is protective against critical illness (odds ratio (OR) = 0.92, 95% CI = 0.89–0.95, P = 1.4 × 10−8) and is in linkage dise- quilibrium with the missense variant rs12329760:C>T (p.Val197Met; r2 = 0.68). SARS-CoV-2 uses the serine protease TMPRSS2 for viral spike protein priming, as well as the previously reported ACE2 for host cell entry which functionally interacts with SLC6A20 (refs. 5,6). Notably, the previously reported association between ABO blood groups and susceptibility could be attributed to the interference of anti-A and anti-B antibodies with the spike protein, potentially interfering with viral entry7. Furthermore, the previously reported SFTPD encodes pul- monary surfactant protein D (SP-D), which contributes dually to the lung’s innate immune molecules and viral-entry response in pulmonary epithelia8,9 along with other genes for airway defence. Second, four loci contain candidate causal genes for entry defence in the airway mucus (Extended Data Fig. 1b), such as previously reported MUC1/THBS3 (1q22) and MUC5B (11p15.5) as well as novel MUC4 (3q29) and MUC16 (19p13.2). We found that the novel lead variants https://doi.org/10.1038/s41586-023-06355-3 Received: 23 December 2022 Accepted: 21 June 2023 Published online: 6 September 2023 Open access Check for updates *A list of authors and their affiliations appears at the end of the paper. The COVID-19 Host Genetics Initiative * arising from: COVID-19 Host Genetics Initiative. Nature https://doi.org/10.1038/s41586-021- 03767-x (2021) E8 | Nature | Vol 621 | 7 September 2023 Matters arising rs2260685:T>C in MUC4 (intronic variant; in linkage disequilibrium (r2 = 0.65) with a missense variant rs2259292:C>T (p.Gly4324Asp)) and rs73005873:G>A in MUC16 (intronic variant) increase the risk of SARS-CoV-2 infection (OR = 1.03 and 1.03, 95% CI = 1.02–1.04 and 1.02–1.04, P = 4.1 × 10−8 and 6.4 × 10−10, respectively). Moreover, the previously reported locus 1q22 contains an intergenic lead variant rs12752585:G>A that decreases the risk of infection (OR = 0.98, 95% CI = 0.97–0.98, P = 1.5 × 10−11) and increases MUC1 expression in the oesophagus mucosa in GTEx v8 (P = 5.2 × 10−9). Notably, the 1q22 locus also contains an independent lead variant, rs35154152:T>C, a missense variant (p.Ser279Gly) of THBS3, that decreases the risk of hospitaliza- tion (OR = 0.88, 95% CI = 0.86–0.90, P = 5.6 × 10−22) but not infection (P = 5.7 × 10−4), suggesting potential distinct mechanisms in the locus. Consistent with these association patterns, MUC1, MUC4 and MUC16 are three known major transmembrane mucins of the respiratory tracts that prevent microbial invasion, whereas previously reported MUC5B, together with nearby MUC5AC, are primary structural components of airways mucus that enable mucociliary clearance of pathogens10. Third, six loci contain candidate causal genes that are linked to the type I interferon pathway (Extended Data Fig. 1c), such as previously reported IFNAR2 (21q22.11), OAS1 (12q24.13) and TYK2 (19p13.2), as well as additionally identified JAK1 (1p31.3), IRF1 (5q31.1) and IFNα-coding genes (9p21.3). Previous studies have reported additional genes in this pathway: TLR7 (refs. 11,12) and DOCK2 (ref. 13). Here we found that the lead variant rs28368148:C>G, a missense variant (p.Trp164Cys) of IFNA10 located within the IFNα gene cluster, increases the risk of critical illness (OR = 1.56, 95% CI = 1.38–1.77, P = 3.7 × 10−12). IFNα is one of the type I interferons that binds specifically to the IFNα receptor consist- ing of IFNAR1–IFNAR2 chains, in which mutations are also known to increase the risk of hospitalization and critical illness. In the genes that enable signalling downstream of IFNAR, we identified that the lead variant rs11208552:G>T, an intronic variant of JAK1, is protective against critical illness and hospitalization (OR = 0.92 and 0.95, 95% CI = 0.89–0.94 and 0.93–0.96, P = 5.5 × 10−10 and 2.2 × 10−9, respectively). This variant was previously reported to decrease lymphocyte counts14 (β = −0.016, P = 5.5 × 10−15) and increase the JAK1 expression in the thy- roid in GTEx15 (P = 6.1 × 10−23). JAK1 and previously reported TYK2 are Janus kinases ( JAKs) that are required for type I interferon-induced JAK–STAT signalling. JAK inhibitors are used to treat patients with severe COVID-19 (ref. 16). Furthermore, downstream of JAK–STAT signalling, we found that the lead variant rs10066378:T>C, located 67 kb upstream of IRF1, increases the risk of critical illness and hospitalization (OR = 1.09 and 1.07, 95% CI = 1.06–1.13 and 1.05–1.09, P = 2.7 × 10−9 and 3.74 × 10−10, respectively). Furthermore, the phenome-wide association analysis identified nine loci previously associated with lung function and respiratory dis- eases. These loci contain genes involved in the upkeep of healthy lung tissue such as previously reported FOXP4 (6p21.1), SFTPD (10q22.3), MUC5B (11p15.5) and DPP9 (19p13.3), as well as additionally identi- fied CIB4 (2p23.3), NPNT (4q24), ZKSCAN1 (7q22.1), ATP11A (13q34) and PSMD3 (17q21.1). For example, we found that three lead variants, rs1662979:G>T (intronic variant of CIB4), rs34712979:G>A (splice region variant of NPNT) and rs2897075:C>T (intronic variant of ZKSCAN1), are significantly associated with hospitalization (OR = 1.05, 0.94 and 1.05, 95% CI = 1.03–1.07, 0.92–0.96 and 1.03–1.07, P = 5.6 × 10−9, 3.8 × 10−8 and 8.9 × 10−9, respectively) and lung function (FEV1/FVC)17, similar to the previously reported lead variant rs3934643:G>A (intronic vari- ant of SFTPD). Notably, whereas the alleles associated with increased risk of COVID-19 severity of rs1662979 and rs3934643 decrease lung function (β = −0.013 and −0.025, P = 5.3 × 10−8 and 6.3 × 10−10), those of rs34712979 and rs2897075 increase lung function (β = 0.068 and 0.023, P = 4.2 × 10−134 and 1.6 × 10−20, respectively). Likewise, we found lead variants that were significantly associated with hospitalization and idiopathic pulmonary fibrosis18,19, such as the aforementioned rs2897075 and rs12585036:C>T (intronic variant of ATP11A; OR = 1.10, 95% CI = 1.08–1.12, P = 3.2 × 10−21), in addition to the previously reported rs35705950:G>T (promoter variant of MUC5B). Whereas the COVID-19 severity risk-increasing alleles of rs2897075 and rs12585036 increase the risk of idiopathic pulmonary fibrosis (OR = 1.12 and 1.27, P = 3.0 × 10−14 and 7.0 × 10−9, respectively), those of rs35705950 decreases the risk (OR = 0.50, P = 3.9 × 10−80). These results highlight the complex pleio- tropic relationships between COVID-19 severity, lung function and respiratory diseases. We used genetic correlations and Mendelian randomization anal- yses to identify potential causal effects of risk factors on COVID-19 phenotypes (Supplementary Fig. 9 and Supplementary Tables 10 and 11). In total, 14 novel genetic correlations and 10 novel robust exposure-COVID-19 trait pairs showed evidence of causal associations PC 2PC 1 PC3 AFR AMR EAS EUR MID SAS i ii iii iv v vi vii i ii iii iv v vi vii Critically ill COVID-19+ Reported SARS-CoV-2 infection Hospitalized COVID-19+ EUR AFR AMR EAS MID OTH Not shown SAS Effective sample size by genetic ancestryAnalysis type Effective sample size 200,000 20,000 2,000 200 16,374 23,812 41,645 53,217 85,496 745,178 669 a b Fig. 1 | Overview of the contributing studies in the HGI data release 7. a, Geographical overview of the studies contributing to the COVID-19 HGI and the composition by major ancestry groups. Populations are defined as Middle Eastern (MID), South Asian (SAS), East Asian (EAS), African (AFR), admixed American (AMR), European (EUR) and other (OTH). b, A principal component analysis (PCA) highlights the population structure and the sample ancestry of the individuals participating in the COVID-19 HGI. Per-cohort PCA results are provided in Supplementary Fig. 2. This figure was reproduced from the original publication by the COVID-19 HGI1 with modifications reflecting the updated analysis from data release 7. Nature | Vol 621 | 7 September 2023 | E9 (Supplementary Note). In particular, smoking initiation and the num- ber of cigarettes per day were positively correlated with severity and susceptibility phenotypes; Mendelian randomization indicated that smoking was causally associated with increased risk of COVID-19, further highlighting the role of the healthy lung tissue in COVID-19 severity. Moreover, genetically instrumented higher glomerular filtra- tion rate (indicative of better kidney function) was associated with a lower risk of COVID-19 critical illness, whereas genetically predi- cted chronic kidney disease was associated with an increased risk of COVID-19 critical illness, suggesting that better kidney function would be beneficial for a lower risk of COVID-19 severity. In summary, we have substantially expanded the current knowl- edge of host genetics for COVID-19 susceptibility and severity by fur- ther doubling the case numbers from the previous data release2 and identifying 28 additional loci. The increased number of loci enables us to map genes to pathways that are involved in viral entry, airway defence and immune system response. Notably, we observed sever- ity loci mapped to type I interferon pathway, while susceptibility loci Suggested phenotypic impact 21 rs9305744 FAM3B TMPRSS2 rs78143111 ATP5PO IFNAR2 IL10RB IL10RB-DT MRPS6 rs9636867 IFNAR2 IL10RB IL10RB-DT 16 rs117169628 AC137932.1 ACSF3 CDH15 LINC02138 SLC22A31 ZNF778 13 rs12585036 AL139384.1 ATP11A 14 rs184781326 FOXG1 12 rs5023077 FBRSL1 rs10850094 OAS1 OAS2 OAS3 11 rs7949972 CAT ELF5 rs35705950 MUC5B 8 rs2875968 CA8 LINC01301 RAB2A COVID-19 phenotypes 19 KCNC3 NAPSA NAPSB NR1H2 POLD1 FUT2 IZUMO1 MAMSTR NTN5 RASIP1 SEC1P FOXA3 AC011529.1 ICAM3 RAVER1 TYK2 MUC16 DPP9 rs1405655 rs516316 rs71175234 rs34536443 rs73005873 rs12610495 Not classied Infection susceptibility Disease severity 17 rs241044 AC005829.2 AC126544.1 AC126544.2 ARHGAP27 ARL17A CRHR1 DND1P1 KANSL1 KANSL1-AS1 LINC02210 LINC02210-CRHR1 LRRC37A2 LRRC37A4P MAPT PLEKHM1 SPPL2C STH rs3744245 CSF3 GSDMA MED24 PSMD3 rs3848456 DLX3 KAT7 TAC4 XYLT2 10 rs7897438 TCF7L2 rs3934643 ANXA11 EIF5AL1 MBL1P NUTM2B NUTM2E PLAC9 SFTPA1 SFTPA2 SFTPD TMEM254 9 rs505922 ABO OBP2B rs60840586 AQP3 rs28368148 IFNA10 IFNA14 IFNA16 IFNA17 IFNA21 IFNA4 IFNA5 IFNA6 IFNA7 KLHL9 rs79611697 CCDC171 PSIP1 SNAPC3 Not genome-wide signicant Closest gene eGenes Gene with highest Variant2Gene Score Genes with coding variants Genes in LD region Type of evidence for gene annotation Critically ill COVID-19+ Hospitalized COVID-19+ Reported SARS-CoV-2 infection 7 rs2897075 AZGP1P1 CNPY4 PILRB TRIM4 ZKSCAN1 rs35866984 HIP1 POM121C rs717624 AC005682.1 SNHG26 TOMM7 6 rs1128175 HLA-C MICA MICB rs2517723 HLA-A HLA-G rs12660421 FOXP4 rs928976 COL11A2 HLA-DPA1 HLA-DPB1rs35122968 HLA-DQA2 5 rs10066378 IRF1 SLC22A5 4 rs13107325 SLC39A8 rs34712979 AC109361.2 NPNT rs7671107 ANAPC4 SLC34A2 rs16877005 PI4K2B 3 rs2260685 MUC20 MUC20-OT1 MUC4 rs11924013 CEP97 NXPE3 PDCL3P4 RPL24 SENP7 ZBTB11 ZBTB11-AS1 rs17713054 CXCR6 LZTFL1 rs73062389 SACM1L SLC6A20 2 rs1123573 BCL11A rs1662979 C2orf70 CIB4 1 rs35154152 THBS3 rs12752585 EFNA1 KRTCAP2 MTX1 MUC1 THBS3 TRIM46 rs7515509 AK5 rs11208552 JAK1 rs10890422 AL450996.1 CYP4B1 X rs190509934 ACE2 BMX PIR Release 7: GWAS hits of COVID-19 a b rs 73 00 58 73 rs 18 47 81 32 6 rs 76 71 10 7 rs 75 15 50 9 rs 16 62 97 9 rs 11 23 57 3 rs 22 60 68 5 rs 16 87 70 05 rs 25 17 72 3 rs 28 97 07 5 rs 71 17 52 34 rs 12 58 50 36 rs 11 20 85 52 rs 10 06 63 78 rs 35 12 29 68 –l og 10 [P ] 1 3 6 10 16 Hospitalized COVID-19+ 2 3 4 5 6 7 8 9 10 13 16 20 25 32 40 50 63 79 100 126 158 200 SARS-CoV-2 reported infection 200 158 126 100 79 63 50 40 32 25 20 16 13 10 9 8 7 6 5 4 3 2 2 54 987 131211 1514 2017 1918 21 X22 –l og 10 [P ] rs 78 14 31 11 rs 28 75 96 8 rs 13 10 73 25 Fig. 2 | GWAS results for COVID-19. a, The results of a GWAS analysis of hospitalized individuals with COVID-19 (n = 49,033 cases and n = 3,393,109 controls) (top), and the results for individuals with reported SARS-CoV-2 infection (n = 219,692 cases and n = 3,001,905 controls) (bottom). The loci highlighted in yellow (top) represent regions that are associated with severity of COVID-19. The loci highlighted in green (bottom) are regions associated with susceptibility to SARS-CoV-2 infection. Lead variants for the loci that were identified in this data release are annotated with their respective rsID. The y axis is on the −log10 (P) scale up to 10, after which it switches to a 10 × log10[−log10(P)] scale to aid presentation. b, Results of gene prioritization using different evidence measures of gene annotation. For the genes in a linkage-disequilibrium (LD) region, genes with coding variants and eGenes (fine-mapped cis-expression quantitative trait locus (cis-eQTL) variant with posterior inclusion probability (PIP) >0.1 in GTEx Lung) are annotated as such if they are in linkage disequilibrium with a COVID-19 lead variant (r2 > 0.6). V2G, the highest gene prioritized by OpenTargetGenetics V2G score. The pink circle indicates SARS-CoV-2 infection susceptibility, the green triangle indicates COVID-19 severity and the blue cross indicates unclassified. This figure was reproduced from the original publication by the COVID-19 HGI1 with modifications reflecting the updated analysis from data release 7. E10 | Nature | Vol 621 | 7 September 2023 Matters arising mapped to viral entry and airway defence pathways, with notable exceptions for severity-classified TMPRSS2 and MUC5B loci. Further investigation of how such susceptibility and severity loci map to dif- ferent pathways would provide mechanistic insights into the human genetic architecture of COVID-19. Online content Any methods, additional references, Nature Portfolio reporting summa- ries, source data, extended data, supplementary information, acknowl- edgements, peer review information; details of author contributions and competing interests; and statements of data and code availability are available at https://doi.org/10.1038/s41586-023-06355-3. Reporting summary Further information on research design is available in the Nature Port- folio Reporting Summary linked to this article. Data availability Summary statistics generated by the COVID-19 HGI are available online, including per-ancestry summary statistics for African, admixed Ameri- can, East Asian, European and South Asian ancestries (https://www. covid19hg.org/results/r7/). The analyses described here used the data release 7. If available, individual-level data can be requested directly from contributing studies, listed in Supplementary Table 1. We used publicly available data from GTEx (https://gtexportal.org/home/), the Neale laboratory (http://www.nealelab.is/uk-biobank/), the Finucane laboratory (https://www.finucanelab.org), the FinnGen Freeze 4 cohort (https://www.finngen.fi/en/access_results) and the eQTL catalogue release 3 (http://www.ebi.ac.uk/eqtl/). Code availability The code for summary statistics lift-over, the projection PCA pipeline including precomputed loadings and meta-analyses (https://github. com/covid19-hg/); for heritability estimation (https://github.com/ AndrewsLabUCSF/COVID19_heritability); for Mendelian randomization and genetic correlation (https://github.com/marcoralab/MRcovid); and subtype analyses (https://github.com/mjpirinen/covid19-hgi_subtypes) are available at GitHub. 1. COVID-19 Host Genetics Initiative. Mapping the human genetic architecture of COVID-19. Nature 600, 472–477 (2021). 2. COVID-19 Host Genetics Initiative. A first update on mapping the human genetic architecture of COVID-19. 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SP-A and SP-D: dual functioning immune molecules with antiviral and immunomodulatory properties. Front. Immunol. 11, 622598 (2020). 10. Chatterjee, M., van Putten, J. P. M. & Strijbis, K. Defensive properties of mucin glycoproteins during respiratory infections-relevance for SARS-CoV-2. mBio 11, e02374-20 (2020). 11. van der Made, C. I. et al. Presence of genetic variants among young men with severe COVID-19. JAMA 324, 663 (2020). 12. Butler-Laporte, G. et al. Exome-wide association study to identify rare variants influencing COVID-19 outcomes: results from the Host Genetics Initiative. PLoS Genet. 18, e1010367 (2022). 13. Namkoong, H. et al. DOCK2 is involved in the host genetics and biology of severe COVID-19. Nature 609, 754–760 (2022). 14. Chen, M.-H. et al. Trans-ethnic and ancestry-specific blood-cell genetics in 746,667 individuals from 5 global populations. Cell 182, 1198–1213 (2020). 15. The GTEx Consortium. The GTEx Consortium atlas of genetic regulatory effects across human tissues. Science 369, 1318–1330 (2020). 16. RECOVERY Collaborative Group. Baricitinib in patients admitted to hospital with COVID-19 (RECOVERY): a randomised, controlled, open-label, platform trial and updated meta-analysis. Lancet 400, 359–368 (2022). 17. Shrine, N. et al. New genetic signals for lung function highlight pathways and chronic obstructive pulmonary disease associations across multiple ancestries. Nat. Genet. 51, 481–493 (2019). 18. Fingerlin, T. E. et al. Genome-wide association study identifies multiple susceptibility loci for pulmonary fibrosis. Nat. Genet. 45, 613–620 (2013). 19. Allen, R. J. et al. Genome-wide association study of susceptibility to idiopathic pulmonary fibrosis. Am. J. Respir. Crit. Care Med. 201, 564–574 (2020). Acknowledgements We acknowledge M. O’Reilly and B. Cooley from the Pattern team at the Broad Institute of MIT and Harvard for designing the geographical map in Fig. 1. A full list of the members of the Genes & Health Research Team is available online (https://www. genesandhealth.org/research/scientific-publications-authorship-and-acknowledgments). A full list of acknowledgements is provided in Supplementary Table 12. Author contributions Detailed author contributions are integrated in the authorship list. Competing interests A full list of competing interests is provided in Supplementary Table 12. Additional information Supplementary information The online version contains supplementary material available at https://doi.org/10.1038/s41586-023-06355-3. Correspondence and requests for materials should be addressed to Andrea Ganna. Reprints and permissions information is available at http://www.nature.com/reprints. Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/. © The Author(s) 2023 Nature | Vol 621 | 7 September 2023 | E11 The COVID-19 Host Genetics Initiative Leadership Masahiro Kanai1, Shea J. Andrews2, Mattia Cordioli3, Christine Stevens1, Benjamin M. Neale1,4, Mark Daly1,3,4 & Andrea Ganna1,3,4 ✉ Writing group Writing group lead Masahiro Kanai1, Shea J. Andrews2, Mattia Cordioli3, Gita A. Pathak5,6 & Andrea Ganna1,3,4 Writing group member Akiko Iwasaki7 Analysis group Manuscript analysis team lead Juha Karjalainen3 & Juha Mehtonen3 Manuscript analysis team members: phenome-wide association study Gita A. Pathak5,6 Manuscript analysis team members: Mendelian randomization Shea J. Andrews2 Manuscript analysis team members: principal component projection, gene prioritization Masahiro Kanai1 Manuscript analysis team members: methods development Mattia Cordioli3 & Matti Pirinen3 Project management group Project management lead Christine Stevens1 Project management support Karolina Chwialkowska8, Amy Trankiem1 & Mary K. Balaconis1 Scientific communication group Scientific communication lead Kumar Veerapen4 & Brooke N. Wolford9 Scientific communication member Hajar Fauzan Ahmad10, Shea Andrews11, Kathrin Aprile von Hohenstaufen Puoti12,13, Cindy Boer1143, Palwende R. Boua14,15, Guillaume Butler-Laporte16, Carmen L. Cadilla17, Karolina Chwiałkowska8, Francesca Colombo18, Venceslas Douillard1143, Nicole Dueker19, Atanu Kumar Dutta20, Yasser M. El-Sherbiny21,22, Madonna M. Eltoukhy23, Sahar Esmaeeli24, Annika Faucon25, Marie-Julie Fave1143, Israel Fernández Cadenas26, Margherita Francescatto27, Laurent Francioli28, Lude Franke29, Macarena Fuentes30, Rocío Gallego Durán1143, David Gómez Cabrero31, Emi N. Harry32, Philip Jansen33, József L. Szentpéteri34, Elżbieta Kaja35, Masahiro Kanai4, Chloe Kirk19, Athanasios Kousathanas36, Jose E. Krieger37, Sanjay K. Patel38, Audrey Lemaçon39, Sophie Limou40, Pietro Lió41, Eirini Marouli42, Minttu M. Marttila43, Carolina Medina-Gómez44, Yael Michaeli1143, Isabelle Migeotte45,46, Soumyajit Mondal47, Andrés Moreno-Estrada48, Leire Moya49, Tomoko Nakanishi50,51,52,53, Jamal Nasir54, Dorote Pasko36, Gita A. Pathak5,6, Nathaniel M. Pearson55, Alexandre C. Pereira56, James Priest57, Vid Prijatelj58, Ivana Prokić59, Alexander Teumer60, Réka Várnai34, Manuel Romero-Gómez61,62, Christina Roos63, Jeffrey Rosenfeld64, Li Ruolin65, Eva C. Schulte66,67,68,69,70, Claudia Schurmann71, Bahar Sedaghati-khayat58, Doaa Shaheen1143, Ilangumaran Shivanathan1143, Csilla Sipeky72, Zhou Sirui50, Pasquale Striano73,74, Yosuke Tanigawa75, Alicia Utrilla Remesal76, Nirmal Vadgama77, Costanza L. Vallerga78, Sander van der Laan79, Ricardo A. Verdugo80,81, Qingbo S. Wang82, Zhou Wei65, Ummu Afeera Zainulabid83 & Ruth N. Zárate84 23andMe Analysis team lead Adam Auton85 Data collection lead Janie F. Shelton85 & Anjali J. Shastri85 Analysis team member Catherine H. Weldon85, Teresa Filshtein-Sonmez85, Daniella Coker85, Antony Symons85, Stella Aslibekyan85 & Jared O’Connell85 Data collection member Chelsea Ye85 & Catherine H. Weldon85 Adolescent brain and cognitive development study Analysis team lead Alexander S. Hatoum86 Admin team lead Arpana Agrawal86 & Ryan Bogdan87 Analysis team member Sarah M. C. Colbert86 Data collection member Wesley K. Thompson88,89,90 & Chun Chieh Fan91 Admin team member Emma C. Johnson86 Armenia_Covid-19hg Data collection lead Lyudmila Niazyan92 & Mher Davidyants92 Admin team lead Arsen Arakelyan93 Admin team member Diana Avetyan93 Assessment of the influence of clinical, functional, immunological and genetic factors on the severity of the course of coronavirus infection with SARS-CoV-2 and post-COVID syndrome Analysis team lead Makhabbat Bekbossynova94 Data collection lead Ainur Tauekelova94 & Mukhtar Tuleutayev95 Admin team lead Aliya Sailybayeva94 Analysis team member Yerlan Ramankulov96, Elena Zholdybayeva96, Jarkyn Dzharmukhanov96 & Kuat Kassymbek96 Data collection member Tatyana Tsechoeva95, Gulsimzhan Turebayeva95 & Zauresh Smagulova95 Admin team member Timur Muratov97 & Sadyk Khamitov94 Avon Longitudinal Study of Parents and Children (ALSPAC) Analysis team lead Alex S. F. Kwong98,99 Data collection lead Nicholas J. Timpson98,99 BelCovid Analysis team lead Mari E. K. Niemi100 Data collection lead Souad Rahmouni101, Julien Guntz102 & Isabelle Migeotte45,46 Admin team lead Yves Beguin103 Analysis team member Mattia Cordioli104, Sara Pigazzini100 & Lindokuhle Nkambule28,105,106 Data collection member Michel Georges101, Michel Moutschen107,108, Benoit Misset107,108, Gilles Darcis107,108, Stéphanie Gofflot103, Youssef Bouysran109, Adeline Busson109, Xavier Peyrassol109, Françoise Wilkin109, Bruno Pichon109, Guillaume Smits109, Isabelle Vandernoot109, Jean-Christophe Goffard110 & Nicky Tiembe109 Biobanque Quebec COVID19 Analysis team lead Tomoko Nakanishi50,51,52,53 Data collection lead David R. Morrison51 & Jonathan Afilalo51 Admin team lead Vincent Mooser111, J. Brent Richards51,112,113, Simon Rousseau114 & Madeleine Durand115,116 Analysis team member Guillaume Butler-Laporte51,117 & Vincenzo Forgetta51 Data collection member Laetitia Laurent51, Zaman Afrasiabi51, Meriem Bouab51, Chris Tselios51, Xiaoqing Xue51, Marc Afilalo118,119, Maureen Oliveira120, Janick St-Cyr111 & Ariane Boisclair111 Admin team member Jiannis Ragoussis111, Daniel Auld111, Daniel E. Kaufmann121,122, G. Mark Lathrop111, Guillaume Bourque111, Simon Décary123, Emilia Liana Falcone124,125, Alexandre Montpetit126, Alain Piché127,128, Christel Renoux129,130,131, Karine Tremblay132,133, Sze Man Tse134 & Ma’n H. Zawati135 BioVU Analysis team lead Lea K. Davis136,137, Nancy J. Cox136,137 & Jennifer E. Below136,137 Analysis team member Julia M. Sealock136,137, Annika B. Faucon136,137, Megan M. Shuey136,137, Hannah G. Polikowsky136,137, Lauren E. Petty136,137, Douglas M. Shaw136,137, Hung-Hsin Chen136,137 & Wanying Zhu136,137 Bonn Study of COVID-19 Genetics Analysis team lead Axel Schmidt138 Data collection lead Kerstin U. Ludwig138 Analysis team member Carlo Maj139 Data collection member Selina Rolker138, Daniella Balla138, Pari Behzad138, Markus M. Nöthen138, Julia Fazaal138, Verena Keitel140, Verena Keitel141, Björn-Erik Ole Jensen140, Torsten Feldt140, Nikolaus Marx142, Michael Dreher143, Isabell Pink144, Markus Cornberg145, Thomas Illig146, Clara Lehmann147,148,149, Philipp Schommers147,148,149, Jan Rybniker147,148,149, Max Augustin147, Lisa Knopp140, Ingo Kurth150, Thomas Eggermann150, Sonja Volland146, Marc M. Berger151, Thorsten Brenner151, Anke Hinney152, Oliver Witzke153, Margarethe J. Konik153, Robert Bals154, Christian Herr154, Nicole Ludwig155, Jörn Walter156, Eicke Latz157 & Susanne V. Schmidt157 CGEn HostSeq—Canadian COVID-19 Human Host Genome Sequencing Databank Analysis team lead Jennifer D. Brooks158, Shelley Bull159,160, Lloyd T. Elliott161, France Gagnon159, Celia M. T. Greenwood16,162, Rayjean J. Hung158,160, Jerry F. Lawless163, Andrew D. Paterson158,164, E12 | Nature | Vol 621 | 7 September 2023 Matters arising Lei Sun158, Michael Rauh165, Laurent Briollais158,166, Anne-Claude Gingras158,160, Yvonne Bombard158,167, Trevor J. Pugh158,168,169, Jared Simpson158,169 & Lee W. Goneau158,170 Data collection lead Anat R. Halevy164, David M. Maslove165, Bjug Borgundvaag158,166, Luke Devine158,166, Erin Bearss158,166, David Richardson171, Saranya Arnoldo158,171, Steven Marc Friedman158,172, Ahmed Taher158,173, Seth Stern173, Marc Dagher158,174, Jovanka Vasilevska-Ristovska164,174, Catherine M. Biggs175,176,177 & Beata Mickiewicz178,179 Admin team lead Lisa J. Strug158,164, Stephen W. Scherer158,164, Naveed Aziz164, Steven J. M. Jones180, Bartha M. Knoppers16, Mark Lathrop16, Stuart E. Turvey175,176, Rae S. M. Yeung158,164, Upton Allen164, Angela M. Cheung158,172, Margaret S. Herridge158,181, Miranda Hunt165, Jordan Lerner-Ellis158,160, Jennifer Taher158,166, Rulan S. Parekh158,164,174, Linda T. Hiraki158,164, Juthaporn Cowan182,183,184, Francine M. Ducharme185,186, Mario Ostrowski158,167, Francois P. Bernier178,179 & Jim Kellner178,179 Analysis team member Elika Garg161, Samantha Yoo164, Caitlyn Vlasschaert165, Erika Frangione166, Monica Chung166, Abdul Noor158,166, Elena Greenfeld158,166, Karen Colwill160, Marc Clausen167, Gary Chao158, FengYun Yue187, Marvin Fritzler178,188, Joseph Whitney164, Bhooma Thiruvahindrapuram164, Jean-Michel Garant189 & Rohan Abraham189 Data collection member Adrienne Davis158,164, Aaron Campigotto164, Jesse Papenburg190, Kissoon Niranjan176, Stephen Betschel167, Manish Sadarangani176, Michelle Barton-Forbes191, Michaela Hanley165, Chun Yiu Jordan Fung166, Elisa Lapadula166, Georgia MacDonald166, Michael Puopolo166, Deepanjali Kaushik171, Konika Nirmalanathan172, Iris Wong173, Zeeshan Khan173, Natasha Zarei173, Maria Michalowska174, Bhavi P. Modi175,176, Pourshahnazari Persia175,177, Antonio Estacio167, Megan Buchholz192, Patti Lou Cheatley167, Miranda Lorenti164, Nowrin F. Aman164,174, Vitaliy Matveev158 & Patrick Budylowski158 Admin team member Julia Upton164, Shaun Morris164, Tracy Boyd165, Sunakshi Chowdhary166, Selina Casalino166, Gregory Morgan158,166, Chloe Mighton158,166,167, Allison McGeer158,166, Tony Mazzulli158,172,193, Shelley L. McLeod158,166, Alexandra Binnie171, Hanna Faghfoury172, Howard Chertkow194, Hilary Racher158,170, Mihaela A. Serbanescu170, Katerina Pavenski167, Michael Esser178,179, Graham Thompson178,179 & Jo-Anne Herbrick164 Colorado Center for Personalized Medicine—CCPM Analysis team lead Christopher R. Gignoux195 Data collection lead Stephen J. Wicks195 & Kristy Crooks195 Admin team lead Kathleen C. Barnes195 Analysis team member Michelle Daya195, Jonathan Shortt195, Nicholas Rafaels195 & Sameer Chavan195 COMRI/Virology Study Analysis team lead Andrea Ganna104 Data collection lead Thomas G. Schulze196,197,198,199 Admin team lead Eva C. Schulte196,200,201,202 Analysis team member Urs Heilbronner196, Sergi Papiol196,200,203, Mattia Cordioli104 & Andrea Corbetta104 Data collection member Clemens M. Wendtner204, Christoph D. Spinner205, Johanna Erber205, Jochen Schneider205, Christof Winter206,207, Jens Wiltfang208,209,210, Monika Budde196, Fanny Senner196,200 & Janos L. Kalman196,200 Admin team member Ulrike Protzer201,211 & Nikola S. Mueller212 Covid19 Ioannina Biobank Analysis team lead Abdou Mousas213 Data collection lead Angelos Liontos214 & Eirini Christaki214 Admin team lead Haralampos Milionis214 & Konstantinos Tsilidis213,215 Analysis team member Alexandros Asimakopoulos213, Afroditi Kanellopoulou213 & Georgios Markozannes213 Data collection member Dimitrios Biros214, Orestis Milionis214, Stavros Tsourlos214, Lazaros Athanasiou214, Nikolaos-Gavriil Kolios214, Christiana Pappa214, Alexandros Papathanasiou214, Eleni Pargana214, Maria Nasiou214, Maria Kosmidou214, Iro Rapti214, Evangelos Ntotsikas213, Konstantinos Chaliasos213, Evangelia Ntzani213,216, Evangelos Evangelou213,215, Konstantina Gartzonika217, Ioannis Georgiou217 & Ioanna Tzoulaki213,215,218 COVID19-Host(a)ge Analysis team lead David Ellinghaus219,220, Frauke Degenhardt219, Mario Cáceres221,222, Simonas Juzenas219,223 & Tobias L. Lenz224,225 Data collection lead Agustín Albillos226,227, Antonio Julià228, Daniele Prati229, Erik Solligård230, Federico Garcia231,232, Florian Tran219,233, Frank Hanses234,235, Guido Baselli229, Heinz Zoller236,237, Jan Cato Holter238,239, Javier Fernández240,241, Jordi Barretina242,243, Luca Valenti229,244, Luis Bujanda227,245, Manuel Romero-Gómez227,246,247,248,249, Maria Buti227,250,251, Mauro D’Amato252,253, Jesus M. Banales227,254, Philip Rosenstiel219, Philipp Koehler255,256,148, Pietro Invernizzi257,258, Rafael de Cid259, Rosanna Asselta260,261, Stefan Schreiber219,233, Stefano Duga260,261 & Ute Hehr262 Admin team lead Andre Franke219,263, Douglas Maya-Miles227,248,249, Johannes R. Hov238,264,265,266, Tom H. Karlsen238,264,265,266 & Trine Folseraas238,264,265,266 Analysis team member Ana Teles224,225, Anja Tanck219, Christoph Gassner219,267, Clinton Azuure224,225, Eike Matthias Wacker219, Florian Uellendahl-Werth219, Georg Hemmrich-Stanisak219, Hesham ElAbd219, Jan Kässens219, Jatin Arora28,268,269,270,271, Jon Lerga-Jaso272, Lars Wienbrandt219, Malte Christoph Rühlemann219,273, Mareike Wendorff219, Maria E. Figuera Basso219, May Sissel Vadla274,275, Michael Wittig219, Nicole Braun219,276, Ole Bernt Lenning274,277, Onur Özer224,225, Ronny Myhre278, Soumya Raychaudhuri28,268,269,270,271,279, Tanja Wesse219, Wolfgang Albrecht219 & Xiaoli Yi219 Data collection member Aaron Blandino Ortiz280, Adolfo de Salazar231,232, Adolfo Garrido Chercoles281, Adriana Palom228,251, Agustín Ruiz282,283, Alba-Estela Garcia-Fernandez284, Albert Blanco-Grau284, Alberto Mantovani260,261, Aleksander Rygh Holten238,285, Alessandra Bandera229,244, Alessandro Cherubini229, Alessandro Protti260,261, Alessio Aghemo260,261, Alessio Gerussi257,258, Alfredo Ramirez256,286,287,288,289, Almut Nebel219, Ana Barreira251, Ana Lleo260,261, Anders Benjamin Kildal290, Andrea Biondi291, Andrea Caballero-Garralda292, Andrea Gori229,293, Andreas Glück233, Andreas Lind239, Anna Carreras Nolla259, Anna Latiano294, Anna Ludovica Fracanzani229,244, Anna Peschuck219, Annalisa Cavallero295, Anne Ma Dyrhol-Riise238,296, Antonella Ruello297, Antonio Muscatello229, Antonio Voza260,261, Ariadna Rando-Segura250,298, Aurora Solier299, Beatriz Cortes259, Beatriz Mateos226,227, Beatriz Nafria-Jimenez281, Benedikt Schaefer236,237, Carla Bellinghausen300, Carlos Ferrando240,301, Carmen de la Horra247,248,249,302,303, Carmen Quereda304, Chiara Scollo305, Christoph Lange306,307,308, Cinzia Hu309, Cinzia Paccapelo229, Claudio Angelini310, Claudio Cappadona261, Cristiana Bianco229, Cristina Cea284, Cristina Sancho311, Dag Arne Lihaug Hoff312,313, Daniela Galimberti229,244, David Haschka314, David Jiménez299, David Pestaña315, David Toapanta240, Eduardo Muñiz-Diaz316, Elena Azzolini260,261, Elena Sandoval240, Eleonora Binatti258,317, Elio Scarpini229,244, Elisabetta Casalone318, Eloisa Urrechaga319,320, Elvezia Maria Paraboschi260,261, Emanuele Pontali321, Enric Reverter240, Enrique J. Calderón247,248,249,302,303, Enrique Navas304, Ernesto Contro322, Eunate Arana-Arri320, Fátima Aziz240, Félix García Sánchez323, Ferruccio Ceriotti229, Filippo Martinelli-Boneschi324,325, Flora Peyvandi326,327, Francesco Blasi328,329, Francesco Malvestiti244, Francisco J. Medrano247,248,302,303,330, Francisco Mesonero226,227, Francisco Rodriguez-Frias227,228,250,331, Fredrik Müller238,239, Giacomo Bellani332,333, Antonio Pesenti329,334, Alberto Zanella329,334, Giacomo Grasselli329,334, Gianni Pezzoli335, Giorgio Costantino229,244, Giovanni Albano297, Giulia Cardamone261, Giuseppe Bellelli336,337, Giuseppe Citerio336,338, Giuseppe Foti336,339, Giuseppe Lamorte229, Giuseppe Matullo318, Hayato Kurihara310, Holger Neb340, Ilaria My260, Isabel Hernández336,341, Itziar de Rojas282,283, Iván Galván-Femenia259, Jan Egil Afset312,342, Jan Heyckendorf306,307,308,343, Jan Kristian Damås344,345, Javier Ampuero227,246,247,248,249, Javier Martín346, Jeanette Erdmann347,348,349, Joan Ramon Badia350, Joaquin Dopazo248,351, Jonas Bergan352, Jose Hernández Quero232,353, Josune Goikoetxea354, Juan Delgado247,248,249,302,303, Juan M. Guerrero247,248,249, Kari Risnes345,355, Karina Banasik356, Karl Erik Müller357, Karoline I. Gaede358,359,360, Koldo Garcia-Etxebarria227,245, Kristian Tonby238,296, Lars Heggelund357,361, Laura Rachele Bettini362, Lauro Sumoy363, Leonardo Terranova229, Lise Tuset Gustad364,365, Lucia Garbarino366, Luigi Santoro229, Luis Téllez226,227, Luisa Roade227,250,251, Mahnoosh Ostadreza229, Maider Intxausti311, Manolis Kogevinas302,367,368,369, Mar Riveiro-Barciela227,250,251, Marco Schaefer370, María A. Gutiérrez-Stampa371, Maria Carrabba309, Maria G. Valsecchi372,373, María Hernandez-Tejero240, Maria J. G. T. Vehreschild374, Maria Manunta229, Marialbert Acosta-Herrera346, Mariella D’Angiò362, Marina Baldini229, Marina Cazzaniga375, Marta Marquié282,283, Massimo Castoldi297, Maurizio Cecconi260,261, Melissa Tomasi229, Mercè Boada282,283, Michael Joannidis376, Michela Mazzocco366, Michele Ciccarelli310, Miguel Rodríguez-Gandía226,227, Monica Bocciolone310, Monica Miozzo244,309, Natale Imaz Ayo320, Natalia Blay259, Natalia Chueca231, Nicola Montano229,244, Nilda Martínez377, Oliver A. Cornely149,255,256,378, Orazio Palmieri294, Paola Faverio336,379, Paoletta Preatoni310, Paolo Bonfanti336,380, Paolo Omodei310, Paolo Tentorio260, Pedro Castro240, Pedro M. Rodrigues227,254, Laura Izquierdo-Sanchez227,245, Pedro Pablo España319, Per Hoffmann381, Petra Bacher219,382, Raúl de Pablo280, Ricard Ferrer383, Roberta Gualtierotti229,244, Rocío Gallego-Durán227,248,249, Rosa Nieto299, Rossana Carpani229, Rubén Morilla247,248,249,302,303, Salvatore Badalamenti260, Sammra Haider384, Sandra Ciesek385,386, Sara Bombace260,261, Sara Marsal228, Sebastian Klein376, Serena Pelusi229,244, Sibylle Wilfling235,262,387, Siegfried Goerg388, Silvano Bosari229,244, Søren Brunak220, Stefanie Heilmann-Heimbach381, Stefano Aliberti260,261, Susanne Dudman238,239, Tenghao Zheng389, Thomas Bahmer233, Tomas Pumarola390,391, Trinidad Gonzalez Cejudo392, Valeria Rimoldi260,261, Valter Monzani229, Vegard Skogen393,394, Vicente Friaza247,248,249,302,303, Victor Andrade286,287, Victor Moreno302,395,396,397, Wolfgang Peter370,398, Xavier Farre259 & Yascha Khodamoradi374 Admin team member Marit M. Grimsrud238,265,266 & Sandra May219 Covid19hg-CL Data collection lead Alicia Colombo399,400, Monardes-Ramírez A. Virginia401, Cristina Dorador402, Macarena Fuentes-Guajardo30, Andrea X. Silva403,404 & Yolanda Espinosa-Parrilla405,406,407 Admin team lead Ricardo A. Verdugo408,409,410 Analysis team member Cristian E. Yáñez409, Rocío M. Retamales-Ortega409 & Juan M. Saez Hidalgo411 Nature | Vol 621 | 7 September 2023 | E13 Data collection member Eduardo A. Tobar-Calfucoy412,413, Laura Carvajal-Silva409, Matías F. Martínez412,413,414, Leslie C. Cerpa415,416, Muñoz A. Christian417, Claudio Cappelli418, Hector Valenzuela-Jorquera418, Daniela Zapata-Contreras405,406, Paula Zuñiga-Pacheco405,406, Estefania A. Nova-Lamperti419, Sergio A. Sanhueza419, Gerardo Donoso400, Pamela Bocchieri399 & Pia Kochifas401 Admin team member Luis A. Quiñones412,413,414 Danish COVID-19 Genomic Consortium Analysis team lead Karina Banasik220 Data collection lead Ole Birger Pedersen420 Analysis team member Frank Geller421, David Westergaard220 & Celia Burgos Sequeros220 Data collection member Janna Nissen422, Susanne Dam Nielsen423, Ulla Feldt-Rasmussen424, Sofie Bliddal424, Kirsten Grønbæk425 & Søren Brunak220 Admin team member Henrik Ullum426, Sisse Rye Ostrowski422 & Bjarke Feenstra421 Egypt hgCOVID hub Analysis team lead Mari E. K. Niemi100 Data collection lead Doaa Shahin22 & Yasser M. El-Sherbiny21,22 Admin team lead Kathrin Aprile von Hohenstaufen Puoti12,13, Ali Sobh427, Madonna M. Eltoukhy23 & Ashraf Shoma428 Analysis team member Mattia Cordioli3, Andrea Corbetta3 & Lindokuhle Nkambul105,429 Data collection member Tamer A. Elhadidy430, Mohamed S. Abd Elghafar431, Jehan J. El-Jawhari21,22, Attia A. S. Mohamed23, Marwa H. Elnagdy432, Amr Samir428, Mahmoud Abdel-Aziz433, Walid T. Khafaga434, Walaa M. El-Lawaty435, Mohamed S. Torky435, Mohamed R. El-Shanshory436, Ahmed Elzeiny22, Amal Rashad437 & Tamer A. Mansour22,438 Admin team member Amr M. Yassen439, Mohamed A. F. Hegazy428, Kamal Okasha440, Mohammed A. Eid441 & Moahmed S. Hanteera435 EraCORE Analysis team lead Carolina Medina-Gomez442 Data collection lead M. Arfan Ikram443 Admin team lead Andre G. Uitterlinden443,444 FinnGen Admin team lead445 Analysis team member Samuli Ripatti1,446,447, Bradley Jermy3 & Sanni Ruotsalainen446 Data collection member Kati Kristiansson448, Sami Koskelainen448, Markus Perola448,449, Kati Donner3, Katja Kivinen3 & Aarno Palotie3 Admin team member Mari Kaunisto3 French hgCOVID Analysis team lead Hanna Julienne450 Data collection lead Hugues Aschard450, Jean-Francois Deleuze451, Stephanie Debette452,453, David-Alexandre Tregouet452 & Laurent Abel454,455 Analysis team member Léo Henches450, Edith Lefloch451 & Dandine-Roulland Claire451 Data collection member Anavaj Sakuntabhai456, Thomas Bourgeron456, Robert Olaso451, Delphine Bacq Daian451, Isabelle Pellegrin457,458, Jean-Laurent Casanova454,455, Aurélie Cobat454, Emmanuelle Jouanguy454, Qian Zhang454,459, Jade Ghosn460, France Mentré460, Cédric Laouénan460, Sarah Tubiana460, Catherine Chirouze461, Luis Quintana-Murci462,463, Etienne Patin464 & Darragh Duffy465 Admin team member Anne Boland451 GCAT. Genomes For Life Analysis team lead Rafael de Cid242 Data collection lead Jordi Barretina243, Anna Carreras242, Victor Moreno466 & Manolis Kogevinas367,368,369,467 Analysis team member Iván Galván-Femenía242, Natalia Blay242, Xavier Farré242 & Lauro Sumoy242 Data collection member Beatriz Cortés242, Josep Maria Mercader468,469,470,471, Marta Guindo-Martinez468, David Torrents468, Judith Garcia-Aymerich367,368,467, Gemma Castaño-Vinyals367,368,369,467 & Carlota Dobaño367,472 Genes & Health Analysis team lead David A. van Heel473 Data collection lead Karen A. Hunt473 Admin team lead Richard C. Trembath474 Analysis team member Benjamin Meir Jacobs475, Qin Qin Huang476 & Hilary C. Martin476 Data collection member Dan Mason477, Bhavi Trivedi473, John Wright477 & Malathy Munisamy477 Admin team member David A. van Heel478, Karen A. Hunt478, Sarah Finer475 & Christopher J. Griffiths475 Genetic influences on severity of COVID-19 illness in Korea Analysis team lead Mark I. McCarthy479 & Carrie Rosenberger479 Data collection lead Jong Eun Lee480 Analysis team member Diana Chang479, Christian Hammer479, Julie Hunkapiller479, Anubha Mahajan479, Rion Pendergrass479, Lara Sucheston-Campbell479 & Brian Yaspan479 Data collection member Hyun Soo Lee480, Eunsoon Shin480, Hye Yoon Jang480, Sunmie Kim481, Sungmin Kym482, Yeon-Sook Kim482, Hyeongseok Jeong482, Ki Tae Kwon483, Shin-Woo Kim483, Jin Yong Kim484, Young Rock Jang484, Hyun ah Kim485, Ji yeon Lee485, Jeong Eun Lee486, Shinwon Lee486, Kang-Won Choe487, Yu Min Kang487, Sun Ha Jee488, Keum Ji Jung488, Hyun-Young Park489, Bong-Jo Kim490, Young Jin Kim490, Mi Yeong Hwang490 & Kyungheon Yoon490 GenOMICC Analysis team lead Erola Pairo-Castineira491,492,493, Konrad Rawlik491, Athanasios Kousathanas36 & Mark J. Caulfield36,494 Data collection lead J. Kenneth Baillie491,492,493,495 Admin team lead Fiona Griffiths493 Analysis team member Andrew D. Bretherick491,492,496, Alex Stuckey36, Christopher A. Odhams36, Susan Walker36, Clark D. Russell493,497, Tomas Malinauskas498, Yang Wu499, Xia Shen500,501, Katherine S. Elliott498, Kirstie Morrice502, Sean Keating495, Bo Wang493, Daniel Rhodes36, Lucija Klaric492, Marie Zechner493, Nick Parkinson493, Angie Fawkes502, Lee Murphy502, Veronique Vitart492, James F. Wilson492,501, Jian Yang503,504, Richard H. Scott36,505, Loukas Moutsianas36 & Andy Law493 Data collection member Colin Begg506, Sara Clohisey Hendry493, Charles Hinds42, Peter Horby507, Julian Knight498, Lowell Ling508, David Maslove509, Danny McAuley510,511, Johnny Millar493, Hugh Montgomery512, Alistair Nichol513, Peter J. M. Openshaw514,515, Alexandre C. Pereira516, Chris P. Ponting492, Kathy Rowan517, Malcolm G. Semple518,519, Manu Shankar-Hari497, Charlotte Summers520, Timothy Walsh495, Jacqueline Pan521, Neus Grau521, Tim Owen Jones521, Rosario Lim521, Martina Marotti521, Christopher Whitton521, Aneta Bociek521, Sara Campos521, Gill Arbane521, Marlies Ostermann521, Mina Cha521, Fabiola DAmato521, Eirini Kosifidou521, Shelley Lorah521, Kyma Morera521, Laura Brady522, Keith Hugill522, Jeremy Henning522, Stephen Bonner522, Evie Headlam522, Jessica Jones522, Abigail List522, Joanne Morley522, Amy Welford522, Bobette Kamangu522, Anitha Ratnakumar522, Abiola Shoremekun522, Zoe Alldis523, Raine Astin-Chamberlain523, Fatima Bibi523, Jack Biddle523, Sarah Blow523, Matthew Bolton523, Catherine Borra523, Ruth Bowles523, Maudrian Burton523, Yasmin Choudhury523, Amber Cox523, Amy Easthope523, Patrizia Ebano523, Stavros Fotiadis523, Jana Gurasashvili523, Rosslyn Halls523, Pippa Hartridge523, Delordson Kallon523, Jamila Kassam523, Ivone Lancoma-Malcolm523, Maninderpal Matharu523, Peter May523, Oliver Mitchelmore523, Tabitha Newman523, Mital Patel523, Jane Pheby523, Irene Pinzuti523, Zoe Prime523, Oleksandra Prysyazhna523, Julian Shiel523, Melanie Taylor523, Carey Tierney523, Olivier Zongo523, Suzanne Wood523, Anne Zak523, David Collier523, Manuela Mundy524, Christopher Thompson524, Lisa Pritchard524, Minnie Gellamucho524, David Cartlidge524, Nageswar Bandla524, Lucy Bailey524, Michelle Davies524, Jane Delaney524, Leanne Scott524, Marwa Abdelrazik525, Frater Alasdair525, David Carter525, Munzir Elhassan525, Arunkumar Ganesan525, Samuel Jenkins525, Zoe Lamond525, Dharam Purohit525, Kumar Rohit525, Malik Saleem525, Alanna Wall525, Kugan Xavier525, Dhanalaksmi Bakthavatsalam525, Kirolos Gehad525, Pakeerathan Gnanapragasam525, Kapil Jain525, Swati Jain525, Abdul Malik525, Naveen Pappachan525, Jeronimo Moreno-Cuesta525, Anne Haldeos525, Rachel Vincent525, Maryjane Oziegb525, Anna Cavazza526, Maeve Cockrell526, Eleanor Corcoran526, Maria Depante526, Clare Finney526, Ellen Jerome526, Abigail Knighton526, Monalisa Nayak526, Evita Pappa526, Rohit Saha526, Sian Saha526, Andrew Dodd526, Kevin O’Reilly526, Mark McPhail526, Emma Clarey526, Harriet Noble526, John Smith526, Phoebe Coghlan527, Stephen Brett527, Anthony Gordon527, Maie Templeton527, David Antcliffe527, Dorota Banach527, Sarah Darnell527, Ziortza Fernandez527, Eleanor Jepson527, Amal Mohammed527, Roceld Rojo527, Sonia Sousa Arias527, Anita Tamang Gurung527, Jenny Wong527, Jaime Fernandez-Roman528, David O. 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Jonathan Whiteside698, Rachael Campbell698, Joanna Matheson698, Deborah McDonald698, Donna Patience698, Polly Rice699, Melanie Clapham699, Rachel Mutch699, Hannah Craig699, Una Poultney699, Karen Burns700, Sophie Twiss701, Janet Barton701, Linsha George701, Clare Harrop701, Sherly Mathew701, David Justin Wright701, Rachel Harrison702, Jordan Toohie702, Ben Chandler702, Alison Turnbull702, Janine Mallinson702, Kerry Elliott702, Rebecca Wolf-Roberts703, Helen Tench703, Maria Hobrok703, Ronda Loosley703, Heather McGuinness703, Tanya Sims703, Deborah Afolabi704, Kathryn Sian Allison704, Taya Anderson704, Rachael Dore704, Dawn Jones704, Naomi Rogers704, Paula Saunderson704, Jennifer Whitbread704, Laura O’Malley704, Laura Rad704, Daniel Hawcutt704, Jonathan Aldridge705, Melanie Tolson705, Sweyn Garrioch705, Joanne Tomlinson706, Michael Grosdenier706, David Loader707, Ritoo Kapoor707, Gemma Hector707, Joslan Scherewode708, Chunda Sri-Chandana708, Lorraine Stephenson708, Sarah Marsh708, Arnold Dela Rosa709, Shaman Jhanji709, Thomas Bemand709, Ryan Howle709, Ravishankar Rao Baikady709, Benjamin Thomas709, Ethel Black709, Kate Tatham709, Sambasivarao Gurram710, Ekaterina Watson710, Sheena Quaid710, Anne Saunderson711, Rachel O’Brien711, Sam Moultrie711, Jen Service711, Clare Cheyne711, Miranda Odam711, Alison Wiliams711, Nicky Barnes712, Peter Csabi712, Joana Da Rocha712, Louika Glynou712, Amy Huffenberger713, Jade Bryant713, Amy Pickard713, Nicholas Roe713, Arianna Bellini713, Anton Mayer713, Amy Burrow713, Natalie Colley713, Jayne Evans713, Alex Howlett713, Zeinab Khalifeh713, Jerldine Pryce714, Claire Gorman714, Rebecca Brady714, Elizabeth Timlick714, Pierre Antoine714, Abhinhav Gupta714, John Hardy715, Henry Houlden715, Eleanor Moncur715, Arianna Tucci715, Ambreen Tariq715, Emma Tagliavini716, Becky Ramsay716, Katy Fidler716, Kevin Donnelly716, Rebecca Hollis716, Jocelyn Barr717, Elizabeth Boyd717, Val Irvine717, Ben Shelley717, Julie Buckley717, Charlene Hamilton717, Kathryn Valdeavella717, Sarah Bircham522, Steven Liggett522 & Anand Patil671 Admin team member Afshan Siddiq36, Peter Goddard36, Sally Donovan36, Tala Zainy36, Fiona Maleady-Crowe36, Linda Todd36, Shahla Salehi36, Greg Elgar36, Georgia Chan36, Prabhu Arumugam36, Christine Patch36, Augusto Rendon36, David Bentley718, Clare Kingsley718, Tom A. Fowler36,719, Paul Elliott720, Albert Tenesa492,493,501, Emma Aitkin493, Latha Aravindan721, Ruth Armstrong493, Heather Biggs722, Ceilia Boz493, Adam Brown493, Primmy Chikowore493, Richard Clark502, Audrey Coutts502, Judy Coyle493, Louise Cullum493, Sukamal Das721, Nicky Day493, Lorna Donnelly502, Esther Duncan493, Paul Finernan493, Max Head Fourman493, Anita Furlong722, James Furniss493, Bernadette Gallagher493, Tammy Gilchrist502, Ailsa Golightly493, Katarzyna Hafezi502, Debbie Hamilton493, Ross Hendry493, Naomi Kearns493, Dawn Law493, Rachel Law493, Sarah Law493, Rebecca Lidstone-Scott493, Christen Lauder502, Louise Macgillivray502, Alan Maclean502, Hanning Mal493, Sarah McCafferty502, Ellie Mcmaster493, Jen Meikle493, Shona C. Moore518, Sheena Murphy721, Hellen Mybaya493, Miranda Odam493, Wilna Oosthuyzen493, Chenqing Zheng723, Jiantao Chen723, Trevor Paterson493, Petra Tucker722, Katherine Schon722, Andrew Stenhouse493, Mihaela Das721, Maaike Swets493,724, Helen Szoor-McElhinney493, Filip Taneski493, Lance Turtle518, Tony Wackett493, Mairi Ward493, Jane Weaver493 & Nicola Wrobel502 Genotek COVID-19 study Analysis team lead Alexander Rakitko725 Admin team lead Valery Ilinsky725 Analysis team member Danat Yermakovich725, Iaroslav Popov725, Alexander Chernitsov725, Elena Kovalenko725, Ekaterina Vergasova725, Alexandr Prokhorov725, Anna Krasnenko725, Nikolay Plotnikov725, Ivan Stetsenko725 & Anna Kim725 Helix Exome+ COVID-19 Phenotypes Analysis team lead Elizabeth T. Cirulli726 Analysis team member Kelly M. Schiabor Barrett726, Alexandre Bolze726, Simon White726, Nicole L. Washington726 & James T. Lu726 Data collection member Stephen Riffle726, Francisco Tanudjaja726, Xueqing Wang726, Jimmy M. Ramirez III726, Nicole Leonetti726, Efren Sandoval726, Iva Neveux727, Shaun Dabe728 & Joseph J. Grzymski727 Host Genetic Factors for COVID-19 Severity and Outcome in Western Indian Population Analysis team lead Madhvi N. Joshi729 Data collection lead Raghav D. Dixit730, Pranay K. Shah731, Kamlesh J. Upadhyay732, Naresh T. Chuhan733, Kairavi J. Desai734, Meenakshi R. Shah735 & Bhavesh Modi736 Admin team lead Chaitanya G. Joshi729 Analysis team member Ramesh J. Pandit729, Indra Singh729 & Afzal I. Ansari729 Data collection member Janvi N. Raval729 & Zarna Z. Patel729 Host Genetics in COVID Cohorts of Mixed ancestry from Mexico Analysis team lead Andres Moreno-Estrada737 Data collection lead Joel Armando Vázquez-Pérez738, Manuel Castillejos-López738, Arnoldo Aq uino-Gálvez738,739, Teresa Shamah-Levy740, María C. Ávila-Arcos741, María Teresa Tusié Luna742 & Alfredo Hidalgo-Miranda743 Analysis team member Carmina Barberena-Jonas737, Consuelo D. Quinto-Cortés737, Marco Antonio Ávila-Arcos740 & Alicia Huerta-Chagoya742 Data collection member Lourdes García-García740, Jesús Martinez Barnetche740, Celia Alpuche-Aranda740, Juan A. Rivera740, Tonatiuh Barrientos Gutiérrez740, Ana Ochoa-Guzmán742, Maria Luisa Ordoñez-Sánchez742, Yayoi Segura-Kato742, Luis Alonso Herrera-Montalvo743, Alfredo Mendoza-Vargas743, Juan Pablo Reyes-Grajeda743, Alonso D. Cruz-Cruz743, Rafael Velázquez-Cruz743, Luz María Torres-Espíndola744, Martha Patricia Sierra-Vargas738, Angel Camarena738, Víctor Ruiz738,745, Jaime Chávez-Alderete738, María Esther Jaime-Capetillo738, Cristóbal Guadarrama-Pérez738, Joaquin Zúñiga-Ramos738,746, Alejandra Olivera-Ramírez738, Miguel Angel López Martínez738 & Ricardo Serna-Muñoz738 Admin team member Adriana Garmendia737 Japan Coronavirus Taskforce Analysis team lead Ho Namkoong747, Yukinori Okada748,749,750 & Seiya Imoto751 Data collection lead Kazuhiko Katayama752, Koichi Fukunaga747, Yuko Kitagawa753, Toshiro Sato754, Naoki Hasegawa755, Atsushi Kumanogoh750,756,757, Akinori Kimura758, Masumi Ai759 & Katsushi Tokunaga760 Admin team lead Takanori Kanai761, Satoru Miyano762 & Seishi Ogawa763,764 Analysis team member Ryuya Edahiro748,756, Kyuto Sonehara748, Yuya Shirai748,756 & Masahiro Kanai270 Data collection member Makoto Ishii747, Hiroki Kabata747, Katsunori Masaki747, Hirofumi Kamata747, Shinnosuke Ikemura747, Shotaro Chubachi747, Satoshi Okamori747, Hideki Terai747, Hiromu Tanaka747, Atsuho Morita747, Ho Lee747, Takanori Asakura747, Junichi Sasaki765, Hiroshi Morisaki766, Yoshifumi Uwamino767, Kosaku Nanki761, Yohei Mikami761, Kazunori Tomono768, Kazuto Kato769, Fumihiko Matsuda770, Meiko Takahashi770, Nobuyuki Hizawa771, Yoshito Takeda756, Haruhiko Hirata756, Takayuki Shiroyama756, Satoru Miyawaki772, Ken Suzuki748, Yuichi Maeda756,773, Takuro Nii756,773, Yoshimi Noda756, Takayuki Niitsu756, Yuichi Adachi756, Takatoshi Enomoto756, Saori Amiya756, Reina Hara756, Kunihiko Takahashi762, Tatsuhiko Anzai762, Takanori Hasegawa762, Satoshi Ito762, Ryuji Koike774, Akifumi Endo775, Yuji Uchimura776, Yasunari Miyazaki777, Takayuki Honda777, Tomoya Tateishi777, Shuji Tohda778, Naoya Ichimura778, Kazunari Sonobe778, Chihiro Sassa778, Jun Nakajima778, Yasuhito Nannya779, Yosuke Omae760, Kazuhisa Takahashi780, Norihiro Harada780, Makoto Hiki781,782, Haruhi Takagi780, Ai Nakamura780, Etsuko Tagaya783, Masatoshi Kawana784, Ken Arimura783, Takashi Ishiguro785, Noboru Takayanagi785, Taisuke Isono785, Yotaro Takaku785, Kenji Takano785, Ryusuke Anan786, Yukiko Nakajima786, Yasushi Nakano786, Kazumi Nishio786, Soichiro Ueda787, Reina Hayashi787, Hiroki Tateno788, Isano Hase788, Shuichi Yoshida788, Shoji Suzuki788, Keiko Mitamura789, Fumitake Saito790, Tetsuya Ueda791, Masanori Azuma791, Tadao Nagasaki791, Yoshinori Yasui792, Yoshinori Hasegawa791, Yoshikazu Mutoh793, Takashi Yoshiyama794, Tomohisa Shoko795, Mitsuaki Kojima795, Tomohiro Adachi795, Motonao Ishikawa796, Kenichiro Takahashi797, Kazuyoshi Watanabe798, Tadashi Manabe799, Fumimaro Ito799, Takahiro Fukui799, Yohei Funatsu799, Hidefumi Koh799, Yoshihiro Hirai800, Hidetoshi Kawashima800, Atsuya Narita800, Kazuki Niwa801, Yoshiyuki Sekikawa801, Fukuki Saito802, Kazuhisa Yoshiya802, Tomoyuki Yoshihara802, Yusuke Suzuki803, Sohei Nakayama803, Keita Masuzawa803, Koichi Nishi804, Masaru Nishitsuji804, Maiko Tani804, Takashi Inoue805, Toshiyuki Hirano805, Keigo Kobayashi805, Naoki Miyazawa806, Yasuhiro Kimura806, Reiko Sado806, Takashi Ogura807, Hideya Kitamura807, Kota Murohashi807, Ichiro Nakachi808, Rie Baba808, Daisuke Arai808, Satoshi Fuke809, Hiroshi Saito809, Naota Kuwahara810, Akiko Fujiwara810, Takenori Okada810, Tomoya Baba811, Junya Noda811, Shuko Mashimo811, Kazuma Yagi812, Tetsuya Shiomi812, Mizuha Hashiguchi812, Toshio Odani813, Takao Mochimaru814,815, Yoshitaka Oyamada814,815, Nobuaki Mori816, Namiki Izumi817, Kaoru Nagata817, Reiko Taki817, Koji Murakami818, Mitsuhiro Yamada818, Hisatoshi Sugiura818, Kentaro Hayashi819, Tetsuo Shimizu819, Yasuhiro Gon819, Shigeki Fujitani820, Tomoya Tsuchida821, Toru Yoshida820, Takashi Kagaya822, Toshiyuki Kita822, Satoru Sakagami822, Yoshifumi Kimizuka823, Akihiko Kawana823, Yoshihiko Nakamura824, Hiroyasu Ishikura824, Tohru Takata825, Takahide Kikuchi826, Daisuke Taniyama826, Morio Nakamura826, Nobuhiro Kodama827, Yasunari Kaneyama827, Shunsuke Maeda827, Yoji Nagasaki828, Masaki Okamoto829, Sayoko Ishihara830, Akihiro Ito831, Yusuke Chihara832, Mayumi Takeuchi832, Keisuke Onoi832, Naozumi Hashimoto833, Keiko Wakahara833, Akira Ando833, Makoto Masuda834, Aya Wakabayashi834, Hiroki Watanabe834, Hisako Sageshima835, Taka-Aki Nakada836, Ryuzo Abe836, Tadanaga Shimada836, Kodai Kawamura837, Kazuya Ichikado837, Kenta Nishiyama837, Masaki Yamasaki838, Satoru Hashimoto838, Yu Kusaka839, Takehiko Ohba839, Susumu Isogai839, Minoru Takada840, Hidenori Kanda840, Yuko Komase841, Fumiaki Sano842, Koichiro Asano843, Tsuyoshi Oguma844, Masahiro Harada845, Takeshi Takahashi845, Takayuki Shibusawa845, Shinji Abe846, Yuta Kono846, Yuki Togashi846, Takehiro Izumo847, Minoru Inomata847, Nobuyasu Awano847, Shinichi Ogawa848, Tomouki Ogata848, Shoichiro Ishihara848, Arihiko Kanehiro849, Shinji Ozaki849, Yasuko Fuchimoto849, Yuichiro Kitagawa850, Shozo Yoshida850, Shinji Ogura850, Kei Nishiyama851, Kousuke Yoshida852, Satoru Beppu852, Satoru Fukuyama853, Yoshihiro Eriguchi854, Akiko Yonekawa854, Yoshiaki Inoue855, Kunihiro Yamagata856, Shigeru Chiba857, Osamu Narumoto858, Hideaki Nagai858, Nobuharu Ooshima858, Mitsuru Motegi859, Hironori Sagara860, Akihiko Tanaka860, Shin Ohta860, Yoko Shibata861, Yoshinori Tanino861, Yuki Sato861, Yuichiro Yamada862, Takuya Hashino862, Masato Shinoki862, Hajime Iwagoe863, Tomonori Imamura864, Akira Umeda865, Hisato Shimada865, Mayu Endo866, Shinichi Hayashi867, Mai Takahashi867, Shigefumi Nakano867, Masakiyo Yatomi868, Toshitaka Maeno868, Tomoo Ishii869, Mitsuyoshi Utsugi870, Akihiro Ono870, Kensuke Kanaoka871, Shoichi Ihara871 & Kiyoshi Komuta871 Nature | Vol 621 | 7 September 2023 | E17 Japan NCGM-COVID19 Analysis team lead Katsushi Tokunaga760 Analysis team member Nao Nishida872, Yosuke Kawai760 & Masaya Sugiyama872 Jordan COVID-19 Host Genomics Initiative (JCHGI) Admin team lead Laith M. Alawneh873 Data collection member Kinda B. Maswadeh874, Fatima S. Alsoub875, Tala J. AlRawashdeh875, Ezaldeen Esawi876, Bayan O. Abu Alragheb875, Ahmad B. Maswadeh875, Abdulfattah Al-Kadash876, Abdallah Al-Ani875, Dana B. Alsafadi875, Mohammad A. Shahin875, Nuseibah Al Qtaish876, Fade M. Alawneh877, Joseph A. Abulail875, Hadi J. Zhlawi873, Tasneem N. Alhousani875, Mohammad R. Alzuraiqi873, Mohammad M. Al-Ja’afreh876, Sara K. Allouzy875, Mothafer ALjalamdeh873, Tareq M. Mansour873, Lina M. Alamer873, Ali Banni Issa878 & Fatima M. Bani younis879 Admin team member Feras I. Hawari873, Wail A. Hayajneh873, Nathir M. Obeidat875, Mamoun Ahram875, Moh’d Wahib J. Zhlawi875, Walhan Alshaer876, Malak M. Al-Kasasbeh875, Randa G. Naffa875, Mohammad A. Ismail876, Mohammed S. Abdullah875 & Said I. Ismail875 Lung eQTL Consortium Data collection member Ma’en Obeidat880, Ana I. Hernández Cordero880, Don D. Sin880,881, Yohan Bossé882, Philippe Joubert882, Ke Hao883, David Nickle884,885, Wim Timens886,887 & Maarten van den Berge887,888 Mass General Brigham—Host Vulnerability to COVID-19 Analysis team lead Yen-Chen Anne Feng889,890,891 & Josep Mercader469,470,471 Data collection lead Scott T. Weiss892, Elizabeth W. Karlson893, Jordan W. Smoller894, Shawn N. Murphy895, James B. Meigs896 & Ann E. Woolley893 Admin team lead Robert C. Green897 Analysis team member Ravi Mandla469,471 & Philip Schroeder469,471 Data collection member Emma F. Perez269 MexGen-COVID Analysis team lead Seung Hyuk T. Lee898 Data collection lead Fernando Daniel Flores-Silva899, Erwin Chiquete899, Carlos Cantú-Brito899, José Sifuentes-Osornio900, Alfredo Ponce de León-Garduño901, Carla Marina Román-Montes901, Emma Chávez-Manzanera902, Donaji Gómez Velasco903, Alexandro Martagón-Rosado903,904,905, Ivette Cruz-Bautista903, Francisco Javier Gómez-Pérez906, Claudia Díaz-Olavarrieta907, Paloma Almeda Valdés906, Linda Liliana Muñóz-Hernández903, Roopa Pravin Mehta906 & Daniel Elías-López903 Admin team lead Teresa Tusié-Luna908,909 & Päivi Pajukanta898,910 Analysis team member Mihir G. Sukhatme898, Hortensia Moreno-Macías911,912, Alicia Huerta-Chagoya913, Johnson Ruth914, Ana Ochoa-Guzmán915 & Sebastián González-Behn-Eschenburg915 Data collection member Miguel García-Grimshaw899, Luz Elizabeth Guillen-Pineda906, María de los Ángeles Vargas-Martínez899, Maribel Rodríguez915, Rosario Rodríguez-Guillén915, María Luisa Ordoñez915, Yayoi Segura-Kato915, Sandra Rajme-López916, María Fernanda González-Lara916, Dalia Luna-Moreno903, Jannet de Jesús Suárez López915, Daniela Mariel Guillén-Quintero915, Juan Vega915, Salvador Duran-Coyote899, María Teresa Guerra-García915, Michelle Durán-Gómez917, Alma Vigueras-Hernández899, Marco Eduardo Ramírez-Jiménez918, Samara Lissete Grajeda-González918, Enrique Manuel Ramos-Galicia918, Martha Fernanda Ramírez-Carrillo919, Juan Fernandez-Ruiz232,920, Amanda Chirino-Pérez899, Ángel Omar Romero-Molina920, David Alejandro Fuentes Zavaleta921, María Andrea Mac Donald Jaramillo918, Katia Jarely Prieto López918, Diego Incontri-Abraham918, Wendy Natalia Rosado918, María Alonso Catalán918, Sonia Evelyn Vital Aguilar915, Ileana Paola Montero Zamudio918, Eduardo Ruíz-Ruíz899, Kevin de Jesús Carrera García918, Cristina Villarreal Guerrero918, Ana Isabel Rodríguez Martínez918, Rogelio Moscoso-Sánchez918, María del Consuelo Rodríguez Mancilla902, Antonio Villa922, Samuel Alejandro Luna-Ramírez907, Sergio Hernández-Hernández915, Erika Frias907, Alejandro Campos903, Paola Alessandra Morales-Sosa915, Luis Enrique Toxqui-Merchant915 & Karla García915 Admin team member Bogdan Pasaniuc898,923,924, Manish J. Butte925,926, Daniel H. Geschwind898,910,927,928,929,930 & Carlos A. Aguilar-Salinas931 Pa-COVID-19 Data collection lead Bettina Heidecker932, Florian Kurth933,934,935 & Leif E. Sander933 Data collection member Carsten Skurk932, Ulf Landmesser932, Zehra Karadeniz932, Xiaomin Wang932, Wolfgang Poller933, Phillip Suwalski932, Stephan Ripke933, Alice Braun933, Julia Kraft933, Alena Mayer933, Lena J. Lippert933, Elisa T. Helbig933, Charlotte Thibeault933 & Swapnil Awasthi933 Qatar Genome Program Analysis team lead Hamdi Mbarek936 Data collection lead Said I. Ismail936 Analysis team member Chadi Saad936 & Yaser Al-Sarraj936 Data collection member Radja Messai Badji936, Wadha Al-Muftah936, Asma Al Thani936 & Nahla Afifi936 SARS-CoV-2 and Host Genome sequencing Data collection lead Fouzia Sadiq937 Admin team lead Mohammad I. Khan938,939 Data collection member Muhammad J. Bhutta940, Hania Khan941, Madeeha Khan937 & Quratul Ain937 Admin team member Ayesha Junaid942 & Ejaz A. Khan940 Search for Genomic Markers Predicting the Severity of the response to COVID-19 Analysis team lead Mateusz Sypniewski943 Data collection lead Adrian Lejman943 Analysis team member Paweł Zawadzki943,944, Elżbieta Kaja943,945 & Paweł Sztromwasser943 Data collection member Zbigniew Król946, Paula Dobosz946,947 & Joanna Stojak946 Spanish COalition to Unlock Research on host GEnetics on COVID-19 (SCOURGE) Analysis team lead Carlos Flores948,949,950,951 & Ángel Carracedo952,953,954,955,956 Data collection lead Carlos Aguilar957, Eunate Arana-Arri958,959, María J. Arranz960, Raúl C. Baptista-Rosas961,962,963, Joao F. Bezerra964, Marcos A. C. Bezerra965, Ramón Brugada966,967,968,969, Matilde Bustos970, Enrique J. Calderón971,972,973, Luis Castano953,958,974,975,976, Rosa Conde-Vicente977, M. Lourdes Cordero-Lorenzana978, Alba De Martino-Rodríguez979,980, Elena Domínguez-Garrido981, María Carmen Fariñas982,983,984, Belén Gil-Fournier985, María José Herrero986,987, María Claudia Lattig988,989, Caridad Martín-López990, Vicente Martín972,991, Oscar Martinez-Nieto989,992, Iciar Martinez-Lopez993,994, Juliana F. Mazzeu995,996,997, Eleuterio Merayo Macías998, Victor Moreno Cuerda999,1000, Vivian N. Silbiger1001, Silviene F. Oliveira995,996,997,1002,1003, Mara Parellada1004,1005,1006, Ney P. C. Santos1007, Patricia Pérez-Matute1008, Patricia Perez1009, Teresa Perucho1010, Gloria L. Porras-Hurtado1011, Aurora Pujol953,1012,1013, Amanda Fernández-Rodríguez1014,1015, María A. Jimenez-Sousa1014,1015, Emilio Rodríguez-Ruiz955,1016, Fernando Rodriguez-Artalejo972,1017,1018,1019, José A. Rodriguez-Garcia1020, Francisco Ruiz-Cabello232,1021,1022, José Manuel Soria1023, Eduardo Tamayo1024,1025, Alejandro Teper1026, Lilian Torres-Tobar1027, Miguel Urioste1028, Zuleima Yáñez1029, Ruth Zarate84, Encarna Guillen-Navarro1030,1031,1032,1033, Carmen Ayuso953,1034, Anna González-Neira1035, José A. Riancho982,983,984, Augusto Rojas-Martinez746 & Pablo Lapunzina953,1036,1037 Analysis team member Raquel Cruz952,953,955,956, Silvia Diz-de Almeida953,956, Francisco C. Ceballos1014, José M. Lorenzo-Salazar948 & Rafaela González-Montelongo948 Data collection member Inés Quintela952, Guillermo Pita1035, Manuela Gago-Domínguez954,955, Marta Sevilla Porras953,1036, Jair Antonio Tenorio-Castaño953,1036,1037, Julian Nevado953,1036,1037, Jose María Aguado1006,1015,1038,1039, Sergio Aguilera-Albesa1040,1041, Virginia Almadana1042, Berta Almoguera953,1034, Nuria Alvarez1035, Álvaro Andreu-Bernabeu1004,1006, Celso Arango1004,1005,1006, Maria-Jesus Artiga1043, María Barreda-Sánchez1030,1044, Moncef Belhassen-Garcia1045,1046, Lucía Boix-Palop1047, María Brion968,1048, Cristina Carbonell1046,1049, Jose E. Castelao1050, Jose L. Cortes-Sanchez746,1051, Marta Corton953,1034, M. Teresa Darnaude1052, Victor del Campo-Pérez1053, Aranzazu Diaz de Bustamante1052, Andre D. Luchessi1001, Rocío Eiros1054, Gladys Mercedes Estigarribia Sanabria1055, Uxía Fernández-Robelo1056, Tania Fernández-Villa1057, Javier Gómez-Arrue979,980, Beatriz González Álvarez979,980, Fernan Gonzalez Bernaldo de Quirós1058, Javier González-Peñas1004,1005,1006, Juan F. Gutiérrez-Bautista1021, Antonio Herrero-Gonzalez1059, Anabel Liger Borja990, Rosario Lopez-Rodriguez953,1034, Esther Mancebo1060,1061, Michel F. Martinez-Resendez746, Angel Martinez-Perez1023, Pablo Minguez953,1034, Eva Ortega-Paino1043, Estela Paz-Artal1060,1061,1062, M. Elena Pérez-Tomás1030, Mel·lina Pinsach-Abuin966,968, Ericka N. Pompa-Mera1063, Soraya Ramiro León985, Salvador Resino1014,1064, Marianne R. Fernandes1007,1065, Javier Ruiz-Hornillos1066,1067,1068, Pablo Ryan1069,1070,1071, Juan Carlos Souto1072, Alvaro Tamayo-Velasco1073, Juan Carlos Taracido-Fernandez1059 & Juan Valencia-Ramos1074 Admin team member Miguel López de Heredia953 The Danish Blood Donor Study Analysis team lead Ole B. Pedersen420, Christian Erikstrup1075, Sisse R. Ostrowski1076, Karina Banasik220, Søren Brunak220, Bjarke Feenstra421 & Kari Stefansson1077 Data collection lead Susan Mikkelsen1075 E18 | Nature | Vol 621 | 7 September 2023 Matters arising Admin team lead Erik Sørensen1076 & Henrik Ullum426 Analysis team member David Westergaard220, Celia Burgos220, Steffen Andersen1078, Kristoffer Brugdorf1076, Thomas F. Hansen1079, Henrik Hjalgrim421,1080, Gregor B. Jemec1081, Mette Nyegaard1082, Thomas Werge1083, Pär I. Johansson1076, Maria Didriksen1076, Frank Geller421, Michael Schwinn1076, Rikke L. Jacobsen1076, Lotte Hindhede1075, Klaus Rostgaard421,1080, Daniel Gudbjartsson1084, Hreinn Stefánsson1084 & Unnur Þorsteinsdóttir1077 Data collection member Kaspar R. Nielsen1085, Mie Topholm Bruun1086, Khoa M. Dinh1075, Margit A. H. Larsen1076 & Susanne G. Sækmose420 The Genetic Predisposition to severe COVID-19 Analysis team lead Mari E. K. Niemi100 Data collection lead Hugo Zeberg1087,1088 Analysis team member Mattia Cordioli104, Sara Pigazzini100 & Lindo Nkambul1,105,1089 Data collection member Robert Frithiof1090, Michael Hultström1090,1091, Miklos Lipcsey1090,1092, Nicolas Tardif1093, Olav Rooyackers1093, Jonathan Grip1093 & Tomislav Maricic1088 Tirschenreuth Study (TiKoco) Analysis team lead Iris M. Heid1094 Data collection lead Ralf Wagner1095 & Klaus Überla1096 Analysis team member Thomas W. Winkler1094, Simon Wiegrebe1094, Mathias Gorski1094 & Klaus J. Stark1094 Data collection member David Peterhoff1095, Sebastian Einhauser1095, Hans-Helmut Niller1095,1097 & Stephanie Beileke1096 UCLA Precision Health COVID-19 Host Genomics Biobank Analysis team lead Ruth Johnson1098 Data collection lead Bogdan Pasaniuc898,923,924 & Manish J. Butte1099,1101 Admin team lead Nelson Freimer1100, Manish J. Butte1099,1101, Daniel H. Geschwind1102 & Bogdan Pasaniuc923,1103 Analysis team member Yi Ding1104, Alec Chiu1104, Timothy S. Chang1105 & Paul Boutros1106 UK Biobank Analysis team lead Tomoko Nakanishi50,51,52,53, Konrad J. Karczewski4,28, Alicia R. Martin4,28, Daniel J. Wilson1107 & Chris C. A. Spencer1108 Data collection lead Derrick W. Crook1109, David H. Wyllie1109,1110 & Anne Marie O’Connell1111 Admin team lead J. Brent Richards51,112,113 Analysis team member Guillaume Butler-Laporte51,117, Vincenzo Forgetta51, Elizabeth G. Atkinson4,28,1112, Masahiro Kanai4,28,1113, Kristin Tsuo4,28,1114, Nikolas Baya4,28, Patrick Turley1115,1116, Rahul Gupta4,28, Raymond K. Walters4,28, Duncan S. Palmer4,28, Gopal Sarma4,28, Matthew Solomonson4,28, Nathan Cheng4,28, Wenhan Lu4,28, Claire Churchhouse4,28, Jacqueline I. Goldstein4,28, Daniel King4,28, Wei Zhou4,28, Cotton Seed4,28, Mark J. Daly1,4,1117, Benjamin M. Neale4,28, Hilary Finucane4,28, Sam Bryant1, F. Kyle Satterstrom4,28, Gavin Band1118, Sarah G. Earle1107, Shang-Kuan Lin1107, Nicolas Arning1107 & Nils Koelling1108 Data collection member Jacob Armstrong1107 & Justine K. Rudkin1107 Admin team member Shawneequa Callier1119, Sam Bryant4,28 & Caroline Cusick28 Variability in Immune Response Genes and Severity of SARS-CoV-2 Infection (INMUNGEN-CoV2 study) Analysis team lead Israel Fernandez-Cadenas26 Data collection lead Anna M. Planas1120,1121 Analysis team member Jordi Perez-Tur1122,1123, Laia Llucià-Carol26,1124, Natalia Cullell26,1125, Elena Muiño26, Jara Cárcel-Márquez26, Marta L. DeDiego1126 & Lara Lloret Iglesias1127 Data collection member Alex Soriano1128, Veronica Rico1129, Daiana Agüero1129, Josep L. Bedini1129, Francisco Lozano1130, Carlos Domingo1129, Veronica Robles1129, Francisca Ruiz-Jaén1131, Leonardo Márquez1132, Juan Gomez1133, Eliecer Coto1133, Guillermo M. Albaiceta1133, Marta García-Clemente1133, David Dalmau1134, Maria J. Arranz1134, Beatriz Dietl1134, Alex Serra-Llovich1134, Pere Soler-Palacin1135, Roger Colobran1135, Andrea Martin-Nalda1135, Alba Parra-Martínez1135, David Bernardo1136, Silvia Rojo1137, Aida Fiz-López1136, Elisa Arribas-Rodriguez1136, Paloma Cal-Sabater1136, Tomás Segura1138, Esther González-Villar1138, Gemma Serrano-Heras1139, Joan Martí-Fàbregas1140, Elena Jiménez-Xarrié1140, Alicia de Felipe Mimbrera1141, Jaime Masjuan1141, Sebastian García-Madrona1141, Anna Domínguez-Mayoral1142, Joan Montaner Villalonga1142 & Paloma Menéndez-Valladares1142 Website development Matthew Solomonson1 1Broad Institute of MIT and Harvard, Cambridge, MA, USA. 2University of California San Francisco, San Francisco, CA, USA. 3Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland. 4Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, MA, USA. 5Department of Psychiatry, School of Medicine, Yale University, New Haven, CT, USA. 6Veteran Affairs Connecticut Healthcare System, West Haven, CT, USA. 7Yale University, New Haven, CT, USA. 8Centre for Bioinformatics and Data Analysis, Medical University of Bialystok, Bialystok, Poland. 9Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, MI, USA. 10Faculty of Industrial Sciences and Technology, Universiti Malaysia Pahang, Pahang, Malaysia. 11Icahn School of Medicine at Mount Sinai, New York, NY, USA. 12Genolier Innovation Network and Hub, Swiss Medical Network, Genolier Healthcare Campus, Genolier, Switzerland. 13Immediate, Independent Multidisciplinary Med & Biotech Research Association, Milan, Italy. 14Clinical Research Unit of Nanoro, Institut de Recherche en Sciences de la Santé, Nanoro, Burkina Faso. 15Sydney Brenner Institute for Molecular Biosciences, University of the Witwatersrand, Johannesburg, South Africa. 16McGill University, Montreal, Quebec, Canada. 17University of Peurto Rico, San Juan, PR, USA. 18Institute for Biomedical Technologies, National Research Council, Milan, Italy. 19University of Miami, Miami, FL, USA. 20Department of Biochemistry, All India Institute of Medical Sciences, Kalyani, India. 21Department of Biosciences, School of Science and Technology, Nottingham Trent University, Nottingham, UK. 22Department of Clinical Pathology, Faculty of Medicine, Mansoura University, Mansoura, Egypt. 23Department of Clinical Pathology, Faculty of Medicine, Tanta University, Tanta, Egypt. 24Neuron23, San Francisco, CA, USA. 25Vanderbilt University, Nasvhille, TN, USA. 26Stroke Pharmacogenomics and Genetics, Institut d’Investigació Biomèdica Sant Pau (IIBSANT PAU), Barcelona, Spain. 27Department of Medicine, Surgery and Health Sciences, University of Trieste, Trieste, Italy. 28Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA. 29Department of Genetics, University Medical Centre Groningen, University of Groningen, Groningen, The Netherlands. 30Departamento de Tecnología Médica, Facultad de Ciencias de la Salud, Universidad de Tarapacá, Arica, Chile. 31King’s College London, London, UK. 32Naina Tech, New York, NY, USA. 33Vrije Universiteit Amsterdam, Amsterdam, The Netherlands. 34Medical School, University of Pécs, Pécs, Hungary. 35MNM Diagnostics, Poznan, Poland. 36Genomics England, London, UK. 37Heart Institute (InCor), University of Sao Paulo Medical School, Sao Paulo, Brazil. 38Department of Physiology, All India Institute of Medical Sciences, Kalyani, India. 39Montreal Heart Institute, Montreal, Quebec, Canada. 40Nantes Université, Ecole Centrale Nantes, INSERM, Center for Research in Transplantation and Translational Immunology, Nantes, France. 41Department of Computer Science and Technology, University of Cambridge, Cambridge, UK. 42William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London, UK. 43University of Helsinki, Helsinki, Finland. 44Department of Internal Medicine, Erasmus MC, University Medical Center Rotterdam, Rotterdam, The Netherlands. 45Fonds de la Recherche Scientifique (FNRS) & Centre de Génétique Humaine, Hôpital Erasme, Université Libre de Bruxelles, Brussels, Belgium. 46Department of Genetics, Hôpital Erasme, Université Libre de Bruxelles, Brussels, Belgium. 47All India Institute of Medical Sciences Kalyani, Kalyani, India. 48National Laboratory of Genomics for Biodiversity, CINVESTAV, Guanajuato, Mexico. 49Queensland University of Technology, Brisbane, Queensland, Australia. 50Department of Human Genetics, McGill University, Montreal, Quebec, Canada. 51Lady Davis Institute, Jewish General Hospital, McGill University, Montreal, Quebec, Canada. 52Kyoto-McGill International Collaborative School in Genomic Medicine, Graduate School of Medicine, Kyoto University, Kyoto, Japan. 53Japan Society for the Promotion of Science, Tokyo, Japan. 54Division of Life Sciences, University of Northampton, Northampton, UK. 55Root Deep, Boston, MA, USA. 56Instituto do Coração, Sao Paulo, Brazil. 57Stanford University, Stanford, CA, USA. 58Department of Internal Medicine, Erasmus MC, University Medical Center Rotterdam, Rotterdam, The Netherlands. 59Department of Epidemiology, Erasmus MC, University Medical Center Rotterdam, Rotterdam, The Netherlands. 60Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany. 61GI and Liver Section, Virgen del Rocío University Hospital and Department of Medicine, University of Seville, Institute of Biomedicine of Seville, Seville, Spain. 62CIBEREHD, Instituto de Salud Carlos III (ISCIII), Seville, Spain. 63Department of Health, Traunstein, Germany. 64Rutgers Cancer Institute of New Jersey, New Brunswick, NJ, USA. 65Erasmus Medical Center, Rotterdam, The Netherlands. 66Institute of Psychiatric Phenomics & Genomics, University of Munich, Munich, Germany. 67Department of Psychiatry, University of Munich, Munich, Germany. 68Institute of Virology, Technical University of Munich/Helmholtz Munich, Munich, Germany. 69Institute of Human Genetics, University Hospital, Faculty of Medicine, University of Bonn, Bonn, Germany. 70Department of Psychiatry, University Hospital, Faculty of Medicine, University of Bonn, Bonn, Germany. 71Digital Health Center, Hasso Plattner Institute, University of Potsdam, Potsdam, Germany. 72Institute of Biomedicine and Cancer Research Laboratories, Western Cancer Centre FICAN West, University of Turku, Turku, Finland. 73IRCCS Istituto Giannina Gaslini, Genova, Italy. 74Department of Neurosciences, Rehabilitation, Ophthalmology, Genetics, Maternal and Child Health, University of Genova, Genova, Italy. 75Department of Biomedical Data Science, Stanford University, Stanford, CA, USA. 76Fundación Pública Andaluza para la Gestión de la Investigación en Salud de Sevilla, Seville, Spain. 77Department of Pediatrics, Stanford University, Stanford, CA, USA. 78Department of Internal Medicine, Erasmus MC, University Medical Center Rotterdam, Rotterdam, The Netherlands. 79University Medical Center Utrecht, Utrecht, The Netherlands. 80Instituto de Investigación Interdisciplinaria & Escuela de Medicina, Universidad de Talca, Talca, Chile. 81Programa de Genética Humana, ICBM and Chile Departamento de Oncología Básico Clínica, Universidad de Chile, Talca, Chile. 82Department of Statistical Genetics, Osaka University Graduate School of Medicine, Osaka, Japan. 83Department of Internal Medicine, International Islamic University Malaysia, Pahang, Nature | Vol 621 | 7 September 2023 | E19 Malaysia. 84Centro para el Desarrollo de la Investigación Científica (CEDIC), Asunción, Paraguay. 8523andMe, Sunnyvale, CA, USA. 86Washington University School Of Medicine, St Louis, MO, USA. 87Department of Psychological & Brain Sciences, Washingon University in St Louis, St Louis, MO, USA. 88Institute of Biological Psychiatry, Mental Health Centre Sct Hans, Mental Health Services Capital Region of Denmark, Copenhagen, Denmark. 89iPSYCH, The Lundbeck Foundation Initiative for Integrative Psychiatric Research, Aarhus, Denmark. 90Laureate Institute for Brain Research, Tulsa, OK, USA. 91Department of Radiology, UCSD, San Diego, CA, USA. 92Nork Infection Clinical Hospital, Yerevan, Armenia. 93Institute of Molecular Biology NAS RA, Yerevan, Armenia. 94National Research Center for Cardiac Surgery, Astana, Kazakhstan. 95City Infectious Disease Center, Astana, Kazakhstan. 96National Center for Biotechnology, Astana, Kazakhstan. 97Health Department of Astana, Astana, Kazakhstan. 98MRC Integrative Epidemiology Unit, University of Bristol, Bristol, UK. 99Division of Psychitary, University of Edinburgh, Edinburgh, UK. 100Institute for Molecular Medicine Finland (FIMM), Helsinki, Finland. 101University of Liege, GIGA-Institute, Liege, Belgium. 102CHC Mont-Légia, Liege, Belgium. 103BHUL (Liège Biobank), CHU of Liège, Liege, Belgium. 104Institute for Molecular Medicine Finland, University of Helsinki, Helsinki, Finland. 105Analytic & Translational Genetics Unit, Massachusetts General Hospital, Boston, MA, USA. 106Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, MA, USA. 107CHU of Liege, Liege, Belgium. 108University of Liege, Liege, Belgium. 109Centre de Génétique Humaine, Hôpital Erasme, Université Libre de Bruxelles, Brussels, Belgium. 110Service de Médecine Interne, Hôpital Erasme, Université Libre de Bruxelles, Brussels, Belgium. 111McGill Genome Centre and Department of Human Genetics, McGill University, Montréal, Quebec, Canada. 112Department of Human Genetics, Epidemiology, Biostatistics and Occupational Health, McGill University, Montreal, Quebec, Canada. 113Department of Twin Research, King’s College London, London, UK. 114The Meakins-Christie Laboratories at the Research Institute of the McGill University Heath Centre Research Institute and Department of Medicine, Faculty of Medicine, McGill University, Montreal, Quebec, Canada. 115Research Centre of the Centre Hospitalier de l’Université de Montréal (CRCHUM), Montreal, Quebec, Canada. 116Centre Hospitalier de l’Université de Montréal (CHUM), Montreal, Quebec, Canada. 117Department of Epidemiology, Biostatistics and Occupational Health, McGill University, Montreal, Quebec, Canada. 118Department of Emergency Medicine, McGill University, Montreal, Quebec, Canada. 119Emergency Department, Jewish General Hospital, McGill University, Montreal, Quebec, Canada. 120McGill AIDS Centre, Department of Microbiology and Immunology, Lady Davis Institute for Medical Research, Jewish General Hospital, McGill University, Montreal, Quebec, Canada. 121Division of Infectious Diseases, Lausanne University Hospital and University of Lausanne, Lausanne, Switzerland. 122Research Centre of the Centre Hospitalier de l’Université de Montréal and Université de Montréal, Montreal, Quebec, Canada. 123School of Rehabilitation, Faculty of Medicine and Health Sciences, Université de Sherbrooke, Sherbrooke, Canada. 124Department of Medicine, Faculty of Medicine, Université de Montréal, Montreal, Quebec, Canada. 125Center for Immunity, Inflammation and Infectious Diseases, Montreal Clinical Research Institute (IRCM), Montreal, Quebec, Canada. 126Génome Québec, Montreal, Quebec, Canada. 127Département de Microbiologie et Infectiologie, Université de Sherbrooke, Sherbrooke, Quebec, Canada. 128Département de Médecine, Service d’Infectiologie, Centre de Recherche Clinique du Centre Hospitalier Universitaire de Sherbrooke, Sherbrooke, Quebec, Canada. 129Department of Epidemiology, Biostatistics and Occupational Health, McGill University, Montreal, Quebec, Canada. 130Center for Clinical Epidemiology, Lady Davis Institute, Jewish General Hospital, Montreal, Quebec, Canada. 131Department of Neurology and Neurosurgery, McGill University, Montreal, Quebec, Canada. 132Centre Intégré Universitaire de Santé et de Services Sociaux (CIUSSS) du Saguenay– Lac-Saint-Jean, Saguenay, Quebec, Canada. 133Department of Pharmacology-Physiology, Medicine and Health Sciences Faculty, Université de Sherbrooke, Sherbrooke, Quebec, Canada. 134Division of Respiratory Medicine, Department of Pediatrics, Sainte-Justine University Hospital Center, University of Montreal, Montreal, Quebec, Canada. 135Centre of Genomics and Policy, McGill University, Montreal, Quebec, Canada. 136Division of Genetic Medicine, Department of Medicine, Vanderbilt University Medical Center, Nashville, TN, USA. 137Vanderbilt Genetics Institute, Vanderbilt University Medical Center, Nashville, TN, USA. 138Institute of Human Genetics, University Hospital Bonn, Medical Faculty University of Bonn, Bonn, Germany. 139Institute of Genomic Statistics and Bioinformatics, University Hospital Bonn, Medical Faculty University of Bonn, Bonn, Germany. 140Department of Gastroenterology, Hepatology and Infectious Diseases, University Hospital Duesseldorf, Medical Faculty Heinrich Heine University, Duesseldorf, Germany. 141Department of Gastroenterology, Hepatology and Infectiology, University Hospital Magdeburg, Magdeburg, Germany. 142Clinic for Cardiology, Angiology and Internal Intensive Medicine, Medical Clinic I, RWTH Aachen University, Aachen, Germany. 143Department of Pneumology and Intensive Care Medicine, Faculty of Medicine, RWTH Aachen University, Aachen, Germany. 144Department of Pneumology, Hannover Medical School, Hannover, Germany. 145Department of Gastroenterology, Hepatology and Endocrinology, Hannover Medical School, Hannover, Germany. 146Hannover Unified Biobank, Hannover Medical School, Hannover, Germany. 147Department I of Internal Medicine, Faculty of Medicine and University Hospital of Cologne, University of Cologne, Cologne, Germany. 148Center for Molecular Medicine Cologne (CMMC), University of Cologne, Cologne, Germany. 149German Center for Infection Research (DZIF), partner site Bonn-Cologne, Cologne, Germany. 150Institute for Human Genetics and Genomic Medicine, Medical Faculty, RWTH Aachen University, Aachen, Germany. 151Department of Anesthesiology and Intensive Care Medicine, University Hospital Essen, University Duisburg-Essen, Essen, Germany. 152Department of Child and Adolescent Psychiatry, University Hospital Essen, University of Duisburg-Essen, Essen, Germany. 153Department of Infectious Diseases, University Hospital Essen, University Duisburg-Essen, Essen, Germany. 154Department of Pneumology, Allergology and Respiratory Medicine, University Hospital Saarland, Homburg, Germany. 155Center of Human and Molecular Biology, Department of Human Genetics, University Hospital Saarland, Homburg, Germany. 156Department of Genetics & Epigenetics, Saarland University, Saarbrücken, Germany. 157Institute of Innate Immunity, University Hospital Bonn, Bonn, Germany. 158University of Toronto, Toronto, Ontario, Canada. 159Dalla Lana School of Public Health, University of Toronto, Toronto, Ontario, Canada. 160Lunenfeld-Tanenbaum Research Institute, Sinai Health, Toronto, Ontario, Canada. 161Simon Fraser University, Burnaby, British Columbia, Canada. 162Lady Davis Institute, Jewish General Hospital, McGill University, Montreal, Quebec, Canada. 163University of Waterloo, Waterloo, Ontario, Canada. 164The Hospital for Sick Children, Toronto, Ontario, Canada. 165Queen’s University, Kingston, Ontario, Canada. 166Mount Sinai Hospital, Sinai Health, Toronto, Ontario, Canada. 167St Michael’s Hospital, Unity Health Toronto, Toronto, Ontario, Canada. 168Princess Margaret Cancer Center, University Health Network, Toronto, Ontario, Canada. 169Ontario Institute for Cancer Research, Toronto, Ontario, Canada. 170Dynacare Laboratory, Toronto, Ontario, Canada. 171William Osler Health System, Brampton, Ontario, Canada. 172University Health Network, Toronto, Ontario, Canada. 173Mackenzie Health, Toronto, Ontario, Canada. 174Women’s College Hospital, Toronto, Ontario, Canada. 175University of British Columbia, Vancouver, British Columbia, Canada. 176BC Children’s Hospital, Vancouver, British Columbia, Canada. 177St Paul’s Hospital, Vancouver, British Columbia, Canada. 178University of Calgary, Calgary, Alberta, Canada. 179Alberta Children’s Hospital, Calgary, Alberta, Canada. 180British Columbia Cancer Research Centre, Vancouver, British Columbia, Canada. 181Toronto General Hospital, Toronto, Ontario, Canada. 182Division of Infectious Diseases, Department of Medicine, University of Ottawa, Ottawa, Ontario, Canada. 183Department of Biochemistry, Microbiology and Immunology, University of Ottawa Centre of Infection, Immunity and Inflammation (CI3), Ottawa, Ontario, Canada. 184The Ottawa Hospital Research Institute, Ottawa, Ontario, Canada. 185University of Montreal, Montreat, Quebec, Canada. 186Ste-Justine University Health Centre, Montreal, Quebec, Canada. 187University of Toronto Ontario, Toronto, Ontario, Canada. 188MitogenDx/Eve Technologies, Toronto, Ontario, Canada. 189Genome Sciences Centre, BC Cancer, Vancouver, British Columbia, Canada. 190Montreal Children’s Hospital, Montreal, Quebec, Canada. 191London Health Sciences Centre, London, Ontario, Canada. 192Kidney and Metabolism Program, St Michael’s Hospital, Unity Health Toronto, Toronto, Ontario, Canada. 193Sinai Health, Mount Sinai Hospital, Toronto, Ontario, Canada. 194Baycrest Health Sciences, Toronto, Ontario, Canada. 195Anschutz Medical Campus, University of Colorado, Aurora, CO, USA. 196Institute of Psychiatric Phenomics and Genomics (IPPG), University Hospital, LMU Munich, Munich, Germany. 197Department of Genetic Epidemiology in Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, University of Heidelberg, Mannheim, Germany. 198Department of Psychiatry and Behavioral Sciences, Johns Hopkins University, Baltimore, MD, USA. 199Department of Psychiatry and Behavioral Sciences, SUNY Upstate Medical University, Syracuse, NY, USA. 200Department of Psychiatry and Psychotherapy, University Hospital, LMU Munich, Munich, Germany. 201Institute of Virology, Technical University Munich/ Helmholtz Zentrum München, Munich, Germany. 202Department of Psychiatry and Psychotherapy, University Hospital Bonn, Medical Faculty University of Bonn, Bonn, Germany. 203CIBER de Salud Mental, Instituto de Salud Carlos III, Barcelona, Spain. 204Munich Clinic Schwabing, Academic Teaching Hospital, Ludwig-Maximilians University of Munich (LMU), Munich, Germany. 205Department of Internal Medicine II, University Hospital rechts der Isar, Technical University of Munich, School of Medicine, Munich, Germany. 206Institute of Clinical Chemistry and Pathobiochemistry, School of Medicine, Technical University of Munich, Munich, Germany. 207TranslaTUM, Center for Translational Cancer Research, Technical University of Munich, Munich, Germany. 208Department of Psychiatry and Psychotherapy, University Medical Center Goettingen, Goettingen, Germany. 209German Center for Neurodegenerative Diseases (DZNE), Goettingen, Germany. 210Neurosciences and Signaling Group, Institute of Biomedicine (iBiMED), Department of Medical Sciences, University of Aveiro, Agra do Crasto, Aveiro, Portugal. 211Munich Site of the German Center for Infection Research (DZIF), Munich, Germany. 212Institute of Computational Biology, Helmholtz Center Munich, Oberschleissheim, Germany. 213Department of Hygiene and Epidemiology, University of Ioannina School of Medicine, Ioannina, Greece. 2141st Department of Internal Medicine and Infectious Diseases Unit, University Hospital of Ioannina, Ioannina, Greece. 215Department of Epidemiology and Biostatistics, School of Public Health, Imperial College London, London, UK. 216Center for Evidence-Based Medicine, Department of Health Services, Policy and Practice, School of Public Health, Brown University, Providence, RI, USA. 217University of Ioannina School of Medicine, Ioannina, Greece. 218Centre for Systems Biology, Biomedical Research Foundation of the Academy of Athens, Athens, Greece. 219Institute of Clinical Molecular Biology, Christian-Albrechts-University, Kiel, Germany. 220Novo Nordisk Foundation Center for Protein Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark. 221Institut de Biotecnologia i de Biomedicina, Universitat Autònoma de Barcelona, Barcelona, Spain. 222ICREA, Barcelona, Spain. 223Institute of Biotechnology, Life Science Centre, Vilnius University, Vilnius, Lithuania. 224Research Group for Evolutionary Immunogenomics, Max Planck Institute for Evolutionary Biology, Plön, Germany. 225Research Unit for Evolutionary Immunogenomics, Department of Biology, University of Hamburg, Hamburg, Germany. 226Department of Gastroenterology, Hospital Universitario Ramón y Cajal, University of Alcalá, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), Madrid, Spain. 227Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII), Madrid, Spain. 228Vall d’Hebron Institut de Recerca (VHIR), Vall d’Hebron Hospital Universitari, Barcelona, Spain. 229Fondazione IRCCS Ca’ Granda Ospedale Maggiore Policlinico, Milan, Italy. 230Department of Medicine, Møre & Romsdal Hospital Trust, Ålesund, Norway Geminicenter for Sepsis Research, Institute of Circulation and Medical Imaging (ISB), NTNU, Trondheim, Norway. 231Microbiology Unit, Hospital Universitario Clinico San Cecilio, Granada, Spain. 232Ibs.Granada Instituto de Investigación Biosanitaria, Granada, Spain. 233Klinik für Innere Medizin I, Universitätsklinikum Schleswig-Holstein, Campus Kiel, Kiel, Germany. 234Emergency Department, University Hospital Regensburg, Regensburg, Germany. 235Department for Infectious Diseases and Infection Control, University Hospital Regensburg, Regensburg, Germany. 236Department of Medicine I, Gastroenterology, Hepatology and Endocrinology, Medical University of Innsbruck, Innsbruck, Austria. 237Christian Doppler Laboratory of Iron and Phosphate Biology at the Department of Medicine I, Medical University of Innsbruck, Innsbruck, Austria. 238Institute of Clinical Medicine, University of Oslo, Oslo, Norway. 239Department of Microbiology, Oslo University Hospital, Oslo, Norway. 240Hospital Clinic, University of Barcelona and IDIBAPS, Barcelona, Spain. 241European Foundation for the Study of Chronic Liver Failure (EF-CLIF), Barcelona, Spain. 242GCAT—Genomes for Life, Germans Trias i Pujol Health Sciences Research Institute (IGTP), Badalona, Spain. 243Genomes for Life—GCAT Laboratory, Germans Trias i Pujol Research Institute (IGTP), Spainermans Trias i Pujol Research Institute (IGTP), Badalona, Spain. 244University of Milan, Milan, Italy. 245Department of Liver and Gastrointestinal Diseases, Biodonostia Health Research Institute, Donostia University Hospital, University of the Basque Country (UPV/EHU), CIBEREHD, Ikerbasque, San Sebastian, Spain. 246Digestive Diseases Unit, Virgen del Rocio University Hospital, Institute of Biomedicine of Seville, University of Seville, Seville, Spain. 247University E20 | Nature | Vol 621 | 7 September 2023 Matters arising of Sevilla, Seville, Spain. 248Instituto de Biomedicina de Sevilla (IBIS), Seville, Spain. 249Hospital Universitario Virgen del Rocío de Seville, Seville, Spain. 250Universitat Autònoma de Barcelona, Bellatera, Spain. 251Liver Unit, Department of Internal Medicine, Hospital Universitari Vall d’Hebron, Vall d’Hebron Barcelona Hospital Campus, Barcelona, Spain. 252Gastrointestinal Genetics Lab, CIC bioGUNE—BRTA, Derio, Spain. 253Ikerbasque, Basque Foundation for Science, Bilbao, Spain. 254Department of Liver and Gastrointestinal Diseases, Biodonostia Health Research Institute—Donostia University Hospital, University of the Basque Country (UPV/EHU), CIBEREHD, Ikerbasque, San Sebastian, Spain Ikerbasque, Basque Foundation for Science, Bilbao, Spain. 255Department I of Internal Medicine, Faculty of Medicine and University Hospital Cologne, University of Cologne, Cologne, Germany. 256University of Cologne, Faculty of Medicine and University Hospital Cologne, Cologne Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD), Cologne, Germany. 257European Reference Network on Hepatological Diseases (ERN RARE-LIVER), Fondazione IRCCS San Gerardo dei Tintori, Monza, Italy. 258Division of Gastroenterology, Center for Autoimmune Liver Diseases, School of Medicine and Surgery, University of Milano-Bicocca, Milan, Italy. 259Genomes for Life—GCAT lab, Germans Trias i Pujol Research Institute (IGTP), Badalona, Spain. 260IRCCS Humanitas Research Hospital, Milan, Italy. 261Department of Biomedical Sciences, Humanitas University, Pieve Emanuele, Milan, Italy. 262Zentrum für Humangenetik Regensburg, Regensburg, Germany. 263University Hospital Schleswig-Holstein (UKSH), Campus Kiel, Kiel, Germany. 264Section for Gastroenterology, Department of Transplantation Medicine, Division for Cancer Medicine, Surgery and Transplantation, Oslo University Hospital Rikshospitalet, Oslo, Norway. 265Research Institute for Internal Medicine, Division of Surgery, Inflammatory Diseases and Transplantation, Oslo University Hospital Rikshospitalet and University of Oslo, Oslo, Norway. 266Norwegian PSC Research Center, Department of Transplantation Medicine, Division of Surgery, Inflammatory Diseases and Transplantation, Oslo University Hospital Rikshospitalet, Oslo, Norway. 267Private University in the Principality of Liechtenstein, Triesen, Liechtenstein. 268Division of Rheumatology, Inflammation and Immunity, Brigham and Women’s Hospital and Harvard Medical School, Boston, MA, USA. 269Division of Genetics, Department of Medicine, Brigham and Women’s Hospital, Boston, MA, USA. 270Department of Biomedical Informatics, Harvard Medical School, Boston, MA, USA. 271Center for Data Sciences, Brigham and Women’s Hospital, Boston, MA, USA. 272Institut de Biotecnologia i de Biomedicina, Universitat Autònoma de Barcelona, Bellaterra, Barcelona, Spain. 273Institute for Medical Microbiology and Hospital Epidemiology, Hannover Medical School, Hannover, Germany. 274Randaberg Municipality, Randaberg, Norway. 275Department of Quality and Health Technology, Faculty of Health Sciences, University of Stavanger, Stavanger, Norway. 276University Hospital Schleswig-Holstein, Campus Kiel, Kiel, Germany. 277Research Department, Stavanger University Hospital, Stavanger, Norway. 278Department of Genetics and Bioinformatics (HDGB), Division of Health Data and Digitalization, Norwegian Institute of Public Health, Oslo, Norway. 279Centre for Genetics and Genomics Versus Arthritis, Centre for Musculoskeletal Research, Manchester Academic Health Science Centre, The University of Manchester, Manchester, UK. 280Department of Intensive Care, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá, Madrid, Spain. 281Clinical Biochemistry Department, Donostialdea Integrated Health Organisation, Osakidetza Basque Health Service, San Sebastian, Spain. 282Research Center and Memory Clinic, Ace Alzheimer Center Barcelona—Universitat Internacional de Catalunya, Barcelona, Spain. 283CIBERNED, Network Center for Biomedical Research in Neurodegenerative Diseases, National Institute of Health Carlos III, Madrid, Spain. 284Department of Biochemistry, University Hospital Vall d’Hebron, Barcelona, Spain. 285Department of Acute Medicine, Oslo University Hospital, Oslo, Norway. 286Division of Neurogenetics and Molecular Psychiatry, Department of Psychiatry and Psychotherapy, Faculty of Medicine and University Hospital Cologne, University of Cologne, Cologne, Germany. 287Department of Neurodegenerative Diseases and Geriatric Psychiatry, Medical Faculty, University Hospital Bonn, Bonn, Germany. 288German Center for Neurodegenerative Diseases (DZNE), Bonn, Germany. 289Department of Psychiatry, Glenn Biggs Institute for Alzheimer’s and Neurodegenerative Diseases, San Antonio, TX, USA. 290Department of Anesthesiology and Intensive Care, University Hospital of North Norway, Tromsø, Norway. 291Pediatrics and Tettamanti Center, Fondazione IRCCS San Gerardo dei Tintori and School of Medicine and Surgery, University of Milano-Bicocca, Monza, Italy. 292Biochemistry Department, Echevarne Laboratory, Sant Cugat del Vallés, Barcelona, Spain. 293Centre for Multidisciplinary Research in Health Science (MACH), University of Milan, Milan, Italy. 294Gastroenterology Unit, Fondazione IRCCS Casa Sollievo della Sofferenza, San Giovanni Rotondo, Italy. 295Microbiology Unit, Fondazione IRCCS San Gerardo, Monza, Italy. 296Department of Infectious Diseases, Oslo University Hospital, Oslo, Norway. 297Humanitas Gavazzeni-Castelli, Bergamo, Italy. 298Microbiology Department, Hospital Universitari Vall d’Hebron, Barcelona, Spain. 299Department of Respiratory Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá, Centro de Investigación Biomédica en Red en Enfermedades Respiratorias (CIBERES), Madrid, Spain. 300Department of Respiratory Medicine and Allergology, University Hospital, Goethe University, Frankfurt am Main, Germany. 301Centro de Investigación Biomédica en Red de Enfermedades Respiratorias (CIBERES), Madrid, Spain. 302Centro de Investigación Biomédica en Red de Epidemiología y Salud Pública (CIBERESP), Madrid, Spain. 303Consejo Superior de Investigaciones Científicas, Seville, Spain. 304Department of Infectious Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá, Madrid, Spain. 305Department of Transfusion Medicine and Haematology Laboratory, San Gerardo Hospital, Monza, Italy. 306Respiratory Medicine & International Health, University of Lübeck, Lübeck, Germany. 307Division of Clinical Infectious Diseases, Research Center Borstel, Borstel, Germany. 308Clinical Tuberculosis Unit, German Center for Infection Research (DZIF), Borstel, Germany. 309Internal Medicine Department, Fondazione IRCCS Ca’ Granda Ospedale Maggiore Policlinico, Milan, Italy. 310Humanitas Clinical and Research Center, IRCCS, Milan, Italy. 311Respiratory Service, Basurto University Hospital, Osakidetza Basque Health Service, Bilbao, Spain. 312Department of Clinical and Molecular Medicine, Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology, Trondheim, Norway. 313Department of Medicine, Møre & Romsdal Hospital Trust, Ålesund, Norway. 314Department of Internal Medicine II, Medical University of Innsbruck, Innsbruck, Austria. 315Department of Anesthesiology and Critical Care, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá, Madrid, Spain. 316Immunohematology Department, Banc de Sang i Teixits, Autonomous University of Barcelona, Barcelona, Spain. 317European Reference Network on Hepatological Diseases (ERN RARE-LIVER), San Gerardo Hospital, Monza, Italy. 318Department of Medical Sciences, Università degli Studi di Torino, Turin, Italy. 319Respiratory Service, Galdakao Hospital, Osakidetza Basque Health Service, Galdakao, Spain. 320Biocruces Bizkaia Health Research Institute, Barakaldo, Spain. 321Department of Infectious Diseases, E.O. Ospedali Galliera, Genova, Italy. 322Accident & Emergency and Emergency Medicine Unit, San Gerardo Hospital, Monza, Italy. 323Histocompatibilidad y Biologia Molecular, Centro de Transfusion de Madrid, Madrid, Spain. 324Dino Ferrari Center, Department of Pathophysiology and Transplantation, University of Milan, Milan, Italy. 325Neurology Unit, IRCCS Fondazione Ca’ Granda Ospedale Maggiore Policlinico, Milan, Italy. 326Department of Pathophysiology and Transplantation, University of Milan, Milan, Italy. 327Angelo Bianchi Bonomi Hemophilia and Thrombosis Center, Fondazione IRCCS Ca’ Granda Ospedale Maggiore Policlinico, Milan, Italy. 328Respiratory Unit, Fondazione IRCCS Ca’ Granda Ospedale Maggiore Policlinico, Milan, Italy. 329Department of Pathophysiology and Transplantation, Università degli Studi di Milano, Milan, Italy. 330Internal Medicine Department, Virgen del Rocio University Hospital, Seville, Spain. 331Biochemistry Department, University Hospital Vall d’Hebron, Barcelona, Spain. 332Centre for Medical Sciences-CISMed, University of Trento, Trento, Italy. 333Anesthesia and Intensive Care, Santa Chiara Hospital, APSS Trento, Trento, Italy. 334Department of Anesthesiology, Intensive Care and Emergency, Fondazione IRCCS Ca’ Granda Ospedale Maggiore Policlinico, Milan, Italy. 335Fondazione Grigioni per il Morbo di Parkinson and Parkinson Institute, ASST Gaetano Pini-CTO, Milan, Italy. 336School of Medicine and Surgery, University of Milano-Bicocca, Milan, Italy. 337Acute Geriatric Unit, IRCCS San Gerardo Foundation, Monza, Italy. 338Neurointensive Care Unit, IRCCS San Gerardo dei Tintori, Monza, Italy. 339Emergency Department, Anesthesia and Intensive Care, IRCCS San Gerardo dei Tintori, Monza, Italy. 340Department of Anesthesiology, Intensive Care Medicine and Pain Therapy, University Hospital Frankfurt, Frankfurt am Main, Germany. 341Acute Geriatric Unit, IRCCS San Gerardo Foundation Monza, Monza, Italy. 342Department of Medical Microbiology, Clinic of Laboratory Medicine, St Olavs Hospital, Trondheim, Norway. 343Department of Internal Medicine I, University Medical Center Schleswig-Holstein, Kiel, Germany. 344Department of Infectious Diseases, St Olavs Hospital, Trondheim University Hospital, Trondheim, Norway. 345Department of Clinical and Molecular Medicine, NTNU, Trondheim, Norway. 346Institute of Parasitology and Biomedicine Lopez-Neyra, CSIC, Granada, Spain. 347Institute for Cardiogenetics, University of Lübeck, Lübeck, Germany. 348German Research Center for Cardiovascular Research, partner site Hamburg–Lübeck–Kiel, Lübeck, Germany. 349University Heart Center Lübeck, Lübeck, Germany. 350Respiratory ICU, Institut Clínic Respiratory, Hospital Clinic, University of Barcelona, and IDIBAPS, Barcelona, Spain. 351Bioinformatics Area, Fundación Progreso y Salud and Institute of Biomedicine of Sevilla (IBIS), Seville, Spain. 352Department of Research, Ostfold Hospital Trust, Gralum, Norway. 353Department of Infectious Diseases, Hospital Universitario Clinico San Cecilio, Granada, Spain. 354Infectious Diseases Service, Osakidetza, Biocruces Bizkaia Health Research Institute, Barakaldo, Spain. 355Department of Research, St Olav Hospital, Trondheim University Hospital, Trondheim, Norway. 356Novo Nordisk Foundation Center for Protein Research, Disease Systems Biology, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark. 357Medical Department, Drammen Hospital, Vestre Viken Hospital Trust, Gjettum, Norway. 358Research Center Borstel, BioMaterialBank Nord, Sülfeld, Germany. 359German Center for Lung Research (DZL), Airway Research Center North (ARCN), Grosshansdorf, Germany. 360Popgen 2.0 Network (P2N), Kiel, Germany. 361Department of Clinical Science, University of Bergen, Bergen, Norway. 362Pediatric Departement and Centro Tettamanti—European Reference Network (ERN) PaedCan, EuroBloodNet, MetabERN- University of Milano-Bicocca-Fondazione MBBM/Ospedale San Gerardo, San Gerardo, Italy. 363Germans Trias i Pujol Research Institute (IGTP), Badalona, Spain. 364Clinic of Medicine and Rehabilitation, Levanger Hospital, Nord-Trondelag Hospital Trust, Levanger, Norway. 365Geminicenter for Sepsis Research, Institute of Circulation and Medical Imaging (ISB), NTNU, Trondheim, Norway. 366HLA Laboratory, E.O. Ospedali Galliera, Genova, Italy. 367ISGlobal, Barcelona, Spain. 368Universitat Pompeu Fabra (UPF), Barcelona, Spain. 369IMIM (Hospital del Mar Medical Research Institute), Barcelona, Spain. 370Stefan-Morsch-Stiftung, Birkenfeld, Germany. 371Osakidetza, OSI Donostialdea, Altza Primary Care, Biodonostia Health Research Institute, San Sebastián, Spain. 372Center of Bioinformatics, Biostatistics and Bioimaging, School of Medicine and Surgery, University of Milano-Bicocca, Milan, Italy. 373Biostatistics and Clinical Epidemiology, Fondazione IRCCS San Gerardo dei Tintori, Monza, Italy. 374Department of Internal Medicine, Infectious Diseases, University Hospital Frankfurt & Goethe University Frankfurt, Frankfurt am Main, Germany. 375Phase 1 Research Centre, ASST Monza, School of Medicine and Surgery, University of Milano-Bicocca, Milan, Italy. 376Division of Intensive Care and Emergency Medicine, Department of Internal Medicine, Medical University Innsbruck, Innsbruck, Austria. 377Department of Anesthesiology, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), Madrid, Spain. 378University of Cologne, Faculty of Medicine and University Hospital Cologne, Clinical Trials Centre Cologne (ZKS Köln), Cologne, Germany. 379Pulmonology Unit, Fondazione IRCCS San Gerardo dei Tintori, Monza, Italy. 380Infectious Diseases Unit, Fondazione IRCCS San Gerardo dei Tintori, Monza, Italy. 381Institute of Human Genetics, University of Bonn School of Medicine & University Hospital Bonn, Bonn, Germany. 382Institute of Immunology, Christian-Albrechts-University of Kiel & UKSH Schleswig-Holstein, Kiel, Germany. 383Intensive Care Department, Vall d’Hebron University Hospital, SODIR-VHIR research group, Barcelona, Spain. 384Department of Medicine, Møre & Romsdal Hospital Trust, Molde, Norway. 385Institute of Medical Virology, University Hospital Frankfurt, Goethe University, Frankfurt am Main, Germany. 386German Centre for Infection Research (DZIF), external partner site Frankfurt, Frankfurt am Main, Germany. 387Department of Neurology, Bezirksklinikum Regensburg, University of Regensburg, Regensburg, Germany. 388Institute of Transfusionsmedicine, University Hospital Schleswig-Holstein (UKSH), Kiel, Germany. 389School of Biological Sciences, Monash University, Clayton, Victoria, Australia. 390Department of Microbiology, University Hospital Vall d’Hebron, Barcelona, Spain. 391Autonoma University of Barcelona, Barcelona, Spain. 392Biochemistry Unit, Hospital Universitario Clinico San Cecilio, Granada, Spain. 393Department of Infectious Diseases, University Hospital of North Norway, Tromsø, Norway. 394Faculty of Health Sciences, UIT The Arctic University of Norway, Tromsø, Norway. 395Catalan Institute of Oncology (ICO), Barcelona, Spain. 396Bellvitge Biomedical Research Institute (IDIBELL), Barcelona, Spain. 397Universitat de Barcelona (UB), Barcelona, Spain. 398Faculty of Medicine and University Nature | Vol 621 | 7 September 2023 | E21 Hospital Cologne, Institute for Transfusion Medicine, University of Cologne, Cologne, Germany. 399Departamento de Anatomía Patológica, Facultad de Medicina, Universidad de Chile, Santiago, Chile. 400Servicio de Anatomía Patológica, Hospital Clínico de la Universidad de Chile, Santiago, Chile. 401Laboratorio Clínico del Área Técnica de Biología Molecular, Hospital del Salvador, Providencia, Chile. 402Departamento de Biotecnología, Universidad de Antofagasta, Antofagasta, Chile. 403AUSTRAL-omics, Vicerrectoría de Investigación, Desarrollo y Creación Artística, Antofagasta, Chile. 404Instituto de Ciencias Ambientales y Evolutivas, Facultad de Ciencias, Universidad Austral de Chile, Valdivia, Chile. 405Escuela de Medicina, Universidad de Magallanes, Punta Arenas, Chile. 406Genómica Evolutiva y Médica de Magallanes (GEMMa), Centro Asistencial, Docente y de Investigación (CADI-UMAG), Punta Arenas, Chile. 407Interuniversity Center on Healthy Aging, Punta Arenas, Chile. 408Instituto de Investigación Interdisciplinaria, Universidad de Talca, Talca, Chile. 409Programa de Genética Humana, ICBM, Universidad de Chile, Santiago, Chile. 410Departamento de Oncología Básico Clínica, Universidad de Chile, Santiago, Chile. 411Departamento de Ciencias de la Computación, Facultadad de Ciencias Físicas y Matemáticas, Universidad de Chile, Santiago, Chile. 412Laboratory of Chemical Carcinogenesis and Pharmacogenetics, Department of Basic and Clinical Oncology (DOBC), Faculty of Medicine, University of Chile, Santiago, Chile. 413Latin American Network for Implementation and Validation of Clinical Pharmacogenomics Guidelines (RELIVAF-CYTED), Santiago, Chile. 414Department of Pharmaceutical Sciences and Technology, Faculty of Chemical and Pharmaceutical Sciences, University of Chile, Santiago, Chile. 415Department of Basic and Clinical Oncology (DOBC), Faculty of Medicine, University of Chile, Santiago, Chile. 416University of Chile & Latin American Network for Implementation and Validation of Clinical Pharmacogenomics Guidelines (RELIVAF-CYTED), Santiago, Chile. 417Departamento de Tecnología Médica, Facultad de Ciencias de la Salud, Universidad de Antofagasta, Antofagasta, Chile. 418AUSTRAL-omics, Vicerrectoría de Investigación, Desarrollo y Creación Artística, Universidad Austral de Chile, Valdivia, Chile. 419Clinical Biochemistry and Immunology Department, Faculty of Pharmacy, Universidad de Concepción, Concepción, Chile. 420Department of Clinical Immunology, Zealand University Hospital, Køge, Denmark. 421Department of Epidemiology Research, Statens Serum Institut, Copenhagen, Denmark. 422Department of Clinical Immunology, Rigshospitalet, University of Copenhagen, Copenhagen, Denmark. 423Department of Infectious Diseases, Copenhagen University Hospital (Rigshospitalet), Copenhagen, Denmark. 424Department of Medical Endocrinology and Metabolism, Copenhagen University Hospital (Rigshospitalet), Copenhagen, Denmark. 425Department of Hematology, Rigshospitalet, Copenhagen, Denmark. 426Statens Serum Institut, Copenhagen, Denmark. 427Department of Pediatrics, Faculty of Medicine, Mansoura University, Mansoura, Egypt. 428Department of Surgery, Faculty of Medicine, Mansoura University, Mansoura, Egypt. 429Stanley Center for Psychiatric Research & Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA. 430Chest Department, Faculty of Medicine, Mansoura University, Mansoura, Egypt. 431Anesthesia, Surgical Intensive Care & Pain Management Department, Faculty of Medicine, Tanta University, Tanta, Egypt. 432Department of Medical Biochemistry, Faculty of Medicine, Mansoura University, Mansoura, Egypt. 433Department of Tropical Medicine, Faculty of Medicine, Mansoura University, Mansoura, Egypt. 434Pediatric and Neonatology, Kafr Elzayat General Hospital, Kafr El-Zayat, Egypt. 435Chest Department, Faculty of Medicine, Tanta University, Tanta, Egypt. 436Pediatrics Department, Faculty of Medicine, Tanta University, Tanta, Egypt. 437Anesthesia, Surgical Intensive Care & Pain Management Departmen, Faculty of Medicine, Mansoura University, Mansoura, Egypt. 438Department of Population Health and Reproduction, School of Veterinary Medicine, University of California, Davis, Davis, CA, USA. 439Department of Anaethesia and Critical Care, Faculty of Medicine, Mansoura University, Mansoura, Egypt. 440Department of Internal Medicine, Faculty of Medicine, Tanta University, Tanta, Egypt. 441Faculty of Science, Tanta University, Tanta, Egypt. 442Department of Internal Medicine, Erasmus MC, University Medical Center, Rotterdam, The Netherlands. 443Department of Epidemiology, Erasmus MC, University Medical Center, Rotterdam, The Netherlands. 444Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands. 445 FinnGen, Helsinki, Finland. 446Institute for Molecular Medicine Finland (FIMM), HiLIFE, University of Helsinki, Helsinki, Finland. 447Public Health, Faculty of Medicine, University of Helsinki, Helsinki, Finland. 448Finnish Institute for Health and Welfare (THL), Helsinki, Finland. 449Clinical and Molecular Metabolism Research Program, Faculty of Medicine, University of Helsinki, Helsinki, Finland. 450Department of Computational Biology, Institut Pasteur, Université de Paris, Paris, France. 451Université Paris-Saclay, CEA, Centre National de Recherche en Génomique Humaine (CNRGH), Evry, France. 452University of Bordeaux, INSERM, Bordeaux Population Health Center, UMR1219, Bordeaux, France. 453Department of Neurology, Institute of Neurodegenerative Diseases, Bordeaux University Hospital, Bordeaux, France. 454Laboratory of Human Genetics of Infectious Diseases, Necker Branch, University of Paris, INSERM U1163, Imagine Institute, Paris, France. 455St Giles Laboratory of Human Genetics of Infectious Diseases, Rockefeller Branch, The Rockefeller University, New York, NY, USA. 456Department of Global Health, Institut Pasteur, Université de Paris, Paris, France. 457Department of Nephrology, Transplantation, Dialysis and Apheresis, Bordeaux University Hospital, Bordeaux, France. 458Immunoconcept CNRS UMR 5164, Bordeaux University, Bordeaux, France. 459St Giles Laboratory of Human Genetics of Infectious Diseases, Rockefeller Branch, The Rockefeller University, New York, NY, USA. 460Université de Paris, INSERM, IAME UMR 1137, Paris, France. 461UMR-CNRS 6249 Chrono-environnement, Université Bourgogne Franche-Comté, Besançon, France. 462Human Evolutionary Genetics, Institut Pasteur, UMR 2000, CNRS, Paris, France. 463Chair of Human Genomics and Evolution, Collège de France, Paris, France. 464Human Evolutionary Genetics Unit, Institut Pasteur, Université Paris Cité, CNRS UMR2000, Paris, France. 465Translational Immunology Laboratory, Institut Pasteur, Paris, France. 466Catalan Institute of Oncology, Bellvitge Biomedical Research Institute, Consortium for Biomedical Research in Epidemiology and Public Health and University of Barcelona, Barcelona, Spain. 467CIBER Epidemiología y Salud Pública (CIBERESP), Madrid, Spain. 468Barcelona Supercomputing Center—Centro Nacional de Supercomputación (BSC-CNS), Life & Medical Sciences, Barcelona, Spain. 469Programs in Metabolism and Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA. 470Department of Medicine, Harvard Medical School, Boston, MA, USA. 471Diabetes Unit and Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA. 472CIBER de Enfermedades Infecciosas (CIBERINFEC), Barcelona, Spain. 473Blizard Institute, Queen Mary University of London, London, UK. 474School of Basic and Medical Biosciences, Faculty of Life Sciences and Medicine, King’s College London, London, UK. 475Wolfson Institute of Population Health, Queen Mary University of London, London, UK. 476Medical and Population Genomics, Wellcome Sanger Institute, Hinxton, UK. 477Bradford Institute for Health Research, Bradford Teaching Hospitals National Health Service (NHS) Foundation Trust, Bradford, UK. 478Genes & Health, Blizard Institute, Queen Mary University of London, London, UK. 479Genentech, San Francisco, CA, USA. 480DNA Link, Seoul, Republic of Korea. 481Seoul National University Hospital Gangnam Center, Seoul, Republic of Korea. 482Division of Infectious Diseases, Department of Internal Medicine, Chungnam National University School of Medicine, Daejeon, Republic of Korea. 483Department of Internal Medicine, School of Medicine, Kyungpook National University, Daegu, Republic of Korea. 484Division of Infectious Diseases, Department of Internal Medicine, Incheon Medical Center, Incheon, Republic of Korea. 485Department of Infectious Diseases, Keimyung University Dongsan Hospital, Keimyung University School of Medicine, Daegu, Republic of Korea. 486Department of Internal Medicine, Pusan National University School of Medicine and Medical Research Institute, Pusan National University Hospital, Busan, Republic of Korea. 487Division of Infectious Diseases, Department of Internal Medicine, Myongji Hospital, Goyang, Republic of Korea. 488Institute for Health Promotion, Graduate School of Public Health, Yonsei University, Seoul, Republic of Korea. 489Department of Precision Medicine, National Institute of Health, Cheongwon-gun, Republic of Korea. 490Division of Genome Science, Department of Precision Medicine, National Institute of Health, Cheongwon-gun, Republic of Korea. 491Baillie Gifford Pandemic Science Hub, Centre for Inflammation Research, The Queen’s Medical Research Institute, University of Edinburgh, Edinburgh, UK. 492MRC Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Western General Hospital, Edinburgh, UK. 493Roslin Institute, University of Edinburgh, Easter Bush, Edinburgh, UK. 494William Harvey Research Institute, Queen Mary University of London, London, UK. 495Intensive Care Unit, Royal Infirmary of Edinburgh, Edinburgh, UK. 496Pain Service, NHS Tayside, Ninewells Hospital and Medical School, Dundee, UK. 497Centre for Inflammation Research, The Queen’s Medical Research Institute, University of Edinburgh, Edinburgh, UK. 498Wellcome Centre for Human Genetics, University of Oxford, Oxford, UK. 499Institute for Molecular Bioscience, The University of Queensland, Brisbane, Queensland, Australia. 500Biostatistics Group, Greater Bay Area Institute of Precision Medicine (Guangzhou), Fudan University, Guangzhou, China. 501Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, Edinburgh, UK. 502Edinburgh Clinical Research Facility, Western General Hospital, University of Edinburgh, Edinburgh, UK. 503School of Life Sciences, Westlake University, Hangzhou, China. 504Westlake Laboratory of Life Sciences and Biomedicine, Hangzhou, China. 505Great Ormond Street Hospital, London, UK. 506Royal Hospital for Children, Glasgow, UK. 507Centre for Tropical Medicine and Global Health, Nuffield Department of Medicine, University of Oxford, Oxford, UK. 508Department of Anaesthesia and Intensive Care, The Chinese University of Hong Kong, Prince of Wales Hospital, Hong Kong, China. 509Department of Critical Care Medicine, Queen’s University and Kingston Health Sciences Centre, Kingston, Ontario, Canada. 510Wellcome-Wolfson Institute for Experimental Medicine, Queen’s University Belfast, Belfast, UK. 511Department of Intensive Care Medicine, Royal Victoria Hospital, Belfast, UK. 512UCL Centre for Human Health and Performance, London, UK. 513Clinical Research Centre at St Vincent’s University Hospital, University College Dublin, Dublin, Ireland. 514National Heart and Lung Institute, Imperial College London, London, UK. 515Imperial College Healthcare NHS Trust: London, London, UK. 516Heart Institute, University of Sao Paulo, Sao Paulo, Brazil. 517Intensive Care National Audit & Research Centre, London, UK. 518NIHR Health Protection Research Unit for Emerging and Zoonotic Infections, Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool, UK. 519Respiratory Medicine, Alder Hey Children’s Hospital, Institute in The Park, University of Liverpool, Liverpool, UK. 520Department of Medicine, University of Cambridge, Cambridge, UK. 521Guys and St Thomas’ Hospital, London, UK. 522James Cook University Hospital, Middlesbrough, UK. 523Barts Health NHS Trust, London, UK. 524Royal Stoke University Hospital, Stoke-on-Trent, UK. 525North Middlesex University Hospital NHS trust, London, UK. 526King’s College Hospital, London, UK. 527Charing Cross Hospital, St Mary’s Hospital and Hammersmith Hospital, London, UK. 528The Royal Liverpool University Hospital, Liverpool, UK. 529John Radcliffe Hospital, Oxford, UK. 530Addenbrooke’s Hospital, Cambridge, UK. 531Nottingham University Hospital, Nottingham, UK. 532St George’s Hospital, London, UK. 533BHRUT (Barking Havering)—Queens Hospital and King George Hospital, London, UK. 534Royal Infirmary of Edinburgh, Edinburgh, UK. 535Kingston Hospital, London, UK. 536Queen Alexandra Hospital, Portsmouth, UK. 537Royal Gwent Hospital, Newport, UK. 538Royal Blackburn Teaching Hospital, Blackburn, UK. 539Stepping Hill Hospital, Stockport, UK. 540Northumbria Healthcare NHS Foundation Trust, North Shields, UK. 541Countess of Chester Hospital, Chester, UK. 542Pinderfields General Hospital, Wakefield, UK. 543Ninewells Hospital, Dundee, UK. 544Croydon University Hospital, Croydon, UK. 545Morriston Hospital, Swansea, UK. 546Queen Elizabeth University Hospital, Glasgow, UK. 547Broomfield Hospital, Chelmsford, UK. 548Heartlands Hospital, Birmingham, UK. 549Royal Sussex County Hospital, Brighton, UK. 550York Hospital, York, UK. 551Queen Elizabeth Hospital, Birmingham, UK. 552Royal Glamorgan Hospital, Pontyclun, UK. 553Barnet Hospital, London, UK. 554Wythenshawe Hospital, Manchester, UK. 555Medway Maritime Hospital, Gillingham, UK. 556Royal Berkshire NHS Foundation Trust, Reading, UK. 557Whiston Hospital, Prescot, UK. 558The Royal Oldham Hospital, Manchester, UK. 559Chesterfield Royal Hospital Foundation Trust, Chesterfield, UK. 560Aberdeen Royal Infirmary, Aberdeen, UK. 561Royal Devon and Exeter Hospital, Exeter, UK. 562Glasgow Royal Infirmary, Glasgow, UK. 563Blackpool Victoria Hospital, Blackpool, UK. 564Southampton General Hospital, Southampton, UK. 565Ashford and St Peter’s Hospital, Chertsey, UK. 566Derriford Hospital, Plymouth, UK. 567East Surrey Hospital, Redhill, UK. 568Poole Hospital, Poole, UK. 569Royal Alexandra Hospital, Paisley, UK. 570St James’s University Hospital and Leeds General Infirmary, Leeds, UK. 571Bedford Hospital, Bedford, UK. 572Southport and Formby District General Hospital, Ormskirk, UK. 573The Tunbridge Wells Hospital and Maidstone Hospital, Kent, UK. 574Queen Elizabeth Hospital, Woolwich, London, UK. 575North Manchester General Hospital, Manchester, UK. 576Royal Victoria Infirmary, Newcastle Upon Tyne, UK. 577Hull Royal Infirmary, Hull, UK. 578Manchester Royal Infirmary, Manchester, UK. 579Royal Derby Hospital, Derby, UK. 580Aintree University Hospital, Liverpool, UK. 581Fairfield General Hospital, Bury, UK. 582Norfolk and Norwich University hospital (NNUH), Norwich, UK. 583Milton Keynes University Hospital, Milton Keynes, UK. 584Good Hope Hospital, Birmingham, UK. 585Queen Elizabeth Hospital Gateshead, Gateshead, UK. 586Royal Bolton Hospital, Bolton, UK. 587Tameside General Hospital, Ashton Under Lyne, UK. 588Salford Royal Hospital, Manchester, UK. 589Great Ormond St Hospital and UCL Great Ormond St Institute of Child E22 | Nature | Vol 621 | 7 September 2023 Matters arising Health NIHR Biomedical Research Centre, London, UK. 590Southmead Hospital, Bristol, UK. 591William Harvey Hospital, Ashford, UK. 592Arrowe Park Hospital, Wirral, UK. 593Royal Hampshire County Hospital, Winchester, UK. 594Bradford Royal Infirmary, Bradford, UK. 595Glan Clwyd Hospital, Bodelwyddan, UK. 596Royal Bournemouth Hospital, Bournemouth, UK. 597Bristol Royal Infirmary, Bristol, UK. 598University Hospital North Durham, Darlington, UK. 599Darlington Memorial Hospital, Darlington, UK. 600Basildon Hospital, Basildon, UK. 601University College Hospital, London, UK. 602Whittington Hospital, London, UK. 603Western General Hospital, Edinburgh, UK. 604Ipswich Hospital, Ipswich, UK. 605Hereford County Hospital, Hereford, UK. 606Sunderland Royal Hospital, Sunderland, UK. 607Queens Hospital Burton, Burton-On-Trent, UK. 608Musgrove Park Hospital, Taunton, UK. 609The Royal Papworth Hospital, Cambridge, UK. 610University Hospital Lewisham, London, UK. 611The Princess Alexandra Hospital, Harlow, UK. 612University Hospital of Wales, Cardiff, UK. 613West Middlesex Hospital, Isleworth, UK. 614Royal Albert Edward Infirmary, Wigan, UK. 615Stoke Mandeville Hospital, Aylesbury, UK. 616Royal Lancaster Infirmary, Lancaster, UK. 617Basingstoke and North Hampshire Hospital, Basingstoke, UK. 618Worthing Hospital, Worthing, UK. 619St Richard’s Hospital, Chichester, UK. 620The Alexandra Hospital, Redditch and Worcester Royal Hospital, Worcester, UK. 621Royal Cornwall Hospital, Truro, UK. 622Watford General Hospital, Watford, UK. 623Macclesfield District General Hospital, Macclesfield, UK. 624Royal Surrey County Hospital, Guildford, UK. 625Rotherham General Hospital, Rotherham, UK. 626Craigavon Area Hospital, Craigavon, UK. 627King’s Mill Hospital, Nottingham, UK. 628Dumfries and Galloway Royal Infirmary, Dumfries, UK. 629Prince Charles Hospital, Merthyr Tydfil, UK. 630Ysbyty Gwynedd, Bangor, UK. 631Royal Preston Hospital, Preston, UK. 632The Great Western Hospital, Swindon, UK. 633Lincoln County Hospital, Lincoln, UK. 634University Hospital of North Tees, Stockton on Tees, UK. 635Glangwili General Hospital, Camarthen, UK. 636Southend University Hospital, Westcliff-on-Sea, UK. 637Lister Hospital, Stevenage, UK. 638Diana Princess of Wales Hospital, Grimsby, UK. 639West Suffolk Hospital, Bury St Edmunds, UK. 640Victoria Hospital, Kirkcaldy, UK. 641Calderdale Royal Hospital, Halifax, UK. 642Huddersfield Royal Infirmary, Huddersfield, UK. 643Dorset County Hospital, Dorchester, UK. 644Russell’s Hall Hospital, Dudley, UK. 645Royal United Hospital, Bath, UK. 646St Mary’s Hospital, Newport, UK. 647George Eliot Hospital NHS Trust, Nuneaton, UK. 648Yeovil Hospital, Yeovil, UK. 649Forth Valley Royal Hospital, Falkirk, UK. 650Frimley Park Hospital, Camberley, UK. 651Chelsea & Westminster NHS Foundation Trust, London, UK. 652Queen Elizabeth the Queen Mother Hospital, Margate, UK. 653Royal Brompton Hospital, London, UK. 654Darent Valley Hospital, Dartford, UK. 655University Hospital Crosshouse, Kilmarnock, UK. 656University Hospital Wishaw, Wishaw, UK. 657University College Dublin, St Vincent’s University Hospital, Dublin, Ireland. 658The Queen Elizabeth Hospital, King’s Lynn, UK. 659Walsall Manor Hospital, Walsall, UK. 660Princess Royal Hospital, Brighton, UK. 661Barnsley Hospital, Barnsley, UK. 662Warrington General Hospital, Warrington, UK. 663Royal Victoria Hospital, Belfast, UK. 664Royal Hallamshire Hospital and Northern General Hospital, Sheffield, UK. 665Harefield Hospital, London, UK. 666Cumberland Infirmary, Carlisle, UK. 667Eastbourne District General Hospital, Eastbourne, UK. 668Conquest Hospital, Saint Leonards-on-sea, UK. 669Salisbury District Hospital, Salisbury, UK. 670Airedale General Hospital, Keighley, UK. 671Leicester Royal Infirmary, Leicester, UK. 672Peterborough City Hospital, Peterborough, UK. 673Hinchingbrooke Hospital, Huntingdon, UK. 674Colchester General Hospital, Colchester, UK. 675Princess Royal Hospital, Telford and Royal Shrewsbury Hospital, Shrewsbury, UK. 676University Hospital Monklands, Airdrie, UK. 677Wrexham Maelor Hospital, Wrexham, UK. 678New Cross Hospital, Wolverhampton, UK. 679University Hospital Hairmyres, East Kilbride, UK. 680Warwick Hospital, Warwick, UK. 681Sandwell General Hospital and City Hospital, Birmingham, UK. 682Royal Manchester Children’s Hospital, Manchester, UK. 683Gloucestershire Royal Hospital, Gloucester, UK. 684University Hospitals Coventry & Warwickshire NHS Trust, Coventry, UK. 685Torbay Hospital, Torquay, UK. 686Pilgrim Hospital, Lincoln, UK. 687Prince Philip Hospital, Lianelli, UK. 688Princess of Wales Hospital, Llantrisant, UK. 689Northampton General Hospital NHS Trust, Northampton, UK. 690The Christie NHS Foundation Trust, Manchester, UK. 691James Paget University Hospital NHS Trust, Great Yarmouth, UK. 692Birmingham Children’s Hospital, Birmingham, UK. 693Withybush General Hospital, Haverfordwest, UK. 694Northwick Park Hospital, London, UK. 695North Devon District Hospital, Barnstaple, UK. 696Scunthorpe General Hospital, Scunthorpe, UK. 697Royal Free Hospital, London, UK. 698Raigmore Hospital, Inverness, UK. 699West Cumberland Hospital, Whitehaven, UK. 700Furness General Hospital, Barrow-in-Furness, UK. 701Liverpool Heart and Chest Hospital, Liverpool, UK. 702Scarborough General Hospital, Scarborough, UK. 703Bronglais General Hospital, Aberystwyth, UK. 704Alder Hey Children’s Hospital, Liverpool, UK. 705Borders General Hospital, Melrose, UK. 706Leighton Hospital, Cheshire, UK. 707Kent & Canterbury Hospital, Canterbury, UK. 708Harrogate and District NHS Foundation Trust, Harrogate, UK. 709The Royal Marsden Hospital, London, UK. 710Ealing Hospital, Southall, UK. 711St John’s Hospital Livingston, Livingston, UK. 712Wexham Park Hospital, Slough, UK. 713Sheffield Children’s Hospital, Sheffield, UK. 714Homerton University Hospital Foundation NHS Trust, London, UK. 715National Hospital for Neurology and Neurosurgery, London, UK. 716The Royal Alexandra Children’s Hospital, Brighton, UK. 717Golden Jubilee National Hospital, Clydebank, UK. 718Illumina Cambridge, Cambridge, UK. 719Test and Trace, The Health Security Agency, Department of Health and Social Care, London, UK. 720Imperial College London, London, UK. 721NIHR Clinical Research Network (CRN), North West London Core Team, London, UK. 722Cambridge University Hospitals NHS Foundation Trust, Cambridge, UK. 723Biostatistics Group, State Key Laboratory of Biocontrol, School of Life Sciences, Sun Yat-sen University, Guangzhou, China. 724Department of Infectious Diseases, Leiden University Medical Center, Leiden, The Netherlands. 725Genotek, Moscow, Russia. 726Helix, San Mateo, CA, USA. 727Center for Genomic Medicine, Desert Research Institute, Reno, NV, USA. 728Renown Health, Reno, NV, USA. 729Gujarat Biotechnology Research Centre (GBRC), Department of Science and Technology (Government of Gujarat), Gandhinagar, India. 730Commissionerate of Health Medical Services and Medical Education Gandhinagar, Gandhinagar, India. 731Department of Microbiology, B.J. Medical College and Civil hospital, Institute of Medical Post-Graduate Studies and Research, Ahmedabad, India. 732Department of Medicine, B.J. Medical College and Civil hospital, Institute of Medical Post-Graduate Studies and Research, Ahmedabad, India. 733Department of Community Medicine, Government Medical College, Surat, India. 734Department of Microbiology, Government Medical College, Bhavnagar, India. 735Department of General Medicine, GMERS Medical College & Hospital, Gotri, Vadodara, India. 736Department of Community & Family Medicine, All India Institute of Medical Sciences, Rajkot, India. 737National Laboratory of Genomics for Biodiversity (LANGEBIO), Advanced Genomics Unit (UGA), CINVESTAV, Irapuato, Guanajuato, Mexico. 738Instituto Nacional de Enfermedades Respiratorias Ismael Cosío Villegas, Tlalpan, Mexico City, Mexico. 739Departamento de Bioquímica, Facultad de Medicina, Universidad Nacional Autónoma de México, Coyoacán, Mexico City, Mexico. 740Instituto Nacional de Salud Pública, Cuernavaca, Mexico. 741International Laboratory for Human Genome Research (LIIGH), UNAM Juriquilla, Queretaro, Mexico. 742Unidad de Biología Molecular y Medicina Genómica, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán/ Instituto de Investigaciones Biomédicas, UNAM, Mexico City, Mexico. 743Instituto Nacional de Medicina Genómica, Mexico City, Mexico. 744Instituto Nacional de Pediatría, Mexico City, Mexico. 745Escuela Superior de Medicina, Instituto Politécnico Nacional, Mexico City, Mexico. 746Tecnologico de Monterrey, Escuela de Medicina y Ciencias de la Salud, Monterrey, Mexico. 747Division of Pulmonary Medicine, Department of Medicine, Keio University School of Medicine, Tokyo, Japan. 748Department of Statiatical Genetics, Osaka University Graduate School of Medicine, Suita, Japan. 749Laboratory of Statistical Immunology, Immunology Frontier Research Center (WPI-IFReC), Osaka University, Suita, Japan. 750Integrated Frontier Research for Medical Science Division, Institute for Open and Transdisciplinary Research Initiatives, Osaka University, Suita, Japan. 751Division of Health Medical Intelligence, Human Genome Center, the Institute of Medical Science, The University of Tokyo, Tokyo, Japan. 752Laboratory of Viral Infection I, Department of Infection Control and Immunology, Ōmura Satoshi Memorial Institute & Graduate School of Infection Control Sciences, Kitasato University, Tokyo, Japan. 753Department of Surgery, Keio University School of Medicine, Tokyo, Japan. 754Department of Organoid Medicine, Keio University School of Medicine, Tokyo, Japan. 755Department of Infectious Diseases, Keio University School of Medicine, Tokyo, Japan. 756Department of Respiratory Medicine and Clinical Immunology, Osaka University Graduate School of Medicine, Suita, Japan. 757Department of Immunopathology, Immunology Frontier Research Center (WPI-IFReC), Osaka University, Suita, Japan. 758Institute of Research, Tokyo Medical and Dental University, Tokyo, Japan. 759Department of Insured Medical Care Management, Tokyo Medical and Dental University Hospital of Medicine, Tokyo, Japan. 760Genome Medical Science Project (Toyama), National Center for Global Health and Medicine, Tokyo, Japan. 761Division of Gastroenterology and Hepatology, Department of Medicine, Keio University School of Medicine, Tokyo, Japan. 762M&D Data Science Center, Tokyo Medical and Dental University, Tokyo, Japan. 763Kyoto University, Department of Pathology and Tumor Biology Institute for the Advanced Study of Human Biology (WPI-ASHBi), Kyoto University, Kyoto, Japan. 764Department of Medicine, Center for Hematology and Regenerative Medicine, Karolinska Institute, Stockholm, Sweden. 765Department of Emergency and Critical Care Medicine, Keio University School of Medicine, Tokyo, Japan. 766Department of Anesthesiology, Keio University School of Medicine, Tokyo, Japan. 767Department of Laboratory Medicine, Keio University School of Medicine, Tokyo, Japan. 768Division of Infection Control and Prevention, Osaka University Hospital, Suita, Japan. 769Department of Biomedical Ethics and Public Policy, Osaka University Graduate School of Medicine, Suita, Japan. 770Center for Genomic Medicine, Kyoto University Graduate School of Medicine, Kyoto, Japan. 771Department of Pulmonary Medicine, Faculty of Medicine, University of Tsukuba, Tsukuba, Japan. 772Department of Neurosurgery, Faculty of Medicine, The University of Tokyo, Tokyo, Japan. 773Laboratory of Immune Regulation, Department of Microbiology and Immunology, Osaka University Graduate School of Medicine, Suita, Japan. 774Medical Innovation Promotion Center, Tokyo Medical and Dental University, Tokyo, Japan. 775Clinical Research Center, Tokyo Medical and Dental University Hospital of Medicine, Tokyo, Japan. 776Department of Medical Informatics, Tokyo Medical and Dental University Hospital of Medicine, Tokyo, Japan. 777Respiratory Medicine, Tokyo Medical and Dental University, Tokyo, Japan. 778Clinical Laboratory, Tokyo Medical and Dental University Hospital of Medicine, Tokyo, Japan. 779Department of Pathology and Tumor Biology, Kyoto University, Kyoto, Japan. 780Department of Respiratory Medicine, Juntendo University Faculty of Medicine and Graduate School of Medicine, Tokyo, Japan. 781Department of Emergency and Disaster Medicine, Juntendo University Faculty of Medicine and Graduate School of Medicine, Tokyo, Japan. 782Department of Cardiovascular Biology and Medicine, Juntendo University Faculty of Medicine and Graduate School of Medicine, Tokyo, Japan. 783Department of Respiratory Medicine, Tokyo Women’s Medical University, Tokyo, Japan. 784Department of General Medicine, Tokyo Women’s Medical University, Tokyo, Japan. 785Department of Respiratory Medicine, Saitama Cardiovascular and Respiratory Center, Saitama, Japan. 786Kawasaki Municipal Ida Hospital, Kawasaki, Japan. 787Internal Medicine, Saitama Medical Center, JCHO (Japan Community Health Care Organization), Saitama, Japan. 788Saitama City Hospital, Saitama, Japan. 789Division of Infection Control, Eiju General Hospital, Tokyo, Japan. 790Department of Pulmonary Medicine, Eiju General Hospital, Tokyo, Japan. 791Department of Respiratory Medicine, Osaka Saiseikai Nakatsu Hospital, Osaka, Japan. 792Department of Infection Control, Osaka Saiseikai Nakatsu Hospital, Osaka, Japan. 793Department of Infectious Diseases, Tosei General Hospital, Seto, Japan. 794Fukujuji Hospital, Kiyose, Japan. 795Department of Emergency and Critical Care Medicine, Tokyo Women’s Medical University Medical Center East, Tokyo, Japan. 796Department of Medicine, Tokyo Women’s Medical University Medical Center East, Tokyo, Japan. 797Department of Pediatrics, Tokyo Women’s Medical University Medical Center East, Tokyo, Japan. 798Japan Community Health Care Organization Kanazawa Hospital, Kanazawa, Japan. 799Division of Pulmonary Medicine, Department of Internal Medicine, Federation of National Public Service Personnel Mutual Aid Associations, Tachikawa Hospital, Tachikawa, Japan. 800Department of Respiratory Medicine, Japan Organization of Occupational Health and Safety, Kanto Rosai Hospital, Kawasaki, Japan. 801Department of General Internal Medicine, Japan Organization of Occupational Health and Safety, Kanto Rosai Hospital, Kawasaki, Japan. 802Department of Emergency and Critical Care Medicine, Kansai Medical Univrsity General Medical Center, Kirakata, Japan. 803Department of Respiratory Medicine, Kitasato University Kitasato Institute Hospital, Tokyo, Japan. 804Ishikawa Prefectural Central Hospital, Kanazawa, Japan. 805Internal Medicine, Sano Kosei General Hospital, Sano, Japan. 806Saiseikai Yokohamashi Nanbu Hospital, Yokohama, Japan. 807Kanagawa Cardiovascular and Respiratory Center, Yokohama, Japan. 808Saiseikai Utsunomiya Hospital, Utsunomiya, Japan. 809Department of Respiratory Medicine, KKR Sapporo Medical Center, Sapporo, Japan. 810Internal Medicine, Internal Medicine Center, Showa University Koto Toyosu Hospital, Tokyo, Japan. 811Department of Respiratory Medicine, Toyohashi Municipal Hospital, Toyohashi, Japan. 812Keiyu Hospital, Yokohama, Japan. 813Department of Rheumatology, National Hospital Organization Hokkaido Medical Center, Sapporo, Japan. 814Department of Respiratory Medicine, National Hospital Organization Tokyo Medical Center, Tokyo, Japan. 815Department of Allergy, National Hospital Organization Tokyo Nature | Vol 621 | 7 September 2023 | E23 Medical Center, Tokyo, Japan. 816Department of General Internal Medicine and Infectious Diseases, National Hospital Organization Tokyo Medical Center, Tokyo, Japan. 817Japanese Red Cross Musashino Hospital, Musashino, Japan. 818Department of Respiratory Medicine, Tohoku University Graduate School of Medicine, Sendai, Japan. 819Department of Internal Medicine, Division of Respiratory Medicine, Nihon University School of Medicine, Tokyo, Japan. 820Department of Emergency and Critical Care Medicine, St Marianna University School of Medicine, Kawasaki, Japan. 821Division of General Internal Medicine, Department of Internal Medicine, St Marianna University School of Medicine, Kawasaki, Japan. 822National Hospital Organization Kanazawa Medical Center, Kanazawa, Japan. 823Division of Infectious Diseases and Respiratory Medicine, Department of Internal Medicine, National Defense Medical College, Saitama, Japan. 824Department of Emergency and Critical Care Medicine, Faculty of Medicine, Fukuoka University, Fukuoka, Japan. 825Department of Infection Control, Fukuoka University Hospital, Fukuoka, Japan. 826Tokyo Saiseikai Central Hospital, Tokyo, Japan. 827Department of Internal Medicine, Fukuoka Tokushukai Hospital, Kasuga, Japan. 828Department of Infectious Disease and Clinical Research Institute, National Hospital Organization Kyushu Medical Center, Fukuoka, Japan. 829Department of Respirology, National Hospital Organization Kyushu Medical Center, Division of Respirology, Rheumatology, and Neurology, Department of Internal Medicine, Kurume University School of Medicine, Fukuoka, Japan. 830Department of Infectious Disease, National Hospital Organization Kyushu Medical Center, Fukuoka, Japan. 831Matsumoto City Hospital, Matsumoto, Japan. 832Uji-Tokushukai Medical Center, Uji, Japan. 833Department of Respiratory Medicine, Nagoya University Graduate School of Medicine, Nagoya, Japan. 834Department of Respiratory Medicine, Fujisawa City Hospital, Fujisawa, Japan. 835Sapporo City General Hospital, Sapporo, Japan. 836Department of Emergency and Critical Care Medicine, Chiba University Graduate School of Medicine, Chiba, Japan. 837Division of Respiratory Medicine, Social Welfare Organization Saiseikai Imperial Gift Foundation, Saiseikai Kumamoto Hospital, Kumamoto, Japan. 838Department of Anesthesiology and Intensive Care Medicine, Kyoto Prefectural University of Medicine, Kyoto, Japan. 839Ome Municipal General Hospital, Ome, Japan. 840Hanwa Daini Hospital, Osaka, Japan. 841Department of Respiratory Internal Medicine, St Marianna University School of Medicine, Yokohama-City Seibu Hospital, Yokohama, Japan. 842Division of Hematology, Department of Internal Medicine, St Marianna University Yokohama-City Seibu Hospital, Yokohama, Japan. 843Division of Pulmonary Medicine, Department of Medicine, Tokai University School of Medicine, Tokai University School of Medicine, Tokyo, Japan. 844Division of Pulmonary Medicine, Department of Medicine, Tokai University School of Medicine, Tokyo, Japan. 845National Hospital Organization Kumamoto Medical Center, Kumamoto, Japan. 846Department of Respiratory Medicine, Tokyo Medical University Hospital, Tokyo, Japan. 847Department of Respiratory Medicine, Japanese Red Cross Medical Center, Tokyo, Japan. 848JA Toride Medical Hospital, Toride, Japan. 849Japan Organization of Occupational Health and Safety Okayama Rosai Hospital, Okayama, Japan. 850Emergency and Disaster Medicine, Graduate School of Medicine, Gifu University School of Medicine, Gifu, Japan. 851Niigata University, Niigata, Japan. 852National Hospital Organization Kyoto Medical Center, Kyoto, Japan. 853Research Institute for Diseases of the Chest, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan. 854Department of Medicine and Biosystemic Science, Kyushu University Graduate School of Medical Sciences, Fukuoka, Japan. 855Department of Emergency and Critical Care Medicine, Tsukuba University, Tsukuba, Japan. 856Department of Nephrology, Faculty of Medicine, University of Tsukuba, Tsukuba, Japan. 857Department of Hematology, Faculty of Medicine, University of Tsukuba, Tsukuba, Japan. 858National Hospital Organization Tokyo Hospital, Tokyo, Japan. 859Fujioka General Hospital, Fujioka, Japan. 860Division of Respiratory Medicine and Allergology, Department of Medicine, School of Medicine, Showa University, Tokyo, Japan. 861Department of Pulmonary Medicine, Fukushima Medical University, Fukushima, Japan. 862Kansai Electric Power Hospital, Osaka, Japan. 863Kumamoto City Hospital, Kumamoto, Japan. 864Department of Emergency and Critical Care Medicine, Tokyo Metropolitan Police Hospital, Tokyo, Japan. 865Department of Respiratory Medicine, International University of Health and Welfare Shioya Hospital, Narita, Japan. 866Department of Clinical Laboratory, International University of Health and Welfare Shioya Hospital, Narita, Japan. 867National Hospital Organization Saitama Hospital, Saitama, Japan. 868Department of Respiratory Medicine, Gunma University Graduate School of Medicine, Maebashi, Japan. 869Department of Orthopedic Surgery, Tokyo Medical University, Ibaraki Medical Center, Tokyo, Japan. 870Department of Internal Medicine, Kiryu Kosei General Hospital, Kiryu, Japan. 871Daini Osaka Police Hospital, Osaka, Japan. 872Genome Medical Science Project (Konodai), National Center for Global Health and Medicine, Chiba, Japan. 873Ministry Of Health, Amman, Jordan. 874School of Dentistry, The University of Jordan, Amman, Jordan. 875School of Medicine, The University of Jordan, Amman, Jordan. 876Cell Therapy Center, The University of Jordan, Amman, Jordan. 877King Hussein Cancer Center, Amman, Jordan. 878King Abdullah University Hospital, Amman, Jordan. 879Gardens Hospital, Amman, Jordan. 880Centre for Heart Lung Innovation, University of British Columbia, Vancouver, British Columbia, Canada. 881Division of Respiratory Medicine, Faculty of Medicine, University of British Columbia, Vancouver, British Columbia, Canada. 882Institut Universitaire de Cardiologie et de Pneumologie de Québec—Université Laval, Quebec City, Quebec, Canada. 883Department of Genetics and Genomic Sciences, Icahn School of Medicine at Mount Sinai, New York, NY, USA. 884Global Health, University of Washington, Seattle, WA, USA. 885Monoceros Bio, San Diego, CA, USA. 886Department of Pathology and Medical Biology Groningen, University Medical Centre Groningen, University of Groningen, Groningen, The Netherlands. 887GRIAC Research Institute, University Medical Centre Groningen, University of Groningen, Groningen, The Netherlands. 888Department of Pulmonary Diseases, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands. 889Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA. 890Institute of Epidemiology and Preventive Medicine, National Taiwan University, Taipei, Taiwan. 891Master of Public Health Program, National Taiwan University, Taipei, Taiwan. 892Channing Division of Network Medicine, Department of Medicine, Brigham and Women’s Hospital, Boston, MA, USA. 893Brigham and Women’s Hospital, Boston, MA, USA. 894Psychiatric and Neurodevelopmental Genetics Unit, Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, USA. 895Department of Neurology, Massachusetts General Hospital, Boston, MA, USA. 896Division of General Internal Medicine, Massachusetts General Hospital and Department of Medicine, Harvard Medical School and Program in Medical and Population GeneticsBroad Institute, Boston, MA, USA. 897Division of Genetics, Department of Medicine, Brigham and Women’s Hospital, Broad Institute of MIT and Harvard, Harvard Medical School, Boston, MA, USA. 898Department of Human Genetics, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA. 899Departamento de Neurología y Psiquiatría, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 900Dirección de Medicina, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 901Infectología, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 902Clínica de Obesidad, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 903Unidad de Investigación en Enfermedades Metabólicas (UIEM), Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 904Tecnologico de Monterrey, The Institute for Obesity Research, Nuevo León, Monterrey, Mexico. 905Tecnologico de Monterrey, Escuela de Medicina y Ciencias de la Salud, Monterrey, Mexico. 906Departamento de Endocrinología y Metabolismo de Lípidos, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 907Departamento de Psiquiatría y Salud Mental, Facultad de Medicina, Universidad Nacional Autónoma de México, Mexico City, Mexico. 908Unidad de Biología Molecular y Medicina Genómica, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán/Instituto de Investigaciones Biomédicas UNAM, Tlalpan, México. 909Universidad Nacional Autónoma de México, Mexico City, Mexico. 910Institute for Precision Health, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA. 911Unidad de Biología Molecular y Medicina Genómica, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, CDMX, Tlalpan, México. 912Universidad Autónoma Metropolitana, Mexico City, Mexico. 913Departamento de Medicina Genómica y Toxicología Ambiental, Instituto de Investigaciones Biomédicas, UNAM, Mexico City, Mexico. 914Department of Computer Science, School of Engineering, UCLA, Los Angeles, CA, USA. 915Unidad de Biología Molecular y Medicina Genómica, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 916Laboratorio de Microbiología, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 917Laboratorio de Neuropsicología y Cognición, Facultad de Psicología, Universidad Nacional Autónoma de México, Mexico City, Mexico. 918Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 919Facultad de Psicología, Universidad Nacional Autónoma de México, Mexico City, Mexico. 920Facultad de Medicina, Universidad Nacional Autónoma de México, Mexico City, Mexico. 921Laboratorio de Neuropsicología, Facultad de Medicina, Universidad Nacional Autónoma de México, UNAM, Coyoacán, Mexico City, Mexico. 922Research Division, School of Medicine, Universidad Nacional Autónoma de México, Mexico City, Mexico. 923Department of Computational Medicine, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA. 924Department of Pathology and Laboratory Medicine, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA. 925Division of Immunology, Allergy and Rheumatology, Department of Pediatrics, UCLA, Los Angeles, CA, USA. 926Department of Microbiology, Immunology, and Molecular Genetics, UCLA, Los Angeles, CA, USA. 927Department of Neurology, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA. 928Department of Psychiatry and Biobehavioral Sciences, Semel Institute, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA. 929Center for Autism Research and Treatment, Semel Institute, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA. 930Institute for Precision Health, Los Angeles, CA, USA. 931Dirección de Nutrición, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Tlalpan, Mexico City, Mexico. 932Deutsches Herzzentrum der Charité, Universitätsmedizin, Berlin, Germany. 933Charite Universitätsmedizin Berlin, Berlin, Germany. 934Department of Tropical Medicine, Bernhard Nocht Institute for Tropical Medicine, Hamburg, Germany. 935Department of Medicine I, University Medical Centre Hamburg-Eppendorf, Hamburg, Germany. 936Qatar Genome Program, Qatar Foundation Research, Development and Innovation, Qatar Foundation, Doha, Qatar. 937Directorate of Research, Shifa Tameer-e-Millat University, Islamabad, Pakistan. 938Shifa Tameer-e-Millat University, Islamabad, Pakistan. 939Department of Vascular Surgery, Shifa International Hospital, Islamabad, Pakistan. 940Department of Infectious Diseases, Shifa International Hospital, Islamabad, Pakistan. 941Shifa International Hospital, Islamabad, Pakistan. 942Department of Pathology, Shifa International Hospital, Islamabad, Pakistan. 943MNM Bioscience, Cambridge, MA, USA. 944Faculty of Physics, Adam Mickiewicz University, Poznań, Poland. 945Department of Medical Chemistry and Laboratory Medicine, Poznan University of Medical Sciences, Poznan, Poland. 946Central Clinical Hospital of Ministry of the Interior and Administration in Warsaw, Warsaw, Poland. 947Institute of Genetics and Biotechnology, Faculty of Biology, University of Warsaw, Warsaw, Poland. 948Genomics Division, Instituto Tecnológico y de Energías Renovables, Santa Cruz de Tenerife, Spain. 949Research Unit, Hospital Universitario N.S. de Candelaria, Santa Cruz de Tenerife, Spain. 950Centre for Biomedical Network Research on Respiratory Diseases (CIBERES), Instituto de Salud Carlos III, Madrid, Spain. 951Facultad de Ciencias de la Salud, Universidad Fernando Pessoa Canarias, Las Palmas de Gran Canaria, Spain. 952Centro Nacional de Genotipado (CEGEN), Universidade de Santiago de Compostela, Santiago de Compostela, Spain. 953Centre for Biomedical Network Research on Rare Diseases (CIBERER), Instituto de Salud Carlos III, Madrid, Spain. 954Fundación Pública Galega de Medicina Xenómica, Sistema Galego de Saúde (SERGAS) Santiago de Compostela, Santiago de Compostela, Spain. 955Instituto de Investigación Sanitaria de Santiago (IDIS), Santiago de Compostela, Spain. 956Centro Singular de Investigación en Medicina Molecular y Enfermedades Crónicas (CIMUS), Universidade de Santiago de Compostela, Santiago de Compostela, Spain. 957Hospital General Santa Bárbara de Soria, Soria, Spain. 958Biocruces Bizkai HRI, Bizkaia, Spain. 959Cruces University Hospital, Osakidetza, Bizkaia, Spain. 960Fundació Docència I Recerca Mutua Terrassa, Barcelona, Spain. 961Hospital General de Occidente, Zapopan, Mexico. 962Centro Universitario de Tonalá, Universidad de Guadalajara, Tonalá, Mexico. 963Centro de Investigación Multidisciplinario en Salud, Universidad de Guadalajara, Tonalá, Mexico. 964Laboratorio de Vigilancia Molecular Aplicada, Escola Tecnica de Saúde, Pará, Brazil. 965Genetics Postgraduate Program, Federal University of Pernambuco, Recife, Brazil. 966Cardiovascular Genetics Center, Institut d’Investigació Biomèdica Girona (IDIBGI), Girona, Spain. 967Medical Science Department, School of Medicine, University of Girona, Girona, Spain. 968Centre for Biomedical Network Research on Cardiovascular Diseases (CIBERCV), Instituto de Salud Carlos III, Madrid, Spain. 969Cardiology Service, Hospital Josep Trueta, Girona, Spain. 970Institute of Biomedicine of Seville (IBiS), Consejo Superior de Investigaciones Científicas (CSIC), University of Seville, Virgen del Rocio University Hospital, Seville, Spain. 971Departamento de Medicina, Hospital Universitario Virgen del Rocío, Universidad de Sevilla, Seville, Spain. 972Centre for Biomedical Network Research on E24 | Nature | Vol 621 | 7 September 2023 Matters arising Epidemiology and Public Health (CIBERESP), Instituto de Salud Carlos III, Madrid, Spain. 973Instituto de Biomedicina de Sevilla, Seville, Spain. 974Osakidetza, Cruces University Hospital, Bizkaia, Spain. 975Centre for Biomedical Network Research on Diabetes and Metabolic Associated Diseases (CIBERDEM), Instituto de Salud Carlos III, Madrid, Spain. 976University of Pais Vasco, UPV/EHU, Bizkaia, Spain. 977Hospital Universitario Río Hortega, Valladolid, Spain. 978Servicio de Medicina intensiva, Complejo Hospitalario Universitario de A Coruña (CHUAC), Sistema Galego de Saúde (SERGAS), A Coruña, Spain. 979Instituto Aragonés de Ciencias de la Salud (IACS), Zaragoza, Spain. 980Instituto Investigación Sanitaria Aragón (IIS-Aragon), Zaragoza, Spain. 981Unidad Diagnóstico Molecular. Fundación Rioja Salud, La Rioja, Spain. 982IDIVAL, Santander, Spain. 983Universidad de Cantabria, Santander, Spain. 984Hospital U M Valdecilla, Santander, Spain. 985Hospital Universitario de Getafe, Servicio de Genética, Madrid, Spain. 986IIS La Fe, Plataforma de Farmacogenética, Valencia, Spain. 987Departamento de Farmacología, Universidad de Valencia, Valencia, Spain. 988Facultad de Ciencias, Universidad de los Andes, Bogotá, Colombia. 989SIGEN Alianza Universidad de los Andes—Fundación Santa Fe de Bogotá, Bogotá, Colombia. 990Hospital General de Segovia, Medicina Intensiva, Segovia, Spain. 991Instituto de Biomedicina (IBIOMED), Universidad de León, León, Spain. 992Departamento Patologia y Laboratorios, Fundación Santa Fe de Bogota, Bogotá, Colombia. 993Unidad de Genética y Genómica Islas Baleares, Islas Baleares, Spain. 994Unidad de Diagnóstico Molecular y Genética Clínica, Hospital Universitario Son Espases, Islas Baleares, Spain. 995Faculdade de Medicina, Universidade de Brasília, Brasília, Brazil. 996Programa de Pós-Graduação em Ciências Médicas, Universidade de Brasília, Brasília, Brazil. 997Programa de Pós-Graduação em Ciências da Saúde, Universidade de Brasília, Brasília, Brazil. 998Unidad Cuidados Intensivos, Hospital El Bierzo, León, Spain. 999Medicina Interna, Hospital Universitario Mostoles, Madrid, Spain. 1000Universidad Francisco de Vitoria, Madrid, Spain. 1001Departamento de Analises Clinicas e Toxicologicas, Universidade Federal do Rio Grande do Norte, Natal, Brazil. 1002Programa de Pós-Graduação em Biologia Animal, Universidade de Brasília, Brasília, Brazil. 1003Programa de Pós-Graduação Profissional em Ensino de Biologia, Universidade de Brasília, Brasília, Brazil. 1004Department of Child and Adolescent Psychiatry, Institute of Psychiatry and Mental Health, Hospital General Universitario Gregorio Marañón (IiSGM), Madrid, Spain. 1005Centre for Biomedical Network Research on Mental Health (CIBERSAM), Instituto de Salud Carlos III, Madrid, Spain. 1006School of Medicine, Universidad Complutense, Madrid, Spain. 1007Núcleo de Pesquisas em Oncologia, Universidade Federal do Pará, Belém, Brazil. 1008Infectious Diseases, Microbiota and Metabolism Unit, Center for Biomedical Research of La Rioja (CIBIR), Logroño, Spain. 1009Inditex, A Coruña, Spain. 1010GENYCA, Madrid, Spain. 1011Clinica Comfamiliar Risaralda, Pereira, Colombia. 1012Neurometabolic Diseases Laboratory, Bellvitge Biomedical Research Institute (IDIBELL), L’Hospitalet de Llobregat, Spain. 1013Catalan Institution of Research and Advanced Studies (ICREA), Barcelona, Spain. 1014Unidad de Infección Viral e Inmunidad, Centro Nacional de Microbiología (CNM), Instituto de Salud Carlos III (ISCIII), Madrid, Spain. 1015Centro de Investigación Biomédica en Red de Enfermedades Infecciosas (CIBERINFEC), Instituto de Salud Carlos III, Madrid, Spain. 1016Unidad de Cuidados Intensivos, Hospital Clínico Universitario de Santiago (CHUS), Sistema Galego de Saúde (SERGAS), Santiago de Compostela, Spain. 1017Department of Preventive Medicine and Public Health, School of Medicine, Universidad Autónoma de Madrid, Madrid, Spain. 1018IdiPaz (Instituto de Investigación Sanitaria Hospital Universitario La Paz), Madrid, Spain. 1019IMDEA-Food Institute, CEI UAM+CSIC, Madrid, Spain. 1020Complejo Asistencial Universitario de León, León, Spain. 1021Servicio de Análisis Clínicos e Inmunología, Hospital Universitario Virgen de las Nieves, Granada, Spain. 1022Departamento Bioquímica, Biología Molecular e Inmunología III, Universidad de Granada, Granada, Spain. 1023Genomics of Complex Diseases Unit, Research Institute of Hospital de la Santa Creu i Sant Pau, IIB Sant Pau, Barcelona, Spain. 1024Servicio de Anestesiologia y Reanimación, Hospital Clinico Universitario de Valladolid, Valladolid, Spain. 1025Departamento de Cirugía, Universidad de Valladolid, Valladolid, Spain. 1026Hospital de Niños Ricardo Gutierrez, Buenos Aires, Argentina. 1027Fundación Universitaria de Ciencias de la Salud, Bogotá, Colombia. 1028Familial Cancer Clinical Unit, Spanish National Cancer Research Centre, Madrid, Spain. 1029Life Science Research Center, Universidad Simón Bolívar, Barranquilla, Colombia. 1030Instituto Murciano de Investigación Biosanitaria (IMIB-Arrixaca), Murcia, Spain. 1031Sección Genética Médica—Servicio de Pediatría, Hospital Clínico Universitario Virgen de la Arrixaca, Servicio Murciano de Salud, Murcia, Spain. 1032Departamento Cirugía, Pediatría, Obstetricia y Ginecología, Facultad de Medicina, Universidad de Murcia (UMU), Murcia, Spain. 1033Grupo Clínico Vinculado, Centre for Biomedical Network Research on Rare Diseases (CIBERER), Instituto de Salud Carlos III, Madrid, Spain. 1034Department of Genetics & Genomics, Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital—Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain. 1035Human Genotyping-CEGEN Unit, Spanish National Cancer Research Centre, Madrid, Spain. 1036Instituto de Genética Médica y Molecular (INGEMM), Hospital Universitario La Paz-IDIPAZ, Madrid, Spain. 1037ERN-ITHACA—European Reference Network, Madrid, Spain. 1038Unit of Infectious Diseases, Hospital Universitario 12 de Octubre, Instituto de Investigación Sanitaria Hospital 12 de Octubre (imas12), Madrid, Spain. 1039Spanish Network for Research in Infectious Diseases (REIPI RD16/0016/0002), Instituto de Salud Carlos III, Madrid, Spain. 1040Pediatric Neurology Unit, Department of Pediatrics, Navarra Health Service Hospital, Pamplona, Spain. 1041Navarra Health Service, NavarraBioMed Research Group, Pamplona, Spain. 1042Neumología, Hospital Universitario Virgen Macarena, Seville, Spain. 1043Spanish National Cancer Research Center, CNIO Biobank, Madrid, Spain. 1044Universidad Católica San Antonio de Murcia (UCAM), Murcia, Spain. 1045Servicio de Medicina Interna-Unidad de Enfermedades Infecciosas, Hospital Universitario de Salamanca-IBSAL, Salamanca, Spain. 1046Universidad de Salamanca, Salamanca, Spain. 1047Hospital Universitario Mutua Terrassa, Barcelona, Spain. 1048Xenética Cardiovascular, Instituto de Investigación Sanitaria de Santiago (IDIS), Santiago de Compostela, Spain. 1049Servicio de Medicina Interna, Hospital Universitario de Salamanca-IBSAL, Salamanca, Spain. 1050Oncology and Genetics Unit, Instituto de Investigacion Sanitaria Galicia Sur, Xerencia de Xestion Integrada de Vigo-Servizo Galego de Saúde, Vigo, Spain. 1051Departament of Microgravity and Translational Regenerative Medicine, Otto von Guericke University, Magdeburg, Germany. 1052Unidad de Genética, Hospital Universitario Mostoles, Madrid, Spain. 1053Preventive Medicine Department, Instituto de Investigacion Sanitaria Galicia Sur, Xerencia de Xestion Integrada de Vigo-Servizo Galego de Saúde, Vigo, Spain. 1054Servicio de Cardiología, Hospital Universitario de Salamanca-IBSAL, Salamanca, Spain. 1055Instituto Regional de Investigación en Salud-Universidad Nacional de Caaguazú, Caaguazú, Paraguay. 1056Urgencias Hospitalarias, Complejo Hospitalario Universitario de A Coruña (CHUAC), Sistema Galego de Saúde (SERGAS), A Coruña, Spain. 1057Grupo de Investigación en Interacciones Gen-Ambiente y Salud (GIIGAS), Instituto de Biomedicina (IBIOMED), Universidad de León, León, Spain. 1058Ministerio de Salud Ciudad de Buenos Aires, Buenos Aires, Argentina. 1059Data Analysis Department, Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital, Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain. 1060Department of Immunology, Hospital Universitario 12 de Octubre, Madrid, Spain. 1061Transplant Immunology and Immunodeficiencies Group, Instituto de Investigación Sanitaria Hospital 12 de Octubre (imas12), Madrid, Spain. 1062Department of Immunology, Ophthalmology and ENT, Universidad Complutense de Madrid, Madrid, Spain. 1063Unidad de Investigación Médica en Enfermedades Infecciosas y Parasitarias, Centro Médico Nacional Siglo XXI, Instituto Mexicano del Seguro Social (IMSS), Mexico City, Mexico. 1064Centro de Investigación Biomédica en Red de Enfermedades Infecciosas (CIBERINFEC), Instituto de Salud Carlos III (ISCIII), Madrid, Spain. 1065Departamento de Ensino e Pesquisa, Hospital Ophir Loyola, Belém, Brazil. 1066Allergy Unit, Hospital Infanta Elena, Madrid, Spain. 1067Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital—Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain. 1068Faculty of Medicine, Universidad Francisco de Vitoria, Madrid, Spain. 1069Hospital Universitario Infanta Leonor, Madrid, Spain. 1070Complutense University of Madrid, Madrid, Spain. 1071Gregorio Marañón Health Research Institute (IiSGM), Madrid, Spain. 1072Haemostasis and Thrombosis Unit, Hospital de la Santa Creu i Sant Pau, IIB Sant Pau, Barcelona, Spain. 1073Servicio de Hematologia y Hemoterapia, Hospital Clinico Universitario de Valladolid, Valladolid, Spain. 1074University Hospital of Burgos, Burgos, Spain. 1075Department of Clinical Immunology, Aarhus University Hospital, Aarhus, Denmark. 1076Department of Clinical Immunology, Copenhagen University Hospital— Rigshospitalet, Copenhagen, Denmark. 1077deCODE genetics, Reykjavik, Iceland. 1078Department of Finance Copenhagen Business School, Copenhagen, Denmark. 1079Danish Headache Center, Department of Neurology, Copenhagen University Hospital, Rigshospitalet—Glostrup, Copenhagen, Denmark. 1080Centre for Cancer Research, Danish Cancer Society, Copenhagen, Denmark. 1081Department of Dermatology, Zealand University hospital, Roskilde, Denmark. 1082Department of Biomedicine, Aarhus University, Aarhus, Denmark. 1083Institute of Biological Psychiatry, Mental Health Centre Sct. Hans, Copenhagen University Hospital, Roskilde, Denmark. 1084deCODE Genetics, Reykjavik, Iceland. 1085Department of Clinical Immunology, Aalborg University Hospital, Aalborg, Denmark. 1086Department of Clinical Immunology, Odense University Hospital, Odense, Denmark. 1087Department of Neuroscience, Karolinska Institutet, Stockholm, Sweden. 1088Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany. 1089Stanley Center for Psychiatric Research & Program in Medical and Population Genetics, Cambridge, UK. 1090Anaesthesiology and Intensive Care Medicine, Department of Surgical Sciences, Uppsala University, Uppsala, Sweden. 1091Integrative Physiology, Department of Medical Cell Biology, Uppsala University, Uppsala, Sweden. 1092Hedenstierna Laboratory, CIRRUS, Anaesthesiology and Intensive Care Medicine, Department of Surgical Sciences, Uppsala University, Uppsala, Sweden. 1093Division Anesthesiology and Intensive Care, CLINTEC, Karolinska Institutet, Stockholm, Sweden. 1094Department of Genetic Epidemiology, University of Regensburg, Regensburg, Germany. 1095Institute of Medical Microbiology and Hygiene, Molecular Microbiology (Virology), University of Regensburg, Regensburg, Germany. 1096Virologisches Institut—Klinische und Molekulare Virologie, Universitätsklinikum Erlangen, Friedrich-Alexander Universität Erlangen-Nürnberg (FAU), Erlangen, Germany. 1097Institute of Clinical Microbiology and Hygiene, University Hospital, Regensburg, Germany. 1098Department of Computer Science, School of Engineering, University of California Los Angeles, Los Angeles, CA, USA. 1099University of California Los Angeles, Los Angeles, CA, USA. 1100Department of Psychiatry and Biobehavioral Sciences, David Geffen School of Medicine at University of California Los Angeles, Los Angeles, CA, USA. 1101Division of Immunology, Allergy and Rheumatology, Department of Pediatrics, Department of Microbiology, Immunology and Molecular Genetics, University of California Los Angeles, Los Angeles, CA, USA. 1102Departments of Neurology, Psychiatry and Human Genetics, Center for Autism Research and Treatment, Institute for Precision Health, Los Angeles, CA, USA. 1103Department of Pathology and Laboratory Medicine, Human Genetics, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA. 1104Bioinformatics IDP, University of California Los Angeles, Los Angeles, CA, USA. 1105Department of Neurology, David Geffen School of Medicine at University of California Los Angeles, Los Angeles, CA, USA. 1106Departments of Human Genetics and Urology, David Geffen School of Medicine at University of California Los Angeles, Los Angeles, CA, USA. 1107Big Data Institute, Nuffield Department of Population Health, University of Oxford, Li Ka Shing Centre for Health Information and Discovery, Oxford, UK. 1108Genomics PLC, Oxford, UK. 1109Nuffield Department of Medicine, Experimental Medicine Division, University of Oxford, John Radcliffe Hospital, Oxford, UK. 1110Public Health England, Field Service, Addenbrooke’s Hospital, Cambridge, UK. 1111Public Health England, Data and Analytical Services, National Infection Service, London, UK. 1112Department of Molecular and Human Genetics, Baylor College of Medicine, Houston, TX, USA. 1113Program in Bioinformatics and Integrative Genomics, Harvard Medical School, Boston, MA, USA. 1114Program in Biological and Biomedical Sciences, Harvard Medical School, Boston, MA, USA. 1115Center for Economic and Social Research, University of Southern California, Los Angeles, CA, USA. 1116Department of Economics, University of Southern California, Los Angeles, CA, USA. 1117Institute for Molecular Medicine Finland (FIMM), Univerisity of Helsinki, Helsinki, Finland. 1118Wellcome Centre for Human Genetics, University of Oxford, Oxford, UK. 1119Department of Clinical Research and Leadership, George Washington University, Washington, DC, USA. 1120Institute for Biomedical Research of Barcelona (IIBB), National Spanish Research Council (CSIC), Madrid, Spain. 1121Institut d’Investigacions Biomèdiques August Pi i Sunyer (IDIBAPS), Barcelona, Spain. 1122Institute of Biomedicine of Valencia (IBV), CSIC, Valencia, Spain. 1123Network Center for Biomedical Research on Neurodegenerative Diseases CIBERNED, Instituto de Salud Carlos III (ISCIII), Valencia, Spain. 1124Institute for Biomedical Research of Barcelona (IIBB), National Spanish Research Council (CSIC), Madrid, Spain. 1125Department of Neurology, Hospital Universitari MútuaTerrassa, Fundació Docència i Recerca MútuaTerrassa, Terrassa, Spain. 1126Department of Molecular and Cell Biology, Centro Nacional de Biotecnología (CNB-CSIC), Campus Universidad Autónoma de Madrid, Madrid, Spain. 1127Instituto de Física de Cantabria (IFCA-CSIC), Santander, Spain. 1128Hospital Clínic, IDIBAPS, Barcelona, Spain. 1129Hospital Clínic, Barcelona, Spain. 1130Hospital Clínic, IDIBAPS, School of Medicine, University of Barcelona Barcelona, Barcelona, Spain. 1131IDIBAPS, Nature | Vol 621 | 7 September 2023 | E25 Barcelona, Spain. 1132IIBB-CSIC, Barcelona, Spain. 1133Servicio de Salud del Principado de Asturias, Oviedo, Spain. 1134Hospital Mutua de Terrassa, Terrassa, Spain. 1135Vall d’Hebron Research Institute, Vall d’Hebron Barcelona Hospital Valle Hebrón Campus, Barcelona, Spain. 1136Unidad de Excelencia Instituto de Biomedicina y Genética Molecular (IBGM, Universidad de Valladolid-CSIC), Valladolid, Spain. 1137Hospital Clínico Universitario de Valladolid (SACYL), Valladolid, Spain. 1138Department of Neurology, University Hospital of Albacete, Albacete, Spain. 1139Research Unit, University Hospital of Albacete, Albacete, Spain. 1140Department of Neurology, Biomedical Research Institute Sant Pau (IIB Sant Pau), Hospital de la Santa Creu i Sant Pau, Barcelona, Spain. 1141Hospital Universitario Ramón Y Cajal, IRYCIS, Madrid, Spain. 1142Institute de Biomedicine of Seville, IBiS/Hospital Universitario Virgen del Rocío/CSIC/University of Seville & Department of Neurology, Hospital Universitario Virgen Macarena, Seville, Spain. 1143Unaffiliated: Cindy Boer, Venceslas Douillard, Marie-Julie Fave, Rocío Gallego Durán, Yael Michaeli, Doaa Shaheen, Ilangumaran Shivanathan. ✉e-mail: andrea.ganna@helsinki.fi E26 | Nature | Vol 621 | 7 September 2023 Matters arising Extended Data Fig. 1 | Major COVID-19 biological pathways mapped with susceptibility and severity GWAS loci. Genome-wide significant variants associated with COVID-19 (white boxes) and the annotated genes (peach boxes) are mapped on to pathways known to be involved in (a) viral entry and innate immunity, (b) entry defence in airway mucus, and (c) type I interferon. The suggested phenotypic impact of the significant variants using the Bayesian approach2 are denoted with shapes; COVID-19 susceptibility (pink circles), COVID-19 disease severity (green triangles), and unclassified variants (blue cross). Other genes known to be involved in the aforementioned pathways are shown using grey boxes. Detailed list of references to studies used to design this pathway can be found in Supplementary Note. 1nature portfolio | reporting sum m ary M arch 2021 Corresponding author(s): Andrea Ganna Last updated by author(s): Apr 20, 2023 Reporting Summary Nature Portfolio wishes to improve the reproducibility of the work that we publish. This form provides structure for consistency and transparency in reporting. For further information on Nature Portfolio policies, see our Editorial Policies and the Editorial Policy Checklist. Statistics For all statistical analyses, confirm that the following items are present in the figure legend, table legend, main text, or Methods section. n/a Confirmed The exact sample size (n) for each experimental group/condition, given as a discrete number and unit of measurement A statement on whether measurements were taken from distinct samples or whether the same sample was measured repeatedly The statistical test(s) used AND whether they are one- or two-sided Only common tests should be described solely by name; describe more complex techniques in the Methods section. A description of all covariates tested A description of any assumptions or corrections, such as tests of normality and adjustment for multiple comparisons A full description of the statistical parameters including central tendency (e.g. means) or other basic estimates (e.g. regression coefficient) AND variation (e.g. standard deviation) or associated estimates of uncertainty (e.g. confidence intervals) For null hypothesis testing, the test statistic (e.g. F, t, r) with confidence intervals, effect sizes, degrees of freedom and P value noted Give P values as exact values whenever suitable. For Bayesian analysis, information on the choice of priors and Markov chain Monte Carlo settings For hierarchical and complex designs, identification of the appropriate level for tests and full reporting of outcomes Estimates of effect sizes (e.g. Cohen's d, Pearson's r), indicating how they were calculated Our web collection on statistics for biologists contains articles on many of the points above. Software and code Policy information about availability of computer code Data collection No code was used to collect data in the study. Data analysis Each individual study that contributed genetic-phenotype association summary statistics to the consortium carried out their association analyses independently of the consortium (study-specific information outlined in Supplementary Table 1). However, the consortium released phenotyping and analysis guidelines as a recommendation (https://www.covid19hg.org/). For quality control of genotype data we recommended using the Ricopili pipeline (PMID: 31393554). For genotype phasing and imputation we recommended the TopMed Imputation Server (PMID: 27571263) or Michigan Imputation Server (PMID: 27571263). For genome-wide association study (GWAS), we recommended SAIGE (PMID: 30104761), but some studies used PLINK (PMID: 17701901). Each study then submitted their GWAS summary statistics to the consortium for meta-analysis. No version was specified in the recommendation—each study used the best available version for them. LD score regression v 1.0.1 [PMID: 25642630] was used for heritability and partitioned heritability analyses. Variants for Mendelian randomization instruments were selected using PLINK version 1.90b6.18 (PMID: 17701901). Exposure and outcome datasets were harmonized, and MR statistical analysis conducted using R version 4.0.3. with the R-package TwoSampleMR version 0.5.5 (PMID: 29846171) (which included a fixed-effect IVW analysis [PMID: 24114802], weighted median estimator [WME] [PMID: 27061298], weighted mode based estimator [WMBE] and MR Egger regression [PMID: 26050253]) and additionally MR-PRESSO version 1.0 (PMID: 29686387). Code availability statement: The code for summary statistics lift-over, the projection PCA pipeline including precomputed loadings and meta- analyses are available at https://github.com/covid19-hg/; heritability estimation at https://github.com/AndrewsLabUCSF/ COVID19_heritability; Mendelian randomization and genetic correlation at https://github.com/marcoralab/MRcovid; and subtype analyses at https://github.com/mjpirinen/covid19-hgi_subtypes. For manuscripts utilizing custom algorithms or software that are central to the research but not yet described in published literature, software must be made available to editors and reviewers. We strongly encourage code deposition in a community repository (e.g. GitHub). See the Nature Portfolio guidelines for submitting code & software for further information. 2nature portfolio | reporting sum m ary M arch 2021 Data Policy information about availability of data All manuscripts must include a data availability statement. This statement should provide the following information, where applicable: - Accession codes, unique identifiers, or web links for publicly available datasets - A description of any restrictions on data availability - For clinical datasets or third party data, please ensure that the statement adheres to our policy Summary statistics generated by COVID-19 HGI are available online, including per-ancestry summary statistics for African, Admixed American, East Asian, European, and South Asian ancestries (https://www.covid19hg.org/results/r7/). The analyses described here utilize the data release 7. If available, individual-level data can be requested directly from contributing studies, listed in Supplementary Table 1. We used publicly available data from GTEx v8 (https://gtexportal.org/home/; dbGaP Accession phs000424.v8.p2), the Neale lab (http://www.nealelab.is/uk-biobank/), Finucane lab (https://www.finucanelab.org), FinnGen Freeze 4 cohort (https:// www.finngen.fi/en/access_results), and eQTL catalogue release 3 (http://www.ebi.ac.uk/eqtl/). Human research participants Policy information about studies involving human research participants and Sex and Gender in Research. Reporting on sex and gender Biological sex was collected by each study. The consortium recommended using biological sex as one of the covariates to correct for potential confounding effects (full recommended covariates are: age + age2 + sex + age*sex + 20 principal components obtained using genetic data). No sex or gender specific analysis was reported in this study. The consortium did not collect gender information. Population characteristics Summary statistics from 82 independent studies were included in consortium meta-analyses. The sample size for genetic ancestry populations for Sars-CoV2 infection was: n=17,898 Middle Eastern; n=53,116 South Asian; 40,580 East Asian; 148,891 African; 129,629 Ad-mixed American; 2,824,391 European. Population characteristics regarding age, sex and exact case and control sample numbers for each contributing study are given in Supplementary Table 1. Recruitment The consortium pre-defined phenotype criteria for cases and controls, but the specific recruitment was carried out independently by each contributing study. COVID-19 disease status (critical illness, hospitalization status) was assessed following the Diagnosis and Treatment Protocol for Novel Coronavirus Pneumonia (PMID: 32358325). The critically ill COVID-19 group included patients who were hospitalized due to symptoms associated with laboratory-confirmed SARSCoV-2 infection and who required respiratory support or whose cause of death was associated with COVID-19. The hospitalized COVID-19 group included patients who were hospitalized due to symptoms associated with laboratory confirmed SARS-CoV-2 infection. The reported infection cases group included individuals with laboratory-confirmed SARSCoV-2 infection or electronic health record, ICD coding or clinically confirmed COVID-19, or self-reported COVID-19 (e.g. by questionnaire), with or without symptoms of any severity. Genetic ancestry-matched controls for the three case definitions were sourced from population-based cohorts, including individuals whose exposure status to SARS-CoV-2 was either unknown or infection- negative for questionnaire/electronic health record based cohorts. Ethics oversight Ethical statements for each contributing study are given in Supplementary Table 1. Note that full information on the approval of the study protocol must also be provided in the manuscript. Field-specific reporting Please select the one below that is the best fit for your research. If you are not sure, read the appropriate sections before making your selection. Life sciences Behavioural & social sciences Ecological, evolutionary & environmental sciences For a reference copy of the document with all sections, see nature.com/documents/nr-reporting-summary-flat.pdf Life sciences study design All studies must disclose on these points even when the disclosure is negative. Sample size The consortium meta-analysed genome-wide association study (GWAS) summary statistics from any individual study that had included a minimum of 100 cases and 100 controls in their analysis. The cutoff was aimed at reducing noise for the meta-analysis, but also to be inclusive of studies that had not yet accumulated large numbers of COVID-19 patient data. The consortium aimed to reach the largest possible sample size for the study and meta-analyzed all the submitted summary statistics. No statistical calculation for adequate sample size was performed, but the results identifying multiple genomic regions at genome-wide significance threshold indicates adequate power for genetic discovery. Data exclusions Individual level phenotype and genotype data exclusions were performed by each individual study, following the consortium analysis plan recommendations (www.covid19hg.org). Possible reasons for sample exclusion included removing genetic ancestry outliers within a study 3nature portfolio | reporting sum m ary M arch 2021 (using principal components analysis), poor quality of genetic data or lack of phenotypic data for a sample. The consortium manually examined GWAS summary statistics data submitted by each study (for each submitted analysis separately), including sample size used for analysis, allele frequency check against Gnomad reference panel, and distribution of test statistics. After meta- analysis, the results were checked for heterogeneity variant effects between contributing studies in Supplementary Table 2 Replication No replication was performed. The consortium meta-analysed GWAS summary statistics, bringing together as many studies as possible to achieve the largest possible sample size and statistical power for association. this meant that the consortium included most large studies of COVID-19 host genetics that have been performed to date, so it was not possible to perform replication analyses in external cohorts. Therefore we performed manual checks on each study contributing summary statistics before entering them into the meta-analysis. In addition, after meta-analysis, we performed a check for heterogeneity between variant association estimates across studies contributing data. This allowed us to better understand whether the variant effects differed much between individual studies. Randomization No randomization was performed because there was no allocation of samples to experimental groups Blinding Blinding was not relevant to the study because no intervention was implemented in the study. The case status and severity of symptoms was evaluated for each sample by investigators from each study respectively. The consortium recommended using covariates to control for confounding: age + age2 + sex + age*sex + 20 principal components (obtained using genetic data) + study specific covariates (if any). The consortium meta-analysed summary statistics from these case/control studies, not individual level data. Details of which variables each study used and how the calculated PCs for their analysis are available in Supplementary Table 1. Reporting for specific materials, systems and methods We require information from authors about some types of materials, experimental systems and methods used in many studies. Here, indicate whether each material, system or method listed is relevant to your study. If you are not sure if a list item applies to your research, read the appropriate section before selecting a response. Materials & experimental systems n/a Involved in the study Antibodies Eukaryotic cell lines Palaeontology and archaeology Animals and other organisms Clinical data Dual use research of concern Methods n/a Involved in the study ChIP-seq Flow cytometry MRI-based neuroimaging