Mapping the human genetic architecture of COVID-19

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472 | Nature | Vol 600 | 16 December 2021
Article
Mapping the human genetic architecture of
COVID-19
COVID-19 Host Genetics Initiative*
The genetic make-up of an individual contributes to the susceptibility and response to
viral infection. Although environmental, clinical and social factors have a role in the
chance of exposure to SARS-CoV-2 and the severity of COVID-191,2, host genetics may
also be important. Identifying host-specic genetic factors may reveal biological
mechanisms of therapeutic relevance and clarify causal relationships of modiable
environmental risk factors for SARS-CoV-2 infection and outcomes. We formed a
global network of researchers to investigate the role of human genetics in SARS-CoV-2
infection and COVID-19 severity. Here we describe the results of three genome-wide
association meta-analyses that consist of up to 49,562 patients with COVID-19 from 46
studies across 19 countries. We report 13 genome-wide signicant loci that are
associated with SARS-CoV-2 infection or severe manifestations of COVID-19. Several of
these loci correspond to previously documented associations to lung or autoimmune
and inammatory diseases3–7. They also represent potentially actionable mechanisms
in response to infection. Mendelian randomization analyses support a causal role for
smoking and body-mass index for severe COVID-19 although not for type II diabetes.
The identication of novel host genetic factors associated with COVID-19 was made
possible by the community of human genetics researchers coming together to
prioritize the sharing of data, results, resources and analytical frameworks. This
working model of international collaboration underscores what is possible for future
genetic discoveries in emerging pandemics, or indeed for any complex human
disease.
The COVID-19 pandemic, caused by infection with SARS-CoV-2, has
resulted in an enormous health and economic burden worldwide. One
of the most remarkable features of SARS-CoV-2 infection is the variation
in consequences, which range from asymptomatic to life-threatening,
viral pneumonia and acute respiratory distress syndrome8. Although
established host factors correlate with disease severity (for example,
increasing age, being a man and higher body-mass index
1
), these risk
factors alone do not explain all of the variability in disease severity
observed across individuals.
Genetic factors contributing to COVID-19 susceptibility and severity
may provide new biological insights into disease pathogenesis and
identify mechanistic targets for therapeutic development or drug
repurposing, as treating the disease remains a highly important goal
despite the recent development of vaccines. Further supporting this
line of inquiry, rare loss-of-function variants in genes involved in the
type I interferon response may be involved in severe forms of COVID-
199–11. At the same time, several genome-wide association studies
that investigate the contribution of common genetic variation
12–15
to
COVID-19 have provided robust support for the involvement of several
genomic loci associated with COVID-19 severity and susceptibility, with
the strongest and most robust finding for severity being at the 3p21.31
locus12–16. However, much remains unknown about the genetic basis of
susceptibility to SARS-CoV-2 and severity of COVID-19.
The COVID-19 Host Genetics Initiative (COVID-19 HGI) (https://www.
covid19hg.org/)
17
is an international, open-science collaboration to
share scientific methods and resources with research groups across the
world with the goal to robustly map the host genetic determinants of
SARS-CoV-2 infection and the severity of the resulting COVID-19 disease.
Here, we report the latest results of meta-analyses of 46 studies from
19 countries (Fig.1) for COVID-19 host genetic effects.
Meta-analyses of COVID-19
Overall, the COVID-19 HGI combined genetic data from 49,562 cases and
2million controls across 46 distinct studies (Fig.1). The data included
studies from populations of different genetic ancestries, including
European, admixed American, African, Middle Eastern, South Asian
and East Asian individuals (Supplementary Table1). An overview of the
study design is provided in Extended Data Fig.1. We performed case–
control meta-analyses in three main categories of COVID-19 disease
according to predefined and partially overlapping phenotypic crite-
ria. These included (1) critically ill cases of COVID-19 defined as those
individuals who required respiratory support in hospital or who died
due to the disease; (2) cases of moderate or severe COVID-19 defined as
those participants who were hospitalized due to symptoms associated
with the infection; and (3) all cases with reported SARS-CoV-2 infection
https://doi.org/10.1038/s41586-021-03767-x
Received: 2 March 2021
Accepted: 23 June 2021
Published online: 8 July 2021
Open access
Check for updates
*Lists of authors and their afiliations appear in the online version of the paper.

Nature | Vol 600 | 16 December 2021 | 473
regardless of symptoms (Methods). Controls for all three analyses
were selected as genetically ancestry-matched samples without known
SARS-CoV-2 infection, if that information was available (Methods).
The average age of the participants with COVID-19 across studies was
55years (Supplementary Table1). We report quantile–quantile plots
in Supplementary Fig.1 and ancestry principal component plots for
contributing studies in Extended Data Fig.2.
Across our three analyses, we reported a total of 13 independent
genome-wide significant loci associated with COVID-19 (the thresh-
old of P<1.67×10
−8
is adjusted for multiple trait testing) (Supple-
mentary Table2), most of which were shared between two or more
COVID-19 phenotypes. Two of these loci are in very close proximity
within the 3p21.31 region, which was previously reported as a single
locus associated with COVID-19 severity12–16 (Extended Data Fig.3).
Overall, we find six genome-wide significant associations for critical
illness due to COVID-19, using data from 6,179 cases and 1,483,780
controls from 16 studies (Extended Data Fig.4). Nine genome-wide
significant loci were detected for moderate to severe hospitalized
COVID-19 (including five of the six critical illness loci) from an analysis
of 13,641 cases of COVID-19 and 2,070,709 controls across 29 studies
(Fig.2a, top). Finally, seven loci reached genome-wide significance
in the analysis using data for all available 49,562 reported cases of
SARS-CoV-2 infection and 1,770,206 controls, using data from a
total of 44 studies (Fig.2a, bottom). The proportion of cases with
non-European genetic ancestry for each of the three analyses was
23%, 29% and 22%. We report the results for the lead variants at the
13 loci in different ancestry-group meta-analyses in Supplementary
Table3. We note that two loci, tagged by lead variants rs1886814
and rs72711165, had higher allele frequencies in southeast Asian
(rs1886814; 15%) and East Asian genetic ancestry (rs72711165; 8%)
whereas the minor allele frequencies in European populations were
less than 3%. This highlights the value of including data from diverse
populations for genetic discovery. We discuss the replication of pre-
vious findings and the new discoveries from these three analyses in
theSupplementary Note.
Variant effects on severity and susceptibility
We found no genome-wide significant sex-specific effects at the 13 loci.
However, we did identify significant heterogeneous effects (P<0.004)
across studies for 3 out of the 13 loci (Methods), which probably reflects
the differential ascertainment of cases (Supplementary Table2). There
was a small number of overlapping samples (n=8,380 European ances-
try; n=745 East Asian ancestry) between controls from the genOMICC
and the UK Biobank studies, but leave-one-out sensitivity analyses
did not reveal any bias in the corresponding effect sizes or P values
(Extended Data Fig.5 and Supplementary Information).
We next wanted to better understand whether the 13 significant loci
were acting through mechanisms that increased the susceptibility to
infection or that affected the progression of symptoms towards more
severe disease. For all 13 loci, we compared the lead variant (strongest
association P value) odds ratios (ORs) for the risk-increasing allele
across our different COVID-19 phenotype definitions.
Focusing on the two better powered analyses: all cases with a reported
SARS-CoV-2 infection and all cases hospitalized due to COVID-19, we find
that four of the loci have similar odds ratios between these two analyses
(Methods and Supplementary Table2). Such consistency suggests a
stronger link to susceptibility to SARS-CoV-2 infection rather than to
the development of severe COVID-19. The strongest susceptibility signal
was the previously reported ABO locus (rs912805253)
12,13,15,16
. Notably,
and in agreement with a previously reported study
15
, we also report
a locus within the 3p21.31 region that was more strongly associated
with susceptibility to SARS-CoV-2 than progression to more severe
COVID-19 phenotypes. rs2271616 showed a stronger association with
a reported SARS-CoV-2 infection (P=1.79×10−3 4; OR (95% confidence
interval (CI))=1.15 (1.13–1.18)) than hospitalization (P=1.05×10−5;
OR (95%CI)=1.12 (1.06–1.19)). For this locus—which contains addi-
tional independent signals—the linkage-disequilibrium (LD) pattern
is discordant with the P-value expectation (Extended Data Fig.6 and
Supplementary Note), pointing to a key missing causal variant or to a
potentially undiscovered multi-allelic or structural variant in this locus.
ab
c
d
e
f
MID 8,875
EAS 5,006
SAS 6,489
AFR 8,876
AMR
EUR
12,841
139,918
Effective sample size
by genetic ancestry
Effective
sample size
d
f
e
c
a
b
120
1,200
12,000
120,000
Analysis type
Reported
SARS-CoV-2
infection
Hospitalized,
SARS-CoV-2
positive
Critically ill,
SARS-CoV-2
positive
Fig. 1 | Geo graphical over view of the cont ributing s tudies to the COV ID-19 HGI and com position by m ajor ancest ry groups . Populations a re defined a s
Africa n (AFR), admixed Ame rican (AMR ), East Asian (EA S), European (EUR), Middle Ea stern (MID) and So uth Asian (SAS).

474 | Nature | Vol 600 | 16 December 2021
Article
By contrast, 9 out of the 13 loci were associated with increased risk
of severe symptoms with significantly larger odds ratios for hospi-
talized COVID-19 compared with the mildest phenotype of reported
SARS-CoV-2 infection (eight loci were below the threshold of P<0.004
(test for effect size difference) and, in addition, the lead variant
rs10774671 had a clear increase in odds ratios despite not passing this
threshold) (Supplementary Table2). We further compared the odds
ratios for these nine loci for critical illness due to COVID-19 versus hos-
pitalized due to COVID-19, and found that these loci exhibited a gen-
eral increase in effect risk for critical illness (Methods, Extended Data
Fig.7a and Supplementary Table4), but the lower power for association
analysis of critically ill COVID-19 means that these results should be
considered as suggestive. Overall, these results indicated that these
nine loci were more likely to be associated with progression of the
disease and worse outcome from SARS-CoV-2 infection compared to
being associated with susceptibility to SARS-CoV-2 infection.
For some of these analyses, the controls were simply existing popula-
tion controls without knowledge of SARS-CoV-2 infection or COVID-19
status, which may bias effect size estimates as some of these individuals
may have either become infected with SARS-CoV-2 or developed COVID-
19. We perform several sensitivity analyses (Extended Data Fig.7b,
Supplementary Note and Supplementary Table4) in which we show
that using population controls can be a valid and powerful strategy
for host genetic discovery of infectious disease, and particularly those
that are widespread and with rare severe outcomes.
Gene prioritization and association with other traits
To better understand the potential biological mechanism of each locus,
we applied several approaches to prioritize candidate causal genes and
explore additional associations with other diseases and traits. Of the 13
genome-wide significant loci, we found that nine loci implicated bio-
logically plausible genes (Supplementary Tables 2, 5). Protein-altering
variants in LD with lead variants implicated genes at six loci, including
TYK2 (chromosome and cytogenetic band (chr.)19p13.2) and PPP1R15A
(chr.19q13.33). The COVID-19 lead variant rs74956615T>A in TYK2,
which confers risk for critical illness (OR (95% CI)=1.43 (1.29–1.59),
P=9.71×10−12) and hospitalization due to COVID-19 (OR (95% CI)=1.27
(1.18–1.36), P=5.05×10−10) is correlated with the missense variant
rs34536443:G>C (p.Pro1104Ala; r2=0.82) . This is consistent with the
primary immunodeficiency described with complete TYK2 loss of func-
tion
3
as this variant is known to reduce function
18,19
. By contrast, this
missense variant was previously reported to be protective against auto-
immune diseases (Extended Data Fig.8 and Supplementary Table6),
including rheumatoid arthritis (OR=0.74, P=3.0×10
−8
; UK Biobank
SAIGE) and hypothyroidism (OR=0.84, P=1.8×10−10; UK Biobank). At
the 19q13.33 locus, the lead variant rs4801778, which was significantly
associated with a reported SARS-CoV-2 infection (OR (95% CI)=0.95
(0.93–0.96), P=2.1×10
−8
), is in LD (r
2
=0.93) with a missense variant
rs11541192:G>A (p.Gly312Ser) in PPP1R15A.
A lung-specific cis-expression quantitative trait loci (cis-eQTLs)
from GTEx v.8
20
(n=515) and the Lung eQTL Consortium
21
(n=1,103)
provided further support for a subset of loci (Supplementary
Table7), including FOXP4 (chr.6p21.1) and ABO (chr.9q34.2),
OAS1/OAS3/OAS2 (chr.12q24.13) and IFNAR2/IL10RB (21q22.11), where
the COVID-19-associated variants modify gene expression in lung.
Furthermore, our phenome-wide association study (PheWAS) analysis
(Supplementary Table6) implicated three additional loci related to lung
function, with modest lung eQTL evidence—that is, the lead variant
was not fine-mapped but significantly associated. An intronic variant
rs2271616
rs11919389
rs912805253
rs4801778
rs13050728
rs2109069
rs74956615
rs1819040
rs77534576
rs10774671
rs72711165
rs1886814
rs10490770
–log
10
(P)–log
10
(P)
Not genome-wide signicant
Closest gene
eGenes
Gene with highest V2G score
Genes with coding variants
Genes in LD region
Type of evidence for gene annotation
Chr. 9
Chr. 12 Chr. 17 Chr. 19 Chr. 21
Chr. 6
Chr. 3 Chr. 8
rs1886814
rs72711165
rs912805253
rs10774671
rs11919389
rs1819040
rs2109069
rs77534576
OAS1
OAS2
OAS3
DPP9
IFNAR2
rs74956615
rs13050728
RAVER1
TYK2
ICAM5
ICAM1
ICAM4
ZGLP1
FDX2
ICAM3
rs4801778
PLEKHA4
TULP2
HSD17B14
PPP1R15A
NUCB1
FOXP4
TMEM65
ABO
RPL24
NXPE3
ZBTB11
CEP97
KANSL1
ARHGAP27
PLEKHM1
LINC02210-CRHR1
CRHR1
SPPL2C
MAPT
STH
LRRC37A
ARL17B
LRRC37A2
ARL17A
NSF
WNT3
TAC4
DLX3
FLJ45513
KAT7
rs10490770
LZTFL1
CXCR6
rs2271616
SLC6A20
CCR3
Critically ill,
SARS-CoV-2 positive
Hospitalized,
SARS-CoV-2 positive
Reported SARS-CoV-2 infection
a b
Hospitalized, SARS-CoV-2 positive
Chromosome
1
79
63
50
40
32
32
40
50
63
79
25
25
20
20
16
16
13
13
10
10
9
9
8
8
7
7
6
6
5
5
4
4
3
3
2
2
2345678910 11 12 13 14 15 16 17 18 20 22 X
Reported SARS-CoV-2 infection
Fig. 2 | Geno me-wide as sociatio n results for COV ID-19. a, Top, results of a
genome-wi de associat ion study of hospi talized cas es of COVID-19 (n=1 3,641
cases an dn=2,070,709 cont rols). Bottom, the res ults of repor ted SARS- CoV-2
infections (n=49,562 cases and n=1,770,206 controls). Loci highlighted in
yellow (top) represent r egions ass ociated with t he severity of th e COVID-19
manifesta tion—that is, incre ased odds of m ore severe COVID-19 phen otypes.
Loci highli ghted in gree n (bottom) are reg ions associ ated with susc eptibility t o
a SARS- CoV-2 infect ion—that is, the ef fect is the sam e across mild and se vere
COVID-19 phenot ypes. We hig hlight in red genom e-wide signif icant vari ants
that had high heterogeneity across contributing studies and that were
therefore exclude d from the list of lo ci found. b, Results o f gene prioriti zation
using diffe rent evidence m easures of gene a nnotation. G enes in the LD re gion,
genes wit h coding variant s and eGene s (fine-mappe d cis-eQTL varian t PIP>0.1
in GTEx Lung ) are annotate d if in LD with a COVID -19 lead variant (r2>0.6). V2G,
highest ge ne prioritiz ed by the V2G score of O pen Target Genetic s.

Nature | Vol 600 | 16 December 2021 | 475
rs2109069:G>A in DPP9 (chr.19p13.3), which is positively associated
with critical illness, was previously reported to be risk-increasing for
interstitial lung disease (tag lead variant rs12610495:A>G (p.Leu8Pro);
OR=1.29, P=2.0×10
−12
)
5
. The COVID-19 lead variant rs1886814:A>C
in the FOXP4 locus is correlated (r
2
=0.64) with a lead variant of lung
adenocarcinoma (tag variant is rs7741164; OR=1.2, P=6.0×10
−13
)
6,22
and similarly with a lead variant reported for subclinical interstitial lung
disease
23
. In severe COVID-19, lung cancer and interstitial lung disease,
the minor, expression-increasing allele is associated with increased
risk. We also found that intronic variants (chr.1q22) and rs1819040:T>A
in KANSL1 (chr.17q21.31), associated with protection against hospi-
talization due to COVID-19, were previously reported for reduced lung
function (for example, tag lead variant rs141942982:G>T; OR (95%
CI)=0.96 (0.95–0.97), P=1.00×10
−20
)
7
. Notably, the 17q21.31 locus
is a well-known locus for structural variants containing a megabase
inversion polymorphism (H1 and inverted H2 forms) and complex
copy-number variations, in which the inverted H2 forms were shown
to be positively selected in European individuals24,25.
Lastly, there are two loci in the 3p21.31 region with varying genes
prioritized by different methods for different independent signals. For
the severity lead variant rs10490770:T>C, we prioritized CXCR6 with
the Variant2Gene (V2G) algorithm26, although LZTFL1 is the closest
gene. The CXCR6 has a role in chemokine signalling27 and LZTFL1 has
been implicated in lung cancer
28
. rs2271616:G>T, which is associated
with susceptibility, tags a complex region including several independ-
ent signals (Supplementary Note) that are all located within the gene
body of SLC6A20, which encodes a protein that is known to function-
ally interact with the SARS-CoV-2 receptor ACE2
29
. However, none of
the lead variants in the 3p21.31 region has been previously associated
with other traits or diseases in our PheWAS analysis. Although these
results provide supporting in silico evidence for candidate causal gene
prioritization, further functional characterization is needed. Detailed
locus descriptions and LocusZoom plots are provided in Supplemen-
tary Fig.2.
Polygenic architecture of COVID-19
To further investigate the genetic architecture of COVID-19, we used
results from meta-analyses including samples from European ancestries
(sample sizes are described in the Methods and Supplementary Table1)
to estimate the heritability explained by common single-nucleotide
polymorphisms—that is, the proportion of variation in the two phe-
notypes that was attributable to common genetic variants—and to
determine whether heritability of COVID-19 phenotypes was enriched
in genes that were specifically expressed in certain tissues30 from the
GTEx dataset
31
. We detected low, but significant, heritability across
all three analyses (<1% on observed scale, all P values were P<0.0001)
(Supplementary Table8). The values are low compared to previously
published studies
14
, but may be explained by differences in the reported
estimate scale (observed versus liability), the specific method used,
disease-prevalence estimates, phenotypic differences between patient
cohorts or ascertainment of controls. Despite the low reported values,
we found that heritability of a reported SARS-CoV-2 infection was sig-
nificantly enriched in genes that were specifically expressed in the
lung (P=5.0×10
−4
) (Supplementary Table9). These findings, together
with the genome-wide significant loci identified in the meta-analyses,
suggest that there is a significant polygenic architecture that can be
better leveraged with future, larger, sample sizes.
Genetic correlation and Mendelian randomization
Genetic correlations (r
g
) between the three COVID-19 phenotypes was
high, although lower correlations were observed between hospitalized
COVID-19 and reported SARS-CoV-2 infection (critical illness versus
hospitalized: rg (95% CI)=1.37 (1.08–1.65), P=2.9×10−21; critical illness
versus reported SARS-CoV-2 infection, rg (95%CI)=0.96 (0.71–1.20),
P=1.1×10−14; hospitalized versus reported SARS-CoV-2 infection: rg
(95%CI)=0.85 (0.68–1.02), P=1.1×10−22). To better understand which
traits are genetically correlated and/or potentially causally associated
with COVID-19 hospitalization, critical illness and reported SARS-CoV-2
infection, we chose a set of 38 disease, health and neuropsychiatric
phenotypes as potential COVID-19 risk factors based on their clinical
correlation with disease susceptibility, severity or mortality (Supple-
mentary Table10).
We found evidence (false-discovery rate (FDR)<0.05) of signifi-
cant genetic correlations between nine traits and hospitalized COVID-
19 and reported SARS-CoV-2 infection (Fig.3, Extended Data Fig.9
and Supplementary Table11). Notably, genetic liability to ischaemic
*
*
*
*
*
*
*
*
*
**
*
*
*
*
*
**
*
*
*
*
*
*
*
Risk factor Biomarker Disease liability
rg
MR
BMI
Cigarettes per day
Depression
Diastolic blood pressure
Height
Insomnia symptoms
Pulse pressure
Risk tolerance
Sleep duration
Smoking initiation
Systolic blood pressure
25-Hydroxyvitamin D
CRP
eGFR
High-density lipoproteins
Low-density lipoproteins
Platelet count
Red-blood-cell count
Total cholesterol
Triglycerides
White-blood-cell count
ADHD
Alzheimer’s disease
Amyotrophic lateral sclerosi
s
Asthma
Autism spectrum disorder
Bipolar disorder
Chronic kidney disease
Coronary artery disease
Diabetes
Heart failure
Idiopathic pul
monary brosis
Ischaemic stroke
Lupus
Multiple sclerosis
Parkinson’s disease
Rheumatoid arthriti
s
Schizophrenia
Reported infection
Hospitalization
Critical illness
Reported infection
Hospitalization
Critical illness
Negative/protective Positive/risk
rg and MR
Fig. 3 | Gene tic correla tions and Me ndelian ran domization c ausal
estima tes betwee n 38 traits and C OVID-19 critic al illness , hospital ization
and repor ted SARS -CoV-2 infection. L arger squares co rrespond to P value s
with high er signific ance, with gen etic correlat ions (rg) or Mendeli an
randomiz ation (MR) caus al estimate s signific antly differen t from zero. The
size of each c oloured square ind icates the mag nitude of the P valu e, with
P<0.05 shown as a full-siz ed square, P=0.05–0.1 as a larg e square, P=0.1–0.5
as a medium sq uare and P>0.5 as a small squ are. Genetic c orrelations or c ausal
estimate s that are signif icantly dif ferent from zero a t an FDR of 5% are marked
with an ast erisk. Two-sided P valu es were calculate d using LDSC for gen etic
correlations and inverse-variance-weighted analysis for Mendelian
randomization. ADHD, attention-deficit hyperactivity disorder; BMI, body
mass index; C RP, C-reac tive protein; eGFR , estimated g lomerular f iltration
rate.

476 | Nature | Vol 600 | 16 December 2021
Article
stroke was only significantly positively correlated with critical illness
or hospitalization due to COVID-19, but not with a higher likelihood of
reported SARS-CoV-2 infection (infection r
g
=0.019 versus hospitaliza-
tion r
g
=0.41, z=2.7, P=0.006; infection r
g
=0.019 versus critical illness
rg=0.40, z=2.49, P=0.013).
We next used two-sample Mendelian randomization to infer poten-
tially causal relationships between these traits. After correcting for
multiple testing (FDR<0.05), eight exposure–COVID-19 trait pairs
showed suggestive evidence of a causal association (Fig.3, Extended
Data Fig.10, Supplementary Table12 and Supplementary Fig.3). Five of
these associations were robust to potential violations of the underlying
assumptions of Mendelian randomization. Corroborating our genetic
correlation results and evidence from epidemiological studies, geneti-
cally predicted higher body-mass index (OR (95%CI)=1.4 (1.3–1.6),
P=8.5×10−11) and smoking (OR (95% CI)=1.9 (1.3–2.8), P=0.0012)
were associated with increased risk of COVID-19 hospitalization,
with body-mass index also being associated with increased risk of
SARS-CoV-2 infection (OR (95% CI)=1.1 (1.1–1.2), P=4.8×10
−7
). Geneti-
cally predicted increased height (OR (95% CI)=1.1 (1–1.1)), P=8.9×10
−4
)
was associated with an increased risk of reported SARS-CoV-2 infection,
whereas a genetically predicted higher red-blood-cell count (OR (95%
CI)=0.93 (0.89–0.96), P=5.7×10
−5
) was associated with a reduced risk
of reported SARS-CoV-2 infection. Despite evidence of a genetic cor-
relation between type II diabetes and COVID-19 outcomes, there was
no evidence of a causal association in the Mendelian randomization
analyses, which suggests that the observed genetic correlations are due
to pleiotropic effects between body-mass index and type 2 diabetes.
Further sensitivity analyses relating to sample overlap are discussed
in theSupplementary Information.
Discussion
The COVID-19 HGI has brought together investigators from across the
world to advance genetic discovery for SARS-CoV-2 infection and severe
COVID-19 disease. We report 13 genome-wide significant loci associated
with some aspect of SARS-CoV-2 infection or COVID-19. Many of these
loci overlap with previously reported associations with lung-related
phenotypes or autoimmune or inflammatory diseases, but some loci
have no obvious candidate gene.
Four out of the thirteen genome-wide significant loci showed similar
effects in the reported SARS-CoV-2 infection analysis (a proxy for dis-
ease susceptibility) and all-hospitalized COVID-19 (a proxy for disease
severity). Of these, one locus was in close proximity to, yet independent
of, the major genetic signal for COVID-19 severity at the 3p21.31 locus.
Notably, this locus was associated with COVID-19 susceptibility rather
than severity. The locus overlaps SLC6A20, which encodes an amino
acid transporter that interacts with ACE2. Nonetheless, we caution that
more data are needed to resolve the nature of the relationship between
genetic variation and COVID-19 at this locus, particularly as the physi-
cal proximity, LD structure and patterns of association suggest that
untagged genetic variation could drive the association signal in the
region. Our findings support the notion that some genetic variants,
most notably at the ABO and PPP1R15A loci, in addition to SLC6A20,
can indeed affect susceptibility to infection rather than progression
to severe COVID-19 once infected.
Several of the loci reported here—as noted in previous publica-
tions
12,14
—intersect with well-known genetic variants that have estab-
lished genetic associations. Examples of these include variants at
DPP9 and FOXP4, which show previous evidence of increasing risk
for interstitial lung disease
5
, and missense variants within TYK2 that
show a protective effect on several autoimmune-related diseases32–35.
Together with the heritability enrichment observed in genes expressed
in lung tissues, these results highlight the involvement of lung-related
biological pathways in the development of severe COVID-19. Several
other loci show no previously documented genome-wide significant
associations, despite the high significance and attractive candidate
genes for COVID-19 (for example, CXCR6, LZTFL1, IFNAR2 and OAS1/
OAS2/OAS3 loci). The previously reported associations for the strongest
association for COVID-19 severity at the 3p21.31 locus and monocytes
count are likely to be due to proximity and not a true co-localization.
Increasing the global representation in genetic studies enhances the
ability to detect novel associations. Two of the loci that affect disease
severity were only discovered by including the four studies of indi-
viduals with East Asian ancestry. One of these loci—close to FOXP4—is
common particularly in East Asian participants (32%) as well as admixed
American participants in the Americas (20%) and Middle Eastern par-
ticipants (7%), but has a low frequency in most European ancestries
(2–3%) in our data. Although we cannot be certain of the mechanism
of action, the FOXP4 association is an attractive biological target, as
it is expressed in the proximal and distal airway epithelium
36
and has
been shown to have a role in controlling epithelial cell fate during lung
development37. The COVID-19 HGI continues to pursue expansion of
the datasets included in the analyses of the consortium to populations
from underrepresented populations in upcoming data releases. We
plan to release ancestry-specific results in full once the sample sizes
allow for a well-powered meta-analysis.
Care should be taken when interpreting the results from a
meta-analysis because of challenges with case and control ascertain-
ment and collider bias (seeSupplementary Note for a more detailed
discussion on study limitations). Drawing a comprehensive and repro-
ducible map of the host genetics factors associated with COVID-19
severity and SARS-CoV-2 requires a sustained international effort to
include diverse ancestries and study designs. To accelerate downstream
research and therapeutic discovery, the COVID-19 HGI regularly pub-
lishes meta-analysis results from periodic data freezes on the website
https://www.covid19hg.org/ and provides an interactive explorer
through which researchers can browse the results and the genomic
loci in more detail. Future work will be required to better understand
the biological and clinical value of these findings. Continued efforts to
collect more samples and detailed phenotypic data should be endorsed
globally, allowing for more thorough investigation of variable, herit-
able symptoms, particularly in light of the newly emerging strains of
SARS-CoV-2, which may provoke different host responses that lead
to disease.
Online content
Any methods, additional references, Nature Research reporting sum-
maries, source data, extended data, supplementary information,
acknowledgements, peer review information; details of author con-
tributions and competing interests; and statements of data and code
availability are available at https://doi.org/10.1038/s41586-021-03767-x.
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Article
COVID-19 Host Genetics Initiative
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Mark Daly1,2,3,1280 ✉ & Andrea Ganna1,2,3,1280 ✉
Writing group
Writing group leaders
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Mohammed Jashim Uddin21, George Davey Smith22, Cristen J. Willer23,24,25 &
Joseph D. Buxbaum26
Analysis group
Manuscript analyses team leader
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Manuscript analyses team member: meta-analysis
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Alicia R. Martin3,29 & Wei Zhou3,29
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Translators
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Kyung Jae Yoon226,227,228 & Nam-Jong Paik229,230
Analysis team members
Woojin Seok231 & Heejun Yoon232
Data collection members
Eun-Jeong Joo233, Yoosoo Chang234,235, Seungho Ryu234,235, Wan Beom Park223, Jeong Su Park236,
Kyoung Un Park236, Sin Young Ham225, Jongtak Jung225, Eu Suk Kim225 & Hong Bin Kim225

Article
COVID-19-Hostage
Analysis team leaders
David Ellinghaus237,238, Frauke Degenhardt237, Mario Cáceres239,240, Simonas Juzenas237 &
Tobias L. Lenz241,242
Data collection leaders
Agustín Albillos243,244, Antonio Julià245, Bettina Heidecker246, Eva C. Schulte9,10,11,
Federico Garcia249,250, Florian Kurth246, Florian Tran237, Frank Hanses251,252, Heinz Zoller253,
Jan C. Holter254,255, Javier Fernández256,257, Leif Erik Sander246, Philip Rosenstiel237,
Philipp Koehler198,259,260, Rafael de Cid261, Rosanna Asselta262,263, Stefan Schreiber237,265,
Ute Hehr266, Daniele Prati285, Guido Baselli285, Luca Valenti285,316, Luis Bujanda244,322,329,
Jesus M. Banales244,321,322, Stefano Duga262,263, Mauro D’Amato321,322,344, Manuel Romero-Góm
ez1241,244, Maria Buti244,292,298 & Pietro Invernizzi286,287
Admin team leaders
Andre Franke237,267, Johannes R. Hov254,268,269,270, Tom H. Karlsen254,268,269,270,
Trine Folseraas254,268,269,270 & Douglas Maya-Miles1244,1245,244
Analysis team members
Ana Teles241,242, Clinton Azuure241, Eike Matthias Wacker237, Florian Uellendahl-Werth237,
Hesham ElAbd237, Jatin Arora29,272,273,274,275, Jon Lerga-Jaso239, Lars Wienbrandt237,
Malte Christoph Rühlemann237, Mareike Wendorff237, May Sissel Vadla277, Ole Bernt Lenning276,
Onur Özer241,242, Ronny Myhre278, Soumya Raychaudhuri29,272,273,274,275,279, Anja Tanck237,
Christoph Gassner237,1240, Georg Hemmrich-Stanisak237, Jan Kässens237, Maria E. Figuera
Basso237, Martin Schulzky237, Michael Wittig237, Nicole Braun237,267, Tanja Wesse237,
Wolfgang Albrecht237 & Xiaoli Yi237
Data collection members
Aaron Blandino Ortiz280, Adolfo Garrido Chercoles281, Agustín Ruiz282,283,
Alberto Mantovani262,263, Aleksander Rygh Holten254,284, Alena Mayer246,
Alessandro Cherubini285, Alessandro Protti262,263, Alessio Aghemo262,263, Alessio Gerussi286,287,
Alfredo Ramirez288,289,290,291, Alice Braun246, Ana Barreira292, Ana Lleo262,263,
Anders Benjamin Kildal293, Andrea Ganna1,2,3,1280, Andreas Glück294, Anna Carreras Nolla261,
Anna Latiano295, Anne Ma Dyrhol-Riise254,296, Antonio Muscatello285, Antonio Voza262,
Ariadna Rando-Segura297,298, Aurora Solier299,1226, Banasik Karina238, Beatriz Cortes261,
Beatriz Mateos243,244, Beatriz Nafria-Jimenez281, Benedikt Schaefer253, Carla Bellinghausen300,
Carlos Ferrando256, Carmen Quereda301, Carsten Skurk246, Charlotte Thibeault246,
Christoph D. Spinner302, Christoph Lange303,304,305, Cinzia Hu285, Claudio Cappadona263,
Cristiana Bianco285, Cristina Sancho306, Dag Arne Lihaug Hoff307,308, Daniela Galimberti285,
David Jiménez299,1226, David Pestaña309, David Toapanta310,313, Elena Azzolini262,263,
Elio Scarpini285, Elisa T. Helbig246, Eloisa Urrechaga311, Elvezia Maria Paraboschi262,263,
Emanuele Pontali312, Enric Reverter256,313, Enrique Navas301, Eunate Arana314, Félix
García Sánchez315, Ferruccio Ceriotti285, Francesco Malvestiti316, Francisco Mesonero243,244,
Gianni Pezzoli317, Giuseppe Lamorte285, Holger Neb318, Ilaria My262, Isabel Hernández282,283,
Itziar de Rojas282,283, Iván Galván-Femenia261, Jan Heyckendorf303,304,305, Jan Rybniker198,260,319,
Joan Ramon Badia256, Jochen Schneider302, Josune Goikoetxea323, Julia Kraft246 ,
Karl Erik Müller324, Karoline I. Gaede325,326,327, Koldo Garcia-Etxebarria244,322,329,
Kristian Tonby254,330, Lars Heggelund324,331, Laura Izquierdo-Sanchez244,322, Lauro Sumoy333,
Lena J. Lippert246, Leonardo Terranova285, Lindokuhle Nkambule3,29,167, Lucia Garbarino312,
Luis Téllez243,244, Luisa Roade298, Mahnoosh Ostadreza285, Maider Intxausti306,
Manolis Kogevinas335,336,337,338, Mari E. K. Niemi1,1280, María A. Gutiérrez-Stampa339,
Maria J. G. T. Vehreschild340, Marta Marquié282,283, Massimo Castoldi341, Mattia Cordioli1,
Maurizio Cecconi262,263, Mercè Boada282,283, Michael J. Seilmaier345, Michela Mazzocco312,
Miguel Rodríguez-Gandía243,244, Natale Imaz Ayo314, Natalia Blay261, Nilda Martínez346,
Norwegian SARS-CoV-2 Study Group*, Oliver A. Cornely198,259,319,349, Orazio Palmieri295,
Paolo Tentorio262, Pedro M. Rodrigues244,322,332, Pedro P. España311, Per Hoffmann351,
Petra Bacher352,353,354, Phillip Suwalski246, Raúl de Pablo280, Rosa Nieto299,1226,
Salvatore Badalamenti262, Sandra Ciesek355,356, Sara Bombace262, Sara Pigazzini1,
Sibylle Willing252,266,357, Søren Brunak238, Stefanie Heilmann-Heimbach351, The Humanitas
COVID-19 Task Force*, The Humanitas Gavazzeni COVID-19 Task Force*, Stephan Ripke246,
Thomas Bahmer294, Ulf Landmesser359, Ulrike Protzer9,3 60, Valeria Rimoldi263,
Vegard Skogen361,362, Victor Andrade289,291, Victor Moreno336,363,364,365, Wolfgang Poller246,
Xavier Farre261, Xiaomin Wang246, Yascha Khodamoradi340, Zehra Karadeniz246, Adolfo de
Salazar249,50 , Adriana Palom245,292, Alba-Estela Garcia-Fernandez1242, Albert Blanco-Grau1242,
Alberto Zanella285,316, Alessandra Bandera285,316, Almut Nebel237, Andrea Biondi1264, Andrea Caba
llero-Garralda1242, Andrea Gori285,316, Andreas Lind254,255, Anna Ludovica Fracanzani285,316,
Anna Peschuck237, Antonio Pesenti285,316, Carmen de la Horra136,1244,1245,336,1246, Chiara Milani286,287,
Cinzia Paccapelo285, Claudio Angelini1247, Cristina Cea1242, Eduardo Muñiz-Diaz1248,
Elena Sandoval1249, Enrique J. Calderón1243,1244,1245,366,1246, Erik Solligård1265,1266, Fátima Aziz1249,
Filippo Martinelli-Boneschi285,316, Flora Peyvandi285,316, Francesco Blasi285,1250,
Francisco J. Medrano1251,1243,1244,366,1246, Francisco Rodriguez-Frias245,298,244,1242, Fredrik Müller254,255,
Giacomo Grasselli285,316, Giorgio Costantino285,316, Giulia Cardamone1252, Giuseppe Foti1253,
Giuseppe Matullo1254, Hayato Kurihara1247, Jan Egil Afset307,1255, Jan Kristian Damås1256,1257,
Javier Ampuero1243,1244,1245,369, Javier Martín1259, Jeanette Erdmann1260,1261,1262, Jonas Bergan1263,
Siegfried Goerg264, Jose Ferrusquía-Acosta1249, Jose Hernández Quero249,250,
Juan Delgado1243,1244,1245,366,1246, Juan M. Guerrero1243,1244,1245, Kari Risnes1257,1258,
Laura Rachele Bettini1257,1258, Leticia Moreira1249, Lise Tuset Gustad1265,1267, Luigi Santoro285,
Luigia Scudeller285, Mar Riveiro-Barciela292,298,244, Marco Schaefer1268, Maria Carrabba285,
Maria G. Valsecchi1269, María Hernandez-Tejero256, Marialbert Acosta-Herrera1259,
Mariella D’Angiò1264, Marina Baldini285, Marina Cazzaniga1270, Michele Ciccarelli1247,
Monica Bocciolone1247, Monica Miozzo285,316, Natalia Chueca249, Nicola Montano285,316,
Paola Faverio1271, Paoletta Preatoni1247, Paolo Bonfanti1272,1273, Paolo Omodei1247, Pedro Castro256,
Ricard Ferrer84,1274, Roberta Gualtierotti285,316, Rocío Gallego-Durán144,1245,244,
Rubén Morilla1243,1244,1245,366,1246, Sammra Haider308, Sara Marsal245, Serena Aneli1254,
Serena Pelusi285,316, Silvano Bosari285,316, Stefano Aliberti285,1250, Susanne Dudman254,255,
Tenghao Zheng344, Tomas Pumarola397, Trinidad Gonzalez Cejudo249, Valter Monzani285,
Vicente Friaza1243,1244,1245,366,1246, Wolfgang Peter1268 & Ximo Dopazo1275
Norwegian SARS-CoV-2 Study Group
Tom H. Karlsen254,268,269,270
Humanitas COVID-19 Task Force
Stefano Duga262,263,341
The Humanitas Gavazzeni COVID-19 Task Force
Stefano Duga262,263,341
Admin team members
Sandra May237 & Marit M. Grimsrud254,269,270
deCODE
Analysis team leader
Daniel F. Gudbjartsson366
Data collection leader
Kari Stefansson366
Analysis team members
Patrick Sulem366, Gardar Sveinbjornsson366, Pall Melsted366, Gudmundur Norddahl366 &
Kristjan Helgi Swerford Moore366
Data collection members
Unnur Thorsteinsdottir366 & Hilma Holm366
Determining the Molecular Pathways & Genetic Predisposition of the Acute Inlammatory
Process Caused by SARS-CoV-2
Analysis team leader
Marta E. Alarcón-Riquelme102
Data collection leader
David Bernardo368,369
Analysis team member
Manuel Martínez-Bueno102
Data collection member
Silvia Rojo Rello370
Estonian Biobank
Analysis team leader
Reedik Mägi371
Data collection leader
Lili Milani371
Admin team leader
Andres Metspalu371
Analysis team members
Triin Laisk371, Kristi Läll371 & Maarja Lepamets371
Data collection members
Tõnu Esko371, Ene Reimann371, Paul Naaber372, Edward Laane373,374, Jaana Pesukova374 ,
Pärt Peterson375, Kai Kisand375, Jekaterina Tabri377, Raili Allos37 7, Kati Hensen377, Joel Starkopf378,
Inge Ringmets379, Anu Tamm380 & Anne Kallaste380
Admin team members
Helene Alavere371, Kristjan Metsalu371 & Mairo Puusepp371
FinnGen
Data collection members
Kati Kristiansson382, Juha Karjalainen1, Sami Koskelainen382, Markus Perola382,383, Kati Donner1,
Katja Kivinen1 & Aarno Palotie1
Admin team member
Mari Kaunisto1
FinnGen Admin team leader
Aarno Palotie381
Functional Host Genomics in Infectious Diseases (FHoGID)
Analysis team leader
Carlo Rivolta384,385

Data collection leaders
Pierre-Yves Bochud386, Stéphanie Bibert386, Noémie Boillat386, Semira Gonseth Nussle388 &
Werner Albrich389
Analysis team members
Mathieu Quinodoz384,385 & Dhryata Kamdar384,385
Data collection members
Noémie Suh390, Dionysios Neofytos391, Véronique Erard392, Cathy Voide393, FHoGID*,
RegCOVID*, P-PredictUs*, SeroCOVID* & CRiPSI*
FHoGID
P. Y. Bochud394,395,398, C. Rivolta394, S. Bibert394, M. Quinodoz394, D. Kamdar394, D. Neofytos394,
V. Erard394, C. Voide394, R. Friolet394, P. Vollenweider394,395, J. L. Pagani394,395, M. Oddo394,
F. Meyer zu Bentrup394, A. Conen394, O. Clerc394, O. Marchetti394, A. Guillet394,
C. Guyat-Jacques394, S. Foucras394, M. Rime394, J. Chassot394, M. Jaquet394, R. Merlet Viollet394,
Y. Lannepoudenx394 & L. Portopena394
RegCOVID
P. Y. Bochud394,395,398, P. Vollenweider394,395, J. L. Pagani394,395, F. Desgranges395, P. Filippidis395,
B. Guéry395, D. Haeliger395, E. E. Kampouri395, O. Manuel395, A. Munting395, M. Papadimitr
iou-Olivgeris395, J. Regina395, L. Rochat-Stettler395, V. Suttels395, E. Tadini395, J. Tschopp395,
M. Van Singer395 & B. Viala395
P-PredictUs
N. Boillat-Blanco396, T. Brahier396, O. Hügli396, J. Y. Meuwly396 & O. Pantet396
SérocoViD
S. Gonseth Nussle397, M. Bochud397, V. D’Acremont397 & S. Estoppey Younes397
CRiPSI
W. C. Albrich398, N. Suh398, A. Cerny398, L. O’Mahony398, C. von Mering, P. Y. Bochud394,395,398,
M. Frischknecht398, G.-R. Kleger398, M. Filipovic398, C. R. Kahlert398, H. Wozniak398,
T. Rochat Negro398, J. Pugin398, K. Bouras398, C. Knapp398 , T. Egger398, A. Perret398, P. Montillier398,
C. di Bartolomeo398 & B. Barda398
GCAT Genomes For Life
Analysis team leader
Rafael de Cid399
Data collection leaders
Anna Carreras399, Victor Moreno400 & Manolis Kogevinas335,336,337,338
Analysis team members
Iván Galván-Femenía399, Natalia Blay399, Xavier Farré399 & Lauro Sumoy399
Data collection members
Beatriz Cortés399, Josep Maria Mercader401,1276,1277,1278,1279, Marta Guindo-Martinez401,
David Torrents401, Judith Garcia-Aymerich335,336,337, Gemma Castaño-Vinyals335,336,337,338 &
Carlota Dobaño335,336
GEN-COVID Multicenter Study
Analysis team leaders
Marco Gori404,405 & Mari E. K. Niemi1,1280
Data collection leaders
Alessandra Renieri63,64,65 , Francesca Mari63,64 ,65, Mario Umberto Mondelli408,409,
Francesco Castelli410, Massimo Vaghi411, Stefano Rusconi412,413, Francesca Montagnani65,414 ,
Elena Bargagli415, Federico Franchi416, Maria Antonietta Mazzei417, Luca Cantarini418,
Danilo Tacconi419, Marco Feri420, Raffaele Scala421, Genni Spargi422, Cesira Nencioni423,
Maria Bandini424, Gian Piero Caldarelli425, Maurizio Spagnesi424, Anna Canaccini426 ,
Agostino Ognibene427, Antonella D’Arminio Monforte428, Massimo Girardis429,
Andrea Antinori430, Daniela Francisci431,432, Elisabetta Schiaroli431,432, Pier Giorgio Scotton433,
Sandro Panese434, Renzo Scaggiante435, Matteo Della Monica436, Mario Capasso437,438,439,
Giuseppe Fiorentino440, Marco Castori441, Filippo Aucella442, Antonio Di Biagio443,
Luca Masucci444,445, Seraina Valente446, Marco Mandalà447, Patrizia Zucchi448,
Ferdinando Giannattasio449, Domenico A. Coviello12,450, Cristina Mussini451, Giancarlo Bosio452,
Luisa Tavecchia453, Lia Crotti454,455,456,457, Marco Rizzi458, Maria Teresa La Rovere459,
Simona Sarzi-Braga460, Maurizio Bussotti461, Sabrina Ravaglia462, Rosangela Artuso463,
Antonio Perrella464, Davide Romani465, Paola Bergomi466, Emanuele Catena466,
Antonella Vincenti1231, Claudio Ferri1232, Davide Grassi1232, Gloria Pessina1233,
Mario Tumbarello65,414, Massimo Di Pietro1234, Ravaglia Sabrina462, Sauro Luchi1235,
Chiara Barbieri1235, Donatella Acquilini1235, Elena Andreucci1235, Francesco Paciosi1235,
Francesco Vladimiro Segala1238, Giusy Tiseo1236, Marco Falcone1236, Mirjam Lista63,65,
Monica Poscente1235, Oreste De Vivo446, Paola Petrocelli1235, Alessandra Guarnaccia444,445,
Silvia Baroni1239 & Valentina Perticaroli63,64,65
Admin team leaders
Simone Furini65 & Simona Dei467
Analysis team members
Elisa Benetti65, Nicola Picchiotti404,468, Maurizio Sanarico469, Stefano Ceri66, Pietro Pinoli66,
Francesco Raimondi470, Filippo Biscarini471, Alessandra Stella471, Mattia Bergomi473,
Kristina Zguro65, Katia Capitani65,475, Mattia Cordioli1, Sara Pigazzini1, Lindokuhle Nkambule3,29,167
& Marco Tanfoni468
Data collection members
Chiara Fallerini63,65, Sergio Daga63,65, Margherita Baldassarri63,65, Francesca Fava63,64,65,
Elisa Frullanti63,65, Floriana Valentino63,65, Gabriella Doddato63,65, Annarita Giliberti63,65,
Rossella Tita64, Sara Amitrano64, Mirella Bruttini63,6 5,64, Susanna Croci63,65, Ilaria Meloni63,65,
Maria Antonietta Mencarelli64, Caterina Lo Rizzo64, Anna Maria Pinto64, Giada Beligni63,65,
Andrea Tommasi63,64,65, Laura Di Sarno63,65, Maria Palmieri63,65, Miriam Lucia Carriero63,65,
Diana Alaverdian63,65, Nicola Iuso63,65, Gabriele Inchingolo63,65, Stefano Busani429,
Raffaele Bruno408,409, Marco Vecchia478, Mary Ann Belli453, Stefania Mantovani478,
Serena Ludovisi408,409, Eugenia Quiros-Roldan410, Melania Degli Antoni410, Isabella Zanella479,480,
Matteo Siano413, Arianna Emiliozzi430, Massimiliano Fabbiani414, Barbara Rossetti414,
Giacomo Zanelli65,414, Laura Bergantini415, Miriana D’Alessandro415, Paolo Cameli415,
David Bennet415, Federico Anedda416, Simona Marcantonio416, Sabino Scolletta416,
Susanna Guerrini417, Edoardo Conticini418, Bruno Frediani418, Chiara Spertilli419, Alice Donati420,
Luca Guidelli421, Marta Corridi422, Leonardo Croci423, Paolo Piacentini424, Elena Desanctis424,
Silvia Cappelli424, Agnese Verzuri426, Valentina Anemoli426, Alessandro Pancrazi427,
Maria Lorubbio427, Esther Merlini428, Federica Gaia Miraglia428, Sophie Venturelli429,
Andrea Cossarizza481, Alessandra Vergori430, Arianna Gabrieli413, Agostino Riva412,413,
Francesco Paciosi432, Francesca Andretta433, Francesca Gatti435, Saverio Giuseppe Parisi482,
Stefano Baratti482, Carmelo Piscopo436, Roberta Russo437,438, Immacolata Andolfo437,438,
Achille Iolascon437,438, Massimo Carella441, Giuseppe Merla437,483, Gabriella Maria Squeo483,
Pamela Raggi484, Carmen Marciano484, Rita Perna484, Matteo Bassetti443,485,
Maurizio Sanguinetti444,445, Alessia Giorli447, Lorenzo Salerni447, Pierpaolo Parravicini448,
Elisabetta Menatti486, Tullio Trotta449, Gabriella Coiro449, Fabio Lena487, Enrico Martinelli452,
Sandro Mancarella453, Chiara Gabbi488, Franco Maggiolo458, Diego Ripamonti458,
Tiziana Bachetti489, Claudia Suardi490, Gianfranco Parati454,455, Giordano Bottà491, Paolo Di
Domenico491, Ilaria Rancan414, Francesco Bianchi65,4 64, Riccardo Colombo466, Chiara Barbieri1235,
Donatella Acquilini1235, Elena Andreucci1235, Francesco Paciosi1235,
Francesco Vladimiro Segala1238, Giusy Tiseo1236, Marco Falcone1236, Mirjam Lista63,65,
Monica Poscente1235, Oreste De Vivo446, Paola Petrocelli1235, Alessandra Guarnaccia444,445,
Silvia Baroni1239 & Valentina Perticaroli63,64,65
Genes & Health
Analysis team leader
David A. van Heel50
Data collection leader
Karen A. Hunt50
Admin team leader
Richard C. Trembath493
Analysis team members
Qin Qin Huang494 & Hilary C. Martin494
Data collection members
Dan Mason495, Bhavi Trivedi50 & John Wright495
Admin team members
Sarah Finer497, Genes & Health Research Team* & Christopher J. Grifiths499
Genes & Health Research Team
Shaheen Akhtar498, Mohammad Anwar498, Elena Arciero498, Samina Ashraf498 , Gerome Breen498,
Raymond Chung498, Charles J. Curtis498, Maharun Chowdhury498, Grainne Colligan498,
Panos Deloukas498, Ceri Durham498, Sarah Finer498, Chris Grifiths498, Qin Qin Huang498,
Matt Hurles498, Karen A. Hunt498, Shapna Hussain498, Kamrul Islam498, Ahsan Khan498,
Amara Khan498, Cath Lavery498, Sang Hyuck Lee498, Robin Lerner498, Daniel MacArthur498,
Bev MacLaughlin498, Hilary Martin498, Dan Mason498, Shefa Miah498, Bill Newman498,
Nishat Safa498, Farah Tahmasebi498, Richard C. Trembath498, Bhavi Trivedi498, David A. van
Heel498 & John Wright498
Genes for Good
Analysis team leader
Albert V. Smith500
Data collection members
Andrew P. Boughton500, Kevin W. Li500, Jonathon LeFaive500 & Aubrey Annis500
Genetic determinants of COVID-19 complications in the Brazilian population
Analysis team leader
Mari E. K. Niemi1,1280
Data collection leader
Cinthia E. Jannes501
Admin team leaders
Jose E. Krieger501 & Alexandre C. Pereira501
Analysis team members
Mariliza Velho501, Emanuelle Marques501, Mattia Cordioli1, Sara Pigazzini1 &
Lindokuhle Nkambule3,29,167

Article
Data collection members
Isabella Ramos Lima501, Mauricio Teruo Tada501 & Karina Valino501
Genetic inluences on severity of COVID-19 illness in Korea
Analysis team leaders
Mark McCarthy502 & Carrie Rosenberger502
Data collection leader
Jong Eun Lee503
Analysis team members
Diana Chang502, Christian Hammer502, Julie Hunkapiller502, Anubha Mahajan502,
Sarah Pendergrass502, Lara Sucheston-Campbell502 & Brian Yaspan502
Data collection members
Hyun Soo Lee503, Eunsoon Shin503, Hye Yoon Jang503, Sunmie Kim504, Sungmin Kym505,
Yeon-Sook Kim505, Hyeongseok Jeong505, Ki Tae Kwon507, Shin-Woo Kim507, Jin Yong Kim508,
Young Rock Jang508, Hyun ah Kim509, Ji Yeon Lee509, Jeong Eun Lee510, Shinwon Lee510,
Kang-Won Choe511, Yu Min Kang511, Sun Ha Jee512 & Keum Ji Jung512
Genomic epidemiology of SARS-CoV-2host genetics in coronavirus disease 2019
Data collection leaders
Victoria Parikh513, Euan Ashley514,515, Matthew Wheeler513, Manuel Rivas516,
Carlos Bustamante515,516, Benjamin Pinksy518, Phillip Febbo519, Kyle Farh519, Gary P. Schroth519 &
Francis deSouza519
Admin team leaders
Karen Dalton513 & Jeff Christle513
Analysis team members
Christopher Deboever520, Sándor Szalma520, Yosuke Tanigawa516, Simone Rubinacci521 &
Olivier Delaneau521
Data collection members
John Gorzynski513, Hannah de Jong513, Shirley Sutton513, Nathan Youlton513, Ruchi Joshi513,
David Jimenez-Morales513, Christopher Hughes513, David Amar513, Alex Ioannidis516,
Steve Hershman513, Anna Kirillova513, Kinya Seo513, Yong Huang513, Massa Shoura518,
Nathan Hammond518, Nathaniel Watson518, Archana Raja513, ChunHong Huang518,
Malaya Sahoo518 & Hannah Wang518
Admin team member
Jimmy Zhen513
Genotek COVID-19 study
Analysis team leader
Alexander Rakitko974
Admin team leader
Valery Ilinsky974
Analysis team members
Danat Yermakovich974, Iaroslav Popov974, Alexander Chernitsov974, Elena Kovalenko974,
Anna Krasnenko974, Nikolay Plotnikov974, Ivan Stetsenko974 & Anna Kim974
Helix & Healthy Nevada Project Exome+ COVID-19 Phenotypes
Analysis team leader
Elizabeth T. Cirulli975
Analysis team members
Kelly M. Schiabor Barrett975, Alexandre Bolze975, Simon White975, Nicole L. Washington975 &
James T. Lu975
Data collection members
Stephen Rifle975, Francisco Tanudjaja975, Xueqing Wang975, Jimmy M. Ramirez III975,
Nicole Leonetti975, Efren Sandoval975, Iva Neveux976 , Shaun Dabe977 & Joseph J. Grzymski976
24Genetics & IdiPaz Genomic Variants associated to COVID-19 infection outcome
Analysis team leader
Juan Ignacio Esteban Miñano978
Data collection leader
Luis A. Aguirre979
Admin team leader
Eduardo López-Collazo979
Analysis team members
Manuel de la Mata Pazos978, Luciano Cerrato978 & Lasse Folkersen978
Data collection members
Roberto Lozano-Rodríguez979, José Avendaño-Ortiz979, Verónica Terrón Arcos979,
Karla Marina Montalbán-Hernández979, Jaime Valentín Quiroga979 &
Alejandro Pascual-Iglesias979
Admin team members
Charbel Maroun-Eid979 & Alejandro Martín-Quirós979
Japan Coronavirus Taskforce
Analysis team leaders
Ho Namkoong980, Yukinori Okada981,982,983 & Seiya Imoto984
Data collection leaders
Kazuhiko Katayama985, Koichi Fukunaga980, Yuko Kitagawa986, Toshiro Sato987,
Naoki Hasegawa988, Atsushi Kumanogoh983,989,990, Akinori Kimura991, Masumi Ai992 &
Katsushi Tokunaga993
Admin team leaders
Takanori Kanai994, Satoru Miyano995 & Seishi Ogawa996,997
Analysis team members
Ryuya Edahiro981,989, Kyuto Sonehara981, Yuya Shirai981,989 & Masahiro Kanai274
Data collection members
Makoto Ishii980, Hiroki Kabata980, Katsunori Masaki980, Hirofumi Kamata980,
Shinnosuke Ikemura980, Shotaro Chubachi980, Satoshi Okamori980, Hideki Terai980,
Hiromu Tanaka980, Atsuho Morita980, Ho Lee980, Takanori Asakura980, Junichi Sasaki999,
Hiroshi Morisaki1000, Yoshifumi Uwamino1001, Kosaku Nanki994, Yohei Mikami994,
Kazunori Tomono1002, Kazuto Kato1003, Fumihiko Matsuda1004, Meiko Takahashi1004,
Nobuyuki Hizawa1005, Yoshito Takeda989, Haruhiko Hirata989, Takayuki Shiroyama989,
Satoru Miyawaki1006, Ken Suzuki981, Yuichi Maeda989,1007, Takuro Nii989,1007, Yoshimi Noda989,
Takayuki Niitsu989, Yuichi Adachi989, Takatoshi Enomoto989, Saori Amiya989, Reina Hara989,
Kunihiko Takahashi995, Tatsuhiko Anzai995, Takanori Hasegawa995, Satoshi Ito995, Ryuji Koike1009,
Akifumi Endo1010, Yuji Uchimura1011, Yasunari Miyazaki1012, Takayuki Honda1012,
Tomoya Tateishi1012, Shuji Tohda1013, Naoya Ichimura1013, Kazunari Sonobe1013, Chihiro Sassa1013,
Jun Nakajima1013, Yasuhito Nannya1014, Yosuke Omae993, Kazuhisa Takahashi1015,
Norihiro Harada1015, Makoto Hiki1016,1017, Haruhi Takagi1015, Ai Nakamura1015, Etsuko Tagaya1018,
Masatoshi Kawana1019, Ken Arimura1018, Takashi Ishiguro1020, Noboru Takayanagi1020,
Taisuke Isono1020, Yotaro Takaku1020, Kenji Takano1020, Ryusuke Anan1021, Yukiko Nakajima1021,
Yasushi Nakano1021, Kazumi Nishio1021, Soichiro Ueda1022, Reina Hayashi1022, Hiroki Tateno1023,
Isano Hase1023, Shuichi Yoshida1023, Shoji Suzuki1023, Keiko Mitamura1024, Fumitake Saito1025,
Tetsuya Ueda1026, Masanori Azuma1026, Tadao Nagasaki1026, Yoshinori Yasui1028,
Yoshinori Hasegawa1026, Yoshikazu Mutoh1029, Takashi Yoshiyama1030, Tomohisa Shoko1031,
Mitsuaki Kojima1031, Tomohiro Adachi1031, Motonao Ishikawa1032, Kenichiro Takahashi1033,
Kazuyoshi Watanabe1034, Tadashi Manabe1035, Fumimaro Ito1035, Takahiro Fukui1035,
Yohei Funatsu1035, Hidefumi Koh1035, Yoshihiro Hirai1036, Hidetoshi Kawashima1036,
Atsuya Narita1036, Kazuki Niwa1037, Yoshiyuki Sekikawa1037, Fukuki Saito1038, Kazuhisa Yoshiya1038,
Tomoyuki Yoshihara1038, Yusuke Suzuki1039, Sohei Nakayama1039, Keita Masuzawa1039,
Koichi Nishi1040, Masaru Nishitsuji1040, Maiko Tani1040, Takashi Inoue1041, Toshiyuki Hirano1041,
Keigo Kobayashi1041, Naoki Miyazawa1042, Yasuhiro Kimura1042, Reiko Sado1042, Takashi Ogura1043,
Hideya Kitamura1043, Kota Murohashi1043, Ichiro Nakachi1044, Rie Baba1044, Daisuke Arai1044,
Satoshi Fuke1045, Hiroshi Saito1045, Naota Kuwahara1046, Akiko Fujiwara1046, Takenori Okada1046,
Tomoya Baba1047, Junya Noda1047, Shuko Mashimo1047, Kazuma Yagi1048, Tetsuya Shiomi1048,
Mizuha Hashiguchi1048, Toshio Odani1049, Takao Mochimaru1050,1051, Yoshitaka Oyamada1050,1051,
Nobuaki Mori1052, Namiki Izumi1053, Kaoru Nagata1053, Reiko Taki1053, Koji Murakami1054,
Mitsuhiro Yamada1054, Hisatoshi Sugiura1054, Kentaro Hayashi1055, Tetsuo Shimizu1055,
Yasuhiro Gon1055, Shigeki Fujitani1056, Tomoya Tsuchida1057, Toru Yoshida1056, Takashi Kagaya1058,
Toshiyuki Kita1058, Satoru Sakagami1058, Yoshifumi Kimizuka1059, Akihiko Kawana1059,
Yoshihiko Nakamura1060, Hiroyasu Ishikura1060, Tohru Takata1061, Takahide Kikuchi1062,
Daisuke Taniyama1062, Morio Nakamura1062, Nobuhiro Kodama1063, Yasunari Kaneyama1063,
Shunsuke Maeda1063, Yoji Nagasaki1064, Masaki Okamoto1065,1027, Sayoko Ishihara1064,
Akihiro Ito1067, Yusuke Chihara1068, Mayumi Takeuchi1068, Keisuke Onoi1068,
Naozumi Hashimoto1069, Keiko Wakahara1069, Akira Ando1069, Makoto Masuda1070,
Aya Wakabayashi1070, Hiroki Watanabe1070, Hisako Sageshima1071, Taka-Aki Nakada1072,
Ryuzo Abe1072, Tadanaga Shimada1072, Kodai Kawamura1073, Kazuya Ichikado1073,
Kenta Nishiyama1073, Masaki Yamasaki1074, Satoru Hashimoto1074, Yu Kusaka1075,
Takehiko Ohba1075, Susumu Isogai1075, Minoru Takada1076, Hidenori Kanda1076, Yuko Komase1077,
Fumiaki Sano1078, Koichiro Asano1079, Tsuyoshi Oguma1080, Masahiro Harada1081,
Takeshi Takahashi1081, Takayuki Shibusawa1081, Shinji Abe1082, Yuta Kono1082, Yuki Togashi1082,
Takehiro Izumo1083, Minoru Inomata1083, Nobuyasu Awano1083, Shinichi Ogawa1084,
Tomouki Ogata1084, Shoichiro Ishihara1084, Arihiko Kanehiro1085, Shinji Ozaki1085,
Yasuko Fuchimoto1085, Yuichiro Kitagawa1086, Shozo Yoshida1086, Shinji Ogura1086,
Kei Nishiyama1087, Kousuke Yoshida1088, Satoru Beppu1088, Satoru Fukuyama1089,
Yoshihiro Eriguchi1090, Akiko Yonekawa1090, Yoshiaki Inoue1091, Kunihiro Yamagata1092,
Shigeru Chiba1093, Osamu Narumoto1094, Hideaki Nagai1 094, Nobuharu Ooshima1094,
Mitsuru Motegi1095, Hironori Sagara1096, Akihiko Tanaka1096, Shin Ohta1096, Yoko Shibata1097,
Yoshinori Tanino1097, Yuki Sato10 97, Yuichiro Yamada1098, Takuya Hashino1098, Masato Shinoki1098,
Hajime Iwagoe1099, Tomonori Imamura1100, Akira Umeda1101, Hisato Shimada1101, Mayu Endo1102,
Shinichi Hayashi1103, Mai Takahashi1103, Shigefumi Nakano1103, Masakiyo Yatomi1104,
Toshitaka Maeno1104, Tomoo Ishii1105, Mitsuyoshi Utsugi1106, Akihiro Ono1106, Kensuke Kanaoka1107,
Shoichi Ihara1107 & Kiyoshi Komuta1107
Lifelines
Analysis team leader
Lude Franke51

Data collection leader
Marike Boezen1109
Analysis team members
Patrick Deelen51,52, Annique Claringbould51, Esteban Lopera51, Robert Warmerdam51,
Judith. M. Vonk1109 & Irene van Blokland51
Data collection members
Pauline Lanting51 & Anil P. S. Ori1112,1113
Lung eQTL Consortium
Data collection members
Ma’en Obeidat1114, Ana I. Hernández Cordero1114, Don D. Sin1114,1115, Yohan Bossé1116,
Philippe Joubert1116, Ke Hao1117, David Nickle1118,1119, Wim Timens1120,1121 &
Maarten van den Berge1121,1122
Mass General Brigham-Host Vulnerability to COVID-19
Analysis team leaders
Yen-Chen Anne Feng1123 & Josep Mercader29,1123
Data collection leaders
Scott T. Weiss1126, Elizabeth W. Karlson1127, Jordan W. Smoller1128, Shawn N. Murphy1129,
James B. Meigs1130,1124,1125 & Ann E. Woolley1127
Admin team leader
Robert C. Green2,273
Data collection member
Emma F. Perez273
Michigan Genomics Initiative (MGI)
Analysis team leader
Brooke Wolford1132
Admin team leader
Sebastian Zöllner500
Analysis team members
Jiongming Wang500 & Andrew Beck500
Mount Sinai Health System COVID-19 Genomics Initiative
Analysis team leader
Laura G. Sloofman26
Data collection leaders
Steven Ascolillo1133, Robert P. Sebra1117,1135, Brett L. Collins26 & Tess Levy26
Admin team leaders
Joseph D. Buxbaum26 & Stuart C. Sealfon7
Analysis team members
Shea J. Andrews7, Daniel M. Jordan1117,1137, Ryan C. Thompson1133,1140,1141, Kyle Gettler1117,
Kumardeep Chaudhary1117,1143, Gillian M. Belbin1144, Michael Preuss1143,1146, Clive Hoggart1147,1148,1142,
Sam Choi1147,1148,1142,1149 & Slayton J. Underwood26
Data collection members
Irene Salib1117, Bari Britvan26, Katherine Keller26, Lara Tang26, Michael Peruggia26,
Liam L. Hiester26, Kristi Niblo26, Alexandra Aksentijevich26, Alexander Labkowsky26,
Avromie Karp26, Menachem Zlatopolsky26 & Marissa Zyndorf1117
Admin team members
Alexander W. Charney1141,1150, Noam D. Beckmann1133, Eric E. Schadt1117,1135,
Noura S. Abul-Husn1144, Judy H. Cho1117,1143, Yuval Itan1117,1143, Eimear E. Kenny1144,
Ruth J. F. Loos1143,1146,1151, Girish N. Nadkarni1133,1143,1153,1154,1155, Ron Do1117,1143, Paul O’Reilly1147,1148,1142,1149
& Laura M. Huckins1147,1148,1142
MyCode Health Initiative
Analysis team leaders
Manuel A. R. Ferreira1157 & Goncalo R. Abecasis1157
Data collection leaders
Joseph B. Leader1158 & Michael N. Cantor1157
Admin team leaders
Anne E. Justice1159 & Dave J. Carey1160
Analysis team members
Geetha Chittoor1159, Navya Shilpa Josyula1159, Jack A. Kosmicki1157, Julie E. Horowitz1157 &
Aris Baras1157
Data collection members
Matthew C. Gass1158 & Ashish Yadav1157
Admin team member
Tooraj Mirshahi1160
Netherlands Twin Register
Analysis team leader
Jouke Jan Hottenga122
Data collection leader
Meike Bartels122
Admin team leader
Eco J. C. de Geus122
Analysis team member
Michel G. Nivard122
Penn Medicine Biobank
Analysis team leaders
Anurag Verma1162 & Marylyn D. Ritchie1162
Admin team leader
Daniel Rader1162
Analysis team members
Binglan Li1163, Shefali S. Verma1162, Anastasia Lucas1162 & Yuki Bradford1162
Population controls
Analysis team leader
Federico Zara12
Analysis team members
Vincenzo Salpietro12, Marcello Scala1172, Michele Iacomino12, Paolo Scudieri12 &
Renata Bocciardi12
Data collection members
Carlo Minetti12, Antonella Riva1172, Maria Stella Vari12, Myriam Mni163, Jean-François Rahier1173,
Elisa Giorgio1174, Federico Zara12 & Diana Carli1175
Data collection leaders
Pasquale Striano12,13, Edouad Louis170, Michel Georges163, Souad Rahmouni163,
Cynthia M. Bulik709,1166,1167, Mikael Landén709,1168, Alfredo Brusco1169 & Giovanni Battista Ferrero1170
Admin team leaders
Francesca Madia12 & Bengt Fundín709
Qatar Genome Program
Analysis team leader
Hamdi Mbarek80
Data collection leader
Said I. Ismail80
Analysis team members
Chadi Saad80 & Yaser Al-Sarraj80
Data collection members
Radja Messai Badji80, Wadha Al-Muftah80, Asma Al Thani80 & Nahla Aii1176
Study of the COVID-19 host genetics in the population of Latvia
Analysis team leader
Janis Klovins1177
Data collection leader
Vita Rovite1177
Analysis team members
Raimonds Rescenko1177 & Raitis Peculis1177
Data collection member
Monta Ustinova1177
The genetic predisposition to severe COVID-19
Analysis team leader
Mari E. K. Niemi1,1280
Data collection leader
Hugo Zeberg1178,1179

Article
Analysis team members
Mattia Cordioli1, Sara Pigazzini1 & Lindokuhle Nkambule3,29,167
Data collection members
Robert Frithiof1180, Michael Hultström1180,1181 & Miklos Lipcsey1180,1182
UCLA Precision Health COVID-19 Host Genomics Biobank
Analysis team leader
Ruth Johnson1183
Data collection leader
UCLA Health ATLAS & Data Mart Working Group*
UCLA Health ATLAS & Data Mart Working Group
Daniel H. Geschwind1184
Admin team leaders
Nelson Freimer1185, Manish J. Butte1186,1171,1184, Daniel H. Geschwind1188,1187,1152 &
Bogdan Pasaniuc1189,1190,1139
Analysis team members
Yi Ding1191, Alec Chiu1191, Timothy S. Chang1192 & Paul Boutros1193,1139
UK 100,000 Genomes Project (Genomics England)
Analysis team leader
Loukas Moutsianas14,95
Data collection leaders
Mark J. Caulield95,695 & Richard H. Scott95,1195,1196
Analysis team members
Athanasios Kousathanas95, Dorota Pasko95, Susan Walker95, Alex Stuckey95,
Christopher A. Odhams95 & Daniel Rhodes95
Data collection members
Tom Fowler95, Augusto Rendon95,1197, Georgia Chan95 & Prabhu Arumugam95
UK Biobank
Analysis team leaders
Tomoko Nakanishi17,56,57, Konrad J. Karczewski3,29, Alicia R. Martin3,29, Daniel J. Wilson1199 &
Chris A. Spencer91
Data collection leaders
Derrick W. Crook1201, David H. Wyllie1201,1202 & Anne Marie O’Connell1203
Admin team leader
J. Brent Richards17,53,55,56
Analysis team members
Guillaume Butler-Laporte17,53, Vincenzo Forgetta17, Elizabeth G. Atkinson3,29,
Masahiro Kanai3,29,1204, Kristin Tsuo3,29,1205, Nikolas Baya3,29, Patrick Turley3,29, Rahul Gupta3,29,
Raymond K. Walters3,29, Duncan S. Palmer3,29, Gopal Sarma3,29, Matthew Solomonson3,29,
Nathan Cheng3,29, Wenhan Lu3,29, Claire Churchhouse3,29, Jacqueline I. Goldstein3,29,
Daniel King3,29, Wei Zhou3,29, Cotton Seed3,29, Mark J. Daly1,2,3, Benjamin M. Neale3,29,
Hilary Finucane3,29, Sam Bryant2, F. Kyle Satterstrom3,29, Gavin Band700, Sarah G. Earle1199,
Shang-Kuan Lin1199, Nicolas Arning1199 & Nils Koelling91
Data collection members
Jacob Armstrong1199 & Justine K. Rudkin1199
Admin team members
Shawneequa Callier1207, Sam Bryant3,29 & Caroline Cusick29
UK Blood Donors Cohort
Analysis team leaders
Nicole Soranzo1208,1209,1210 & Jing Hua Zhao1211
Data collection leaders
John Danesh1211,1212,1213,1214,1215 & Emanuele Di Angelantonio1211,1212,1213,1214
Analysis team member
Adam S. Butterworth1211,1212,1213,1214
VA Million Veteran Program (MVP)
Analysis team leaders
Yan V. Sun1216,1217 & Jennifer E. Huffman1218
Data collection leader
Kelly Cho1219
Admin team leaders
Christopher J. O’Donnell1218, Phil Tsao1220,1221 & J. Michael Gaziano1219
Analysis team member
Gina Peloso1218,1222
Data collection member
Yuk-Lam Ho1219
Val Gardena
Analysis team leader
Christian Fuchsberger209
Data collection leader
Michael Mian1223
Data collection member
Federica Scaggiante1224
Admin team members
Cristian Pattaro209 & Peter Pramstaller209
CHOP_CAG
Xiao Chang1227, Joseph R. Glessner1227,1228 & Hakon Hakonarson1227,1228,1229
GenOMICC/ISARIC4C
Data collection leaders
J. Kenneth Baillie60,61,62, Peter J. McGuigan523, Luke Stephen Prockter Moore524,
Marcela Paola Vizcaychipi524, Kathryn Hall525, Andy Campbell526, Ailstair Nichol527,
Geraldine Ward528, Valerie Joan Page529, Malcolm G. Semple530, Kayode Adeniji531,
Daniel Agranoff532, Ken Agwuh533, Dhiraj Ail534, Erin L. Aldera535, Ana Alegria536,506,
Brian Angus537, Abdul Ashish538, Dougal Atkinson539, Shahedal Bari540, Gavin Barlow541,
Stella Barnass542, Nicholas Barrett543, Christopher Bassford544, Sneha Basude54 5,
David Baxter546, Michael Beadsworth547, Jolanta Bernatoniene548, John Berridge549,
Nicola Best550, Pieter Bothma551, David Chadwick552, Robin Brittain-Long553, Naomi Bulteel554,
Tom Burden555, Andrew Burtenshaw556, Vikki Caruth557, David Chadwick552,
Duncan Chambler558, Nigel Chee559, Jenny Child560, Srikanth Chukkambotla56 1, Tom Clark562,
Paul Collini563, Catherine Cosgrove564, Jason Cupitt565, Maria-Teresa Cutino-Moguel566,
Paul Dark567, Chris Dawson568, Samir Dervisevic569, Phil Donnison837, Sam Douthwaite543,
Andrew Drummond572,773, Ingrid DuRand573, Ahilanadan Dushianthan574 , Tristan Dyer575,
Cariad Evans563, Chi Eziefula532, Christopher Fegan576, Adam Finn57 7, Duncan Fullerton578,
Sanjeev Garg579, Atul Garg580, Effrossyni Gkrania-Klotsas581, Jo Godden582, Arthur Goldsmith583,
Clive Graham584, Elaine Hardy585, Stuart Hartshorn586, Daniel Harvey587, Peter Havalda588,
Daniel B. Hawcutt589, Maria Hobrok590, Luke Hodgson591, Anil Hormis592, Michael Jacobs593,
Susan Jain594, Paul Jennings595, Agilan Kaliappan596, Vidya Kasipandian5 97, Stephen Kegg598,
Michael Kelsey599, Jason Kendall1281, Caroline Kerrison600, Ian Kerslake601, Oliver Koch602,
Gouri Koduri603, George Koshy604, Shondipon Laha605, Steven Laird606, Susan Larkin607,
Tamas Leiner604, Patrick Lillie608, James Limb609, Vanessa Linnett610, Jeff Little611, Mark Lyttle612,
Michael MacMahon1281, Emily MacNaughton613, Ravish Mankregod614, Huw Masson615,
Elijah Matovu578, Katherine McCullough616, Ruth McEwen617, Manjula Meda618, Gary H. Mills563,
Jane Minton620, Karl Ward620, Mariyam Mirfenderesky621, Kavya Mohandas622, Quen Mok623,
James Moon624, Elinoor Moore581, Patrick Morgan625, Craig Morris626, Katherine Mortimore604,
Samuel Moses627, Mbiye Mpenge628, Rohinton Mulla629, Michael Murphy630, Megan Nagel631,
Thapas Nagarajan632, Mark Nelson633, Matthew K. O’Shea634, Igor Otahal635,
Marlies Ostermann543, Mark Pais636, Selva Panchatsharam637, Danai Papakonstantinou638,
Hassan Paraiso639, Brij Patel640, Natalie Pattison641, Justin Pepperell642, Mark Peters1281,
Mandeep Phull643, Stefania Pintus644, Jagtur Singh Pooni645, Frank Post646, David Price647,
Rachel Prout648, Nikolas Rae649, Henrik Reschreiter650, Tim Reynolds651, Neil Richardson652,
Mark Roberts653, Devender Roberts654, Alistair Rose655, Guy Rousseau656, Brendan Ryan657,
Taranprit Saluja658, Aarti Shah659, Prad Shanmuga660, Anil Sharma661, Anna Shawcross662,
Jeremy Sizer663, Manu Shankar-Hari543, Richard Smith664, Catherine Snelson665 , Nick Spittle666,
Nikki Staines667, Tom Stambach668, Richard Stewart669, Pradeep Subudhi670,
Tamas Szakmany671, Kate Tatham672, Jo Thomas673, Chris Thompson674, Robert Thompson1281,
Ascanio Tridente675, Darell Tupper-Carey551, Mary Twagira676, Andrew Ustianowski572,
Nick Vallotton677, Lisa Vincent-Smith678, Shico Visuvanathan667, Alan Vuylsteke679,
Sam Waddy680, Rachel Wake681, Andrew Walden682, Ingeborg Welters547, Tony Whitehouse665,
Paul Whittaker683, Ashley Whittington684, Padmasayee Papineni685, Meme Wijesinghe686,
Martin Williams1281, Lawrence Wilson617, Sarah Cole966, Stephen Winchester687,
Martin Wiselka688, Adam Wolverson689, Daniel G. Wooton690, Andrew Workman588,
Bryan Yates691 & Peter Young692
Analysis team members
J. Kenneth Baillie60,61,62, Rupert Beale693, Andrew D. Bretherick62, Mark J. Caulield95,695,
Sara Clohisey60, Max Head Fourman60, James Furniss60, Elvina Gountouna696,
Graeme Grimes62, Chris Haley60, David Harrison697, Caroline Hayward62,696, Sean Keating61,
Lucija Klaric62, Paul Klenerman700, Athanasios Kousathanas95, Andy Law60, Alison M. Meynert62,
Jonathan Millar60, Loukas Moutsianas14,95, Erola Pairo-Castineira60,6 2, Nicholas Parkinson60,
Dorota Pasko95, Chris P. Ponting62, David J. Porteous696, Konrad Rawlik60, Anne Richmond62,
Kathy Rowan697, Clark D. Russell60,705, Richard H. Scott95,706, Xia Shen707,708,709, Barbara Shih60,
Albert Tenesa60,62,708, Veronique Vitart62, Susan Walker95, Bo Wang60, James F. Wilson62,708,
Yang Wu710, Jian Yang711,712, Zhijian Yang707, Marie Zechner60, Ranran Zhai7 07, Chenqing Zheng707,
Lisa Norman714, Riinu Pius714, Thomas M. Drake714, Cameron J. Fairield714, Stephen R. Knight714,
Kenneth A. Mclean714, Derek Murphy714, Catherine A. Shaw714, Jo Dalton715, Michelle Girvan715,
Egle Saviciute715, Stephanie Roberts715, Janet Harrison715, Laura Marsh715, Marie Connor715,

Sophie Halpin715, Clare Jackson715, Carrol Gamble715, Gary Leeming716, Andrew Law60,
Murray Wham717, Sara Clohisey60, Ross Hendry60 & James Scott-Brown718
Data collection members
Colin Begg719, Sara Clohisey60, Charles Hinds695, Antonia Ying Wai Ho721, Peter W. Horby722,
Julian Knight700, Lowell Ling724, David Maslove725, Danny McAuley726,727, Jonathan Millar60,
Hugh Montgomery728, Alistair Nichol729, Peter J. M. Openshaw730,731, Chris P. Ponting62,
Kathy Rowan697, Malcolm G. Semple732,733, Manu Shankar-Hari734, Charlotte Summers735,
Timothy Walsh61, Lisa Armstrong736, Hayley Bates736, Emma Dooks736, Fiona Farquhar736,
Brigid Hairsine736, C. McParland736, Sophie Packham736, Zoe Alldis737, Raine Astin-Chamber
lain737, Fatima Bibi737, Jack Biddle737 , Sarah Blow737, Matthew Bolton737, Catherine Borra737,
Ruth Bowles737, Maudrian Burton737, Yasmin Choudhury737, David Collier7 37, Amber Cox737,
Amy Easthope737, Patrizia Ebano737, Stavros Fotiadis737, Jana Gurasashvili737 , Rosslyn Halls737,
Pippa Hartridge737, Delordson Kallon737, Jamila Kassam7 37, Ivone Lancoma-Malcolm737,
Maninderpal Matharu737, Peter May737, Oliver Mitchelmore737, Tabitha Newman737, Mital Patel737,
Jane Pheby737, Irene Pinzuti737, Zoe Prime737, Oleksandra Prysyazhna737, Julian Shiel737,
Melanie Taylor737, Carey Tierney737, Suzanne Wood737, Anne Zak737, Olivier Zongo737,
Miranda Forsey738, Agilan Kaliappan738, Anne Nicholson738, Joanne Riches738, Mark Vertue738,
Christopher Wasson523, Stephanie Finn523, Jackie Green523, Erin Collins523, Bernadette King523,
Lina Grauslyte739, Musarat Hussain739, Mandeep Phull739, Tatiana Pogreban739, Lace Rosaroso739,
Erika Salciute739, George Franke739, Joanna Wong739, Aparna George739, Louise Akeroyd740,
Shereen Bano740, Matt Bromley74 0, Lucy Gurr740 , Tom Lawton740, James Morgan740 ,
Kirsten Sellick740, Deborah Warren740 , Brian Wilkinson740 , Janet McGowan740,
Camilla Ledgard740, Amelia Stacey74 0, Kate Pye740 , Ruth Bellwood740 , Michael Bentley740,
Maria Hobrok741, Ronda Loosley741 , Heather McGuinness741 , Helen Tench741,
Rebecca Wolf-Roberts741, Sian Gibson742, Amanda Lyle742, Fiona McNeela742,
Jayachandran Radhakrishnan742, Alistair Hughes74 2, Asifa Ali74 3, Megan Brady74 3, Sam Dale743 ,
Annalisa Dance743, Lisa Gledhill74 3, Jill Greig74 3, Kathryn Hanson74 3, Kelly Holdroyd743,
Marie Home743, Diane Kelly74 3, Ross Kitson743 , Lear Matapure743 , Deborah Melia743 ,
Samantha Mellor743, Tonicha Nortcliffe743, Jez Pinnell743, Matthew Robinson743 , Lisa Shaw743,
Ryan Shaw743, Lesley Thomis74 3, Alison Wilson743 , Tracy Wood743, Lee-Ann Bayo743,
Ekta Merwaha743, Tahira Ishaq743 , Sarah Hanley743, David Antcliffe744, Dorota Banach744,
Stephen Brett744, Phoebe Coghlan74 4, Ziortza Fernandez74 4, Anthony Gordon744 , Roceld Rojo744,
Sonia Sousa Arias744, Maie Templeton744 , Rajeev Jha745, Vinodh Krishnamurthy745, Lai Lim745 ,
Rehana Bi746, Barney Scholeield74 6, Lydia Ashton746 , Alison Williams747, Claire Cheyne747,
Anne Saunderson747, Angela Allan74 8, Felicity Anderson748 , Callum Kaye748 , Jade Liew748,
Jasmine Medhora748, Teresa Scott748, Erin Trumper748, Adriana Botello748, Petra Polgarova749,
Katerina Stroud749, Eoghan Meaney74 9, Megan Jones74 9, Anthony Ng74 9, Shruti Agrawal749 ,
Nazima Pathan749, Deborah White74 9, Esther Daubney74 9, Kay Elston74 9, Robert Parker750,
Amie Reddy750, Ian Turner-Bone750, Laura Wilding750, Peter Harding750, Reni Jacob752,
Cathy Jones752, Craig Denmade752, Maria Croft753, Ian White753, Rajeev Jha745,
Vinodh Krishnamurthy745, Li Lim74 5, Denise Grifin754, Nycola Muchenje754,
Mcdonald Mupudzi754, Richard Partridge754, Jo-Anna Conyngham754, Rachel Thomas754 ,
Mary Wright754, Maria Alvarez Corral754, Victoria Bastion663, Daphene Clarke663, Beena David663,
Harriet Kent663, Rachel Lorusso663, Gamu Lubimbi663, Sophie Murdoch663 ,
Melchizedek Penacerrada663, Alastair Thomas663, Jennifer Valentine663, Ana Vochin663,
Retno Wulandari663, Brice Djeugam663, Joy Dawson755, Sweyn Garrioch755, Melanie Tolson755,
Jonathan Aldridge755, Laura Gomes de Almeida Martins524 , Jaime Carungcong524,
Sarah Beavis756, Katie Dale756, Rachel Gascoyne756, Joanne Hawes756, Kelly Pritchard756,
Lesley Stevenson756, Amanda Whileman756, Anne Cowley757, Judith Highgate757,
Rikki Crawley758, Abigail Crew758, Mishell Cunningham758, Allison Daniels758, Laura Harrison758,
Susan Hope758, Ken Inweregbu758, Sian Jones758, Nicola Lancaster758, Jamie Matthews758,
Alice Nicholson758, Gemma Wray758, Leonie Benham759, Zena Bradshaw759, Joanna Brown759,
Melanie Caswell759, Jason Cupitt759, Sarah Melling759, Stephen Preston759, Nicola Slawson759,
Emma Stoddard759, Scott Warden759, Edward Combes760, Teishel Joeield760, Sonja Monnery760,
Valerie Beech760, Sallyanne Trotman760, Bridget Hopkins761, James Scriven761,
Laura Thrasyvoulou761, Heather Willis761, Susan Anderson762, Janine Birch762, Emma Collins762,
Kate Hammerton762, Ryan O’Leary762, Caroline Abernathy763, Louise Foster763, Andrew Gratrix763,
Vicky Martinson763, Priyai Parkinson763, Elizabeth Stones763 , Llucia Carbral-Ortega763,
Ritoo Kapoor765, David Loader765, Karen Castle765, Craig Brandwood766, Lara Smith766,
Richard Clark766, Katie Birchall766, Laurel Kolakaluri766, Deborah Baines766, Anila Sukumaran766,
Isheunesu Mapfunde525, Megan Meredith767, Lucy Morris767, Lucy Ryan767, Amy Clark767,
Julia Sampson767, Cecilia Peters767, Martin Dent767, Margaret Langley767 , Saima Ashraf767,
Shuying Wei767, Angela Andrew767, Manish Chablani768, Amy Kirkby768, Kimberley Nether ton768,
Michelle Bates769, Jo Dasgin769, Jaspret Gill769, Annette Nilsson769, James Scriven769,
Elena Apetri770, Cathrine Basikolo770, Bethan Blackledge770, Laura Catlow770, Bethan Charles770,
Paul Dark770, Reece Doonan770, Jade Harris770, Alice Harvey770, Daniel Horner770,
Karen Knowles770, Stephanie Lee770, Diane Lomas770, Chloe Lyons770, Tracy Marsden770,
Danielle McLaughlan770, Liam McMorrow770, Jessica Pendlebury770, Jane Perez770,
Maria Poulaka770, Nicola Proudfoot770, Melanie Slaughter770, Kathryn Slevin770 , Melanie Taylor770,
Vicky Thomas770, Danielle Walker770, Angiy Michael770, Matthew Collis770 , Martyn Clark771,
Martina Coulding771, Edward Jude771, Jacqueline McCormick771, Oliver Mercer771, Darsh Potla771,
Haiz Rehman771, Heather Savill771, Victoria Turner771, Miriam Davey772, David Golden772,
Rebecca Seaman772, Jodie Hunt773, Joy Dearden773, Emma Dobson773, Andrew Drummond572,773,
Michelle Mulcahy773, Sheila Munt773, Grainne O’Connor773, Jennifer Philbin773, Chloe Rishton773,
Redmond Tully773, Sarah Winnard773, Lenka Cagova774, Adama Fofano774 , Lucie Garner774,
Helen Holcombe774, Sue Mepham7 74, Alice Michael Mitchell774, Lucy Mwaura774, K. Praman774,
Alain Vuylsteke774, Julie Zamikula774 , Miriam Davey772, David Golden772, Rebecca Seaman772,
Georgia Bercades775, David Brealey775, Ingrid Hass775, Niall MacCallum775, Gladys Martir775,
Eamon Raith775, Anna Reyes775, Deborah Smyth775, Abigail Taylor776, Rachel Anne Hughes776,
Helen Thomas776, Alun Rees776, Michaela Duskova776, Janet Phipps7 76, Suzanne Brooks776,
Michelle Edwards776, Peter Alexander777, Schvearn Allen777, Joanne Bradley-Potts777,
Craig Brantwood777, Jasmine Egan777, Timothy Felton777, Grace Padden777, Luke Ward777,
Stuart Moss777, Susannah Glasgow777, Kate Beesley778, Sarah Board778,
Agnieszka Kubisz-Pudelko778, Alison Lewis778, Jess Perry778, Lucy Pippard778, Di Wood778,
Clare Buckley778, Alison Brown779, Jane Gregory779, Susan O’Connell779, Tim Smith779,
Zakaula Belagodu780, Bridget Fuller780, Anca Gherman780, Olumide Olufuwa780,
Remi Paramsothy780, Carmel Stuart780, Naomi Oakley780, Charlotte Kamundi780, David Tyl780,
Katy Collins780, Pedro Silva780, June Taylor780, Laura King780, Charlotte Coates780 ,
Maria Crowley780, Phillipa Wakeield780, Jane Beadle780, Laura Johnson780, Janet Sargeant780,
Madeleine Anderson780, Catherine Jardine781, Dewi Williams781, Victoria Parris782,
Sheena Quaid782, Ekaterina Watson782, Julie Melville783, Jay Naisbitt783, Rosane Joseph783,
Maria Lazo783, Olivia Walton783, Alan Neal783, Michaela Hill784, Thogulava Kannan784,
Laura Wild784, Elizabeth Allan785, Kate Darlington785 , Ffyon Davies785, Jack Easton785,
Sumit Kumar785, Richard Lean785, Daniel Menzies785, Richard Pugh785, Xinyi Qiu785,
Llinos Davies785, Hannah Williams785, Jeremy Scanlon785, Gwyneth Davies785, Callum Mackay785,
Joannne Lewis785, Stephanie Rees785, Samantha Coetzee786, Alistair Gales786, Igor Otahal786,
Meena Raj786, Craig Sell786, Helen Langton787 , Rachel Prout787, Malcolm Watters787,
Catherine Novis787, Gill Arbane788, Aneta Bociek788 , Sara Campos788, Neus Grau788,
Tim Owen Jones788, Rosario Lim788, Martina Marotti788, Marlies Ostermann788,
Manu Shankar-Hari788, Christopher Whitton788, Anthony Barron789, Ciara Collins789 ,
Sundeep Kaul789, Heather Passmore789, Claire Prendergast789, Anna Reed789, Paula Rogers789,
Rajvinder Shokkar789, Meriel Woodruff789, Hayley Middleton789, Oliver Polgar789, Claire Nolan78 9,
Vicky Thwaites789, Kanta Mahay789, Chunda Sri-Chandana790, Joslan Scherewode790,
Lorraine Stephenson790, Sarah Marsh790, Hollie Bancroft638, Mary Bellamy638,
Margaret Carmody638, Jacqueline Daglish638, Faye Moore638, Joanne Rhodes638,
Mirriam Sangombe638, Salma Kadiri638, James Scriven638, Amanda Ayers792, Wendy Harrison792,
Julie North792, Anna Cavazza646, Maeve Cockrell646, Eleanor Corcoran646, Maria Depante646,
Clare Finney646, Ellen Jerome646, Mark McPhail646, Monalisa Nayak646, Harriet Noble646,
Kevin O’Reilly646, Evita Pappa646, Rohit Saha646, Sian Saha646, John Smith646, Abigail Knighton646,
Mandy Gill794, Paul Paul794, Valli Ratnam794, Sarah Shelton794 , Inez Wynter794, David Baptista795,
Rebecca Crowe795, Rita Fernandes795, Rosaleen Herdman-Grant795, Anna Joseph795,
Adam Loveridge795, India McKenley795, Eriko Morino795, Andres Naranjo795, Richard Simms795,
Kathryn Sollesta795, Andrew Swain795, Harish Venkatesh795, Jacyntha Khera795, Jonathan Fox795,
Russell Barber796, Claire Hewitt796, Annette Hilldrith796, Karen Jackson-Lawrence796,
Sarah Shepardson796, Maryanne Wills796, Susan Butler796, Silvia Tavares796, Amy Cunningham796,
Julia Hindale796, Sarwat Arif796, Linsha George797, Sophie Twiss797, David Wright797,
Maureen Holland798, Natalie Keenan798, Marc Lyons798, Helen Wassall798, Chris Marsh798,
Mervin Mahenthran798, Emma Carter798, Thomas Kong798, Oluronke Adanini799, Nikhil Bhatia799,
Maines Msiska799, Miranda Forsey738, Agilan Kaliappan738, Anne Nicholson738, Joanne Riches738,
Mark Vertue738, Louise Mew800, Esther Mwaura800, Richard Stewart800, Felicity Williams800,
Lynn Wren800, Sara-Beth Sutherland800, Ceri Battle801, Elaine Brinkworth801, Rachel Har ford801,
Carl Murphy801, Luke Newey801, Tabitha Rees801, Marie Williams801, Sophie Arnold801,
David Brealey802, John Hardy802, Henry Houlden802, Eleanor Moncur80 2, Eamon Raith802,
Ambreen Tariq802, Arianna Tucci802, Karen Convery803, Deirdre Fottrell-Gould803, Lisa Hudig803,
Jocelyn Keshet-Price803, Georgina Randell803, Katie Stammers803, Marwa Abdelrazik804 ,
Dhanalakshmi Bakthavatsalam804, Munzir Elhassan804, Arunkumar Ganesan804,
Anne Haldeos804, Jeronimo Moreno-Cuesta804, Dharam Purohit804, Rachel Vincent804,
Kugan Xavier804, Kumar Rohit804, Frater Alasdair804, Malik Saleem804, Carter David804,
Samuel Jenkins804, Zoe Lamond804, Alanna Wall804, Bryan Yates806, Jessica Reynolds806,
Helen Campbell806, Maria Thompsom806, Steve Dodds806, Stacey Duffy806, Deborah Butcher807,
Susie O’Sullivan807, Nicola Butterworth-Cowin807, Bethan Deacon808, Meg Hibbert808,
Carla Pothecary808, Dariusz Tetla808, Christopher Woodford808, Latha Durga808,
Gareth Kennard-Holden808, Laura Ortiz-Ruiz de Gordoa809, Emily Peasgood809, Claire Phillips809,
Denise Skinner810, Jane Gaylard810, Dee Mullan810, Julie Newman810, Ellie Davies811, Lisa Roche811,
Sonia Sathe811, Lutece Brimield812, Zoe Daly812, David Pogson812, Steve Rose812, Amy Collins813,
Waqas Khaliq813,853, Estefania Treus Gude813, Louise Allen536, Eva Beranova536, Nikki Crisp536,
Joanne Deery536, Tracy Hazelton536, Alicia Knight536, Carly Price536, Sorrell Tilbey536,
Salah Turki536, Sharon Turney536, Julian Giles815, Simon Booth815, Gillian Bell816, Katy English816,
Amro Katary816, Louise Wilcox816, Rachael Campbell817, Noreen Clarke817, Jonathan Whiteside817,
Mairi Mascarenhas817, Avril Donaldson817, Joanna Matheson817, Fiona Barrett817,
Marianne O’Hara817, Laura O’Keefe817, Clare Bradley817, Dawn Collier818, Anil Hormis818,
Rachel Walker818, Victoria Maynard818, Tahera Patel819, Matthew Smith819,
Srikanth Chukkambotla819, Aayesha Kazi819, Janice Hartley819, Joseph Dykes819,
Muhammad Hijazi819, Sarah Keith819, Meherunnisa Khan819, Janet Ryan-Smith819,
Philippa Springle819, Jacqueline Thomas819, Nick Truman819, Samuel Saad819 ,
Dabheoc Coleman819, Christopher Fine819, Roseanna Matt819, Bethan Gay819, Jack Dalziel819,
Syamlan Ali819, Drew Goodchild819, Rhiannan Harling819, Ravi Bhatterjee819, Wendy Goddard819,
Chloe Davison819, Stephen Duberly819, Jeanette Hargreaves819, Rachel Bolton819,
Shondipon Laha820, Mark Verlander820, Alexandra Williams820, Helen Blackman821,
Ben Creagh-Brown821, Sinead Donlon821, Natalia Michalak-Glinska821, Sheila Mtuwa821,
Veronika Pristopan821, Armorel Salberg821, Eleanor Smith821, Sarah Stone821, Charles Piercy821,
Jerik Verula821, Dorota Burda821, Rugia Montaser821, Lesley Harden821, Irving Mayangao821,
Cheryl Marriott821, Paul Bradley821, Celia Harris821, Joshua Cooper822, Cheryl Finch822,
Sarah Liderth822, Alison Quinn822, Natalia Waddington822, Katy Fidler823, Emma Tagliavini823,
Kevin Donnelly823, Lynn Abel824, Michael Brett824, Brian Digby8 24, Lisa Gemmell824,
James Hornsby824, Patrick MacGoey824, Pauline O’Neil824, Richard Price824, Natalie Rodden824,
Kevin Rooney824, Radha Sundaram824, Nicola Thomson824 , Rebecca Flanagan825,
Gareth Hughes825, Scott Latham825, Emma McKenna825, Jennifer Anderson825, Robert Hull825,
Kat Rhead825, Debbie Branney826, Jordan Frankham826, Sally Pitts826, Nigel White826,
Daniele Cristiano827, Natalie Dormand827, Zohreh Farzad827, Mahitha Gummadi827,
Kamal Liyanage827, Brijesh V. Patel828, Sara Salmi827, Geraldine Sloane827, Vicky Thwaites827,
Mathew Varghese827, Anelise C. Zborowski827, Sarah Bean829, Karen Burt82 9, Michael Spivey829,
Christine Eastgate-Jackson830, Helder Filipe830, Daniel Martin830, Amitaa Maharajh830,
Sara Mingo Garcia830, Mark De Neef830, Bethan Deacon831, Ceri Lynch831, Carla Pothecary831,
Lisa Roche831, Gwenllian Sera Howe831, Jayaprakash Singh831, Keri Turner831, Hannah Ellis831,
Natalie Stroud831, Shiney Cherian832, Sean Cutler832, Anne Emma Heron832,
Anna Roynon-Reed832, Tamas Szakmany832, Gemma Williams832, Owen Richards832,
Yusuf Cheema832, Norfaizan Ahmad563,833, Joann Barker563,833, Kris Bauchmuller563,833,
Sarah Bird563,833, Kay Cawthron563,833, Kate Harrington563,833, Yvonne Jackson563,833,
Faith Kibutu563,833, Becky Lenagh563,833, Shamiso Masuko563,833, Gary H. Mills563,833,
Ajay Raithatha563,833, Matthew Wiles563,833, Jayne Willson563,833, Helen Newell563,833,
Alison Lye563,833, Lorenza Nwafor563,833, Claire Jarman563,833, Sarah Rowland-Jones563,833,
David Foote563,833, Joby Cole563,833, Roger Thompson563,833, James Watson563,833,

Article
Lisa Hesseldon563,833, Irene Macharia563,833, Luke Chetam563,833, Jacqui Smith563,833,
Amber Ford563,833, Samantha Anderson563,833, Kathryn Birchall563,833, Kay Housley563,833,
Sara Walker563,833, Leanne Milner563,833, Helena Hanratty563,833, Helen Trower563,833,
Patrick Phillips563,833, Simon Oxspring563,833, Ben Donne563,833, Emily Bevan834, Jane Martin834,
Dawn Trodd834, Geoff Watson834, Caroline Wrey Brown834, Lara Bunni835, Claire Jennings835,
Monica Latif835, Rebecca Marshall835, Gayathri Subramanian835, Nageswar Bandla836,
Minnie Gellamucho836, Michelle Davies836, Christopher Thompson836, Laura Ortiz-Ruiz de
Gordoa809, Emily Peasgood809, Claire Phillips809, Denise Skinner810, Jane Gaylard810,
Dee Mullan810, Julie Newman810, Phil Donnison837, Fiona Trim837, Beena Eapen837,
Cecilia Ahmed838, Balvinder Baines838, Sarah Clamp838, Julie Colley838, Risna Haq838,
Anne Hayes838, Jonathan Hulme838, Samia Hussain838, Sibet Joseph838, Rita Kumar838,
Zahira Maqsood838, Manjit Purewal838, Ben Chandler839, Kerry Elliott839, Janine Mallinson839,
Alison Turnbull839, Kathy Dent840, Elizabeth Horsley840, Muhmmad Nauman Akhtar840,
Sandra Pearson840, Dorota Potoczna840, Sue Spencer840, Hayley Blakemore841,
Borislava Borislavova841, Beverley Faulkner841, Emma Gendall841, Elizabeth Goff841, Kati Hayes841,
Matt Thomas841, Ruth Worner841, Kerry Smith841, Deanna Stephens841, Carlos Castro Delgado842,
Deborah Dawson842, Lijun Ding842, Georgia Durrant842, Obiageri Ezeobu842 ,
Sarah Farnell-Ward842, Abiola Harrison842, Rebecca Kanu842, Susannah Leaver842,
Elena Maccacari842, Soumendu Manna842, Romina Pepermans Saluzzio842, Joana Queiroz842,
Tinashe Samakomva842, Christine Sicat842, Joana Texeira842, Edna Fernandes Da Gloria842,
Ana Lisboa842, John Rawlins842, Jisha Mathew842, Ashley Kinch842, William James Hurt842,
Nirav Shah842, Victoria Clark842, Maria Thanasi842, Nikki Yun842, Kamal Patel842, Alison Brown843,
Vikki Crickmore843, Gabor Debreceni843, Joy Wilkins843, Liz Nicol843, Iona Burn844,
Geraldine Hambrook844, Katarina Manso844, Ruth Penn844, Pradeep Shanmugasundaram844,
Julie Tebbutt844, Danielle Thornton844, Anthony Rostron845, Alistair Roy845, Lindsey Woods845,
Sarah Cornell845, Fiona Wakinshaw845, Kimberley Rogerson845, Jordan Jarmain845,
Peter Anderson846, Katie Archer846, Karen Austin846, Caroline Davis846, Alison Durie84 6,
Olivia Kelsall846, Jessica Thrush846, Charlie Vigurs846, Laura Wild846, Hannah-Louise Wood846,
Helen Tranter846, Alison Harrison846, Nicholas Cowley846, Michael McAlindon846,
Andrew Burtenshaw846, Stephen Digby846, Emma Low846, Aled Morgan846, Naiara Cother846,
Tobias Rankin846, Sarah Clayton846, Alex McCurdy846, Suzanne Allibone847,
Roman Mary-Genetu847, Vidya Kasipandian847, Amit Patel847, Ainhi Mac8 47, Anthony Murphy847,
Parisa Mahjoob847, Roonak Nazari847, Lucy Worsley847, Andrew Fagan847, Inthakab Ali Mohamed
Ali848, Karen Beaumont848, Mark Blunt848, Zoe Coton848, Hollie Curgenven848,
Mohamed Elsaadany848, Kay Fernandes848, Sameena Mohamed Ally848, Harini Rangarajan848,
Varun Sarathy848, Sivarupan Selvanayagam848, Dave Vedage848, Matthew White848,
Jaime Fernandez-Roman849, David O. Hamilton849, Emily Johnson849, Brian Johnston84 9,
Maria Lopez Martinez849, Suleman Mulla849, David Shaw849, Alicia A. C. Waite849,
Victoria Waugh849, Ingeborg D. Welters849, Karen Williams849, Thomas Bemand850,
Ethel Black850, Arnold Dela Rosa850, Ryan Howle850, Shaman Jhanji850,
Ravishankar Rao Baikady850, Kate Colette Tatham850, Benjamin Thomas850, Matthew Halkes851,
Pauline Mercer851, Lorraine Thornton851, Joe West852, Tracy Baird852, Jim Ruddy852,
Rosie Reece-Anthony853, Mark Birt854, Amanda Cowton854, Andrea Kay854, Melanie Kent854,
Kathryn Potts854, Ami Wilkinson854, Suzanne Naylor854, Ellen Brown854, Michele Clark855,
Sarah Purvis855, Jade Cole856, Michelle Davies856, Rhys Davies856, Donna Dufin856, Helen Hill856,
Ben Player856, Emma Thomas856, Angharad Williams856, Claire Marie Beith857, Karen Black857 ,
Suzanne Clements857, Alan Morrison857, Dominic Strachan857, Margaret Taylor857,
Michelle Clarkson857, Stuart D’Sylva857, Kathryn Norman857 , Tina Coventry858, Susan Fowler858,
Michael MacMahon858, Amanda McGregor858, Ailbhe Brady859, Rebekah Chan859, Jeff Little859,
Shane McIvor859, Helena Prady859, Helen Whittle859, Bijoy Mathew859 , Melanie Clapham860,
Rosemary Harper860, Una Poultney860, Polly Rice860, Tim Smith860, Rachel Mutch860,
Yolanda Baird861, Aaron Butler861, Indra Chadbourn861, Linda Folkes861, Heather Fox861,
Amy Gardner861, Raquel Gomez861, Gillian Hobden861, Luke Hodgson861 , Kirsten King861,
Michael Margarson861, Tim Martindale861, Emma Meadows861, Dana Raynard861,
Yvette Thirlwall861, David Helm861, Jordi Margalef861, Sandra Greer862, Karen Shuker862,
Ascanio Tridente862, Sara Smuts526, Joseph Dufield526, Oliver Smith526, Lewis Mallon526,
Watkins Claire526, Isobel Birkinshaw863, Joseph Carter863, Kate Howard863, Joanne Ingham863,
Rosie Joy863, Harriet Pearson863, Samantha Roche863, Zoe Scott863, Ellen Knights864,
Alicia Price864, Alice Thomas864, Chris Thorpe864, Azmerelda Abraheem865, Peter Bamford865,
Kathryn Cawley865, Charlie Dunmore865, Maria Faulkner865, Rumanah Girach865, Helen Jeffrey865,
Rhianna Jones865, Emily London865, Imrun Nagra865, Farah Nasir865, Hannah Sainsbury865,
Clare Smedley865, Reena Khade866, Ashok Sundar866, George Tsinaslanidis866, Teresa Behan867,
Caroline Burnett867, Jonathan Hatton867, Elaine Heeney867, Atideb Mitra867, Maria Newton867,
Rachel Pollard867, Rachael Stead867, Jenny Birch868, Laura Bough868, Josie Goodsell868,
Rebecca Tutton868, Patricia Williams868, Sarah Williams868, Barbara Winter-Goodwin868,
Anne Cowley757, Judith Highgate757, Fiona Auld869, Joanne Donnachie869, Ian Edmond869,
Lynn Prentice869, Nikole Runciman869, Dario Salutous869, Lesley Symon869, Anne Todd869,
Patricia Turner869, Abigail Short869, Laura Sweeney869, Euan Murdoch869,
Dhaneesha Senaratne869, Karen Burns870, Andrew Higham870,905 , Taya Anderson871,
Dan Hawcutt871, Laura O’Malley871, Laura Rad871 , Naomi Rogers871, Paula Saunderson871 ,
Kathryn Sian Allison871, Deborah Afolabi871, Jennifer Whitbread871 , Dawn Jones871,
Rachael Dore871, Liana Lankester872, Nikitas Nikitas872, Colin Wells872, Bethan Stowe872,
Kayleigh Spencer872, Susanne Cathcart873, Katharine Duffy873, Alex Puxty873, Kathryn Puxty873,
Lynne Turner873, Jane Ireland873, Gary Semple873, Peter Barry874 , Paula Hilltout875, Jayne Evitts87 5,
Amanda Tyler875, Joanne Waldron875, Val Irvine876, Benjamin Shelley876,
Olugbenga Akinkugbe877, Alasdair Bamford877, Emily Beech877 , Holly Belield877, Michael Bell87 7,
Charlene Davies877, Gareth A. L. Jones877, Tara McHugh877, Hamza Meghari877 , Lauran O’Neill877,
Mark J. Peters877, Samiran Ray877, Ana Luisa Tomas877, Amy Easthope878, Claire Gorman878,
Abhinav Gupta878, Elizabeth Timlick878, Rebecca Brady878, Stephen Bonner879, Keith Hugill879,
Jessica Jones879, Steven Liggett879, Archana Bashyal880, Neil Davidson880, Paula Hutton880,
Stuart McKechnie880, Jean Wilson880, Neil Flint881, Patel Rekha881, Dawn Hales881, Carina Cruz882,
Natalie Pattison882, Shameer Gopal883, Nichola Harris883 , Victoria Lake883, Stella Metherell883,
Elizabeth Radford883, Ian Clement884, Bijal Patel884, A. Gulati884, Carole Hays884, K. Webster884,
Anne Hudson884, Andrea Webster884, Elaine Stephenson884, Louise McCormack884,
Victoria Slater884, Rachel Nixon884, Helen Hanson884, Maggie Fearby884, Sinead Kelly884,
Victoria Bridgett884, Philip Robinson884, Christine Almaden-Boyle885, Pauline Austin885,
Louise Cabrelli885, Stephen Cole885, Matt Casey885, Susan Chapman885, Stephen Cole885,
Clare Whyte885, Adam Brayne886, Emma Fisher886, Jane Hunt886, Peter Jackson886,
Duncan Kaye886, Nicholas Love886, Juliet Parkin886, Victoria Tuckey886, Lynne van Koutrik886,
Sasha Carter886, Benedict Andrew886, Louise Findlay886, Katie Adams886, Michelle Bruce887,
Karen Connolly887, Tracy Duncan887, Helen T.-Michael887, Gabriella Lindergard887,
Samuel Hey887, Claire Fox887, Jordan Alfonso887, Laura Jayne Durrans887, Jacinta Guerin887,
Bethan Blackledge887, Jade Harris887, Martin Hruska887, Ayaa Eltayeb887, Thomas Lamb887,
Tracey Hodgkiss887, Lisa Cooper887, Joanne Rothwell887, Catherine Dennis888,
Alastair McGregor888, Victoria Parris888, Sinduya Srikaran888, Anisha Sukha888, Kim Davies889,
Linda O’Brien889, Zohra Omar889, Igor Otahal889, Emma Perkins889, Tracy Lewis889,
Isobel Sutherland889, Hollie Brooke890, Sarah Buckley890, Jose Cebrian Suarez890,
Ruth Charlesworth890, Karen Hansson890, John Norris890, Alice Poole890, Alastair Rose890,
Rajdeep Sandhu890, Brendan Sloan890, Elizabeth Smithson890, Muthu Thirumaran890,
Veronica Wagstaff890, Alexandra Metcalfe890, Julie Camsooksai891, Charlotte Humphrey891,
Sarah Jenkins891, Henrik Reschreiter891, Beverley Wadams891, Yasmin DeAth891, Colene Adams892,
Anita Agasou892, Tracie Arden893, Amy Bowes892, Pauline Boyle892, Mandy Beekes893,
Heather Button893, Nigel Capps624,893, Mandy Carnahan892, Anne Carter892, Danielle Childs892,
Denise Donaldson893, Kelly Hard892, Fran Hurford893, Yasmin Hussain892, Ayesha Javaid893,
James Jones893, Sanal Jose624,893, Michael Leigh892, Terry Martin893, Helen Millward893,
Nichola Motherwell893, Rachel Rikunenko892, Jo Stickley892, Julie Summers893, Louise Ting893,
Helen Tivenan892, Louise Tonks893, Rebecca Wilcox892, Maria Bokhari899, Vanessa Linnett899,
Rachael Lucas899, Wendy McCormick899, Jenny Ritzema899, Amanda Sanderson899,
Helen Wild899, Nicola Baxter900, Steven Henderson900, Sophie Kennedy-Hay900,
Christopher McParland900, Laura Rooney900, Malcolm Sim900, Gordan McCreath900,
Mark Brunton901, Jess Caterson901, Holly Coles901, Matthew Frise901, Sabi Gurung Rai901,
Nicola Jacques901, Liza Keating901, Emma Tilney901, Shauna Bartley901, Parminder Bhuie901,
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Deepak Subramanian902, Priya Daniel902, Letizia Zitter903, Sarah Benyon903, Suzie Marriott903,
Linda Park903, Samantha Keenan903, Elizabeth Gordon903, Helen Quinn903, Kizzy Baines903,
Gillian Andrew904, J. Kenneth Baillie904, Lucy Barclay904, Marie Callaghan904,
Rachael Campbell904, Sarah Clark904, Dave Hope9 04, Lucy Marshall904, Corrienne McCulloch904 ,
Kate Briton904, Jo Singleton904, Sophie Birch904, Andrew Higham905, Kerry Simpson905,
Jayne Craig905, Carrie Demetriou906, Charlotte Eckbad906, Sarah Hierons906, Lucy Howie906,
Sarah Mitchard906, Lidia Ramos906, Alfredo Serrano-Ruiz906, Katie White906, Fiona Kelly906,
Vishal Amin907, Elena Anastasescu907 , Vikram Anumakonda907, Komala Karthik907,
Rizwana Kausar907, Karen Reid907, Jacqueline Smith907, Janet Imeson-Wood907, Arianna Bellini908,
Jade Bryant908, Anton Mayer908, Amy Pickard908, Nicholas Roe908, Jason Sowter908,
Alex Howlett908, Kristine Criste909, Rebecca Cusack909, Kim Golder909, Hannah Golding909,
Oliver Jones909, Samantha Leggett909, Michelle Male909, Martyna Marani909, Kirsty Prager909,
Toran Williams909, Belinda Roberts909, Karen Salmon909, Prisca Gondo910, B. Hadebe910,
Abdul Kayani910, Bridgett Masunda910, Ashar Ahmed911, Anna Morris911, Srinivas Jakkula911,
Kate Long912, Simon Whiteley912, Elizabeth Wilby912, Bethan Ogg912, Sam Moultrie747, M. Odam747,
Jeremy Bewley913, Zoe Garland913, Lisa Grimmer913, Bethany Gumbrill913, Rebekah Johnson913,
Katie Sweet913, Denise Webster913, Georgia Efford913, Sara Bennett914, Emma Goodwin914,
Matthew Jackson914, Alissa Kent914, Clare Tibke914, Wiesia Woodyatt914, Ahmed Zaki9 14,
Amelia Daniel915, Joanne Finn915, Rajnish Saha915, Nikki Staines915, Amy Easthope915,
Pamela Bremmer528, J. Allan916, T. Geary916, Gordon Houston916, A. Meikle916, P. O’Brien916,
Dina Bell917, Rosalind Boyle917, Katie Douglas917, Lynn Glass917, Emma Lee917, Liz Lennon917,
Austin Rattray917, Rob Charnock918, Denise McFarland918, Denise Cosgrove918, Ben Attwood919,
Penny Parsons919, Siobhain Carmody529, Metod Oblak920, Monica Popescu920,
Mini Thankachen920, Rosie Baruah602, Sheila Morris602, Susie Ferguson602, Amy Shepherd602,
Abdelhakim Altabaibeh922, Ana Alvaro922, Kayleigh Gilbert922, Louise Ma922, Loreta Mostoles922,
Chetan Parmar922, Kathryn Simpson922, Champa Jetha922, Lauren Booker922, Anezka Pratley922,
Tracey Cosier923, Gemma Millen923, Neil Richardson923, Natasha Schumacher923,
Heather Weston923, James Rand923, Beatrice Alex718, Benjamin Bach718, Wendy S. Barclay932,
Debby Bogaert705, Meera Chand933, Graham S. Cooke934, Annemarie B. Docherty714,
Jake Dunning935, Ana da Silva Filipe936, Tom Fletcher937, Christoper A. Green634,
Ewen M. Harrison714, Julian A. Hiscox938, Samreen Ijaz940, Saye Khoo941, Paul Klenerman942,
Andrew Law60, Wei Shen Lim944, Alexander J. Mentzer945, Laura Merson946, Alison M. Meynert62,
Mahdad Noursadeghi947, Shona C. Moore948, Massimo Palmarini936, William A. Paxton948,
Georgios Pollakis948, Nicholas Price949, Andrew Rambaut950, David L. Robertson936,
Clark D. Russell705, Vanessa Sancho-Shimizu951, Janet T. Scott936, Thushan de Silva952,
Louise Sigfrid946, Tom Solomon530, Shiranee Sriskandan934, David Stuart953, Richard S. Tedder955,
Emma C. Thomson936, A. A. Roger Thompson956, Ryan S. Thwaites730, Lance C. W. Turtle530,732,
Rishi K. Gupta957, Carlo Palmieri958, Olivia V. Swann959, Maria Zambon935,
Marc-Emmanuel Dumas960, Julian L. Grifin960, Zoltan Takats960, Kanta Chechi961,
Petros Andrikopoulos960, Anthonia Osagie960, Michael Olanipekun960, Sonia Liggi960,
Matthew R. Lewis962, Gonçalo dos Santos Correia962, Caroline J. Sands962, Panteleimon Takis962,
Lynn Maslen962, William Greenhalf963, Victoria Shaw964, Sarah E. McDonald936, Seán Keating965,
Katie A. Ahmed966, Jane A. Armstrong966, Milton Ashworth966, Innocent G. Asiimwe966,
Siddharth Bakshi966, Samantha L. Barlow966, Laura Booth966, Benjamin Brennan967,
Katie Bullock966, Benjamin W. A. Catterall966, Jordan J. Clark966, Emily A. Clarke966,
Louise Cooper966, Helen Cox966, Christopher Davis966, Oslem Dincarslan9 66, Chris Dunn966,
Philip Dyer966, Angela Elliott966, Anthony Evans966, Lorna Finch966, Lewis W. S. Fisher966,
Terry Foster966, Isabel Garcia-Dorival966, William Greenhalf966, Philip Gunning966,
Catherine Hartley966, Rebecca L. Jensen966, Christopher B. Jones966, Trevor R. Jones966,
Shadia Khandaker966, Katharine King966, Robyn T. Kiy966, Chrysa Koukorava966, Annette Lake967,
Suzannah Lant966, Diane Latawiec966, Lara Lavelle-Langham966, Daniella Lefteri967, Lauren Lett966,
Lucia A. Livoti966, Maria Mancini966, Sarah McDonald966, Laurence McEvoy966,
John McLauchlan967, Soeren Metelmann966, Nahida S. Miah966, Joanna Middleton966,
Joyce Mitchell966, Shona C. Moore966, Ellen G. Murphy966, Rebekah Penrice-Randal966,
Jack Pilgrim966, Tessa Prince966, Will Reynolds966, P. Matthew Ridley966, Debby Sales966,
Victoria E. Shaw966, Rebecca K. Shears966, Benjamin Small966, Krishanthi S. Subramaniam966,
Agnieska Szemiel966, Aislynn Taggart967, Jolanta Tanianis-Hughes966, Jordan Thomas966,
Erwan Trochu966, Libby van Tonder966, Eve Wilcock966, J. Eunice Zhang966, Lisa Flaherty966 ,
Nicole Maziere966, Emily Cass966, Alejandra Doce Carracedo966, Nicola Carlucci966,
Anthony Holmes966, Hannah Massey966, Lee Murphy554, Nicola Wrobel554, Sarah McCafferty554,
Kirstie Morrice554 & Alan MacLean554

Admin team members
Ruth Armstrong60, J. Kenneth Baillie60,61,62, Ceilia Boz60, Adam Brown60, Richard Clark968,
Sara Clohisey60, Audrey Coutts968, Louise Cullum60, Nicky Day60, Lorna Donnelly968,
Esther Duncan60, Angie Fawkes968, Paul Finernan60, Max Head Fourman60, James Furniss60,
Tammy Gilchrist968, Ailsa Golightly60, Katarzyna Hafezi968, Ross Hendry60, Andy Law60,
Dawn Law60, Rachel Law60, Sarah Law60, Louise Macgillivray968, Alan Maclean968,
Hanning Mal60, Sarah McCafferty968, Ellie Mcmaster60, Jen Meikle60 , Shona C. Moore732,
Kirstie Morrice968, Lee Murphy968, Wilna Oosthuyzen60, Nicholas Parkinson60, Trevor Paterson60,
Andrew Stenhouse60, Maaike Swets60,970, Helen Szoor-McElhinney60, Filip Taneski60,
Lance C. W. Turtle530,732, Tony Wackett60, Mairi Ward60, Jane Weaver60, Nicola Wrobel968,
Marie Zechner60, Judy Coyle60, Bernadette Gallagher60, Rebecca Lidstone-Scott60,
Debbie Hamilton60, Katherine Schon971, Anita Furlong971, Heather Biggs971, Fiona Grifiths60,
Eleanor Andrews762, Kathy Brickell527, Michelle Smyth527, Lorna Murphy527, Gail Carson946,
Hayley Hardwick530 & Chloe Donohue715
COVID-19 HGI corresponding authors
Benjamin M. Neale4,1280 ✉, Mark Daly1,2,3,1280 ✉ & Andrea Ganna1,2,3,1280 ✉
1Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland.
2Broad Institute of MIT and Harvard, Cambridge, MA, USA. 3Analytic and Translational
Genetics Unit, Massachusetts General Hospital, Boston, MA, USA. 4Massachusetts General
Hospital, Broad Institute of MIT and Harvard, Cambridge, MA, USA. 6Yale University, New
Haven, CT, USA. 7Icahn School of Medicine at Mount Sinai, New York, NY, USA. 8Stroke
Pharmacogenomics and Genetics, Biomedical Research Institute Sant Pau (IIB Sant Pau), Sant
Pau Hospital, Inmungen-CoV2, Barcelona, Spain. 9Institute of Virology, Technical University
Munich and Helmholtz Zentrum München, Munich, Germany. 10Institute of Psychiatric
Phenomics and Genomics, Medical Center of the University of Munich, Munich, Germany.
11Department of Psychiatry, Medical Center of the University of Munich, Munich, Germany.
12IRCCS, Istituto Giannina Gaslini, Genova, Italy. 13Department of Neurosciences,
Rehabilitation, Ophthalmology, Genetics, Maternal and Child Health, University of Genova,
Genova, Italy. 14Queen Mary University of London, London, UK. 15Open Targets, Wellcome
Genome Campus, Hinxton, UK. 16Department of Complex Trait Genetics, Center for
Neurogenomics and Cognitive Research, Amsterdam Neuroscience, Vrije Universiteit
Amsterdam, Amsterdam, The Netherlands. 17Lady Davis Institute, Jewish General Hospital,
McGill University, Montreal, Quebec, Canada. 18Medical Research Institute, Kangbuk
Samsung Hospital, Sungkyunkwan University School of Medicine, Suwon, Republic of Korea.
19Osaka University Graduate School of Medicine, Osaka, Japan. 20Baylor College of Medicine,
Houston, TX, USA. 21Mohammed Bin Rashid University of Medicine and Health Sciences,
Dubai, United Arab Emirates. 22MRC Integrative Epidemiology Unit (IEU), University of Bristol,
Bristol, UK. 23Department of Internal Medicine, Division of Cardiovascular Medicine, Michigan
Medicine, Ann Arbor, MI, USA. 24Department of Human Genetics, University of Michigan
Medical School, Ann Arbor, MI, USA. 25Department of Computational Medicine and
Bioinformatics, University of Michigan Medical School, Ann Arbor, MI, USA. 26Seaver Autism
Center for Research and Treatment, Department of Psychiatry, Icahn School of Medicine at
Mount Sinai, New York, NY, USA. 29Program in Medical and Population Genetics, Broad
Institute of MIT and Harvard, Cambridge, MA, USA. 32David Geffen School of Medicine at
UCLA, Los Angeles, CA, USA. 33Institut Pasteur, Paris, France. 34Harvard School of Public
Health, Boston, MA, USA. 35Institute for Molecular Bioscience, The University of Queensland,
Brisbane, Queensland, Australia. 38Wellcome Sanger Institute, Wellcome Genome Campus,
Hinxton, UK. 40European Molecular Biology Laboratory, European Bioinformatics Institute
(EMBL-EBI), Wellcome Genome Campus, Hinxton, UK. 42Centre for Bioinformatics and Data
Analysis, Medical University of Bialystok, Bialystok, Poland. 43Trieste University, Trieste, Italy.
44Vanderbilt University Medical Center, Nashville, TN, USA. 45University of California San
Francisco, San Francisco, CA, USA. 46Stanford University, Stanford, CA, USA. 47University of
Siena, Siena, Italy. 49Boston Children’s Hospital, Broad Institute of MIT and Harvard,
Cambridge, MA, USA. 50Blizard Institute, Queen Mary University of London, London, UK.
51Department of Genetics, University Medical Centre Groningen, Groningen, The Netherlands.
52Department of Genetics, University Medical Centre Utrecht, Utrecht, The Netherlands.
53Department of Epidemiology, Biostatistics and Occupational Health, McGill University,
Montreal, Quebec, Canada. 55Department of Twin Research, King’s College London, London,
UK. 56Department of Human Genetics, McGill University, Montreal, Quebec, Canada.
57Kyoto-McGill International Collaborative School in Genomic Medicine, Graduate School of
Medicine, Kyoto University, Kyoto, Japan. 59National Institutes of Health, Bethesda, MD, USA.
60The Roslin Institute, University of Edinburgh, Edinburgh, UK. 61Intensive Care Unit, Royal
Inirmary of Edinburgh, Edinburgh, UK. 62MRC Human Genetics Unit, Institute of Genetics and
Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, UK.
63Medical Genetics, University of Siena, Siena, Italy. 64Genetica Medica, Azienda
Ospedaliero-Universitaria Senese, Siena, Italy. 65Med Biotech Hub and Competence Center,
Department of Medical Biotechnologies, University of Siena, Siena, Italy. 66Department of
Electronics, Information and Bioengineering (DEIB), Politecnico di Milano, Milano, Italy.
67Politecnico di Milano, Milan, Italy. 68University of Michigan, Ann Arbor, MI, USA. 69Vanderbilt
School of Medicine, Nashville, TN, USA. 70All India Institute of Medical Sciences Kalyani,
Kalyani, India. 71Hasso Plattner Institute, New York, NY, USA. 72Naina Tech, Hyderabad, India.
73EMBL-European Bioinformatics Institute, Hinxton, UK. 74University of Northampton,
Northampton, UK. 75University of Helsinki, Helsinki, Finland. 76University of Miami, Miami, FL,
USA. 78Ecole Centrale de Nantes, Inserm, Centre de Recherche en Transplantation et
Immunologie, Nantes University, UMR1064, ITUN, Nantes, France. 79University of Liège, Liège,
Belgium. 80Qatar Genome Program, Qatar Foundation Research, Development and
Innovation, Qatar Foundation, Doha, Qatar. 82Medical and Population Genetics and
Cardiovascular Disease Initiative, Broad Institute of Harvard and MIT, Cambridge, Cambridge,
MA, USA. 83Cardiovascular Research Center, Massachusetts General Hospital, Boston, MA,
USA. 84Intensive Care Unit, Vall d’Hebron Hospital, Barcelona, Spain. 85Institut de Biomedicina
de València - CSIC, València, Spain. 86Centro de Investigación Biomédica en Red en
Enfermedades Neurodegenerativas (CIBERNED), València, Spain. 87Unidad Mixta de
Neurología y Genética, Instituto de Investigación Sanitaria La Fe, València, Spain. 88Erasmus
Medical Center, Rotterdam, The Netherlands. 89National Genome Center, Copenhagen,
Denmark. 90University of Copenhagen, Copenhagen, Denmark. 91Genomics PLC, Oxford, UK.
93Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.
94Department of Population Medicine and Lifestyle Diseases Prevention, Medical University of
Bialystok, Bialystok, Poland. 95Genomics England, London, UK. 96Junta de Andalucía, Seville,
Spain. 97Human Genetics Program of ICBM and Department of Basic-Clinical Oncology,
University of Chile, Santiago, Chile. 98Center for the Development of Scientiic Research
(CEDIC), Asunción, Paraguay. 100Translational Bioinformatics Unit, Navarrabiomed, Complejo
Hospitalario de Navarra (CHN), Universidad Pública de Navarra (UPNA), IdiSNA, Pamplona,
Spain. 101Mucosal & Salivary Biology Division, King’s College London Dental Institute, London,
UK. 102GENYO, Center for Genomics and Oncological Research Pizer, University of Granada,
Andalusian Regional Government, Granada, Spain. 103University of Puerto Rico, San Juan,
Puerto Rico. 104National Laboratory of Genomics for Biodiversity (LANGEBIO), Advanced
Genomics Unit, CINVESTAV, Irapuato, Mexico. 105Queensland University of Technology,
Brisbane, Queensland, Australia. 106Clinical Research Unit of Nanoro, Institut de Recherche en
Sciences dela Santé, CNRST, Ouagadougou, Burkina Faso. 107McGill University, Montreal,
Quebec, Canada. 108Université de Montréal, Montreal, Quebec, Canada. 109Fonds dela
Recherche Scientiique (FNRS) & Centre de Génétique Humaine, Hôpital Erasme, Université
Libre de Bruxelles, Brussels, Belgium. 112University of Pecs Medical School, Pécs, Hungary.
113Institute of Biomedicine and Cancer Research Laboratories, Western Cancer Centre FICAN
West, University of Turku, Turku, Finland. 114Institute of Biomedical Technologies, National
Research Council, Segrate, Italy. 115Immediate, Milan, Italy. 116University of Cambridge,
Cambridge, UK. 117Genome Opinion, Seoul, Republic of Korea. 119University of Groningen,
Groningen, The Netherlands. 120Universiti Malaysia Pahang, Gambang, Malaysia. 122Vrije
Universiteit Amsterdam, Amsterdam, The Netherlands. 123University Medical Centre
Groningen, University of Groningen, Groningen, The Netherlands. 124MNM DIAGNOSTICS,
Pozna?, Poland. 125Institute for Systems Biology, Seattle, WA, USA. 126Sultan Idris Education
University, Tanjung Malim, Malaysia. 127Hospital Kulim, Kedah, Malaysia. 128AbbVie, Lake Buff,
IL, USA. 129Root Deep Insight, Boston MA, USA. 13023andMe, Sunnyvale, CA, USA. 131GSK,
Stevenage, UK. 132Department of Pharmacology, Feinberg School of Medicine, Northwestern
University, Chicago, IL, USA. 133Department of Medicine, Northwestern University, Chicago, IL,
USA. 134Washington DC Veterans Affairs Medical Center, Hospital Medicine, Washington, DC,
USA. 135Department of Medicine, George Washington University, Washington, DC, USA.
136Section of Hospital Medicine, Department of Medicine, University of Chicago, Chicago, IL,
USA. 137Section of Hematology and Oncology, Department of Medicine, University of Chicago,
Chicago, IL, USA. 138College of Pharmacy, University of Illinois at Chicago, Chicago, IL, USA.
144Department of Pharmacology, George Washington University, Washington, DC, USA.
148Department of Neurology, Amsterdam UMC, Amsterdam Neuroscience, Amsterdam, The
Netherlands. 149Department of Intensive Care, Amsterdam UMC, Amsterdam, The
Netherlands. 150Department of Infectious Diseases, Amsterdam UMC, Amsterdam, The
Netherlands. 151Department of Clinical Epidemiology, Biostatistics and Bioinformatics,
Amsterdam UMC, Amsterdam, The Netherlands. 152Experimental Immunology, Amsterdam
UMC, Amsterdam, The Netherlands. 153Department of Pulmonology, Amsterdam UMC,
Amsterdam, The Netherlands. 154Department of Pathology, Amsterdam UMC, Amsterdam, The
Netherlands. 155Department of Anesthesiology, Amsterdam UMC, Amsterdam, The
Netherlands. 156Amsterdam UMC Biobank Core Facility, Amsterdam UMC, Amsterdam, The
Netherlands. 157Department of Radiology, Amsterdam UMC, Amsterdam, The Netherlands.
158Department of Medical Microbiology, Amsterdam UMC, Amsterdam, The Netherlands.
159Department of Clinical Chemistry, Amsterdam UMC, Amsterdam, The Netherlands.
160Amsterdam UMC Biobank, Amsterdam UMC, Amsterdam, The Netherlands. 161Core Facility
Genomics, Amsterdam UMC, Amsterdam, The Netherlands. 162Ancestry, Lehi, UT, USA.
163GIGA-Institute, University of Liège, Liège, Belgium. 164CHC Mont-Légia, Liège, Belgium.
165BHUL (Liège Biobank), CHU of Liège, Liège, Belgium. 167Stanley Center for Psychiatric
Research, Broad Institute of MIT and Harvard, Cambridge, MA, USA. 168Centre de Génétique
Humaine, Hôpital Erasme, Université Libre de Bruxelles, Brussels, Belgium. 169Service de
Médecine Interne, Hôpital Erasme, Université Libre de Bruxelles, Brussels, Belgium. 170CHU of
Liège, University of Liège, Liège, Belgium. 174McGill Genome Centre, McGill University,
Montréal, Québec, Canada. 177Department of Emergency Medicine, McGill University,
Montreal, Quebec, Canada. 178Emergency Department, Jewish General Hospital, McGill
University, Montreal, Quebec, Canada. 179McGill AIDS Centre, Department of Microbiology
and Immunology, Lady Davis Institute for Medical Research, Jewish General Hospital, McGill
University, Montreal, Quebec, Canada. 180McGill Centre for Viral Diseases, Department of
Infectious Disease, Lady Davis Institute, Jewish General Hospital, Montreal, Quebec, Canada.
181Research Centre of the Centre Hospitalier de l’Université de Montréal, Montreal, Quebec,
Canada. 182Department of Medicine, Research Centre of the Centre Hospitalier de l’Université
de Montréal, Montreal, Quebec, Canada. 183Department of Medicine, Université de Montréal,
Montreal, Quebec, Canada. 184Department of Medicine and Human Genetics, McGill
University, Montreal, Quebec, Canada. 185Department of Intensive Care, Research Centre of
the Centre Hospitalier de l’Université de Montréal, Montreal, Quebec, Canada. 186Division of
Infectious Diseases, Research Centre of the Centre Hospitalier de l’Université de Montréal,
Montréal, Quebec, Canada. 187Division of Genetic Medicine, Department of Medicine,
Vanderbilt University Medical Center, Nashville, TN, USA. 188Vanderbilt Genetics Institute,
Vanderbilt University Medical Center, Nashville, TN, USA. 189Institute of Human Genetics,
University Hospital Bonn, Medical Faculty University of Bonn, Bonn, Germany. 190Institute of
Genomic Statistics and Bioinformatics, University Hospital Bonn, Medical Faculty University
of Bonn, Bonn, Germany. 191Department of Gastroenterology, Hepatology and Infectious
Diseases, University Hospital Düsseldorf, Medical Faculty Heinrich Heine University,
Düsseldorf, Germany. 192Institute of Human Genetics, Medical Faculty, RWTH Aachen
University, Aachen, Germany. 193Clinic for Cardiology, Angiology and Internal Intensive
Medicine, Medical Clinic I, RWTH Aachen University, Aachen, Germany. 194Department of
Pneumology and Intensive Care Medicine, Faculty of Medicine, RWTH Aachen University,
Aachen, Germany. 195Department of Pneumology, Hannover Medical School, Hannover,
Germany. 196Department of Gastroenterology, Hepatology and Endocrinology, Hannover
Medical School, Hannover, Germany. 197Hannover Uniied Biobank, Hannover Medical School,
Hannover, Germany. 198Department I of Internal Medicine, Faculty of Medicine and University
Hospital of Cologne, University of Cologne, Cologne, Germany. 199Center for Molecular
Medicine Cologne (CMMC), University of Cologne, Cologne, Germany. 200German Center for
Infection Research (DZIF), Partner Site Bonn-Cologne, Cologne, Germany. 202Cologne Center

Article
for Genomics (CCG), University of Cologne, Cologne, Germany. 203Department of
Anesthesiology and Intensive Care Medicine, University Hospital Essen, University
Duisburg-Essen, Essen, Germany. 204Department of Child and Adolescent Psychiatry,
University Hospital Essen, University of Duisburg-Essen, Essen, Germany. 205Department of
Infectious Diseases, University Hospital Essen, University Duisburg-Essen, Essen, Germany.
206Department of Pneumology, Allergology and Respiratory Medicine, University Hospital
Saarland, Homburg/Saar, Germany. 207Center of Human and Molecular Biology, Department
of Human Genetics, University Hospital Saarland, Homburg/Saar, Germany. 208Department of
Genetics & Epigenetics, Saarland University, Saarbrücken, Germany. 209Eurac Research,
Institute for Biomedicine (afiliated to the University of Lübeck), Bolzano, Italy. 210University of
Colorado Anschutz Medical Campus, Aurora, CO, USA. 211Department of Genetics and
Development, Institute for Genomic Medicine, Columbia University, New York, NY, USA.
212Department of Medicine, Institute for Genomic Medicine, Columbia University, New York,
NY, USA. 213Department of Biomedical Informatics, Columbia University, New York, NY, USA.
214Department of Pediatrics, Columbia University, New York, NY, USA. 215Department of
Medicine, Columbia University, New York, NY, USA. 216Institute for Genomic Medicine,
Columbia University, New York, NY, USA. 217Department of Biostatistics, Mailman School of
Public Health, Columbia University, New York, NY, USA. 218Department of Pathology and Cell
Biology, Columbia University, New York, NY, USA. 219Medical Research Institute, Kangbuk
Samsung Hospital, Sungkyunkwan University School of Medicine, Seoul, Republic of Korea.
220Department of Clinical Research Design and Evaluation, SAIHST, Sungkyunkwan University,
Seoul, Republic of Korea. 221Division of Gastroenterology, Department of Medicine, Kangbuk
Samsung Hospital, Sungkyunkwan University, School of Medicine, Seoul, Republic of Korea.
222Department of Biochemistry, College of Medicine, Ewha Womans University, Seoul,
Republic of Korea. 223Department of Internal Medicine, Seoul National University Hospital,
Seoul National University College of Medicine, Seoul, Republic of Korea. 224Department of
Periodontology, Section of Dentistry, Seoul National University Bundang Hospital, Seongnam,
Republic of Korea. 225Department of Internal Medicine, Seoul National University Bundang
Hospital, Seoul National University College of Medicine, Seongnam, Republic of Korea.
226Department of Physical & Rehabilitation Medicine, Kangbuk Samsung Hospital,
Sungkyunkwan University School of Medicine, Seoul, Republic of Korea. 227Department of
Clinical Research Design & Evaluation, SAIHST, Sungkyunkwan University, Seoul, Republic of
Korea. 228Biomedical Institute for Convergence at SKKU, Sungkyunkwan University School of
Medicine, Suwon, Republic of Korea. 229Department of Public Health Service, Seoul National
University Bundang Hospital, Seongnam, Republic of Korea. 230Department of Rehabilitation
Medicine, Seoul National University College of Medicine, Seoul, Republic of Korea. 231Korea
Research Environment Open NETwork, Korea Institute of Science and Technology
Information, Daejeon, Republic of Korea. 232Global Science Experimental Data Hub Center,
Korea Institute of Science and Technology Information, Daejeon, Republic of Korea.
233Division of Infectious Diseases, Department of Medicine, Kangbuk Samsung Hospital,
Sungkyunkwan University School of Medicine, Seoul, Republic of Korea. 234Center for Cohort
Studies, Kangbuk Samsung Hospital, Sungkyunkwan University School of Medicine, Seoul,
Republic of Korea. 235Department of Occupational and Environmental Medicine,
Sungkyunkwan University School of Medicine, Seoul, Republic of Korea. 236Department of
Laboratory Medicine, Seoul National University Bundang Hospital, Seoul National University
College of Medicine, Seongnam, Republic of Korea. 237Institute of Clinical Molecular Biology,
Christian-Albrechts-University, Kiel, Germany. 238Novo Nordisk Foundation Center for Protein
Research, Disease Systems Biology, Faculty of Health and Medical Sciences, University of
Copenhagen, Copenhagen, Denmark. 239Institut de Biotecnologia i de Biomedicina,
Universitat Autònoma de Barcelona, Barcelona, Spain. 240ICREA, Barcelona, Spain. 241Research
Group for Evolutionary Immunogenomics, Max Planck Institute for Evolutionary Biology, Plön,
Germany. 242Research Unit for Evolutionary Immunogenomics, Department of Biology,
University of Hamburg, Hamburg, Germany. 243Department of Gastroenterology, Hospital
Universitario Ramón y Cajal, University of Alcalá, Instituto Ramón y Cajal de Investigación
Sanitaria (IRYCIS), Madrid, Spain. 244Centro de Investigación Biomédica en Red en
Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII),
Madrid, Spain. 245Vall d’Hebron Institut de Recerca (VHIR), Vall d’Hebron Hospital Universitari,
Barcelona, Spain. 246Charite Universitätsmedizin Berlin, Berlin, Germany. 249Hospital
Universitario Clinico San Cecilio, Granada, Spain. 250Instituto de Investigación Ibs.Granada,
Granada, Spain. 251Emergency Department, University Hospital Regensburg, Regensburg,
Germany. 252Department for Infectious Diseases and Infection Control, University Hospital
Regensburg, Regensburg, Germany. 253Medical University of Innsbruck, Department of
Medicine and Christian Doppler Laboratory on Iron and Phosphate Biology, Innsbruck,
Austria. 254Institute of Clinical Medicine, University of Oslo, Oslo, Norway. 255Department of
Microbiology, Oslo University Hospital, Oslo, Norway. 256Hospital Clinic, University of
Barcelona and IDIBAPS, Barcelona, Spain. 257European Foundation for the Study of Chronic
Liver Failure (EF-CLIF), Barcelona, Spain. 259Cologne Excellence Cluster on Cellular Stress
Responses in Aging-Associated Diseases (CECAD), University of Cologne, Cologne, Germany.
260Center for Molecular Medicine Cologne (CMMC), University of Cologne, Cologne,
Germany. 261Genomes for Life-GCAT labGermans Trias i Pujol Research Institute (IGTP),
Badalona, Spain. 262IRCCS Humanitas Research Hospital, Milan, Italy. 263Department of
Biomedical Sciences, Humanitas University, Pieve Emanuele, Milan, Italy. 264Institute of
Transfusionsmedicine, University Hospital Schleswig-Holstein (UKSH), Kiel, Germany. 265Klinik
für Innere Medizin I, Universitätsklinikum Schleswig-Holstein, Kiel Campus, Kiel, Germany.
266Zentrum für Humangenetik Regensburg, Regensburg, Germany. 267University Hospital
Schleswig-Holstein (UKSH), Kiel Campus, Kiel, Germany. 268Section for Gastroenterology,
Department of Transplantation Medicine, Division for Cancer Medicine, Surgery and
Transplantation, Oslo University Hospital Rikshospitalet, Oslo, Norway. 269Research Institute
for Internal Medicine, Division of Surgery, Inlammatory Diseases and Transplantation, Oslo
University Hospital Rikshospitalet and University of Oslo, Oslo, Norway. 270Norwegian PSC
Research Center, Department of Transplantation Medicine, Division of Surgery, Inlammatory
Diseases and Transplantation, Oslo University Hospital Rikshospitalet, Oslo, Norway.
272Division of Rheumatology, Inlammation and Immunity, Brigham and Women’s Hospital and
Harvard Medical School, Boston, MA, USA. 273Division of Genetics, Department of Medicine,
Brigham and Women’s Hospital, Boston, MA, USA. 274Department of Biomedical Informatics,
Harvard Medical School, Boston, MA, USA. 275Center for Data Sciences, Brigham and Women’s
Hospital, Boston, MA, USA. 276Randaberg Municipality, Randaberg, Norway. 277Department of
Quality and Health Technology, Faculty of Health Sciences, University of Stavanger,
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. 281Osakidetza Basque
Health Service, Donostialdea Integrated Health Organisation, Clinical Biochemistry
Department, San Sebastian, Spain. 282Research Center and Memory Clinic, Fundació ACE,
Institut Català de Neurociències Aplicades, Universitat Internacional de Catalunya, Barcelona,
Spain. 283Networking Research Center on Neurodegenerative Diseases (CIBERNED), Instituto
de Salud Carlos III, Madrid, Spain. 284Department of Acute Medicine, Oslo University Hospital,
Oslo, Norway. 285Fondazione IRCCS Ca’ Granda Ospedale Maggiore Policlinico, Milan, Italy.
286European Reference Network on Hepatological Diseases (ERN RARE LIVER), San Gerardo
Hospital, Monza, Italy. 287Division of Gastroenterology, Center for Autoimmune Liver Diseases,
Department of Medicine and Surgery, University of Milan Bicocca, Milan, Italy. 288German
Center for Neurodegenerative Diseases (DZNE Bonn), Bonn, Germany. 289Division of
Neurogenetics and Molecular Psychiatry, Department of Psychiatry and Psychotherapy,
Medical Faculty, University of Cologne, Cologne, Germany. 290Department of Psychiatry,
Glenn Biggs Institute for Alzheimer’s and Neurodegenerative Diseases, San Antonio, TX, USA.
291Department of Neurodegenerative Diseases and Geriatric Psychiatry, University Hospital
Bonn, Bonn, Germany. 292Liver Unit, Department of Internal Medicine, Hospital Universitari
Vall d’Hebron, Vall d’Hebron Barcelona Hospital Campus, Barcelona, Spain. 293Department of
Anesthesiology and Intensive Care, University Hospital of North Norway, Tromsø, Norway.
294Klinik für Innere Medizin I, Universitätsklinikum Schleswig-Holstein, Kiel Campus, Kiel,
Germany. 295Gastroenterology Unit, Fondazione IRCCS Casa Sollievo della Sofferenza, San
Giovanni Rotondo, Italy. 296Department of Infectious Diseases, Oslo University Hospital, Oslo,
Norway. 297Microbiology Department, Hospital Universitari Vall d’Hebron, Barcelona, Spain.
298Universitat Autònoma de Barcelona, Bellaterra, Spain. 299Department of Respiratory
Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación
Sanitaria (IRYCIS), Madrid, Spain. 300Department of Respiratory Medicine and Allergology,
University Hospital, Goethe University, Frankfurt am Main, Germany. 301Department of
Infectious Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de
Investigación Sanitaria (IRYCIS), University of Alcalá, Madrid, Spain. 302Department of Internal
Medicine II, Technical University of Munich, School of Medicine, University Hospital rechts der
Isar, Munich, Germany. 303Division of Clinical Infectious Diseases, Research Center Borstel,
Borstel, Germany. 304German Center for Infection Research (DZIF) Clinical Tuberculosis Unit,
Borstel, Germany. 305Respiratory Medicine & International Health, University of Lübeck,
Lübeck, Germany. 306Osakidetza Basque Health Service, Basurto University Hospital,
Respiratory Service, Bilbao, Spain. 307Department of Clinical and Molecular Medicine, Faculty
of Medicine and Health Science, Norwegian University of Science and Technology,
Trondheim, Norway. 308Clinic Ålesund Hospital, Department of Medicine, Møre & Romsdal
Hospital Trust, Ålesund, Norway. 309Department of Anesthesiology, Hospital Universitario
Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of
Alcalá, Madrid, Spain. 310Spain Hospital Clinic, University of Barcelona and IDIBAPS,
Barcelona, Spain. 311Osakidetza Basque Health Service, Galdakao Hospital, Respiratory
Service, Galdakao, Spain. 312IBMDR - E.OOspedali Galliera, Genova, Italy. 313Liver ICU, Hospital
Clinic Barcelona, Barcelona, Spain. 314Biocruces Bizkaia Health Research Institute, Barakaldo,
Spain. 315Histocompatibilidad y Biologia Molecular, Centro de Transfusion de Madrid, Madrid,
Spain. 316University of Milan, Milan, Italy. 317Fondazione Grigioni per il Morbo di Parkinson,
Milan, Italy. 318Department of Anesthesiology, Intensive Care Medicine and Pain Therapy,
University Hospital Frankfurt, Frankfurt am Main, Germany. 319German Center for Infection
Research (DZIF), Medical Faculty and University Hospital Cologne, University of Cologne,
Partner Site Bonn-Cologne, Cologne, Germany. 321Ikerbasque, Basque Foundation for
Science, Bilbao, Spain. 322Department of Liver and Gastrointestinal Diseases, Biodonostia
Health Research Institute, Donostia University Hospital, University of the Basque Country
(UPV/EHU), San Sebastian, Spain. 323Infectious Diseases Service, Osakidetza, Biocruces
Bizkaia Health Research Institute, Barakaldo, Spain. 324Medical Department, Drammen
Hospital, Vestre Viken Hospital Trust, Drammen, Norway. 325Research Center Borstel,
BioMaterialBank Nord, Borstel, Germany. 326German Center for Lung Research (DZL), Airway
Research Center North (ARCN), Giessen, Germany. 327Popgen 2.0 Network (P2N), Kiel,
Germany. 329Department of Liver and Gastrointestinal Diseases, Biodonostia Health Research
Institute, Donostia University Hospital, University of the Basque Country (UPV/EHU),
CIBERehd, San Sebastian, Spain. 330Department of Infectious Diseases, Oslo University
Hospital, Oslo, Norway. 331Department of Clinical Science, University of Bergen, Bergen,
Norway. 332Biodonostia Health Research Institute, Donostia University Hospital, San Sebastian,
Spain. 333Germans Trias i Pujol Research Institute (IGTP), Badalona, Spain. 335ISGlobal,
Barcelona, Spain. 336CIBER Epidemiología y Salud Pública (CIBERESP), Madrid, Spain.
337Universitat Pompeu Fabra (UPF), Barcelona, Spain. 338Hospital del Mar Medical Research
Institute (IMIM), Barcelona, Spain. 339Osakidetza Basque Health Service, Donostialdea
Integrated Health Organization, San Sebastian, Spain. 340Department of Internal Medicine,
Infectious Diseases, University H.ospital Frankfurt and Goethe University Frankfurt, Frankfurt
am Main, Germany. 341Humanitas Gavazzeni-Castelli, Bergamo, Italy. 344School of Biological
Sciences, Monash University, Clayton, Victoria, Australia. 345Munich Clinic Schwabing,
Academic Teaching Hospital, Ludwig-Maximilians-University (LMU), Munich, Germany.
346Department of Anesthesiology, Hospital Universitario Ramón y Cajal, Instituto Ramón y
Cajal de Investigación Sanitaria (IRYCIS), Madrid, Spain. 349Clinical Trials Centre Cologne, ZKS
Köln, Cologne, Germany. 351Institute of Human Genetics, University of Bonn School of
Medicine, University Hospital Bonn, Bonn, Germany. 352Institute of Clinical Molecular Biology,
Christian-Albrechts-University of Kiel, Kiel, Germany. 353UKSH Schleswig-Holstein, Kiel,
Germany. 354Institute of Immunology, Christian-Albrechts-University of Kiel, Kiel, Germany.
355Institute of Medical Virology, University Hospital Frankfurt, Goethe University, Frankfurt am
Main, Germany. 356German Centre for Infection Research (DZIF), External Partner Site
Frankfurt, Frankfurt am Main, Germany. 357Department of Neurology, Bezirksklinikum
Regensburg, University of Regensburg, Regensburg, Germany. 359Charite Universitätsmedizin
Berlin, Berlin Institute of Health, Berlin, Germany. 360German Center for Infection Research
(DZIF), Partner Site Munich, Munich, Germany. 361Department of Infectious Diseases,

University Hospital of North Norway, Tromsø, Norway. 362Faculty of Health Sciences, UIT The
Arctic University of Norway, Tromsø, Norway. 363Catalan Institute of Oncology (ICO),
Barcelona, Spain. 364Bellvitge Biomedical Research Institute (IDIBELL), Barcelona, Spain.
365Universitat de Barcelona (UB), Barcelona, Spain. 366deCODE genetics, Reykjavik, Iceland.
368Mucosal Immunology Lab, Unidad de Excelencia Instituto de Biomedicina y Genética
Molecular de Valladolid (IBGM), Universidad de Valladolid-CSIC, Valladolid, Spain. 369Centro
de Investigaciones Biomédicas en Red de Enfermedades Hepáticas y Digestivas (CIBERehd),
Madrid, Spain. 370Valladolid University Hospital, Valladolid, Spain. 371Estonian Genome Centre,
Institute of Genomics, University of Tartu, Tartu, Estonia. 372SYNLAB Estonia, University of
Tartu, Tartu, Estonia. 373University of Tartu, Tartu, Estonia. 374Kuressaare Hospital, Kuressaare,
Estonia. 375Institute of Biomedicine and Translational Medicine, University of Tartu, Tartu,
Estonia. 377West Tallinn Central Hospital, Tallinn, Estonia. 378University of Tartu, Tartu University
Hospital, Tartu, Estonia. 379Estonian Health Insurance Fund, Tallinn, Estonia. 380Tartu University
Hospital, Tartu, Estonia. 381FinnGen, Helsinki, Finland. 382Finnish Institute for Health and
Welfare (THL), Helsinki, Finland. 383University of Helsinki, Faculty of Medicine, Clinical and
Molecular Metabolism Research Program, Helsinki, Finland. 384Institute of Molecular and
Clinical Ophthalmology Basel (IOB), Basel, Switzerland. 385Department of Ophthalmology,
University of Basel, Basel, Switzerland. 386Infectious Diseases Service, Department of
Medicine, University Hospital and University of Lausanne, Lausanne, Switzerland. 388Centre
for Primary Care and Public Health, University of Lausanne, Lausanne, Switzerland. 389Division
of Infectious Diseases and Hospital Epidemiology, Cantonal Hospital St Gallen, St Gallen,
Switzerland. 390Division of Intensive Care, Geneva University Hospitals and the University of
Geneva Faculty of Medicine, Geneva, Switzerland. 391Infectious Disease Service, Department
of Internal Medicine, Geneva University Hospital, Geneva, Switzerland. 392Clinique de
Médecine et spécialités, Infectiologie, HFR-Fribourg, Fribourg, Switzerland. 393Infectious
Diseases Division, University Hospital Centre of the Canton of Vaud, Hospital of Valais, Sion,
Switzerland. 394Functional Host Genomics of Infectious Diseases, University Hospital and
University of Lausanne, Lausanne, Switzerland. 395Registry COVID, University Hospital and
University of Lausanne, Lausanne, Switzerland. 396Pneumonia Prediction using Lung
Ultrasound, University Hospital and University of Lausanne, Lausanne, Switzerland. 397Center
for Primary Care and Public Health (Unisanté), University of Lausanne, Lausanne, Switzerland.
398COVID-19 Risk Prediction in Swiss ICUs-Trial, Division of Infectious Diseases and Hospital
Epidemiology, Cantonal Hospital St Gallen, St Gallen, Switzerland. 399GCAT-Genomes for Life,
Germans Trias i Pujol Health Sciences Research Institute (IGTP), Badalona, Spain. 400Catalan
Institute of Oncology, Bellvitge Biomedical Research Institute, Consortium for Biomedical
Research in Epidemiology and Public Health, University of Barcelona, Barcelona, Spain.
401Barcelona Supercomputing Center, Centro Nacional de Supercomputación (BSC-CNS), Life
& Medical Sciences, Barcelona, Spain. 404University of Siena, DIISM-SAILAB, Siena, Italy.
405Université Côte d’Azur, Inria, CNRS, I3S, Maasai, Nice, France. 408Division of Infectious
Diseases and Immunology, Department of Medical Sciences and Infectious Diseases,
Fondazione IRCCS Policlinico San Matteo, Pavia, Italy. 409Department of Internal Medicine and
Therapeutics, University of Pavia, Pavia, Italy. 410Department of Infectious and Tropical
Diseases, University of Brescia and ASST Spedali Civili Hospital, Brescia, Italy. 411Chirurgia
Vascolare, Ospedale Maggiore di Crema, Crema, Italy. 412III Infectious Diseases Unit,
ASST-FBF-Sacco, Milan, Italy. 413Department of Biomedical and Clinical Sciences Luigi Sacco,
University of Milan, Milan, Italy. 414Department of Specialized and Internal Medicine, Tropical
and Infectious Diseases Unit, Azienda Ospedaliera Universitaria Senese, Siena, Italy. 415Unit of
Respiratory Diseases and Lung Transplantation, Department of Internal and Specialist
Medicine, University of Siena, Siena, Italy. 416Department of Emergency and Urgency,
Medicine, Surgery and Neurosciences, Unit of Intensive Care Medicine, Siena University
Hospital, Siena, Italy. 417Department of Medical, Surgical and Neurosciences and Radiological
Sciences, Unit of Diagnostic Imaging, University of Siena, Siena, Italy. 418Rheumatology Unit,
Department of Medicine, Surgery and Neurosciences, University of Siena, Policlinico Le
Scotte, Siena, Italy. 419Department of Specialized and Internal Medicine, Infectious Diseases
Unit, San Donato Hospital Arezzo, Arezzo, Italy. 420Department of Emergency, Anesthesia Unit,
San Donato Hospital, Arezzo, Italy. 421Department of Specialized and Internal Medicine,
Pneumology Unit and UTIP, San Donato Hospital, Arezzo, Italy. 422Department of Emergency,
Anesthesia Unit, Misericordia Hospital, Grosseto, Italy. 423Department of Specialized and
Internal Medicine, Infectious Diseases Unit, Misericordia Hospital, Grosseto, Italy.
424Department of Preventive Medicine, Azienda USL Toscana Sud Est, Arezzo, Italy. 425Clinical
Chemical Analysis Laboratory, Misericordia Hospital, Grosseto, Italy. 426Territorial Scientiic
Technician Department, Azienda USL Toscana Sud Est, Arezzo, Italy. 427Clinical Chemical
Analysis Laboratory, San Donato Hospital, Arezzo, Italy. 428Department of Health Sciences,
Clinic of Infectious Diseases, ASST Santi Paolo e Carlo, University of Milan, Milan, Italy.
429Department of Anesthesia and Intensive Care, University of Modena and Reggio Emilia,
Modena, Italy. 430HIV/AIDS Department, National Institute for Infectious Diseases, IRCCS,
Lazzaro Spallanzani, Rome, Italy. 431Infectious Diseases Clinic, Department of Medicine,
Azienda Ospedaliera di Perugia, Perugia, Italy. 432Infectious Diseases Clinic, Santa Maria
Hospital, University of Perugia, Perugia, Italy. 433Department of Infectious Diseases, Treviso
Hospital, Treviso, Italy. 434Clinical Infectious Diseases, Mestre Hospital, Venezia, Italy.
435Infectious Diseases Clinic, ULSS1, Belluno, Italy. 436Medical Genetics and Laboratory of
Medical Genetics Unit, A.O.R.N “Antonio Cardarelli”, Naples, Italy. 437Department of Molecular
Medicine and Medical Biotechnology, University of Naples Federico II, Naples, Italy. 438CEINGE
Biotecnologie Avanzate, Naples, Italy. 439IRCCS SDN, Naples, Italy. 440Unit of Respiratory
Physiopathology, AORN dei Colli, Monaldi Hospital, Naples, Italy. 441Division of Medical
Genetics, Fondazione IRCCS Casa Sollievo della Sofferenza Hospital, San Giovanni Rotondo,
Italy. 442Department of Medical Sciences, Fondazione IRCCS Casa Sollievo della Sofferenza
Hospital, San Giovanni Rotondo, Italy. 443Infectious Diseases Clinic, Policlinico San Martino
Hospital, IRCCS for Cancer Research, Genova, Italy. 444Microbiology, Fondazione Policlinico
Universitario Agostino Gemelli IRCCS, Catholic University of Medicine, Rome, Italy.
445Department of Laboratory Sciences and Infectious Diseases, Fondazione Policlinico
Universitario AGemelli IRCCS, Rome, Italy. 446Department of Cardiovascular Diseases,
University of Siena, Siena, Italy. 447Otolaryngology Unit, University of Siena, Siena, Italy.
448Department of Internal Medicine, ASST Valtellina e Alto Lario, Sondrio, Italy. 449First Aid
Department, Luigi Curto Hospital, Polla, Italy. 450U.O.C. Laboratorio di Genetica Umana,
Genova, Italy. 451Infectious Diseases Clinics, University of Modena and Reggio Emilia, Modena,
Italy. 452Department of Respiratory Diseases, Azienda Ospedaliera di Cremona, Cremona,
Italy. 453U.O.C. Medicina, ASST Nord Milano, Ospedale Bassini, Milan, Italy. 454Department of
Cardiovascular, Neural and Metabolic Sciences, Istituto Auxologico Italiano, IRCCS, San Luca
Hospital, Milan, Italy. 455Department of Medicine and Surgery, University of Milano-Bicocca,
Milan, Italy. 456Center for Cardiac Arrhythmias of Genetic Origin, Istituto Auxologico Italiano,
IRCCS, Milan, Italy. 457Laboratory of Cardiovascular Genetics, Istituto Auxologico Italiano,
IRCCS, Milan, Italy. 458Unit of Infectious Diseases, ASST Papa Giovanni XXIII Hospital,
Bergamo, Italy. 459Department of Cardiology, Institute of Montescano, Istituti Clinici Scientiici
Maugeri, IRCCS, Pavia, Italy. 460Department of Cardiac Rehabilitation, Institute of Tradate (VA),
Istituti Clinici Scientiici Maugeri, IRCCS, Pavia, Italy. 461Cardiac Rehabilitation Unit,
Fondazione Salvatore Maugeri, IRCCS, Scientiic Institute of Milan, Milan, Italy. 462IRCCS
CMondino Foundation, Pavia, Italy. 463Medical Genetics Unit, Meyer Children’s University
Hospital, Florence, Italy. 464Department of Medicine, Pneumology Unit, Misericordia Hospital,
Grosseto, Italy. 465Department of Preventive Medicine, Azienda USL Toscana Sud Est, Arezzo,
Italy. 466Department of Anesthesia and Intensive Care Unit, ASST Fatebenefratelli Sacco, Luigi
Sacco Hospital, Polo Universitario, University of Milan, Milan, Italy. 467Health Management,
Azienda USL Toscana Sudest, Arezzo, Italy. 468Department of Mathematics, University of Pavia,
Pavia, Italy. 469Independent researcher, Milan, Italy. 470Scuola Normale Superiore, Pisa, Italy.
471CNR-Consiglio Nazionale delle Ricerche, Istituto di Biologia e Biotecnologia Agraria (IBBA),
Milano, Italy. 473Veos Digital, Milan, Italy. 475Core Research Laboratory, ISPRO, Florence, Italy.
478Division of Infectious Diseases and Immunology, Fondazione IRCCS Policlinico San Matteo,
Pavia, Italy. 479Department of Molecular and Translational Medicine, University of Brescia,
Brescia, Italy. 480Clinical Chemistry Laboratory, Cytogenetics and Molecular Genetics Section,
Diagnostic Department, ASST Spedali Civili di Brescia, Brescia, Italy. 481Department of Medical
and Surgical Sciences for Children and Adults, University of Modena and Reggio Emilia,
Modena, Italy. 482Department of Molecular Medicine, University of Padova, Padua, Italy.
483Laboratory of Regulatory and Functional Genomics, Fondazione IRCCS Casa Sollievo della
Sofferenza, San Giovanni Rotondo, Italy. 484Clinical Trial Ofice, Fondazione IRCCS Casa
Sollievo della Sofferenza Hospital, San Giovanni Rotondo, Italy. 485Department of Health
Sciences, University of Genova, Genova, Italy. 486Oncologia Medica e Uficio Flussi Sondrio,
Sondrio, Italy. 487Local Health Unit, Pharmaceutical Department of Grosseto, Toscana Sud Est
Local Health Unit, Grosseto, Italy. 488Independent researcher, Milan, Italy. 489Direzione
Scientiica, Istituti Clinici Scientiici Maugeri IRCCS, Pavia, Italy. 490Fondazione per la ricerca
Ospedale di Bergamo, Bergamo, Italy. 491Allelica, New York, NY, USA. 493School of Basic and
Medical Biosciences, Faculty of Life Sciences and Medicine, King’s College London, London,
UK. 494Medical and Population Genomics, Wellcome Sanger Institute, Hinxton, UK. 495Bradford
Institute for Health Research, Bradford Teaching Hospitals National Health Service (NHS)
Foundation Trust, Bradford, UK. 497Institute of Population Health Sciences, Queen Mary
University of London, London, UK. 498Genes & Health, Blizard Institute, Queen Mary University
of London, London, UK. 499Institute of Population Health Sciences, Queen Mary University of
London, London, UK. 500Department of Biostatistics, University of Michigan, Ann Arbor, MI,
USA. 501Heart Institute (InCor), University of Sao Paulo Med School, São Paulo, Brazil.
502Genentech, San Francisco, CA, USA. 503DNA Link Inc., Seoul, Republic of Korea. 504Seoul
National University Hospital Gangnam Center, Seoul, Republic of Korea. 505Division of
Infectious Diseases, Department of Internal Medicine, Chungnam National University School
of Medicine, Daejeon, Republic of Korea. 506East Kent Hospitals NHS Foundation Trust,
Canterbury, UK. 507Department of Internal Medicine, School of Medicine, Kyungpook National
University, Daegu, Republic of Korea. 508Division of Infectious Diseases, Department of
Internal Medicine, Incheon Medical Center, Incheon, Republic of Korea. 509Department of
Infectious Diseases, Keimyung University Dongsan Hospital, Keimyung University School of
Medicine, Daegu, Republic of Korea. 510Department of Internal Medicine, Pusan National
University School of Medicine and Medical Research Institute, Pusan National University
Hospital, Busan, Republic of Korea. 511Division of Infectious Diseases, Department of Internal
Medicine, Myongji Hospital, Goyang, Republic of Korea. 512Institute for Health Promotion,
Graduate School of Public Health, Yonsei University, Seoul, Republic of Korea. 513Division of
Cardiovascular Medicine, Stanford University, Stanford, CA, USA. 514Department of Medicine,
Stanford University, Stanford, CA, USA. 515Department of Genetics, Stanford University,
Stanford, CA, USA. 516Department of Biomedical Data Science, Stanford University, Stanford,
CA, USA. 518Department of Pathology, Stanford University, Stanford, CA, USA. 519Illumina, San
Diego, CA, USA. 520Computational Biology, Drug Discovery Sciences, Takeda
Pharmaceuticals, Boston, MA, USA. 521Department of Computational Biology, Swiss Institute
of Bioinformatics (SIB), University of Lausanne, Lausanne, Switzerland. 523Royal Victoria
Hospital, Belfast, UK. 524Chelsea & Westminster NHS Foundation Trust, London, UK.
525Northampton General Hospital NHS Trust, Northampton, UK. 526Wrexham Maelor Hospital,
Wrexham, UK. 527University College Dublin, St Vincent’s University Hospital, Dublin, Ireland.
528University Hospitals Coventry & Warwickshire NHS Trust, Coventry, UK. 529Watford General
Hospital, Watford, UK. 530NIHR Health Protection Research Unit, Institute of Infection,
Veterinary and Ecological Sciences, Faculty of Health and Life Sciences, University of
Liverpool, Liverpool, UK. 531Queen Alexandra Hospital (Hampshire), Portsmouth Hospital
Trust, Portsmouth, UK. 532Princess Royal Hospital, Brighton & Sussex Universities Hospitals
NHS Trust, Brighton, UK. 533Bassettlaw Hospital, Doncaster and Bassetlaw, Worksop, UK.
534Darent Valley Hospital, Dartford & Gravesham NHS Trust, Dartford, UK. 535High Containment
Laboratories, University of Birmingham, Birmingham, UK. 536Queen Elizabeth the Queen
Mother Hospital, Margate, UK. 537John Radcliffe Hospital, Oxford University Hospitals NHS
Foundation Trust, Oxford, UK. 538Royal Albert Edward Inirmary (Wigan), Wrightington, Wigan
and Leigh, Wigan, UK. 539Manchester Royal Inirmary, Manchester University Hospitals NHS
Foundation Trust, Manchester, UK. 540Furness General Hospital, Morecambe Bay NHS
Foundation Trust, Barrow-in-Furness, UK. 541Castle Hill Hospital, Hull University Teaching
Hospital Trust, Hull, UK. 542Hillingdon Hospital, Hillingdon Hospital, London, UK. 543St Thomas
Hospital, Guys and St Thomas Foundation Trust, London, UK. 544University Hospitals Coventry
and Warwickshire, Coventry, UK. 545St Michaels Hospital (Bristol), University Hospitals Bristol
and Weston NHS Foundation Trust, Bristol, UK. 546Stepping Hill Hospital, Stockport NHS
Foundation Trust, Manchester, UK. 547Royal Liverpool Hospital, Liverpool University Hospitals
NHS Foundation Trust, Liverpool, UK. 548Bristol Royal Hospital (Children’s), University
Hospitals Bristol and Weston NHS Foundation Trust, Bristol, UK. 549Scarborough Hospital, York
Teaching Hospitals NHS Foundation Trust, York, UK. 550Liverpool Heart & Chest Hospital,
Liverpool Heart & Chest NHS Foundation Trust, Liverpool, UK. 551James Paget University
Hospital, James Paget University Hospitals NHS Foundation Trust, Great Yarmouth, UK. 552The

Article
James Cook University Hospital, South Tees NHS Foundation Trust, Middlesbrough, UK.
553Aberdeen Royal Inirmary, Grampian, Aberdeen, UK. 554University of Edinburgh, Edinburgh,
UK. 555Royal Devon and Exeter Hospital, Royal Devon and Exeter NHS Foudation Trust, Exeter,
UK. 556Worcestershire Royal Hospital, Worcestershire Acute Hospitals NHS Trust, Worcester,
UK. 557Conquest Hospital, Hastings, East Sussex Healthcare NHS Trust, Seaford, UK. 558Dorset
County Hospital, Dorset County Hospital NHS Foundation Trust, Dorchester, UK. 559Royal
Bournemouth General Hospital, University Hospitals Dorset NHS Foundation Trust,
Bournemouth, UK. 560Harrogate Hospital, Harrogate and District NHS Foundation Trust,
Harrogate, UK. 561Burnley General Teaching Hospital, East Lancashire Hospitals NHS
Hospitals, Burnley, UK. 562Torbay Hospital, Torbay & South Devon NHS Foundation Trust,
Torquay, UK. 563Royal Hallamshire Hospital, Shefield Teaching Hospitals NHS Foundation
Trust, Shefield, UK. 564St Georges Hospital (Tooting), St Georges University Hospitals NHS
Foundation Trust, London, UK. 565Blackpool Victoria Hospital, Blackpool Teaching Hospitals
NHS Foundation Trust, Blackpool, UK. 566The Royal London Hospital, Barts Health NHS Trust,
London, UK. 567Salford Royal NHS Foundation Trust, Salford Royal NHS Foundation Trust,
Manchester, UK. 568University Hospital of North Durham, County Durham and Darlington
Foundation Trust, Durham, UK. 569Norfolk and Norwich University Hospital, Norfolk and
Norwich University Hospital NHS Foundation Trust, Norwich, UK. 572Fairield General Hospital,
Pennine Acute Hospitals NHS Trust, Manchester, UK. 573Hereford County Hospital, Wye Valley
NHS Trust, Hereford, UK. 574Southampton General Hospital, University Hospital Southampton
NHS Foundation Trust, Southampton, UK. 575Northampton General Hospital, Northampton
General Hospital NHS Trust, Northampton, UK. 576University Hospital of Wales, Cardiff and
Vale University Health Board, Cardiff, UK. 577University of Bristol, Bristol, UK. 578Leighton
Hospital, Mid Cheshire Hospitals NHS Foundation Trust, Crewe, UK. 579Diana Princess of Wales
Hospital (Grimsby), North Lincolnshire & Goole, Grimsby, UK. 580Manor Hospital, Walsall
Healthcare NHS Trust, Walsall, UK. 581Addenbrookes Hospital, Cambridge University Hospital
NHS Foundation Trust, Cambridge, UK. 582West Suffolk Hospital, West Suffolk Hospital NHS
Foundation Trust, Bury St Edmunds, UK. 583Basingstoke and North Hampshire Hospital,
Hampshire Hospitals NHS Foundation Trust, Basingstoke, UK. 584North Cumbria Integrated
Care NHS Foundation Trust, Carlisle, UK. 585Warwick Hospital, South Warwickshire NHS
Foundation Trust, Warwick, UK. 586Birmingham Women’s and Children’s Hospital, Birmingham
Women’s and Children’s Hospital NHS Foundation Trust, Birmingham, UK. 587Nottingham City
Hospital, Nottingham University Hospitals NHS Trust, Nottingham, UK. 588Glangwili Hospital
Child Health Section, Hywel Dda University Health Board, Carmarthen, UK. 589Alder Hey
Children’s Hospital, Alder Hey Children’s NHS Foundation Trust, Liverpool, UK. 590Bronglais
General Hospital, Hywel Dda University Health Board, Aberystwyth, UK. 591Worthing Hospital,
Western Sussex Hospitals NHS Foundation Trust, Worthing, UK. 592Rotheram District General
Hospital, The Rotheram NHS Foundation Trust, Rotherham, UK. 593Royal Free Hospital, Royal
Free London NHS Foundation Trust, London, UK. 594Homerton Hospital, Homerton University
Hospital NHS Foundation Trust, London, UK. 595Airedale Hospital, Airedale NHS Foundation
Trust, Keighley, UK. 596Basildon Hospital, Basildon and Thurrock University Hospitals NHS
Foundation Trust, Basildon, UK. 597The Christie NHS Foundation Trust, Manchester, UK.
598Queen Elizabeth Hospital (Greenwich), Lewisham and Greenwich NHS Trust, London, UK.
599The Whittington Hospital, Whittington Health NHS Trust, London, UK. 600Shefield
Children’s Hospital, Shefield Children’s NHS Foundation Trust, Shefield, UK. 601Royal United
Hospital, Bath, Royal United Hospitals Bath NHS Foundation Trust, Bath, UK. 602Western
General Hospital, Edinburgh, UK. 603Mid and South Essex NHS Foundation Trust, Basildon, UK.
604Hinchingbrooke Hospital, North West Anglia NHS Foundation Trust, Peterborough, UK.
605Royal Preston Hospital, Lancashire Teaching Hospitals NHS Foundation Trust, Preston, UK.
606University Hospital (Coventry), University Hospitals Coventry and Warwickshire, Coventry,
UK. 607The Walton Centre, The Walton Centre, Liverpool, UK. 608Hull Royal Inirmary, Hull
University Teaching Hospital Trust, Hull, UK. 609Darlington Memorial Hospital, County Durham
and Darlington Foundation Trust, Darlington, UK. 610Queen Elizabeth Hospital (Gateshead),
Gateshead NHS Foundation Trust, Newcastle, UK. 611Warrington Hospital, Warrington & Halton
Hospitals NHS Foundation Trust, Warrington, UK. 612University Hospitals Bristol and Weston
NHS Foundation Trust, Bristol, UK. 613St Mary’s Hospital (Isle of Wight), Isle of Wight NHS Trust,
Isle of Wight, UK. 614The Maidstone Hospital, Maidstone & Tunbridge Wells NHS Trust,
Maidstone, UK. 615Huddersield Royal, Calderdale and Huddersield NHS Foundation Trust,
Huddersield, UK. 616Royal Surrey County Hospital, Guildford, UK. 617Countess of Chester
Hospital, Countess of Chester Hospital NHS Foundation Trust, Chester, UK. 618Frimley Park
Hospital, Frimley Health Foundation Trust, Frimley, UK. 620Leeds General Inirmary, Leeds
Teaching Hospitals, Leeds, UK. 621North Middlesex Hospital, North Middlesex University
Hospital NHS Trust, London, UK. 622Arrowe Park Hospital, Wirral University Teaching Hospital
NHS Foundation Trust, Wirral, UK. 623Great Ormond Street Hospital, Great Ormond Street
Hospital for Children NHS Foundation Trust, London, UK. 624Royal Shrewsbury Hospital,
Shrewsbury and Telford Hospital NHS Trust, Shrewsbury, UK. 625East Surrey Hospital (Redhill),
Surrey & Sussex Healthcare, Redhill, UK. 626Burton Hospital, University Hospitals of Derby &
Burton NHS Foundation Trust, Burton-on-Trent, UK. 627Kent and Canterbury Hospital, East Kent
Hospitals NHS Foundation Trust, Canterbury, UK. 628Weston Area General Trust, University
Hospitals Bristol and Weston NHS Foundation Trust, Bristol, UK. 629Luton and Dunstable
University Hospital, Luton, UK. 630Glasgow Royal Inirmary, Greater Glasgow and Clyde,
Glasgow, UK. 631Derbyshire Healthcare, Derbyshire Healthcare NHS Foundation Trust, Derby,
UK. 632Macclesield General Hospital, East Cheshire NHS Foundation Trust, Macclesield, UK.
633Chelsea and Westminster Hospital, Chelsea and Westminster NHS Trust, London, UK.
634Institute of Microbiology and Infection, University of Birmingham, Birmingham, UK.
635Prince Philip Hospital, Hwyel Dda University Health Board, Llanelli, UK. 636George Eliot
Hospital - Acute Services, George Eliot Hospital, Nuneaton, UK. 637Kettering General Hospital,
Kettering General Hospital NHS Foundation Trust, Kettering, UK. 638Birmingham Heartlands
Hospital, Birmingham, UK. 639Russells Hall Hospital, The Dudley Group NHS Foundation Trust,
Dudley, UK. 640Hareield Hospital, Royal Brompton & Hareield Trust, London, UK. 641Lister
Hospital, East and North Hertfordshire NHS Trust, Stevenage, UK. 642Musgrove Park Hospital
(Taunton & Somerset), Somerset NHS Foundation Trust, Taunton, UK. 643Queen’s Hospital,
Havering (Romford), Barking, Havering and Redbridge University Hospitals NHS Trust,
London, UK. 644Southport & Formby District General Hospital, Southport and Ormskirk
Hospital NHS Trust, Southport, UK. 645New Cross Hospital, The Royal Wolverhampton NHS
Trust, Wolverhampton, UK. 646King’s College Hospital, London, UK. 647The Royal Victoria
Inirmary, Newcastle Hospitals NHS Trust, Newcastle, UK. 648The Great Western Hospital,
Great Western Hospitals NHS Foundation Trust, Swindon, UK. 649Ninewells Hospital, Tayside,
Dundee, UK. 650Poole Hospital NHS Trust, Poole, UK. 651Burton Hospital, University Hospitals of
Derby & Burton NHS Foundation Trust, Derby, UK. 652William Harvey Hospital, Ashford, East
Kent Hospitals NHS Foundation Trust, Willesborough, UK. 653King’s Mill Hospital, Sherwood
Forest Hospitals NHS Foundation Trust, Sutton-in-Ashield, UK. 654Liverpool Women’s NHS
Foundation Trust, Liverpool, UK. 655Dewsbury Hospital, Mid Yorkshire Hospitals NHS Trust,
Dewsbury, UK. 656Northern Devon District Hospital, Northern Devon Healthcare NHS Trust,
Barnstaple, UK. 657Tameside General Hospital, Tameside and Glossop Integrated Care NHS
Foundation Trust, Manchester, UK. 658Sandwell General Hospital, Sandwell and West
Birmingham Hospitals NHS Trust, Birmingham, UK. 659Broomield Hospital, Mid and South
Essex University Hospitals Group, Broomield, UK. 660Wycombe Hospital, Buckingham
Healthcare NHS Trust, Wycombe, UK. 661University Hospital of North Tees, North Tees and
Hartlepool NHS Trust, Stockton-on-Tees, UK. 662Royal Manchester Children’s Hospital,
Manchester University Hospitals NHS Foundation Trust, Manchester, UK. 663Bedford Hospital,
Bedford, UK. 664Colchester General Hospital, East Suffolk and North Essex Foundation Trust,
Colchester, UK. 665Queen Elizabeth Hospital (Birmingham) and Heartlands, University Hospital
Birmingham NHS Foundation Trust, Birmingham, UK. 666Chesterield Royal Hospital,
Chesterield Royal Hospital NHS Foundation Trust, Chesterield, UK. 667Princess Alexandra
Hospital, The Princess Alexandra Hospital NHS Trust, Harlow, UK. 668Watford General Hospital,
West Hertfordshire Hospitals NHS Trust, Watford, UK. 669Milton Keynes Hospital, Milton
Keynes University Hospital NHS Foundation Trust, Milton Keynes, UK. 670Royal Bolton General
Hospital, Bolton Foundation Trust, Bolton, UK. 671Royal Gwent (Newport), Aneurin Bevan
University Health Board, Newport, UK. 672The Royal Marsden Hospital (London), The Royal
Marsden NHS Foundation Trust, London, UK. 673Queen Victoria Hospital (East Grinstead),
Queen Victoria Hospital NHS Foundation Trust, East Grinstead, UK. 674County Hospital
(Stafford), University Hospitals of North Midlands NHS Trust, Stafford, UK. 675Whiston Hospital,
St Helen’s & Knowlsey Hospitals NHS Trust, Prescot, UK. 676Croydon University Hospital,
London, UK. 677Gloucester Royal, Gloucestershire Hospitals NHS Foundation Trust,
Gloucester, UK. 678Medway Maritime Hospital, Medway Maritime NHS Trust, Gillingham, UK.
679Royal Papworth Hospital Everard, Royal Papworth Hospital NHS Foundation Trust,
Cambridge, UK. 680Derriford (Plymouth), University Hospital Plymouth NHS Trust, Plymouth,
UK. 681St Helier Hospital, Epsom and St Helier University Hospital NHS Trust, London, UK.
682Royal Berkshire Hospital, Royal Berkshire Foundation Trust, London, UK. 683Bradford Royal
Inirmary, Bradford Teaching Hospitals NHS Foundation Trust, Bradford, UK. 684Northwick Park,
London North West University Hospital Trust, London, UK. 685Ealing Hospital, London North
West University Hospital Trust, London, UK. 686Royal Cornwall Hospital (Tresliske), Royal
Cornwall NHS Trust, Truro, UK. 687Ashford Hospital, Ashford & St Peter’s Hospital, Stanwell,
UK. 688Leicester Royal Inirmary (includes Glenield Site), University Hospitals of Leicester,
Leicester, UK. 689Grantham and District Hospital, United Lincolnshire Hospitals NHS Trust,
Grantham, UK. 690University Hospital Aintree, Liverpool University Hospitals NHS Foundation
Trust, Liverpool, UK. 691North Tyneside General Hospital, Northumbria Healthcare NHS Trust,
North Shields, UK. 692Queen Elizabeth Hospital (King’s Lynn), Queen Elizabeth Hospital King’s
Lynn NHS Foundation Trust, King’s Lynn, UK. 693The Crick Institute, London, UK. 695William
Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen
Mary University of London, London, UK. 696Centre for Genomic and Experimental Medicine,
Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General
Hospital, Edinburgh, UK. 697Intensive Care National Audit & Research Centre, London, UK.
700Wellcome Centre for Human Genetics, University of Oxford, Oxford, UK. 705Centre for
Inlammation Research, The Queen’s Medical Research Institute, University of Edinburgh,
Edinburgh, UK. 706Great Ormond Street Hospital for Children NHS Foundation Trust, London,
UK. 707Biostatistics Group, School of Life Sciences, Sun Yat-sen University, Guangzhou, China.
708Centre for Global Health Research, Usher Institute of Population Health Sciences and
Informatics, Edinburgh, UK. 709Department of Medical Epidemiology and Biostatistics,
Karolinska Institutet, Stockholm, Sweden. 710Institute for Molecular Bioscience, The University
of Queensland, Brisbane, Queensland, Australia. 711School of Life Sciences, Westlake
University, Hangzhou, China. 712Westlake Laboratory of Life Sciences and Biomedicine,
Westlake University, Hangzhou, China. 714Centre for Medical Informatics, The Usher Institute,
University of Edinburgh, Edinburgh, UK. 715Liverpool Clinical Trials Centre, University of
Liverpool, Liverpool, UK. 716Centre for Health Informatics, Division of Informatics, Imaging and
Data Science, School of Health Sciences, Faculty of Biology, Medicine and Health, University
of Manchester, Manchester Academic Health Science Centre, Manchester, UK. 717MRC Human
Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh,
Edinburgh, UK. 718School of Informatics, University of Edinburgh, Edinburgh, UK. 719Royal
Hospital for Children, Glasgow, UK. 721MRC-University of Glasgow Centre for Virus Research,
Institute of Infection, Immunity and Inlammation, College of Medical, Veterinary and Life
Sciences, University of Glasgow, Glasgow, UK. 722Centre for Tropical Medicine and Global
Health, Nufield Department of Medicine, University of Oxford, Oxford, UK. 724Department of
Anaesthesia and Intensive Care, The Chinese University of Hong Kong, Prince of Wales
Hospital, Hong Kong, China. 725Department of Critical Care Medicine, Queen’s University and
Kingston Health Sciences Centre, Kingston, Ontario, Canada. 726Wellcome-Wolfson Institute
for Experimental Medicine, Queen’s University Belfast, Belfast, UK. 727Department of Intensive
Care Medicine, Royal Victoria Hospital, Belfast, UK. 728UCL Centre for Human Health and
Performance, London, UK. 729Clinical Research Centre at St Vincent’s University Hospital,
University College Dublin, Dublin, Ireland. 730National Heart and Lung Institute, Imperial
College London, London, UK. 731Imperial College Healthcare NHS Trust London, London, UK.
732NIHR Health Protection Research Unit for Emerging and Zoonotic Infections, Institute of
Infection, Veterinary and Ecological Sciences University of Liverpool, Liverpool, UK.
733Respiratory Medicine, Alder Hey Children’s Hospital, Institute in The Park, University of
Liverpool, Alder Hey Children’s Hospital, Liverpool, UK. 734Department of Intensive Care
Medicine, Guy’s and St Thomas NHS Foundation Trust, London, UK. 735Department of
Medicine, University of Cambridge, Cambridge, UK. 736Airedale General Hospital, Keighley,
UK. 737Barts Health NHS Trust, London, UK. 738Basildon Hospital, Basildon, UK. 739BHRUT
(Barking Havering) - Queens Hospital and King George Hospital, Romford, UK. 740Bradford
Royal Inirmary, Bradford, UK. 741Bronglais General Hospital, Aberystwyth, UK. 74 2Broomield
Hospital, Chelmsford, UK. 743Calderdale Royal Hospital, Halifax, UK. 744Charing Cross Hospital,
St Mary’s Hospital and Hammersmith Hospital, London, UK. 745Barnet Hospital, London, UK.
746Birmingham Children’s Hospital, Birmingham, UK. 747St John’s Hospital Livingston,

Livingston, UK. 748Aberdeen Royal Inirmary, Aberdeen, UK. 749Addenbrooke’s Hospital,
Cambridge, UK. 750Aintree University Hospital, Liverpool, UK. 752Arrowe Park Hospital, Wirral,
UK. 753Ashford and St Peter’s Hospital, Lyne, UK. 754Basingstoke and North Hampshire Hospital,
Basingstoke, UK. 755Borders General Hospital, Melrose, UK. 756Chesterield Royal Hospital
Foundation Trust, Chesterield, UK. 757Eastbourne District General Hospital, East Sussex, UK
and Conquest Hospital, Eastbourne, UK. 758Barnsley Hospital, Barnsley, UK. 759Blackpool
Victoria Hospital, Blackpool, UK. 760East Surrey Hospital, Redhill, UK. 761Good Hope Hospital,
Birmingham, UK. 762Hereford County Hospital, Hereford, UK. 763Hull Royal Inirmary, Hull, UK.
765Kent & Canterbury Hospital, Canterbury, UK. 766Manchester Royal Inirmary, Manchester,
UK. 767Nottingham University Hospital, Nottingham, UK. 768Pilgrim Hospital, Lincoln, UK.
769Queen Elizabeth Hospital, Birmingham, UK. 770Salford Royal Hospital, Manchester, UK.
771Tameside General Hospital, Ashton-under-Lyne, UK. 772The Tunbridge Wells Hospital and
Maidstone Hospital, Maidstone, UK. 773The Royal Oldham Hospital, Manchester, UK. 774The
Royal Papworth Hospital, Cambridge, UK. 775University College Hospital, London, UK.
776Withybush General Hospital, Haverfordwest, UK. 777Wythenshawe Hospital, Manchester, UK.
778Yeovil Hospital, Yeovil, UK. 779Cumberland Inirmary, Carlisle, UK. 780Darent Valley Hospital,
Dartford, UK. 781Dumfries and Galloway Royal Inirmary, Dumfries, UK. 782Ealing Hospital,
London, UK. 783Fairield General Hospital, Bury, UK. 784George Eliot Hospital NHS Trust,
Nuneaton, UK. 785Glan Clwyd Hospital, Bodelwyddan, UK. 786Glangwili General Hospital,
Camarthen, UK. 787The Great Western Hospital, Swindon, UK. 788Guys and St Thomas’ Hospital,
London, UK. 789Hareield Hospital, London, UK. 790Harrogate and District NHS Foundation
Trust, Harrogate, UK. 792James Paget University Hospital NHS Trust, Great Yarmouth, UK.
794King’s Mill Hospital, Nottingham, UK. 795Kingston Hospital, Kingston, UK. 796Lincoln County
Hospital, Lincoln, UK. 797Liverpool Heart and Chest Hospital, Liverpool, UK. 798Macclesield
District General Hospital, Macclesield, UK. 799Medway Maritime Hospital, Gillingham, UK.
800Milton Keynes University Hospital, Milton Keynes, UK. 801Morriston Hospital, Swansea, UK.
802National Hospital for Neurology and Neurosurgery, London, UK. 803Norfolk and Norwich
University hospital (NNUH), Norwich, UK. 804North Middlesex University Hospital NHS Trust,
London, UK. 806Northumbria Healthcare NHS Foundation Trust, North Shields, UK.
807Peterborough City Hospital, Peterborough, UK. 808Prince Charles Hospital, Merthyr Tydil,
UK. 809Royal Sussex County Hospital, Brighton, UK. 810Princess Royal Hospital, Haywards
Heath, UK. 811Princess of Wales Hospital, Llantrisant, UK. 812Queen Alexandra Hospital,
Portsmouth, UK. 813Queen Elizabeth Hospital, London, UK. 815Queen Victoria Hospital, East
Grinstead, UK. 816Queen’s Hospital Burton, Burton-On-Trent, UK. 817Raigmore Hospital,
Inverness, UK. 818Rotherham General Hospital, Rotherham, UK. 819Royal Blackburn Teaching
Hospital, Blackburn, UK. 820Royal Preston Hospital, Preston, UK. 821Royal Surrey County
Hospital, Guildford, UK. 822Royal Albert Edward Inirmary, Wigan, UK. 823The Royal Alexandra
Children’s Hospital, Brighton, UK. 824Royal Alexandra Hospital, Paisley, UK. 825Royal Bolton
Hospital, Bolton, UK. 826University Hospitals Dorset NHS Foundation Trust, Dorchester, UK.
827Royal Brompton Hospital, London, UK. 828Imperial College London, London, UK. 829Royal
Cornwall Hospital, Truro, UK. 830Royal Free Hospital, London, UK. 831Royal Glamorgan Hospital,
Pontyclun, UK. 832Royal Gwent Hospital, Newport, UK. 833Northern General Hospital, Shefield,
UK. 834Royal Hampshire County Hospital, Winchester, UK. 835Royal Manchester Children’s
Hospital, Manchester, UK. 836Royal Stoke University Hospital, Stoke-on-Trent, UK. 837Salisbury
District Hospital, Salisbury, UK. 838Sandwell General Hospital, Birmingham, UK.
839Scarborough General Hospital, Scarborough, UK. 840Scunthorpe General Hospital,
Scunthorpe, UK. 841Southmead Hospital, Bristol, UK. 842St George’s Hospital, London, UK. 843St
Mary’s Hospital, Newport, UK. 844Stoke Mandeville Hospital, Aylesbury, UK. 845Sunderland
Royal Hospital, Sunderland, UK. 846Alexandra Hospital, Redditch and Worcester Royal
Hospital, Worcester, UK. 847The Christie NHS Foundation Trust, Manchester, UK. 848The Queen
Elizabeth Hospital, King’s Lynn, UK. 849The Royal Liverpool University Hospital, Liverpool, UK.
850The Royal Marsden NHS Foundation Trust, London, UK. 851Torbay Hospital, Torquay, UK.
852University Hospital Monklands, Airdrie, UK. 853University Hospital Lewisham, London, UK.
854University Hospital North Durham, Darlington, UK. 855University Hospital of North Tees,
Stockton-on-Tees, UK. 856University Hospital of Wales, Cardiff, UK. 857University Hospital
Wishaw, Wishaw, UK. 858Victoria Hospital, Kirkcaldy, UK. 859Warrington General Hospital,
Warrington, UK. 860West Cumberland Hospital, Whitehaven, UK. 861Western Sussex Hospitals,
Chichester, UK. 862Whiston Hospital, Prescot, UK. 863York Hospital, York, UK. 864Ysbyty
Gwynedd, Bangor, UK. 865Countess of Chester Hospital, Chester, UK. 866Croydon University
Hospital, Croydon, UK. 867Diana Princess of Wales Hospital, Grimsby, UK. 868Dorset County
Hospital, Dorchester, UK. 869Forth Valley Royal Hospital, Falkirk, UK. 870Furness General
Hospital, Barrow-in-Furness, UK. 871Alder Hey Children’s Hospital, Liverpool, UK. 872Derriford
Hospital, Plymouth, UK. 873Glasgow Royal Inirmary, Glasgow, UK. 874Glenield Hospital,
Leicester, UK. 875Gloucestershire Royal Hospital, Gloucester, UK. 876Golden Jubilee National
Hospital, Clydebank, UK. 877Great Ormond St Hospital and UCL Great Ormond St Institute of
Child Health NIHR Biomedical Research Centre, London, UK. 878Homerton University Hospital
Foundation NHS Trust, London, UK. 879James Cook University Hospital, Middlesbrough, UK.
880John Radcliffe Hospital, Oxford, UK. 881Leicester Royal Inirmary, Leicester, UK. 882Lister
Hospital, Stevenage, UK. 883New Cross Hospital, Wolverhampton, UK. 884Royal Victoria
Inirmary, Newcastle Upon Tyne, UK. 885Ninewells Hospital, Dundee, UK. 886North Devon
District Hospital, Barnstaple, UK. 887North Manchester General Hospital, Manchester, UK.
888Northwick Park Hospital, London, UK. 889Prince Philip Hospital, Lianelli, UK. 890Pinderields
General Hospital, Wakeield, UK. 891Poole Hospital, Poole, UK. 892Royal Shrewsbury Hospital,
Shrewsbury, UK. 893Princess Royal Hospital, Telford, UK. 899Queen Elizabeth Hospital
Gateshead, Gateshead, UK. 900Queen Elizabeth University Hospital, Glasgow, UK. 901Royal
Berkshire NHS Foundation Trust, Reading, UK. 902Royal Derby Hospital, Derby, UK. 903Royal
Devon and Exeter Hospital, Exeter, UK. 904Royal Inirmary of Edinburgh, Edinburgh, UK.
905Royal Lancaster Inirmary, Lancaster, UK. 906Royal United Hospital, Bath, UK. 907Russells Hall
Hospital, Dudley, UK. 908Shefield Children’s Hospital, Shefield, UK. 909Southampton General
Hospital, Southampton, UK. 910Southend University Hospital, Westcliff-on-Sea, UK.
911Southport and Formby District General Hospital, Ormskirk, UK. 912St James’s University
Hospital and Leeds General Inirmary, Leeds, UK. 913Bristol Royal Inirmary, Bristol, UK.
914Stepping Hill Hospital, Stockport, UK. 915The Princess Alexandra Hospital, Harlow, UK.
916University Hospital Crosshouse, Kilmarnock, UK. 917University Hospital Hairmyres, East
Kilbride, UK. 918Craigavon Area Hospital, Craigavon, UK. 919Warwick Hospital, Warwick, UK.
920West Middlesex Hospital, Isleworth, UK. 922Whittington Hospital, London, UK. 923William
Harvey Hospital, Ashford, UK. 932Section of Molecular Virology, Imperial College London,
London, UK. 933Antimicrobial Resistance and Hospital Acquired Infection Department, Public
Health England, London, UK. 934Department of Infectious Disease, Imperial College London,
London, UK. 935National Infection Service, Public Health England, London, UK.
936MRC-University of Glasgow Centre for Virus Research, Glasgow, UK. 937Liverpool School of
Tropical Medicine, Liverpool, UK. 938Institute of Infection and Global Health, University of
Liverpool, Liverpool, UK. 940Virology Reference Department, National Infection Service, Public
Health England, London, UK. 941Department of Pharmacology, University of Liverpool,
Liverpool, UK. 942Nufield Department of Medicine, University of Oxford, Oxford, UK.
944Nottingham University Hospitals NHS Trust, Nottingham, UK. 945Nufield Department of
Medicine, John Radcliffe Hospital, Oxford, UK. 946ISARIC Global Support Centre, Centre for
Tropical Medicine and Global Health, Nufield Department of Medicine, University of Oxford,
Oxford, UK. 947Division of Infection and Immunity, University College London, London, UK.
948Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool,
UK. 949Centre for Clinical Infection and Diagnostics Research, Department of Infectious
Diseases, School of Immunology and Microbial Sciences, King’s College London, London, UK.
950Institute of Evolutionary Biology, University of Edinburgh, Edinburgh, UK. 951Department of
Pediatrics and Virology, Imperial College London, London, UK. 952The Florey Institute for
Host-Pathogen Interactions, Department of Infection, Immunity and Cardiovascular Disease,
University of Shefield, Shefield, UK. 953Division of Structural Biology, The Wellcome Centre
for Human Genetics, University of Oxford, Oxford, UK. 955Blood Borne Virus Unit, Virus
Reference Department, National Infection Service, Public Health England, London, UK.
956Department of Infection, Immunity and Cardiovascular Disease, University of Shefield,
Shefield, UK. 957Institute for Global Health, University College London, London, UK.
958Molecular and Clinical Cancer Medicine, Institute of Systems, Molecular and Integrative
Biology, University of Liverpool, Liverpool, UK. 959Department of Child Life and Health,
University of Edinburgh, Edinburgh, UK. 960Section of Biomolecular Medicine, Division of
Systems Medicine, Department of Metabolism, Digestion and Reproduction, Imperial College
London, London, UK. 961Department of Epidemiology and Biostatistics, School of Public
Health, Faculty of Medicine, Imperial College London, London, UK. 962National Phenome
Centre, Department of Metabolism, Digestion and Reproduction, Imperial College London,
London, UK. 963Department of Molecular and Clinical Cancer Medicine, University of
Liverpool, Liverpool, UK. 964Institute of Translational Medicine, University of Liverpool,
Liverpool, UK. 965Intensive Care Unit, Royal Inirmary Edinburgh, Edinburgh, UK. 966University
of Liverpool, Liverpool, UK. 967University of Glasgow, Glasgow, UK. 968Edinburgh Clinical
Research Facility, Western General Hospital, University of Edinburgh, Edinburgh, UK.
970Department of Infectious Diseases, Leiden University Medical Center, Leiden, The
Netherlands. 971Cambridge University Hospitals NHS Foundation Trust, Cambridge, UK.
974Genotek, Moscow, Russia. 975Helix, San Mateo, CA, USA. 976Center for Genomic Medicine,
Desert Research Institute, Reno, NV, USA. 977Renown Health, Reno, NV, USA. 97824Genetics,
Boston, MA, USA. 979Hospital La Paz Institute for Health Research, Madrid, Spain. 980Division of
Pulmonary Medicine, Department of Medicine, Keio University School of Medicine, Tokyo,
Japan. 981Department of Statistical Genetics, Osaka University Graduate School of Medicine,
Suita, Japan. 982Laboratory of Statistical Immunology, Immunology Frontier Research Center
(WPI-IFReC), Osaka University, Suita, Japan. 983Integrated Frontier Research for Medical
Science Division, Institute for Open and Transdisciplinary Research Initiatives, Osaka
University, Suita, Japan. 984Division of Health Medical Intelligence, Human Genome Center,
the Institute of Medical Science, The University of Tokyo, Tokyo, Japan. 985Laboratory of Viral
Infection I, Department of Infection Control and Immunology, Ōmura Satoshi Memorial
Institute & Graduate School of Infection Control Sciences, Kitasato University, Tokyo, Japan.
986Department of Surgery, Keio University School of Medicine, Tokyo, Japan. 987Department of
Organoid Medicine, Keio University School of Medicine, Tokyo, Japan. 988Department of
Infectious Diseases, Keio University School of Medicine, Tokyo, Japan. 989Department of
Respiratory Medicine and Clinical Immunology, Osaka University Graduate School of
Medicine, Suita, Japan. 990Department of Immunopathology, Immunology Frontier Research
Center (WPI-IFReC), Osaka University, Suita, Japan. 991Institute of Research, Tokyo Medical and
Dental University, Tokyo, Japan. 992Department of Insured Medical Care Management, Tokyo
Medical and Dental University Hospital of Medicine, Tokyo, Japan. 993Genome Medical
Science Project (Toyama), National Center for Global Health and Medicine, Chiba, Japan.
994Division of Gastroenterology and Hepatology, Department of Medicine, Keio University
School of Medicine, Tokyo, Japan. 995M&D Data Science Center, Tokyo Medical and Dental
University, Tokyo, Japan. 996Department of Pathology and Tumor Biology Institute for the
Advanced Study of Human Biology (WPI-ASHBi), Kyoto University, Kyoto, Japan.
997Department of Medicine, Center for Hematology and Regenerative Medicine, Karolinska
Institute, Stockholm, Sweden. 999Department of Emergency and Critical Care Medicine, Keio
University School of Medicine, Tokyo, Japan. 1000Department of Anesthesiology, Keio
University School of Medicine, Tokyo, Japan. 1001Department of Laboratory Medicine, Keio
University School of Medicine, Tokyo, Japan. 1002Division of Infection Control and Prevention,
Osaka University Hospital, Suita, Japan. 1003Department of Biomedical Ethics and Public
Policy, Osaka University Graduate School of Medicine, Suita, Japan. 1004Center for Genomic
Medicine, Kyoto University Graduate School of Medicine, Kyoto, Japan. 1005Department of
Pulmonary Medicine, Faculty of Medicine, University of Tsukuba, Tsukuba, Japan.
1006Department of Neurosurgery, Faculty of Medicine, The University of Tokyo, Tokyo, Japan.
1007Laboratory of Immune Regulation, Department of Microbiology and Immunology, Osaka
University Graduate School of Medicine, Suita, Japan. 1009Medical Innovation Promotion
Center, Tokyo Medical and Dental University, Tokyo, Japan. 1010Clinical Research Center, Tokyo
Medical and Dental University Hospital of Medicine, Tokyo, Japan. 1011Department of Medical
Informatics, Tokyo Medical and Dental University Hospital of Medicine, Tokyo, Japan.
1012Respiratory Medicine, Tokyo Medical and Dental University, Tokyo, Japan. 1013Clinical
Laboratory, Tokyo Medical and Dental University Hospital of Medicine, Tokyo, Japan.
1014Department of Pathology and Tumor Biology, Kyoto University, Kyoto, Japan.
1015Department of Respiratory Medicine, Graduate School of Medicine, Faculty of Medicine,
Juntendo University, Tokyo, Japan. 1016Department of Emergency and Disaster Medicine,
Graduate School of Medicine, Faculty of Medicine, Juntendo University, Tokyo, Japan.
1017Department of Cardiovascular Biology and Medicine, Graduate School of Medicine,
Faculty of Medicine, Juntendo University, Tokyo, Japan. 1018Department of Respiratory
Medicine, Tokyo Women’s Medical University, Tokyo, Japan. 1019Department of General
Medicine, Tokyo Women’s Medical University, Tokyo, Japan. 1020Department of Respiratory

Article
Medicine, Saitama Cardiovascular and Respiratory Center, Saitama, Japan. 1021Kawasaki
Municipal Ida Hospital, Kanagawa, Japan. 1022Saitama Medical Center, Internal Medicine,
Japan Community Healthcare Organization (JCHO), Saitama, Japan. 1023Saitama City Hospital,
Saitama, Japan. 1024Division of Infection Control, Eiju General Hospital, Tokyo, Japan.
1025Department of Pulmonary Medicine, Eiju General Hospital, Tokyo, Japan. 1026Department of
Respiratory Medicine, Osaka Saiseikai Nakatsu Hospital, Osaka, Japan. 1027Division of
Respirology, Rheumatology, and Neurology, Department of Internal Medicine Kurume
University School of Medicine, Fukuoka, Japan. 1028Department of Infection Control, Osaka
Saiseikai Nakatsu Hospital, Osaka, Japan. 1029Department of Infectious Diseases, Tosei General
Hospital, Aichi, Japan. 1030Fukujuji Hospital, Kiyose, Japan. 1031Department of Emergency and
Critical Care Medicine, Tokyo Women’s Medical University Medical Center East, Tokyo, Japan.
1032Department of Medicine, Tokyo Women’s Medical University Medical Center East, Tokyo,
Japan. 1033Department of Pediatrics, Tokyo Women’s Medical University Medical Center East,
Tokyo, Japan. 1034Japan Community Healthcare Organization Kanazawa Hospital, Kanazawa,
Japan. 1035Division of Pulmonary Medicine, Department of Internal Medicine, Federation of
National Public Service Personnel Mutual Aid Associations, Tachikawa Hospital, Tachikawa,
Japan. 1036Department of Respiratory Medicine, Japan Organization of Occupational Health
and Safety, Kanto Rosai Hospital, Kawasaki, Japan. 1037Department of General Internal
Medicine, Japan Organization of Occupational Health and Safety, Kanto Rosai Hospital,
Kawasaki, Japan. 1038Department of Emergency and Critical Care Medicine, Kansai Medical
University General Medical Center, Kirakata, Japan. 1039Department of Respiratory Medicine,
Kitasato University, Kitasato Institute Hospital, Tokyo, Japan. 1040Ishikawa Prefectural Central
Hospital, Kanazawa, Japan. 1041Internal Medicine, Sano Kosei General Hospital, Sano, Japan.
1042Saiseikai Yokohamashi Nanbu Hospital, Yokohama, Japan. 1043Kanagawa Cardiovascular
and Respiratory Center, Yokohama, Japan. 1044Saiseikai Utsunomiya Hospital, Utsunomiya,
Japan. 1045Department of Respiratory Medicine, KKR Sapporo Medical Center, Sapporo, Japan.
1046Internal Medicine, Internal Medicine Center, Showa University Koto Toyosu Hospital, Tokyo,
Japan. 1047Department of Respiratory Medicine, Toyohashi Municipal Hospital, Toyohashi,
Japan. 1048Keiyu Hospital, Yokohama, Japan. 1049Department of Rheumatology, National
Hospital Organization Hokkaido Medical Center, Sapporo, Japan. 1050Department of
Respiratory Medicine, National Hospital Organization Tokyo Medical Center, Tokyo, Japan.
1051Department of Allergy, National Hospital Organization Tokyo Medical Center, Tokyo, Japan.
1052Department of General Internal Medicine and Infectious Diseases, National Hospital
Organization Tokyo Medical Center, Tokyo, Japan. 1053Japanese Red Cross Musashino
Hospital, Musashino, Japan. 1054Department of Respiratory Medicine, Tohoku University
Graduate School of Medicine, Sendai, Japan. 1055Division of Respiratory Medicine, Department
of Internal Medicine, Nihon University School of Medicine, Tokyo, Japan. 1056Department of
Emergency and Critical Care Medicine, St Marianna University School of Medicine, Kawasaki,
Japan. 1057Division of General Internal Medicine, Department of Internal Medicine, St Marianna
University School of Medicine, Kawasaki, Japan. 1058National Hospital Organization Kanazawa
Medical Center, Kanazawa, Japan. 1059Division of Infectious Diseases and Respiratory
Medicine, Department of Internal Medicine, National Defense Medical College, Tokorozawa,
Japan. 1060Department of Emergency and Critical Care Medicine, Faculty of Medicine,
Fukuoka University, Fukuoka, Japan. 1061Department of Infection Control, Fukuoka University
Hospital, Fukuoka, Japan. 1062Tokyo Saiseikai Central Hospital, Tokyo, Japan. 1063Department of
Internal Medicine, Fukuoka Tokushukai Hospital, Kasuga, Japan. 1064Department of Infectious
Disease and Clinical Research Institute, National Hospital Organization Kyushu Medical
Center, Fukuoka, Japan. 1065Department of Respirology, National Hospital Organization
Kyushu Medical Center, Fukuoka, Japan. 1067Matsumoto City Hospital, Matsumoto, Japan.
1068Uji-Tokushukai Medical Center, Uji, Japan. 1069Department of Respiratory Medicine, Nagoya
University Graduate School of Medicine, Nagoya, Japan. 1070Department of Respiratory
Medicine, Fujisawa City Hospital, Fujisawa, Japan. 1071Sapporo City General Hospital, Sapporo,
Japan. 1072Department of Emergency and Critical Care Medicine, Chiba University Graduate
School of Medicine, Chiba, Japan. 1073Division of Respiratory Medicine, Social Welfare
Organization Saiseikai Imperial Gift Foundation, Saiseikai Kumamoto Hospital, Kumamoto,
Japan. 1074Department of Anesthesiology and Intensive Care Medicine, Kyoto Prefectural
University of Medicine, Kyoto, Japan. 1075Ome Municipal General Hospital, Ome, Japan.
1076Hanwa Daini Hospital, Osaka, Japan. 1077Department of Respiratory Internal Medicine, St
Marianna University School of Medicine, Yokohama-City Seibu Hospital, Yokohama, Japan.
1078Division of Hematology, Department of Internal Medicine, St Marianna University
Yokohama-City Seibu Hospital, Yokohama, Japan. 1079Division of Pulmonary Medicine,
Department of Medicine, Tokai University School of Medicine, Tokai University School of
Medicine, Tokyo, Japan. 1080Division of Pulmonary Medicine, Department of Medicine, Tokai
University School of Medicine, Tokyo, Japan. 1081National Hospital Organization Kumamoto
Medical Center, Kumamoto, Japan. 1082Department of Respiratory Medicine, Tokyo Medical
University Hospital, Tokyo, Japan. 1083Department of Respiratory Medicine, Japanese Red
Cross Medical Center, Tokyo, Japan. 1084JA Toride Medical Hospital, Toride, Japan. 1085Japan
Organization of Occupational Health and Safety Okayama Rosai Hospital, Okayama, Japan.
1086Emergency and Disaster Medicine, Graduate School of Medicine, Gifu University School of
Medicine, Gifu, Japan. 1087Niigata University, Niigata, Japan. 1088National Hospital Organization
Kyoto Medical Center, Kyoto, Japan. 1089Research Institute for Diseases of the Chest, Graduate
School of Medical Sciences, Kyushu University, Fukuoka, Japan. 1090Department of Medicine
and Biosystemic Science, Kyushu University Graduate School of Medical Sciences, Fukuoka,
Japan. 1091Department of Emergency and Critical Care Medicine, Tsukuba University, Tsukuba,
Japan. 1092Department of Nephrology, Faculty of Medicine, University of Tsukuba, Tsukuba,
Japan. 1093Department of Hematology, Faculty of Medicine, University of Tsukuba, Tsukuba,
Japan. 1094National Hospital Organization Tokyo Hospital, Tokyo, Japan. 1095Fujioka General
Hospital, Fujioka, Japan. 1096Division of Respiratory Medicine and Allergology, Department of
Medicine, School of Medicine, Showa University, Tokyo, Japan. 1097Department of Pulmonary
Medicine, Fukushima Medical University, Fukushima, Japan. 1098Kansai Electric Power Hospital,
Osaka, Japan. 1099Kumamoto City Hospital, Kumamoto, Japan. 1100Department of Emergency
and Critical Care Medicine, Tokyo Metropolitan Police Hospital, Tokyo, Japan. 1101Department
of Respiratory Medicine, International University of Health and Welfare, Shioya Hospital,
Narita, Japan. 1102Department of Clinical Laboratory, International University of Health and
Welfare, Shioya Hospital, Narita, Japan. 1103National Hospital Organization Saitama Hospital,
Saitama, Japan. 1104Department of Respiratory Medicine, Gunma University Graduate School
of Medicine, Maebashi, Japan. 1105Department of Orthopedic Surgery, Tokyo Medical
University, Ibaraki Medical Center, Tokyo, Japan. 1106Department of Internal Medicine, Kiryu
Kosei General Hospital, Kiryu, Japan. 1107Daini Osaka Police Hospital, Osaka, Japan.
1109Department of Epidemiology, University Medical Centre Groningen, University of
Groningen, Groningen, The Netherlands. 1112Department of Psychiatry, University Medical
Center Groningen, Groningen, The Netherlands. 1113Department of Genetics, University
Medical Center Groningen, Groningen, The Netherlands. 1114Centre for Heart Lung Innovation,
University of British Columbia, Vancouver, British Columbia, Canada. 1115Division of Respiratory
Medicine, Faculty of Medicine, University of British Columbia, Vancouver, British Columbia,
Canada. 1116Institut Universitaire de Cardiologie et de Pneumologie de Québec, Université
Laval, Quebec, Quebec, Canada. 1117Department of Genetics and Genomic Sciences, Icahn
School of Medicine at Mount Sinai, New York, NY, USA. 1118University of Washington, Global
Health, Seattle, WA, USA. 1119Gossamer Bio, San Diego, CA, USA. 1120Department of Pathology
and Medical Biology, University Medical Centre Groningen, University of Groningen,
Groningen, The Netherlands. 1121GRIAC Research Institute, University Medical Centre
Groningen, University of Groningen, Groningen, The Netherlands. 1122Department of
Pulmonary Diseases, University Medical Centre Groningen, University of Groningen,
Groningen, The Netherlands. 1123Center for Genomic Medicine, Massachusetts General
Hospital, Boston, MA, USA. 1124Harvard Medical School, Cambridge, MA, USA. 1125Program in
Medical and Population Genetics, Broad Institute, Boston, MA, USA. 1126Channing Division of
Network Medicine, Department of Medicine, Brigham and Women’s Hospital, Boston, MA,
USA. 1127Brigham and Women’s Hospital, Boston, MA, USA. 1128Psychiatric and
Neurodevelopmental Genetics Unit, Center for Genomic Medicine, Massachusetts General
Hospital, Boston, MA, USA. 1129Department of Neurology, Massachusetts General Hospital,
Boston, MA, USA. 1130Division of General Internal Medicine, Massachusetts General Hospital
and Department of Medicine, Boston, MA, USA. 1132Department of Human Genetics, University
of Michigan, Ann Arbor, MI, USA. 1133Mount Sinai Clinical Intelligence Center, Department of
Genetics and Genomic Sciences, Icahn School of Medicine at Mount Sinai, New York, NY,
USA. 1135Sema4, a Mount Sinai venture, Stamford, CT, USA. 1137Mount Sinai Clinical Intelligence
Center, Charles Bronfman Institute for Personalized Medicine, New York, NY, USA.
1139Department of Human Genetics, David Geffen School of Medicine at UCLA, Los Angeles,
CA, USA. 1140Icahn Institute of Data Science and Genomics Technology, Icahn School of
Medicine, New York, NY, USA. 1141Mount Sinai Clinical Intelligence Center, Icahn School of
Medicine, New York, NY, USA. 1142Department of Genetic and Genomic Sciences, Icahn School
of Medicine at Mount Sinai, New York, NY, USA. 1143Charles Bronfman Institute for Personalized
Medicine, Icahn School of Medicine at Mount Sinai, New York, NY, USA. 1144Institute for
Genomic Health, Icahn School of Medicine at Mount Sinai, New York, NY, USA. 1146The Mindich
Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York,
NY, USA. 1147Pamela Sklar Division of Psychiatric Genomics, Icahn School of Medicine at Mount
Sinai, New York, NY, USA. 1148Department of Psychiatry, Icahn School of Medicine at Mount
Sinai, New York, NY, USA. 1149Icahn School of Medicine at Mount Sinai, New York, NY, USA.
1150Department of Psychiatry, Department of Genetic and Genomic Sciences, Icahn School of
Medicine at Mount Sinai, New York, NY, USA. 1151Department of Environmental Medicine and
Public Health, Icahn School of Medicine at Mount Sinai, New York, NY, USA. 1152Department of
Human Genetics, Center for Autism Research and Treatment, Institute for Precision Health,
University of California Los Angeles, Los Angeles, CA, USA. 1153The Hasso Plattner Institute of
Digital Health at Mount Sinai, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
1154BioMe Phenomics Center, >Icahn School of Medicine at Mount Sinai, New York, NY, USA.
1155Department of Medicine, Icahn School of Medicine at Mount Sinai, New York, NY, USA.
1157Regeneron Genetics Center, Tarrytown, NY, USA. 1158Phenomic Analytics & Clinical Data
Core, Geisinger Health System, Danville, PA, USA. 1159Department of Population Health
Sciences, Geisinger Health System, Danville, PA, USA. 1160Department of Molecular and
Functional Genomics, Geisinger Health System, Danville, PA, USA. 1162Department of
Genetics, University of Pennsylvania Perelman School of Medicine, Philadelphia, PA, USA.
1163Department of Biomedical Data Science, Stanford University, Stanford, CA, USA.
1166Department of Psychiatry, University of North Carolina at Chapel Hill, Chapel Hill, USA.
1167Department of Nutrition, University of North Carolina at Chapel Hill, Chapel Hill, USA.
1168Institute of Neuroscience and Physiology, University of Gothenburg, Gothenburg, Sweden.
1169Department of Medical Sciences, University of Turin, Turin, Italy. 1170Department of Clinical
and Biological Sciences, University of Turin, Orbassano, Italy. 1171Department of Pediatrics,
Department of Microbiology, Immunology and Molecular Genetics, University of California
Los Angeles, Los Angeles, CA, USA. 1172University of Genova, Genova, Italy. 1173Hopital
Mont-Godinne, Yvoir, Belgium. 1174Department of Molecular Medicine, University of Pavia,
Pavia, Italy. 1175Department of Public Health and Pediatric Sciences, University of Turin, Turin,
Italy. 1176Qatar Biobank for Medical Research, Qatar Foundation Research, Development and
Innovation, Qatar Foundation, Doha, Qatar. 1177Latvian Biomedical Research and Study Centre,
Riga, Latvia. 1178Department of Neuroscience, Karolinska Institutet, Stockholm, Sweden.
1179Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany. 1180Anaesthesiology
and Intensive Care Medicine, Department of Surgical Sciences, Uppsala University, Uppsala,
Sweden. 1181Integrative Physiology, Department of Medical Cell Biology, Uppsala University,
Uppsala, Sweden. 1182Hedenstierna Laboratory, CIRRUS, Anaesthesiology and Intensive Care
Medicine, Department of Surgical Sciences, Uppsala University, Uppsala, Sweden.
1183Department of Computer Science, School of Engineering, University of California Los
Angeles, Los Angeles, CA, USA. 1184University of California Los Angeles, Los Angeles, CA, USA.
1185Department of Psychiatry and Biobehavioral Sciences, David Geffen School of Medicine at
University of California Los Angeles, Los Angeles, CA, USA. 1186Division of Immunology,
Allergy, and Rheumatology, University of California Los Angeles, Los Angeles, CA, USA.
1187Department of Psychiatry, University of California Los Angeles, Los Angeles, CA, USA.
1188Department of Neurology, University of California Los Angeles, Los Angeles, CA, USA.
1189Department of Computational Medicine, University of California Los Angeles, Los Angeles,
CA, USA. 1190Department of Pathology and Laboratory Medicine, University of California Los
Angeles, Los Angeles, CA, USA. 1191Bioinformatics IDP, UCLA, Los Angeles, CA, USA.
1192Department of Neurology, David Geffen School of Medicine at UCLA, Los Angeles, CA,
USA. 1193Department of Urology, David Geffen School of Medicine at UCLA, Los Angeles, CA,
USA. 1195Queen Mary University, London, UK. 1196UCL Great Ormond Street Institute of Child
Health, London, UK. 1197University of Cambridge, Cambridge, UK. 1199Big Data Institute, Nufield
Department of Population Health, Li Ka Shing Centre for Health Information and Discovery,
University of Oxford, Oxford, UK. 1201Experimental Medicine Division, Nufield Department of

Medicine, John Radcliffe Hospital, University of Oxford, Oxford, UK. 1202Public Health England,
Field Service, Addenbrooke’s Hospital, Cambridge, UK. 1203Public Health England, Data and
Analytical Services, National Infection Service, London, UK. 1204Program in Bioinformatics and
Integrative Genomics, Harvard Medical School, Boston, MA, USA. 1205Program in Biological
and Biomedical Sciences, Harvard Medical School, Boston, MA, USA. 1207Department of
Clinical Research and Leadership, George Washington University, Washington, DC, USA.
1208Department of Human Genetics, The Wellcome Sanger Institute, Wellcome Genome
Campus, Hinxton, Cambridge, UK. 1209Strangeways Research Laboratory, The National
Institute for Health Research Blood and Transplant Unit in Donor Health and Genomics,
University of Cambridge, Cambridge, UK. 1210Department of Haematology, University of
Cambridge, Cambridge Biomedical Campus, Cambridge, UK. 1211British Heart Foundation
Cardiovascular Epidemiology Unit, Department of Public Health and Primary Care, University
of Cambridge, Cambridge, UK. 1212British Heart Foundation Centre of Research Excellence,
University of Cambridge, Cambridge, UK. 1213The National Institute for Health Research Blood
and Transplant Research Unit in Donor Health and Genomics, University of Cambridge,
Cambridge, UK. 1214Health Data Research UK Cambridge, Wellcome Genome Campus and
University of Cambridge, Cambridge, UK. 1215Department of Human Genetics, Wellcome
Sanger Institute, Hinxton, UK. 1216Department of Epidemiology, Emory University Rollins
School of Public Health, North Druid Hills, GA, USA. 1217Atlanta CA Health Care System, North
Druid Hills, GA, USA. 1218Center for Population Genomics, MAVERIC, VA Boston Healthcare
System, Boston, MA, USA. 1219MAVERIC, VA Boston Healthcare System, Boston, MA, USA.
1220Stanford University, Stanford, CA, USA. 1221Palo Alto VA Healthcare System, Stanford, CA,
USA. 1222Department of Biostatistics, Boston University School of Public Health, Boston, MA,
USA. 1223Department of Haematology, Central Hospital of Bolzano (SABES-ASDAA), Bolzano,
Italy. 1224Laboratory of Clinical Pathology, Hospital of Bressanone (SABES-ASDAA),
Bressanone, Italy. 1226University of Alcalá, Centro de Investigación Biomédica en Red en
Enfermedades Respiratorias (CIBERES), Madrid, Spain. 1227Center for Applied Genomics, The
Children’s Hospital of Philadelphia, Philadelphia, PA, USA. 1228Division of Human Genetics,
Department of Pediatrics, The Perelman School of Medicine, University of Pennsylvania,
Philadelphia, PA, USA. 1229Faculty of Medicine, University of Iceland, Reykjavik, Iceland.
1231Infectious Disease Unit, Hospital of Massa, Massa, Italy. 1232Department of Clinical Medicine,
Public Health, Life and Environment Sciences, University of L’Aquila, L’Aquila, Italy. 1233UOSD
Laboratorio di Genetica Medica - ASL Viterbo, San Lorenzo, Italy. 1234Unit of Infectious
Diseases, S. M. Annunziata Hospital, Florence, Italy. 1235Infectious Disease Unit, Hospital of
Lucca, Lucca, Italy. 1236Department of Clinical and Experimental Medicine, Infectious Diseases
Unit, University of Pisa, Pisa, Italy. 1238Clinic of Infectious Diseases, Catholic University of the
Sacred Heart, Rome, Italy. 1239Department of Diagnostic and Laboratory Medicine, Institute of
Biochemistry and Clinical Biochemistry, Fondazione Policlinico Universitario A. Gemelli
IRCCS, Catholic University of the Sacred Heart, Rome, Italy. 1240Private University in the
Principality of Liechtenstein, Triesen, Liechtenstein. 1241Digestive Diseases Unit, Virgen del
Rocio University Hospital, Institute of Biomedicine of Seville, University of Seville, Seville,
Spain. 1242Department of Biochemistry, University Hospital Vall d’Hebron, Barcelona, Spain.
1243University of Sevilla, Sevilla, Spain. 1244Instituto de Biomedicina de Sevilla, Sevilla, Spain.
1245Hospital Universitario Virgen del Rocío de Sevilla, Sevilla, Spain. 1246Consejo Superior de
Investigaciones cientíicas, Madrid, Spain. 1247Humanitas Clinical and Research Center,
IRCCS, Milan, Italy. 1248Immunohematology Department, Banc de Sang i Teixits,
Autonomous University of Barcelona, Barcelona, Spain. 1249August Pi i Sunyer Biomedical
Research Institute, Hospital Clinic, University of Barcelona, Barcelona, Spain.
1250Department of Pathophysiology and Transplantation, Università degli Studi di Milano,
Milan, Italy. 1251Internal Medicine Department, Virgen del Rocio University Hospital, Sevilla,
Spain. 1252Department of Biomedical Sciences, Humanitas University, Milan, Italy.
1253Department Emergency, Anesthesia and Intensive Care, University Milano-Bicocca,
Monza, Italy. 1254Department of Medical Sciences, Università degli Studi di Torino, Turin,
Italy. 1255Department of Medical Microbiology, Clinic of Laboratory Medicine, St Olav’s
Hospital, Trondheim, Norway. 1256Department of Infectious Diseases, St Olav’s Hospital,
Trondheim University Hospital, Trondheim, Norway. 1257Department of Clinical and
Molecular Medicine, NTNU, Trondheim, Norway. 1258Department of Research, St Olav’s
Hospital, Trondheim University Hospital, Trondheim, Norway. 1259Institute of Parasitology
and Biomedicine Lopez-Neyra, Granada, Spain. 1260Institute for Cardiogenetics, University of
Lübeck, Lübeck, Germany. 1261German Research Center for Cardiovascular Research,
partner site Hamburg-Lübeck-Kiel, Lübeck, Germany. 1262University Heart Center Lübeck,
Lübeck, Germany. 1263Department of Research, Ostfold Hospital Trust, Gralum, Norway.
1264Pediatric Departement, Centro Tettamanti- European Reference Network (ERN) PaedCan,
EuroBloodNet, MetabERN-University of Milano-Bicocca-Fondazione MBBM/Ospedale San
Gerardo, Milan, Italy. 1265Geminicenter for Sepsis Research, Institute of Circulation and
Medical Imaging (ISB), NTNU, Trondheim, Norway. 1266Clinic of Anesthesia and Intensive
Care, St Olav’s Hospital, Trondheim University Hospital, Trondheim, Norway. 1267Clinic of
Medicine and Rehabilitation, Levanger Hospital, Nord-Trondelag Hospital Trust, Levanger,
Norway. 1268Stefan-Morsch-Stiftung, Birkenfeld, Germany. 1269Center of Bioinformatics,
Biostatistics, and Bioimaging, School of Medicine and Surgery, University of Milano
Bicocca, Milan, Italy. 1270Phase 1 Research Centre, ASST Monza, School of Medicine and
Surgery, University of Milano-Bicocca, Milan, Italy. 1271Pneumologia ASST-Monza, University
of Milano-Bicocca, Milano, Italy. 1272School of Medicine and Surgery, University of
Milano-Bicocca, Milano, Italy. 1273Infectious Diseases Unit, San Gerardo Hospital, Monza,
Italy. 1274SODIR-VHIR research group, Barcelona, Spain. 1275Bioinformatics area, Fiundación
progreso y Salud, Andalucia, Spain. 1276Present address: Program in Metabolism, Broad
Institute of MIT and Harvard, Cambridge, MA, USA. 1277Present address: Program in Medical
and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA.
1278Present address: Diabetes Unit, Center for Genomic Medicine, Massachusetts General
Hospital, Boston, MA, USA. 1279Present address: Harvard Medical School, Boston, MA, USA.
1280These authors contributed equally: Mari E. K. Niemi, Juha Karjalainen, Benjamin M. Neale,
Mark Daly, Andrea Ganna. 1281Unafiliated: Sangyoon Im, Jason Kendall, Michael MacMahon,
Mark Peters, Robert Thompson, Martin Williams. ✉e-mail: bneale@broadinstitute.org;
mark.daly@helsinki.i; andrea.ganna@helsinki.i

Article
Methods
Contributing studies
All of the participants were recruited following protocols approved
by local Institutional Review Boards; this information is collected in
Supplementary Table1 for all 46 studies. All protocols followed local
ethics recommendations and informed consent was obtained when
required. Information about sample numbers, sex and age from for
each contributing study is given in Supplementary Table1. In total, 16
studies contributed data to the analysis of critical illness due to COVID-
19, 29 studies contributed data to hospitalized COVID-19 analysis and
44 studies contributed to the analysis of all cases of COVID-19. Each
individual study that contributed data to a particular analysis met a
minimum threshold of 50 cases, as defined by the phenotypic criteria,
for statistical robustness. The effective sample sizes for each ancestry
group shown in Fig.1 were calculated for display using the formula:
((4×N
case
×N
control
)/(N
case
+N
control
)). Details of contributing research
groups are provided in Supplementary Table1.
Phenotype definitions
COVID-19 disease status (critical illness and hospitalization status) was
assessed following the Diagnosis and Treatment Protocol for Novel
Coronavirus Pneumonia
38
. The critically ill COVID-19 group included
patients who were hospitalized owing to symptoms associated with
laboratory-confirmed SARS-CoV-2 infection and who required respira-
tory support or whose cause of death was associated with COVID-19.
The hospitalized COVID-19 group included patients who were hos-
pitalized owing to symptoms associated with laboratory-confirmed
SARS-CoV-2 infection.
The reported SARS-CoV-2 infection group included individu-
als with laboratory-confirmed SARS-CoV-2 infection or electronic
health record, ICD coding or clinically confirmed COVID-19, or
self-reported COVID-19 (for example, by questionnaire), with or
without symptoms of any severity. Genetic-ancestry-matched con-
trol individuals 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. Additional
information regarding individual studies contributing to the consor-
tium are described in Supplementary Table1.
Genome-wide association studies and meta-analyses
Each contributing study genotyped the samples and performed quality
controls, data imputation and analysis independently, but following
the consortium recommendations (information is available at https://
www.covid19hg.org/). We recommended that genome-wide association
study (GWAS) analyses were run using Scalable and Accurate Implemen-
tation of GEneralized mixed model (SAIGE)39 on chromosomes 1–22 and
X. The recommended analysis tool was SAIGE, but studies also used
other software such as PLINK
40
. The suggested covariates were age,
age
2
, sex, age×sex and the 20 first principal components. Any other
study-specific covariates to account for known technical artefacts could
be added. SAIGE automatically accounts for sample relatedness and
case–control imbalances. Quality-control and analysis approaches for
individual studies are reported in Supplementary Table1.
Study-specific summary statistics were then processed for
meta-analysis. Potential false positives, inflation and deflation were
examined for each submitted GWAS. Allele frequency plots against
gnomAD 3.0 genomes were manually inspected for each study. Stand-
ard error values as a function of the effective sample size were used to
find studies that deviated from the expected trend. Summary statistics
passing this manual quality control were included in the meta-analysis.
Variants with an allele frequency of >0.1% and an imputation INFO score
of >0.6 were carried forward from each study. Variants and alleles were
lifted over to genome build GRCh38, if needed, and harmonized to
gnomAD 3.0 genomes
41
by finding matching variants by strand flipping
or switching the ordering of alleles. If multiple matching variants were
included, the best match was chosen according to the minimum fold
change in absolute allele frequency. Meta-analysis was performed
using the inverse-variance-weighted (IVW) method on variants that
were present in at least two-thirds of the studies contributing to the
phenotype analysis. The method summarizes effect sizes across the
multiple studies by computing the mean of the effect sizes weighted
by the inverse variance in each individual study.
We report 13 meta-analysis variants that pass the genome-wide
significance threshold after adjusting the threshold for multiple
traits tested (P<5×10−8/3). We report the unadjusted P values for
each variant. We tested for heterogeneity between estimates from
contributing studies using Cochran’s Q-test42,43. This is calculated
for each variant as the weighted sum of squared differences between
the effects sizes and their meta-analysis effect, the weights being the
inverse variance of the effect size. Q is distributed as a χ
2
statistic with
k (number of studies) minus one degrees of freedom. Two loci reached
genome-wide significance but were excluded from the significant
results in Supplementary Table2 due to heterogeneity between esti-
mates from contributing studies and missingness between studies at
chr.6: 31057940–31380334 and chr.7: 54671568–54759789; however,
these regions are not excluded from the corresponding summary
statistics in data release 5 (COVID-19 HGI (https://www.covid19hg.
org/results/r5/) and GWAS Catalog (study code GCST011074)). For
each of the lead variants reported in Supplementary Table2, we aimed
to find loci specific to susceptibility or severity by testing whether
there was heterogeneity between the effect sizes associated with
hospitalized COVID-19 (progression to severe disease) and reported
SARS-CoV-2 infection. We used the Cochran’s Q measure
42,43
, calculated
for each variant as the weighted sum of squared differences between
the two analysis effect sizes and their meta-analysis effect with the
weights being the inverse variance of the effect size. A significant P
value of P<0.004 ((0.05/13 loci) for multiple tests) indicates that
the effect sizes for a particular variant are significantly different in
the two analyses (Supplementary Table2). For the nine loci, in which
the lead variant effect size was significantly higher for hospitalized
COVID-19, we carried out the same test again but comparing effect sizes
from hospitalized COVID-19 with critically ill COVID-19 (Supplemen-
tary Table4). Furthermore, we carried out the same test comparing
meta-analysed hospitalized COVID-19 (population as controls) and
hospitalized COVID-19 (SARS-CoV-2-positive but non-hospitalized
as controls) (Supplementary Table4). For these pairs of phenotype
comparisons, we generated new meta-analysis summary statistics to
use; including only those studies that could contribute data to both
phenotypes that were under comparison.
Principal component projection
To project every GWAS participant into the same principal compo-
nent (PC) space, we used pre-computed PC loadings and reference
allele frequencies. For reference, we used unrelated samples from
the 1000 Genomes Project and the Human Genome Diversity Pro-
ject and computed PC loadings and allele frequencies for the 117,221
single-nucleotide polymorphisms (SNPs) that (1) are available in every
cohort; (2) have a minor allele frequency of >0.1% in the reference; and
(3) are LD-pruned (r
2
<0.8; 500-kb window). We then asked each cohort
to project their samples using our automated script provided at https://
github.com/covid19-hg/. It internally uses the PLINK244 --score function
with the variance-standardize option and reference allele frequencies
(--read-freq); so that each cohort-specific genotype/dosage matrix is
mean-centred and variance-standardized with respect to reference
allele frequencies, but not cohort-specific allele frequencies. We fur-
ther normalized the projected PC scores by dividing the values by a
square root of the number of variants used for projection to account
for a subtle difference due to missing variants.

Gene prioritization
To prioritize candidate causal genes reported in full in Supplementary
Table2, we used various gene prioritization approaches using both
locus-based and similarity-based methods. Because we only describe
the in silico gene prioritization results without characterizing the actual
functional activity invitro or invivo, we aimed to provide a systematic
approach to nominate potential causal genes in a locus using the fol-
lowing criteria.
(1) The closest gene: a gene that is closest to a lead variant by distance
to the gene body.
(2) Genes in the LD region: genes that overlap with a genomic range
containing any variants in LD (r
2
>0.6) with a lead variant. For LD compu-
tation, we retrieved LD matrices provided by gnomAD v.2.1.141 for each
population analysed in this study (except for admixed American, Middle
Eastern and South Asian genetic ancestry populations, for whom data
are not available). We then constructed a weighted-average LD matrix
by per-population sample sizes in each meta-analysis, which we used
as a LD reference.
(3) Genes with coding variants: genes with at least one loss-of-function
or missense variant (annotated by VEP
45
v.95 with GENCODE v.29) that
is in LD with a lead variant (r2>0.6).
(4) eGenes: genes with at least one fine-mapped cis-eQTL variant
(PIP>0.1) that is in LD with a lead variant (r
2
>0.6) (Supplementary
Table 5). We retrieved fine-mapped variants from the GTEx v.820 (https://
www.finucanelab.org/) and eQTL catalogue46. In addition, we looked
up significant associations in the Lung eQTL Consortium21 (n=1,103)
to further support our findings in lung with a larger sample size (Sup-
plementary Table7). We note that, in contrast to the GTEx or eQTL
catalogue, we only looked at associations and did not fine-map our
data to the Lung eQTL Consortium data.
(5) V2G: a gene with the highest overall V2G score based on Open
Targets Genetics (OTG)26. For each variant, the overall V2G score
aggregates differentially weighted evidence of variant–gene associa-
tions from several data sources, including molecular cis-QTL data (for
example, cis-protein QTLs from ref. 47, cis-eQTLs from GTEx v.7 and so
on), interaction-based datasets (for example, promoter capture Hi-C),
genomic distance and variant effect predictions (VEP) from Ensembl. A
detailed description of the evidence sources and weights used is pro-
vided in the OTG documentation (https://genetics-docs.opentargets.
org/our-approach/data-pipeline)26.
Phenome-wide association study
To investigate the evidence of shared effects of 15 index variants
for COVID-19 and previously reported phenotypes, we performed a
phenome-wide association study. We considered phenotypes in OTG
obtained from the GWAS catalogue (this included studies with and
without full summary statistics, n=300 and 14,013, respectively)
48
and from the UK Biobank. Summary statistics for UK Biobank traits
were extracted from SAIGE39 for binary outcomes (n=1,283 traits)
and Neale v.2 (n=2,139 traits) for both binary and quantitative traits
(http://www.nealelab.is/uk-biobank/) and FinnGen Freeze 4 cohort
(https://www.finngen.fi/en/access_results). We report PheWAS results
for phenotypes for which the lead variants were in high LD (r2>0.8) with
the 13 genome-wide significant lead variants from our main COVID-19
meta-analysis (Supplementary Table6). This conservative approach
allowed spurious signals primarily driven by proximity rather than
actual colocalization to be removed (seeMethods).
To remove plausible spurious associations, we retrieved phenotypes
for GWAS lead variants that were in LD (r
2
>0.8) with COVID-19 index
variants.
Heritability
LD score regression v.1.0.1
49
was used to estimate the SNP heritability of
the phenotypes from the meta-analysis summary statistic files. As this
method depends on matching the LD structure of the analysis sample
to a reference panel, the summary statistics of European ancestry only
were used. Sample sizes were n=5,101 critically ill cases of COVID-19
and n=1,383,241 control participants, n=9,986 hospitalized cases of
COVID-19 and n=1,877,672 control participants, and n=38,984 cases
and n=1,644,784 control participants for the analysis of all cases—
all including the 23andMe cohort. Pre-calculated LD scores from the
1000 Genomes European reference population were obtained online
(https://data.broadinstitute.org/alkesgroup/LDSCORE/). Analyses
were conducted using the standard program settings for variant filter-
ing (removal of non-HapMap3 SNPs, the HLA region on chromosome
6, non-autosomal, χ2>30, minor allele frequency of <1%, or allele mis-
match with reference). We additionally report SNP heritability estimates
for the all-ancestries meta-analyses, calculated using European panel
LD scores, in Supplementary Table8.
Partitioned heritability
We used partitioned LD score regression50 to partition COVID-19 SNP
heritability in cell types in our summary statistics for European ancestry
only. We ran the analysis using the baseline model LD scores calculated
for European populations and regression weights that are available
online (https://github.com/bulik/ldsc). We used the COVID-19 summary
statistics for European ancestry only for the analysis.
Genome-wide association summary statistics
We obtained genome-wide association summary statistics for 43
complex-disease, neuropsychiatric, behavioural or biomarker phe-
notypes (Supplementary Table10). These phenotypes were selected
based on their putative relevance to COVID-19 susceptibility, severity
or mortality, with 19 selected based on the Centers for Disease Control
list of underlying medical conditions associated with COVID-19 sever-
ity51 or traits reported to be associated with increased risk of COVID-19
mortality by OpenSafely52. Summary statistics generated from GWAS
using individuals of European ancestry were preferentially selected if
available. These summary statistics were used in subsequent genetic
correlation and Mendelian randomization analyses.
Genetic correlation
LD score regression50 was also used to estimate the genetic correlations
between our COVID-19 meta-analysis phenotypes reported using sam-
ples of only European ancestry, and between these and the curated set
of 38 summary statistics. Genetic correlations were estimated using the
same LD score regression settings as for heritability calculations. Differ-
ences between the observed genetic correlations of SARS-CoV-2 infec-
tion and COVID-19 severity were compared using a z-score method53.
Mendelian randomization
Two-sample Mendelian randomization was used to evaluate the poten-
tial for causal association of the 38 traits on COVID-19 hospitalization,
on COVID-19 severity and reported SARS-CoV-2 infection using samples
of only European ancestry. Independent genome-wide significant SNPs
robustly associated with the exposures of interest (P<5×10
−8
) were
selected as genetic instruments by performing LD clumping using
PLINK40. We used a strict r2 threshold of 0.001, a 10-Mb clumping win-
dow, and the European reference panel from the 1000 Genomes Pro-
ject
54
to discard SNPs in LD with another variant with a smaller P-value
association. For genetic variants that were not present in the hospital
-
ized COVID-19 analysis, PLINK was used to identify proxy variants that
were in LD (r
2
>0.8). Next, the exposure and outcome datasets were
harmonized using the R package TwoSampleMR
55
. Namely, we ensured
that the effect of a variant on the exposure and outcome corresponded
to the same allele, we inferred positive-strand alleles and dropped
palindromes with ambiguous allele frequencies, as well as incompat-
ible alleles. Supplementary Table10 includes the harmonized datasets
used in the analyses.

Article
The global test from Mendelian randomization pleiotropy residual
sum and outlier (MR-PRESSO)
56
software was used to investigate over-
all horizontal pleiotropy. In brief, the standard IVW meta-analytic
framework was used to calculate the average causal effect by excluding
each genetic variant used to instrument the analysis. A global statistic
was calculated by summing the observed residual sum of squares, that
is, the difference between the effect predicted by the IVW slope exclud
-
ing the SNP, and the observed effect of the SNP on the outcome. Over-
all horizontal pleiotropy was subsequently analysed by comparing the
observed residual sum of squares, with the residual sum of squares
expected under the null hypothesis of no pleiotropy. The MR-PRESSO
global test was shown to perform well when the outcome and exposure
GWASs are not disjoint (although the power to detect horizontal plei-
otropy is slightly reduced by complete sample overlap). We also used
the regression intercept in MR-Egger
57
to evaluate potential bias due
to directional pleiotropic effects. This additional check was used in
Mendelian randomization analyses with an
IGX
2
index surpassing the
recommended threshold (
I
>9
0%
GX
2
)58. Contingent on the MR-PRESSO
global test results we analysed the causal effect of each exposure on
COVID-19 hospitalization by using a fixed-effect IVW meta-analysis as
the primary analysis, or, if pleiotropy was present, the MR-PRESSO
outlier-corrected test. The IVW approach estimates the causal effect
by aggregating the single-SNP causal effects (obtained using the ratio
of coefficients method—that is, the ratio of the effect of the SNP on the
outcome over the effect of the SNP on the exposure) in a fixed-effects
meta-analysis. The SNPs were assigned weights based on their inverse
variance. The IVW method confers the greatest statistical power for
estimating causal associations
59
, but assumes that all variants are valid
instruments and can produce biased estimates if the average pleio-
tropic effect differs from zero. Alternatively, when horizontal pleiot-
ropy was present, we used the MR-PRESSO outlier-corrected method
to correct the IVW test by removing outlier SNPs. We conducted further
sensitivity analyses using alternative Mendelian randomization meth-
ods that provide consistent estimates of the causal effect even when
some instrumental variables are invalid, at the cost of reduced statisti-
cal power including: (1) Weighted median estimator (WME); (2)
weighted mode-based estimator (WMBE); and (3) MR-Egger regression.
Robust causal estimates were defined as those that were significant at
an FDR of 5% and either (1) showed no evidence of heterogeneity
(MR-PRESSO global test P>0.05) or horizontal pleiotropy (Egger inter-
cept P>0.05); or (2) in the presence of heterogeneity or horizontal
pleiotropy, the WME-, WMBE-, MR-Egger- or MR-PRESSO-corrected
estimates were significant (P<0.05). All statistical analyses were con-
ducted using R v.4.0.3. Mendelian randomization analysis was per-
formed using the ‘TwoSampleMR’ v.0.5.5 package55.
Website and data distribution
In anticipation of the need to coordinate many international partners
around a single meta-analysis effort, we created the COVID-19 HGI
website (https://covid19hg.org). We were able to centralize informa-
tion, recruit partner studies, rapidly distribute summary statistics
and present preliminary interpretations of the results to the public.
Open meetings are held on a monthly basis to discuss future plans and
new results; video recordings and supporting documents are shared
(https://covid19hg.org/meeting-archive). This centralized resource
provides a conceptual and technological framework for organizing
global academic and industry groups around a shared goal. The website
source code and additional technical details are available at https://
github.com/covid19-hg/covid19hg.
To recruit new international partner studies, we developed a work-
flow in which new studies are registered and verified by a curation
team (https://covid19hg.org/register). Users can explore the registered
studies using a customized interface to find and contact studies with
similar goals or approaches (https://covid19hg.org/partners). This
helps to promote organic assembly around focused projects that are
adjacent to the centralized effort (https://covid19hg.org/projects). Visi-
tors can query study information, including study design and research
questions. Registered studies are visualized on a world map and are
searchable by institutional affiliation, city and country.
To encourage data sharing and other forms of participation, we
created a rolling acknowledgements page (https://covid19hg.org/
acknowledgements) and directions on how to contribute data to the
central meta-analysis effort (https://covid19hg.org/data-sharing).
Upon the completion of each data freeze, we post summary statistics,
plots and sample size breakdowns for each phenotype and contributing
cohort (https://covid19hg.org/results). The results can be explored
using an interactive web browser (https://app.covid19hg.org). Several
computational research groups carry out follow-up analyses, which
are made available for download (https://covid19hg.org/in-silico). To
enhance scientific communication to the public, preliminary results
are described in blog posts by the scientific communications team and
shared on Twitter. The first post was translated to 30 languages with
the help of 85 volunteer translators. We compile publications and pre
-
prints submitted by participating groups and summarize genome-wide
significant findings from these publications (https://covid19hg.org/
publications).
Reporting summary
Further information on research design is available in theNature
Research Reporting Summary linked to this paper.
Data availability
Summary statistics generated by the COVID-19 HGI are available at
https://www.covid19hg.org/results/r5/ and are available in the GWAS
Catalog (study code GCST011074). The analyses described here include
the freeze-5 data. COVID-19 HGI continues to regularly release new
data freezes. Summary statistics for non-European ancestry samples
are not currently available due to the small individual sample sizes of
these groups, but results for lead variants of 13 loci are reported in
Supplementary Table3. 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 lab (http://www.nealelab.is/uk-biobank/), Finucane lab (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 are available on
GitHub (https://github.com/covid19-hg/) and the code for the Men-
delian randomization and genetic correlation pipeline is available on
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Acknowledgements We thank the entire COVID-19 HGI community for their contributions and
continued collaboration. The work of the contributing studies was supported by numerous
grants from governmental and charitable bodies. Acknowledgements speciic to contributing
studies are provided in Supplementary Table 13. We thank G. Butler-Laporte, G. Wojcik,
M.-G. Hollm-Delgado, C. Willer and G. Davey Smith for their extensive feedback and discussion.
Author contributions Author contributions are provided within the author list.
Competing interests A full list of competing interests is supplied as Supplementary Table 13.
Additional information
Supplementary information The online version contains supplementary material available at
https://doi.org/10.1038/s41586-021-03767-x.
Correspondence and requests for materials should be addressed to Benjamin M. Neale, Mark
Daly, Andrea Ganna, Benjamin M. Neale or Mark Daly.
Peer review information Nature thanks Samira Asgari, Paul McLaren and Neneh Sallah for their
contribution to the peer review of this work. Peer reviewer reports are available.
Reprints and permissions information is available at http://www.nature.com/reprints.

Article
Extende d Data Fig. 1 | An alytical su mmary of the C OVID-19 HGI
meta-analysis. Usi ng the analyti cal plan set by the COV ID-19 HGI, each
individua l study runs thei r analyses and upl oads the result s to the Initiati ve,
who then run s the meta-anal ysis. There are th ree main analyse s that each stud y
can contr ibute summar y statistic s to: critically ill C OVID-19, hospitalized
COVID-19 and repo rted SARS -CoV-2 infection. The ph enotypic cr iteria used t o
define c ases are list ed in the dark grey boxes , along with the nu mbers of cas es
(N) included in t he final all-anc estries me ta-analysis. Co ntrols were def ined in
the same way ac ross all three anal yses as everyb ody that is not a c ase—for
example, population controls (light grey box). Sensitivity analyses—not
reporte d in this extende d data fig ure—also include d mild and/or asymp tomatic
cases of COV ID-19 as control ind ividuals. Sam ple number (N) of co ntrol
individua ls differed bet ween the analys es due to the dif ference in the numb er
of studies contributing data to these.

Extende d Data Fig. 2 | Pr ojection o f contribut ing studie s samples in to the
same PC sp ace. We asked par ticipating stu dies to perfor m a PC projectio n
using the 100 0 Genomes P roject and Hum an Genome Diver sity Projec t as a
reference, w ith a common set o f variants. For e ach panel (except for the
reference), coloured points correspond to contributed samples from each
cohort, wh ereas grey poin ts correspo nd to the reference s amples from th e
1000 Geno mes Projec t. Colour repres ents a geneti c population tha t each
cohort sp ecified . As 23andMe , Genomics Eng land 100,000 Ge nomes Projec t
(Genomics England100kg p), and Million Veterans Pro gram (MVP) only
submitte d PCA images, we overl aid their submit ted transpare nt images using
the same co ordinates, ins tead of direct ly plotting the m. Populations a re
define d as African ( AFR), admixed Ame rican (AMR), Ea st Asian (EA S), European
(EUR), Middle Ea stern (MID) and Sou th Asian (SAS), Oceani an (OCE).

Article
Extende d Data Fig. 3 | Loc us-zoom plots o f the 3p21.31 re gion for repo rted
SARS- CoV-2 infection. a, A stan dard plot without exc lusion. Here, th e severity
lead varian t rs10490770 (chr.3:45823240T:C) is shown as a lea d variant.
b, Additional in dependent su sceptibilit y signal(s) after exclud ing variants w ith
r2>0.05 with rs10490770. Th e susceptibi lity lead varia nt rs2271616
(chr.3:45796521G:T) is highlighted .

Extende d Data Fig. 4 | Ge nome-wide m eta-analys is associ ation result s for
critica l illness du e to COVID-19. The locus o n chromosome 6 is th e HLA locus,
which was rem oved from the list of re ported loc i in Supplement ary Table2 due
to the high he terogeneit y in effect size e stimated be tween studie s included in
the analysis . The locus on c hromosome 7 was al so not report ed in
Supplemen tary Table2 due to missing ness acros s studies—that is, t he high
number of st udies in the met a-analysis that di d not report sum mary stati stics
for this regio n. There are two a ssociation p eaks on chromo some 19.

Article
Extende d Data Fig. 5 | Se nsitivit y analyses for overl apping cont rols in
genomiCC and UK Biobank. Comparison o f the beta effe ct sizes (top) and
unadjusted P values (bot tom) of the 13 lead var iants, using da ta from the
COVID-19 criti cal illness me ta-analysis in all th e cohorts to le aving out
genomiCC (cas es, n=4,354; controls, n=1,474,655; tot al, n=1,479,009), leaving
out the UK Bio bank (UKBB; cas es, n=5,870; controls, n=1,155,203; tot al,
n=1,161,073) and leaving ou t both genomiCC an d UK Biobank (case s, n=4,04 5;
controls, n=1,146,078; to tal, n=1,150,12 3) (from left to right, re spectivel y).
Top, dots and grey bars rep resent the be ta effect si ze estimates ± s tandard
error from th e correspond ing GWAS meta-analys is. Bottom, d ots represen t
two-side d P values from the corre sponding GWAS met a-analysis. Fill ed dots
indicate var iants that showe d genome-wide s ignific ance in the full
meta-anal ysis of critical i llness due to COVI D-19, and empty dots repre sent
variants t hat were not signif icant for cri tical illness b ut were signif icant for
either hos pitalizati on due to COVID-19 or repor ted SARS -CoV-2 infection. Red
dots repre sent variant s that showed genom e-wide signif icance in th e
leave-one -out analysis for gen omiCC, UK Biob ank or genomiCC and U K
Biobank.

Extende d Data Fig. 6 | Com parison of χ2 stati stics and r2 values to the lead
variant in th e 3p21.31 re gion. a–c, Data are sho wn for critical ill ness (a),
hospitalization (b) and repor ted SARS- CoV-2 infection (c). The left blue p eak in
c, which is unco rrelated wit h the lead variant s in the region, i ndicates tha t there
are independent signals.

Article
Extende d Data Fig. 7 | Com parison of th e effect s izes of lead var iants
betwee n pairs of COVID -19 meta-analys es. Compar ison of effec t sizes for the
nine variant s associat ed with severit y of COVID-19 disea se. a, Comparin g
hospital ized cases of C OVID-19 versus popula tion controls (n=10,428 case s
and n=1,483,270 contr ols) and critically il l cases of COVID -19 versus
population controls (n=6,179 ca ses and n=1,483,780 controls). b, Hospitalized
cases of COV ID-19 versus popula tion controls (n=5,806 c ases and n=1 ,144, 263
controls) and ho spitalized c ases of COVID -19 versus non-hospita lized cases o f
COVI D-19 ( n=5,773 cases and n=15,497 cont rols). Sample sizes for ho spitalized
cases of COV ID-19 versus popula tion controls dif fer betwee n a and b due to
differenc es in the samplin g of studies sel ected for the ana lysis. This se lection
included all s tudies that were a ble to contribu te data to the resp ective analy ses
that the dat a were compared to (show n on the y axis) in each pa nel. Dots
represen t the effect si ze beta esti mates, bars repr esent the 95% co nfidence
interval o f the estimate s. Effect siz e estimates a nd P values for hetero geneity
tests (Cochran’s Q, two-tailed tes t) are reported in Su pplementar y Table3.

Extende d Data Fig. 8 | PheWAS for ge nome-wide s ignif icant lead var iants.
Select ed phenoty pes associ ated with geno me-wide signi ficant COVI D-19
variants (se e Supplement ary Table6 for a complete list). We repor t those
associa tions for which a lea d variant from a prev ious GWAS result was i n high
LD (r2>0.8) with the index COV ID-19 variants. T he colour repre sents the
z-scores of c orrelated ris k increasing alle les for the trait . The total numb er of
associa tions for each COVI D-19 variant is highli ghted in the grey b ox.

Article
Extende d Data Fig. 9 | Ge netic corre lation with COV ID-19 phenot ypes. Eac h
column shows th e genetic corre lation result s for the three COVID -19
phenoty pes (Europea n-ancestry a nalyses only): crit ical illness, h ospitaliza tion
and repor ted SARS- CoV-2 infect ion. The trait s that the genet ic correlation i s
run agains t are listed on the le ft. Signif icant corre lations (FDR<0.05) are
shown with their 95% confidence intervals in red, nominally significant
correlations (P<0.05) are in black and non-sig nifican t correlations a re in grey.
Two -s ide d P values we re calculated us ing LDSC for genet ic correlation s and
exact est imates, unadj usted stand ard errors and two -sided P values are
available in Supplementary Table11.

Extended Data Fig. 10 | Mendelian randomization sensitivity analyses.
Genetic c orrelations a nd Forest plots dis playing the cau sal estimate s for each
of the sensi tivity anal yses used in th e Mendelian rand omization an alysis for
trait pairs t hat were signif icant at an FDR of 5% . Two-sided P values were
estimate d using IVW, WM E, WMBE and MR-PR ESSO analyse s. RBC, red b lood
cell count.

1
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Corresponding author(s): Andrea Ganna
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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
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release 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
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LD score regression v 1.0.1 [PMID: 25642630] was used for heritability and partitioned heritability analyses. Variants for Mendelian
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European ancestry samples are not currently available due to the small individual sample sizes of these groups, but results for 13 loci lead variants are reported in
Supplementary Table 3. 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 lab (http://www.nealelab.is/uk-biobank/), Finucane lab (https://www.finucanelab.org), FinnGen Freeze 4
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Sample size The consortium meta-analysed genome-wide association study (GWAS) summary statistics from any individual study that had included a
minimum of n=50 cases and n=50 controls in their analysis. The cutoff at n=50 cases and n=50 controls was aimed at reducing noise to the
meta-analysis, but also to be inclusive of studies that had not yet accumulated large numbers of COVID-19 patient data. No statistical
calculation for adequate sample size was performed, but the results identifying multiple genomic regions at genome-wide significance
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Data exclusions Individual level phenotype and genotype data exclusions were performed by each individual study, following the consortium analysis plan
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summary statistics (with heterogeneity test values).
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. 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.

3
nature research | reporting summary April 2020
Materials & experimental systems
n/a Involved in the study
Antibodies
Eukaryotic cell lines
Palaeontology and archaeology
Animals and other organisms
Human research participants
Clinical data
Dual use research of concern
Methods
n/a Involved in the study
ChIP-seq
Flow cytometry
MRI-based neuroimaging
Human research participants
Policy information about studies involving human research participants
Population characteristics Summary statistics from 46 independent studies were included in consortium meta-analyses. Mean age of cases across
studies was 55.3 years. The effective sample size for genetic ancestry populations was: n=11,598 Middle Eastern; n=28,918
South Asian; 43,332 East Asian; 48,714 African; 70,902 Ad-mixed American; 738,538 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 SARS-
CoV-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 SARS-
CoV-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.