BRIEF COMMUNICATION OPEN
Defining he causes o spo adic Pa kinson’s disease in he
global Pa kinson’s gene ics p og am (GP2)
Clodagh Towns
1
, Madeleine Riche
1
, Simona Jasai y e
1
, Eleano J. S a o d
1,2
, Julie Joube
1
, Ta ek An a
3
,
Alejand o Ma inez-Ca asco
1,2
, Ma y B. Maka ious
1,3,4
, B ad o d Casey
5,6
, Dan Vi ale
4,7,8
, K is in Le ine
7,8
,
Hamp on Leona d
3,7,8,9
, Ca oline B. Pan azis
4,7
, Lau el A. Sc e en
7
, Dena G. He nandez
3
, Clai e E. Wegel
10
, Jus in Solle
5
,
Mike A. Nalls
3,4,7,8
, Co nelis Blauwend aa
4,7,11
, And ew B. Single on
3,7,12
, Manuela M. X. Tan
13
, Hi o aka Iwaki
3,7,8
,
Huw R. Mo is
1,2
✉and he Global Pa kinson’s Gene ics P og am (GP2)*
The Global Pa kinson’s Gene ics P og am (GP2) will geno ype o e 150,000 pa icipan s om a ound he wo ld, and in eg a e
gene ic and clinical da a o use in la ge-scale analyses o d ama ically expand ou unde s anding o he gene ic a chi ec u e o PD.
This epo de ails he wo kflow o coho in eg a ion in o he complex a m o GP2, and oge he wi h ou ou line o he
monogenic hub in a companion pape , p o ides a gene alizable bluep in o es ablishing la ge scale collabo a i e esea ch
conso ia.
npj Pa kinson’s Disease (2023) 9:131 ; h ps://doi.o g/10.1038/s41531-023-00533-w
INTRODUCTION
Pa kinson’s disease (PD) is a mul i ac o ial diso de wi h complex
e iology. The la ges genome-wide associa ion s udy (GWAS) o-
da e included 37,688 cases, 18,618 p oxy cases (una ec ed fi s -
deg ee ela i es), and 1.4 million con ols om Eu opean ances y,
and iden ified 90 independen isk signals ac oss 78 genomic
egions; 38 o which we e no el signals
1
. Despi e hese ad ances,
PD GWAS a e cu en ly limi ed by scale, a ocus on Eu opean
popula ions, and limi ed in eg a ion wi h clinical pheno ype da a.
A powe calcula ion based on he 2019 GWAS da a indica es ha
inclusion o an addi ional ~99,000 cases would enable a ian s o
smalle e ec size ha con ibu e o polygenic isk (p- alue cu o :
1.35 × 10
−3
) o each genome-wide significance. The e o e, expand-
ing PD GWAS o a leas his size will esul in iden ifica ion o
addi ional isk loci and imp o e gene ic p edic ion o PD occu -
ence
1
. The he i abili y o PD can be es ima ed using win s udies o
s a is ical gene ic me hods and is hough o lie be ween 22% and
40% in Eu opean popula ions. Known genome wide significan loci
cu en ly explain ~16% o he he i abili y o PD
1
.Theuseo
polygenic isk sco e analysis (including loci ha do no each
genome wide significance) indica es ha he e a e likely o be a
subs an ial numbe o loci con ibu ing o PD isk ha ha e no ye
been defined. Ou powe analysis indica es ha 99,000 PD cases will
be needed o define loci wi h 80% powe , a mino allele equency o
0.21 and simila e ec s o he cu en s a e-o - he-a analysis. The
a iabili y o pheno ypes obse ed in PD a e likely o ha e a gene ic
basis
2–4
. Knowledge o associa ions be ween geno ype and clinical
ou comes will enable clinicians o p o ide pa ien s wi h a mo e
accu a e p ognosis. Unde s anding he gene- o-pheno ype pa h-
ways esponsible o specific PD ea u es would p o ide an
oppo uni y o de elop ea men s a ge ing pheno ype-specific
disease pa hways, esul ing in mo e e ficien and pe sonalized
ea men wi h ewe side e ec s. We aim o cap u e he di e si y o
PD ou comes, including Pa kinson’s i sel bu also ela ed condi ions
such as p od omal Pa kinson’s, demen ia wi h Lewy bodies and
o he Pa kinson’s plus synd omes, and o pe o m la ge-scale
analyses o clinical-gene ic da a wi h su ficien powe o gene
disco e y. This will comp ise eg ession and ime- o-e en analysis
o he pheno ype o in e es (e.g., demen ia, dyskinesias, mo o
p og ession). I is likely ha his will be limi ed o a ound 25% o
samples included in his s udy wi h in dep h longi udinal da a, and
u he la ge scale longi udinal coho s will be needed o explo e he
biology o p og ession and di e se pheno ypes. The ocus o PD
GWAS on indi iduals o Eu opean ances y has le gaps in ou
knowledge o PD-associa ed gene ic a ian s in unde ep esen ed
popula ions and limi ed ou abili y o esol e GWAS loci
5
.Toad ance
unde s anding o he gene ic de e minan s o Pa kinsonism on a
global scale, we need o ensu e ep esen a ion o di e se ances ies
wi h sample sizes su ficien o de ec ances y-specific signals. We
aim o include a leas 15,000 pa icipan s o A ican, Sou h Asian,
and Eas Asian ances y, espec i ely.
The Global Pa kinson’s Gene ics P og am (GP2, h p://gp2.o g/),
unded by he Aligning Science Ac oss Pa kinson’s ini ia i e (ASAP,
h ps://pa kinsons oadmap.o g/) will ec ui PD, P od omal Pa -
kinson’s and Pa kinson’s Plus (including Demen ia wi h Lewy
bodies, P og essi e Sup anuclea Palsy, Mul iple Sys em A ophy
and Co ico-basal synd ome) coho s om ac oss he wo ld
6
.We
will geno ype >150,000 pa icipan s and in eg a e gene ic da a
wi h ha monized clinical da a o use in case-con ol and
geno ype-pheno ype associa ion s udies. This figu e will include
a minimum o 50,000 indi iduals om ances ies cu en ly
unde ep esen ed in Pa kinson’s esea ch: including Black
1
Depa men o Clinical and Mo emen Neu osciences, Queen Squa e Ins i u e o Neu ology, Uni e si y College London, London, UK.
2
Uni e si y College London, London, UK.
3
Molecula Gene ics Sec ion, Labo a o y o Neu ogene ics, Na ional Ins i u e on Aging, Na ional Ins i u es o Heal h, Be hesda, MD, USA.
4
Na ional Ins i u es o Heal h, Be hesda,
MD, USA.
5
Depa men o Clinical Resea ch, Michael J. Fox Founda ion o Pa kinson’s Resea ch, New Yo k Ci y, NY, USA.
6
The Michael J. Fox Founda ion o Pa kinson’s
Resea ch, New Yo k, NY, USA.
7
Cen e o Alzheime ’s and Rela ed Demen ias (CARD), Na ional Ins i u e on Aging and Na ional Ins i u e o Neu ological Diso de s and S oke,
Na ional Ins i u es o Heal h, Be hesda, MD, USA.
8
Da a Tecnica In e na ional, Washing on, DC, USA.
9
Na ional Ins i u e on Aging/Na ional Ins i u es o Heal h, Be hesda, MD, USA.
10
Depa men o Medical and Molecula Gene ics, Indiana Uni e si y School o Medicine, Indianapolis, IN, USA.
11
In eg a i e Genomics Uni , Labo a o y o Neu ogene ics,
Na ional Ins i u e on Aging, Na ional Ins i u es o Heal h, Be hesda, MD, USA.
12
Na ional Ins i u e on Aging, Be hesda, MD, USA.
13
Depa men o Neu ology, Oslo Uni e si y
Hospi al, Oslo, No way. *A lis o au ho s and hei a filia ions appea a he end o he pape . ✉email: [email p o ec ed]
www.na u e.com/npjpa kd
Published in pa ne ship wi h he Pa kinson’s Founda ion
1234567890():,;
Ame ican, A ican, Middle Eas e n, Cen al/ Eas /Sou h Asian,
Indian, Ca ibbean and Cen al/Sou h Ame ican. The emaining
pa icipan s a e expec ed o be o Eu opean ances y
6
. Cleaned
gene ic and clinical da a will be ha monized ac oss coho s and
made a ailable o Pa kinson’s, P od omal and Pa kinson’s Plus
esea che s ia a con olled-access online eposi o y wi h a unified
use ag eemen . Con ibu ing in es iga o s a e encou aged o
play an ac i e ole in he p ojec by p oposing and leading
analyses, and will ecei e au ho ship on GP2 publica ions as well
as suppo o hei analyses. GP2 also p o ides comp ehensi e
aining oppo uni ies o esea che s om con ibu ing ins i u-
ions ia an online lea ning managemen sys em. The ou come o
GP2 will be an open access esou ce which will in eg a e clinical
and gene ic da a om a e y la ge, di e se sample o cases, and
acili a e disco e y o new gene ic de e minan s o Pa kinson’s,
P od omal and Pa kinson’s Plus occu ence and pheno ypic
a ia ion in mul iple ances ies. A global ne wo k o PD
esea che s will be es ablished, acili a ing u u e collabo a ion
and ad ancemen o PD esea ch. The p ojec began in Janua y
2020 and cu en unding ex ends o 2029.
Coo dina ing coho s a ound he wo ld o include 150,000
pa icipan s is a conside able logis ical challenge. As o May 2022,
we ha e es ablished a wo kflow o coho in eg a ion, a
s anda dized se o clinical da a elemen s, a ecommended
consen empla e, a p o ocol o e alua ing coho s joining he
s udy, and a p ocess o in eg a ing and ha monizing he
incoming clinical da a. Mo e b oadly, we ha e es ablished he
complex hub amewo k, which c ea ed a code o conduc and
publica ion policy, c ea ed a DNA quali y and shipping p o ocol,
and c ea ed he Neu oBoos e A ay (NBA) geno yping wo kflow.
The size and dep h o he da ase which is being gene a ed by
GP2 will p o ide a majo oppo uni y o disco e new geno ype-
pheno ype associa ions. Co e analyses, analyses ha will be
con inuously upda ed by he analysis eams a GP2 wi h he
inclusion o new samples, will include case-con ol, age a onse ,
and p og ession GWAS, as well as wi hin-ances y analyses o
p e iously unde ep esen ed popula ions. Beyond his, GP2 is
suppo ing addi ional analy ical p ojec s p oposed by con ibu ing
collabo a o s. The GP2 ne wo k will also acili a e collabo a ion on
auxilia y s udies be ween in es iga o s sha ing he same a ea o
in e es (e.g., bioma ke s o di e en modali ies) o add ess
ou s anding ques ions in PD esea ch.
GP2 is conduc ed acco ding o o e a ching p inciples o
democ a iza ion o da a, collabo a ion and coope a ion, sa e and
esponsible da a sha ing, commi men o di e si y in esea ch,
anspa ency and ep oducibili y, and p oduc ion o an ac ionable
esou ce in acco dance wi h he Findable, Accessible, In e ope -
able and Reusable (FAIR) scien ific da a managemen p inciples
7
.I
is GP2 policy ha local esea che s a e included in publica ions
ha use hei da a/samples. He e, we epo he specific s eps
unde aken by he Coho In eg a ion Wo king G oup (CIWG) o
iden i y, ec ui , and ha monize clinical coho s wi hin he complex
disease a m o GP2. GP2 also has a monogenic a m (h ps://
gp2.o g/wo king-g oups/) which ocuses on cases wi h po en ial
monogenic causes o PD, i.e., hose wi h ea ly age a onse o a
amily his o y o PD (Re e ence: PMID:37369645).
METHODS
PD and Pa kinson’s Plus in es iga o s and coho s a e iden ified
h ough ele an publica ions, o h ough exis ing conso ia. We
welcome con ac om in e es ed in es iga o s a ound he wo ld
(Email: [email p o ec ed]). P ospec i e collabo a o s comple e a Si e
In e es Fo m (SIF) (Supplemen a y Ma e ial) o p o ide a b ie
o e iew o hei coho [Supplemen a y ma e ial]. Da a a ail-
abili y, ype o coho (e.g., b ain bank, d ug s udy e c.), e hnic
di e si y and sample/da a sha ing es ic ions a e all conside ed
by he CIWG when e iewing SIF submissions. Coho s’consen
documen s a e e iewed by he Ope a ions and Compliance
Wo king G oup (OCWG) o ensu e consen o sample/da a
sha ing was ob ained, and compliance wi h egional and coho -
specific da a sha ing es ic ions. The OCWG assis s he in es i-
ga o i e isions a e needed, and empla e consen language is
p o ided online (h ps://gp2.o g/ esou ces/consen -guidelines/
[Supplemen a y Ma e ial]). We ha e an inclusi e app oach; o-
da e >95% o coho s e alua ed ha e been accep ed. Following
coho app o al, a concise Collabo a ion Ag eemen is signed by
he collabo a ing PI and he Michael J. Fox Founda ion (MJFF), and
he necessa y ma e ial and da a ans e ag eemen s a e
comple ed. The coho ’s samples a e hen ans e ed o one o
GP2’s geno yping cen e s, and da a a e ans e ed h ough secu e
upload o a cloud se e .
Once he coho ’s clinical da a ha e been ans e ed, ha mo-
niza ion and quali y con ol (QC) a e pe o med by he CIWG,
ollowing a s anda d amewo k
8
. Co e clinical da a we e decided
on he basis o clinical scales a ailable in exis ing PD coho s
(p ima ily he T acking Pa kinson’s
9
, PPMI
10
, PDBP
11
, NET PD-LS1
12
,
and GEoPD
13
coho s) as well as on he basis o ecen p oposals
o a modula se o assessmen c i e ia o cha ac e ize long-
i udinal PD coho s
14,15
. The co e clinical da a elemen s a e he
clinical da a ca ego ies o in e es which will be ha monized ac oss
coho s and used o analysis (Table 1). We ha e gene a ed a
common da ase o b ain banks, and ha e added da a elemen s
o ime- o-e en analysis (Supplemen a y Table 1). We a e also
collec ing in o ma ion on a ailabili y o o he da a and samples,
including, plasma, se um, RNA, fib oblas s, CSF, skin biopsy, PBL,
iPS o expand awa eness o he esou ces ha a e a ailable a
con ibu ing si es (Supplemen a y Ma e ial). The common da ase
is defined in a common da a dic iona y (Supplemen a y Table 2).
The ha moniza ion o aw da a uses a se o coding ules o e-
coding and handling o missing da a, using a modified cus om
sc ip o each coho .
DNA samples om each coho a e geno yped on he Neu o Boos e
A ay (NBA; h ps://gi hub.com/GP2code/Neu o_Boos e _A ay), de el-
oped in collabo a ion be ween Da a Tecnica In e na ional, NIA, NINDS,
Illumina Inc and GP2. The NBA consis s o he Illumina Infinium Global
Di e si y A ay (GDA; h ps://www.illumina.com/p oduc s/by- ype/
mic oa ay-ki s/infinium-global-di e si y.h ml), a high-densi y (1.9 M
o al a ian s) global backbone op imized o c oss-popula ion
impu a ion co e age o he human genome, and addi ional cus om
con en (>95,000 a ian s) which includes known causal a ian s o
a ious neu odegene a i e diseases and impu a ion boos e s o
unde ep esen ed popula ions. QC o geno ype da a by he Da a
Analysis Wo king G oup (DAWG) ollows a s anda d pipeline. A cus om
geno ype clus e ing file is used o ensu e ep esen a ion o he di e se
gene ic ances ies wi hin he GP2 da a. The clus e ing file used o he
la es da a elease is based on 2793 samples ac oss 6 ances y g oups,
and includes 420 Gauche disease cases o cap u e a ian s o in e es
in he GBA1 isk gene. This file is a ailable o download ia he GP2
Gi hub eposi o y (h ps://gi hub.com/GP2code). Cleaned gene ic da a
a e e u ned o he con ibu ing in es iga o o use as hey wish, and
a e uploaded o he GP2 eposi o y o use in combined analyses (Fig.
1). GP2 co e s he cos s o geno yping and sample shipmen o all
con ibu ing coho s, and can assis in es iga o s wi h he analysis o
hei coho ’s da a.
The da ase which we a e cu en ly agg ega ing and ha moniz-
ing con ains a wide a ie y o demog aphic and clinical ac o s,
hanks o he di e si y o con ibu ing coho s. As o Ma ch 2023,
he CIWG has app o ed 145 coho s o inclusion in GP2, o which
128 ha e been app o ed by he OCWG, and 74 ha e comple ed all
necessa y ag eemen s and a e in he p ocess o ans e ing
samples and clinical da a. Samples ha ha e been ans e ed a e
cu en ly being geno yped and passed h ough he da a QC
pipeline. So a , he app o ed coho s span o e 50 di e en
coun ies and e i o ies. A map showing he geog aphic
dis ibu ion o hese coho s, as well as cu en expec ed and
C. Towns e al.
2
npj Pa kinson’s Disease (2023) 131 Published in pa ne ship wi h he Pa kinson’s Founda ion
1234567890():,;
Fig. 1 Coho in eg a ion wo kflow. Da a a e con ibu ed and e u ned o he local PI (g een); new gene ic da a, da a cleaning and
ha moniza ion a e ca ied ou by GP2 and made a ailable o analysis wi hin he GP2 conso ium (blue); and da a a e eleased o qualified
in es iga o s ia AMP-PD (o ange). AMP-PD Accele a ing Medicines Pa ne ship - Pa kinson’s Disease (h ps://amp-pd.o g).
Table 1. Recommended co e clinical da a.
MINIMUM MINIMUM PLUS CORE EXTENDED
Demog aphics (e.g., NINDS CDE Gene al Co e)
16
✓✓ ✓✓
Rec ui men ca ego y (Case/Con ol) ✓✓ ✓✓
Family His o y ✓✓ ✓✓
Diagnos ic checklis (MDS clinical diagnosis c i e ia
17
o UK Pa kinson’s Disease Socie y B ain Bank
Diagnos ic C i e ia)
18
✓✓✓
P ima y diagnosis and PD ce ain y (PPMI P ima y Clinical Diagnosis)
10
✓✓✓
PD his o y (e.g., NINDS CDE o Pa kinson’s Disease)
16
✓✓✓
Global PD se e i y (CISI-PD)
19
✓✓✓
Beha io al and En i onmen al His o y (PD-RFQ-U)
16
✓✓
Medical His o y ✓✓
Cu en Medica ion S a us ✓✓
nM-EDL (MDS-UPDRS Pa I [o UPDRS])
20,21
✓✓
M-EDL (MDS-UPDRS Pa II [o UPDRS])
20,21
✓✓
Mo o Assessmen (MDS-UPDRS Pa III [o UPDRS])
20,21
✓✓
Complica ions (MDS-UPDRS Pa IV [o UPDRS])
20,21
✓✓
Cogni i e Assessmen (MMSE, MoCA)
22,23
✓✓
Mo o (Hoehn and Yah s age)
24
✓✓
Au onomic unc ion assessmen (SCOPA-AUT)
25
✓✓
pRBD (RBDSQ)
26
✓✓
Day ime sleepiness (ESS)
27
✓
Dep ession (GDS-15)
28
✓
O hos a ic hypo ension (S anding/si ing/supine HR and BP) ✓
Ol ac o y unc ion (UPSIT o Sni fin-s ick o BSIT)
29–31
✓
Gene al ADL (Schwab & England ADL)
32
✓
PD EQL (PDQ-8)
33
✓
Pain (Kings PD Pain Scale)
34
✓
C. Towns e al.
3
Published in pa ne ship wi h he Pa kinson’s Founda ion npj Pa kinson’s Disease (2023) 131
comple ed sample numbe s, can be ound on he GP2 websi e
(h ps://gp2.o g/coho -dashboa d/ [Supplemen a y Ma e ial]).
Repo ing summa y
Fu he in o ma ion on esea ch design is a ailable in he Na u e
Resea ch Repo ing Summa y linked o his a icle.
DATA AVAILABILITY
GP2 has pa ne ed wi h he Accele a ing Medicines Pa ne ship - Pa kinson’s Disease
(AMP-PD; h ps://amp-pd.o g) o sha e da a gene a ed by GP2, and in Decembe
2021 he fi s GP2 geno yping da a we e eleased on he AMP-PD pla o m. As o
2023, he da a consis o 14,902 samples (8190 PD cases), ep esen ing a b oad ange
o di e se ances ies defined di ec ly om he geno yping da a om hese coho s.
Geno yping and da a QC is ongoing, and he e will be egula da a eleases (2–4
imes pe yea ) as he p ojec p og esses. All con ibu ing in es iga o s ha e access o
GP2 da a, and ex e nal esea che s can also gain access by ollowing ins uc ions on
he GP2 websi e (h ps://gp2.o g/applying- o -gp2-da a-access-on- he-amp-pd-
pla o m/). The e a e wo ie s o da a access. Tie 1 consis s o summa y s a is ics
and any esea che can gain access by comple ing an online applica ion. Tie 2 access
includes deiden ified, indi idual le el gene ic da a, and o gain access esea che s
mus sign a Da a Usage Ag eemen co-signed by hei ins i u ion.
CODE AVAILABILITY
Code used in GP2 co e analyses and he cus om clus e ing files a e deposi ed in he
GP2 Gi Hub eposi o y (h ps://gi hub.com/GP2code). Code c ea ed and used by he
CIWG can be ound on he “GP2-Wo kingG oups/CD-Coho -In eg a ion”subpage o
he GP2 Gi Hub eposi o y (h ps://gi hub.com/GP2code/GP2-Wo kingG oups/ ee/
main/CD-Coho -In eg a ion). The Da a and Code Dissemina ion Wo king G oup
(DCD-WG) a GP2 is commi ed o making hese da a, esou ces, and aining
ma e ials accessible o he scien ific communi y. The DCD-WG ac i ely suppo s
e o s o consolida e analysis sc ip s wi h he necessa y analy ical ools and decisions
ia Gi Hub (h ps://gi hub.com/GP2code), consolida e aining ma e ials wi h se e al
ansla ions in collabo a ion wi h he T aining and Ne wo king Wo king G oup
(h ps://gp2.o g/ aining/), and consolida e addi ional analy ical pipelines on Te a
(h ps://amp-pd.o g/ ools).
Recei ed: 25 No embe 2022; Accep ed: 22 May 2023;
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ACKNOWLEDGEMENTS
This esea ch is suppo ed by he Aligning Science Ac oss Pa kinson’s Ini ia i e, he
In amu al Resea ch P og am, Na ional Ins i u e on Aging, Na ional Ins i u es o
Heal h, Depa men o Heal h and Human Se ices, p ojec ZO1 AG000949, and he
Michael J. Fox Founda ion o Pa kinson’s Resea ch. Da a used in he p epa a ion o
his a icle we e ob ained om Global Pa kinson’s Gene ics P og am (GP2). GP2 is
unded by he Aligning Science Ac oss Pa kinson’s (ASAP) ini ia i e and implemen ed
by The Michael J. Fox Founda ion o Pa kinson’s Resea ch (h ps://gp2.o g). Fo a
comple e lis o GP2 membe s see h ps://gp2.o g.
AUTHOR CONTRIBUTIONS
C.T., M.R., S.J., E.J.S., J.J., T.A., A.C.M., M.M.X.T., H.I., and H.R.M. a e membe s o he GP2
Coho In eg a ion Wo king G oup (CIWG), o which H.R.M. is he lead, and M.M.X.T.
and H.I. a e co-leads. C.T. was he p ima y con ibu o in he d a ing o his
manusc ip , assis ed by M.R. and J.J. The d a was e iewed by all CIWG membe s
p io o ci cula ion o o he con ibu ing au ho s o e iew. M.B.M. is he co-lead o
he GP2 Da a and Code Dissemina ion Wo king G oup and d a ed he da a
a ailabili y sec ion o his manusc ip , oge he wi h B.C. who is senio associa e
C. Towns e al.
4
npj Pa kinson’s Disease (2023) 131 Published in pa ne ship wi h he Pa kinson’s Founda ion
di ec o a he Michael J. Fox Founda ion’s esea ch di ision. D.V., K.L., H.L., and M.A.N.
a e membe s o he GP2 Complex Disease Da a Analysis Wo king G oup (DAWG), o
which M.A.N. is he lead and H.L. is a co-lead. DAWG a e esponsible o he da a
cleaning and analysis desc ibed in his manusc ip . DGH di ec s he Genomic
Technologies G oup wi hin he Labo a o y o Neu ogene ics a NIH, which conduc s
he majo i y o sample geno yping o he Complex Ne wo k. C.E.W. and J.S. a e
membe s o he GP2 Ope a ions and Compliance Wo king G oup o which J.S. is a co-
lead. C.B.P. and L.A.S. a e scien ific p og am manage s o he GP2 Complex Disease
Ne wo k. A.B.S. is he lead o he GP2 Complex Disease Ne wo k and C.B. is he co-
lead. All lis ed au ho s e iewed he manusc ip and p o ided commen s and
e isions p io o submission.
COMPETING INTERESTS
A.B.S. and C.B. a e suppo ed by he In amu al Resea ch P og am o he Na ional
Ins i u e on Aging and ha e ecei ed g an suppo om he Michael J. Fox
Founda ion o Pa kinson’s Resea ch and he Aligning Science Ac oss Pa kinson’s
Ini ia i e. A.B.S. has ecei ed oyal y paymen s ela ed o a diagnos ic o s oke. A.B.S.
is an edi o o npj Pa kinson’s Disease. A.B.S. was no in ol ed in he jou nal’s e iew
o , o decisions ela ed o, his manusc ip . H.L., H.I., D.V., K.L., and M.A.N. a e
consul an s employed by Da a Tecnica In e na ional. Da a Tecnica is engaged in a
consul ing ag eemen wi h he US Na ional Ins i u es o Heal h. H.R.M. is employed
by UCL. In he las 24 mon hs, he epo s paid consul ancy om Biogen, Bioha en,
Lundbeck; lec u e ees/hono a ia om Wellcome T us , Mo emen Diso de s Socie y;
Resea ch G an s om Pa kinson’s UK, Cu e Pa kinson’s T us , PSP Associa ion, CBD
Solu ions, D ake Founda ion, Medical Resea ch Council, Michael J. Fox Founda ion.
H.R.M. is also a co-applican on a pa en applica ion ela ed o C9ORF72—Me hod o
diagnosing a neu odegene a i e disease (PCT/GB2012/052140). BC and JS a e
employed by he Michael J. Fox Founda ion o Pa kinson’s Resea ch. All o he
au ho s decla e no financial o non-financial compe ing in e es s.
ADDITIONAL INFORMATION
Supplemen a y in o ma ion The online e sion con ains supplemen a y ma e ial
a ailable a h ps://doi.o g/10.1038/s41531-023-00533-w.
Co espondence and eques s o ma e ials should be add essed o Huw R. Mo is.
Rep in s and pe mission in o ma ion is a ailable a h p://www.na u e.com/
ep in s
Publishe ’s no e Sp inge Na u e emains neu al wi h ega d o ju isdic ional claims
in published maps and ins i u ional a filia ions.
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© The Au ho (s) 2023
THE GLOBAL PARKINSON’S GENETICS PROGRAM (GP2)
Emilia M. Ga o14, Ma celo Kau man15, Samson Khacha yan16, Za uhi Ta adyan16, Clai e E. Shephe d17, Julie Hun e 18,
Kisho e Kuma 19, Melina Ellis20, Miguel E. Ren e ía21, Sule Koks22, Alexande Zimp ich23, A u F. Schumache -Schuh24, Ca los Riede 25,
Paula Sa fie Awad26, Vi o Tumas27, Sa ah Cama gos28, Edwa d A. Fon29, Ou y Monchi30, Ted Fon31, Benjamin Piza o Galleguillos32,
Ma celo Mi anda33, Ma ia Leono Bus aman e34, Pa icio Olguin32, Ped o Chana35, Beisha Tang36, Hui ang Shang37, Ji eng Guo38,
Piu Chan39, Wei Luo40, Gonzalo A boleda41, Jo ge O ozco42, Ma lene Jimenez del Rio43, Al a o He nandez44, Mohamed Salama45,
Walaa A. Kamel46, Ya ed Z. Zewde47, Alexis B ice48, Jean-Ch is ophe Co ol49, Ana Wes enbe ge 50, Anas asia Illa iono a51,
B i Mollenhaue 52, Ch is ine Klein50, E a-Juliane Volls ed 50, F anziska Hop ne 53, Gün e Höglinge 53, Ha u yun Madoe 50,
Joanne T inh50, Johanna Junke 50, Ka ja Lohmann50, La a M. Lange54, Manu Sha ma55, Se giu G oppa56, Thomas Gasse 55,
Zih-Hua Fang57, Albe Akpalu58, Geo gia Xi ome isiou59, Geo gios Hadjigo giou59, Ioannis Dagklis60, Ioannis Ta nanas61,
Leonidas S e anis62, Ma ia S amelou63, E hymios Dadio is59, Alex Medina64, Ge maine Hiu-Fai Chan65, Nancy Ip66,
Nelson Yuk-Fai Cheung65, Phillip Chan66, Xiaopu Zhou66, Asha Kisho e67, Di ya KP68, P amod Pal69, P ashan h Lingappa Kukkle70,
Roopa Rajan71, Rupam Bo gohain72, Meh i Sala i73, And ea Qua one74, Enza Ma ia Valen e75, Lucilla Pa ne i76, Micol A enali75,
Tommaso Schi inzi77, Manabu Funayama78, Nobu aka Ha o i79, Tomo aka Shi aishi80, Al ynay Ka imo a81, Gulnaz Kaishibaye a81,
Cholpon Shambe o a82, Rejko K üge 83, Ai Huey Tan84, Azlina Ahmad-Annua 84, Mohamed Ib ahim No linah85,
No Azian Abdul Mu ad86, Shah ul Azmin87, Shen-Yang Lim84, Wael Mohamed88, Yi Wen Tay84, Daniel Ma inez-Rami ez89,
Mayela Rod iguez-Violan e90, Paula Reyes-Pé ez91, Bayasgalan Tse ensodnom92, Rajee Ojha93, Tim J. Ande son94, Toni L. Pi che 94,
A inola Sanyaolu95, Njideka Okubadejo95, Oluwadamilola Ojo96, Jan O. Aasly97, Lasse Pihls øm98, Manuela Tan98, Shoaib U -Rehman99,
Ma io Co nejo-Oli as100, Ma ia Leila Doquenia101, Raymond Rosales101, Angel Vinuela102, Elena Iako enko103, Bashaye Al Muba ak104,
Muhammad Umai 105, Eng-King Tan106, Jia Nee Foo107, Fe zana Amod108, Jona han Ca 109, So aya Ba dien110, Beomseok Jeon111,
Yun Joong Kim112, Es he Cubo113, Ignacio Al a ez114, Jane Hoenicka115, Ka in Beye 116, Ma ia Te esa Pe iñan117, Pau Pas o 118,
Sa ah El-Sadig119, Ch is iane Zweie 120, Paul K ack120, Chin-Hsien Lin121, Hsiu-Chuan Wu122, Pin-Jui Kung123, Ruey-Meei Wu121,
Yih u Wu122, Rim Amou i124, Samia Ben Sassi125, A. NazlıBaşak126, Gence Genc127, Özgü Öz op Çakmak126, Sibel E an126,
Alas ai Noyce128, Alejand o Ma inez-Ca asco 1,2, Ane e Sch ag2, An hony Schapi a2, Camille Ca oll129, Clai e Bale130,
Donald G osse 131, Eleano J. S a o d 1,2, Hen y Houlden2, Huw R. Mo is 1,2 ✉, John Ha dy2, Kin Ying Mok2, Mie Rizig2,
Nicholas Wood2, Nigel Williams132, Olai an Okunoye2, Pa ick Al yn Lewis133, Rauan Kaiy zhano 2, Rimona Weil2, Se h Lo e134,
Simon S o 135, Simona Jasai ye2, Sumi Dey128, Vida Obese2, Albe o Espay136, Alyssa O’G ady6, And ew B. Single on 3,7,12,
And ew K. Sobe ing137, Be nade e Siddiqi6, B ad o d Casey 5,6, B ian Fiske6, Cabell Jonas138, Ca los C uchaga139,
Ca oline B. Pan azis 4,7, Cha isse Coma 6, Clai e Wegel140, Co nelis Blauwend aa 4,7,11, Dan Vi ale4,7,8, Debo ah Hall141,
Dena He nandez4, Ejaz Shiamim142, Ekemini Riley143, Fa az Fagh i4, Geidy E. Se ano144, Hamp on Leona d3,7,8,9, Hi o aka Iwaki3,7,8,
Honglei Chen145, Ignacio F. Ma a146, Ignacio Juan Kelle Sa mien o147, Ja ed Williamson142, Jonggeol Je Kim4, Joseph Janko ic148,
Joshua Shulman149, Jus in C. Solle6, Kaileigh Mu phy6, Ka en Nuy emans150, Ka l Kiebu z151, Ka e ina Ma kopoulou152,
Kenne h Ma ek153, K is in S. Le ine8, Lana M. Chahine154, Lau a Ibanez139, Lau el Sc e en12, Lau en Ru age155, Lisa Shulman156,
Luca Ma sili136, Maggie Kuhl6, Ma issa Dean155, Ma y B. Maka ious 1,3,4, Ma hew Ko e sky4, Megan J. Puckelwa z157,
C. Towns e al.
5
Published in pa ne ship wi h he Pa kinson’s Founda ion npj Pa kinson’s Disease (2023) 131

Miguel Inca-Ma inez146, Mike A. Nalls3,4,7,8, Naomi Louie6, Niccolò Emanuele Mencacci147, Roge Albin158, Roy Alcalay159,
Ru h Walke 160, Sa a Band es-Ciga4, Sohini Chowdhu y6, Sonya Dumanis161, S e en Lubbe157, Tao Xie162, Ta iana Fo oud163,
Thomas Beach164, Todd She e 6, Yeajin Song4, Duan Nguyen165, Toan Nguyen165 and Masha ip A adzhano 166
14
Sana o io de la T inidad Mi e- INEBA, Buenos Ai es, A gen ina.
15
Hospi al JM Ramos Mejia, Buenos Ai es, A gen ina.
16
Somnus Neu ology Clinic, Ye e an, A menia.
17
Neu oscience Resea ch Aus alia, Sydney, NSW, Aus alia.
18
ANZAC Resea ch Ins i u e, Conco d, NSW, Aus alia.
19
Ga an Ins i u e o Medical Resea ch and Conco d Repa ia ion
Gene al Hospi al, Da linghu s , NSW, Aus alia.
20
Conco d Hospi al, Conco d, NSW, Aus alia.
21
QIMR Be gho e Medical Resea ch Ins i u e, He s on, QLD, Aus alia.
22
Mu doch
Uni e si y, Pe h, Aus alia.
23
Medical Uni e si y Vienna Aus ia, Vienna, Aus ia.
24
Uni e sidade Fede al do Rio G ande do Sul / Hospi al de Clínicas de Po o Aleg e, Po o Aleg e,
B azil.
25
Fede al Uni e si y o Heal h Sciences o Po o Aleg e, Po o Aleg e, B azil.
26
Uni e sidade Fede al do Rio G ande do Sul, Po o Aleg e, B azil.
27
Uni e si y o São Paulo, São
Paulo, B azil.
28
Uni e sidade Fede al de Minas Ge ais, Belo Ho izon e, B azil.
29
Mon eal Neu ological Ins i u e, Mon eal, QC, Canada.
30
Ins i u uni e si ai e de gé ia ie de
Mon éal, Mon eal, QC, Canada.
31
McGill Uni e si y, Mon eal, QC, Canada.
32
Uni e sidad de Chile, San iago, Chile.
33
Fundación Diagnosis, San iago, Chile.
34
Facul y o Medicine
Uni e sidad de Chile, San iago, Chile.
35
CETRAM, San iago, Chile.
36
Cen al Sou h Uni e si y, Changsha, China.
37
Wes China Hospi al Sichuan Uni e si y, Chengdu, China.
38
Xiangya Hospi al, Changsha, China.
39
Capi al Medical Uni e si y, Beijing, China.
40
Zhejiang Uni e si y, Hangzhou, China.
41
Uni e sidad Nacional de Colombia, Bogo á, Colombia.
42
Fundación Valle del Lili, San iago De Cali, Colombia.
43
Uni e si y o An ioquia, Medellin, Colombia.
44
Uni e si y o Cos a Rica, San Jose, Cos a Rica.
45
The Ame ican Uni e si y in
Cai o, Cai o, Egyp .
46
Beni-Sue Uni e si y, Beni Sue , Egyp .
47
Addis Ababa Uni e si y, Addis Ababa, E hiopia.
48
Pa is B ain Ins i u e, Pa is, F ance.
49
So bonne Uni e si é, Pa is,
F ance.
50
Uni e si y o Lübeck, Lübeck, Ge many.
51
Deu sches Zen um ü Neu odegene a i e E k ankungen, Gö ingen, Ge many.
52
Uni e si y Medical Cen e
Gö ingen, Gö ingen, Ge many.
53
Depa men o Neu ology, Uni e si y Hospi al, LMU Munich, Munich, Ge many.
54
Uni e si y o Lübeck and Uni e si y Medical Cen e
Schleswig-Hols ein, Lübeck, Ge many.
55
Uni e si y o Tubingen, Tübingen, Ge many.
56
Uni e si y o Mainz, Mainz, Ge many.
57
The Ge man Cen e o Neu odegene a i e
Diseases, Gö ingen, Ge many.
58
Uni e si y o Ghana Medical School, Acc a, Ghana.
59
Uni e si y o Thessaly, Volos, G eece.
60
A is o le Uni e si y o Thessaloniki, Thessaloniki,
G eece.
61
Ionian Uni e si y, Co u, G eece.
62
Biomedical esea ch Founda ion o he Academy o A hens, A hens, G eece.
63
Diagnos ic and The apeu ic Cen e HYGEIA
Hospi al, Ma ousi, G eece.
64
Hospi al San Felipe, Tegucigalpa, Hondu as.
65
Queen Elizabe h Hospi al, Kowloon, Hong Kong.
66
The Hong Kong Uni e si y o Science and
Technology, Kowloon, Hong Kong.
67
As e Medci y, Kochi, India.
68
S ee Chi a Ti unal Ins i u e o Medical Sciences and Technology, Thi u anan hapu am, India.
69
Na ional
Ins i u e o Men al Heal h & Neu osciences, Bengalu u, India.
70
Manipal Hospi al, Delhi, India.
71
All India Ins i u e o Medical Sciences, Delhi, India.
72
Nizam’s Ins i u e O Medical
Sciences, Hyde abad, India.
73
Shahid Behesh i Uni e si y o Medical Science, Teh an, I an.
74
Magna G æcia Uni e si y o Ca anza o, Ca anza o, I aly.
75
Uni e si y o Pa ia, Pa ia,
I aly.
76
Uni e si y o Pe ugia, Pe ugia, I aly.
77
Uni e si y o Rome To Ve ga a, Rome, I aly.
78
Jun endo Uni e si y, Tokyo, Japan.
79
Jun endo Uni e si y acul y o medicine, Tokyo,
Japan.
80
Jikei Uni e si y School o Medicine, Tokyo, Japan.
81
Ins i u e o Neu ology and Neu o ehabili a ion, Alma y, Kazakhs an.
82
Ky gyz S a e Medical Academy, Bishkek,
Ky gyzs an.
83
Uni e si y o Luxembou g, Luxembou g, Luxembou g.
84
Uni e si y o Malaya, Kuala Lumpu , Malaysia.
85
Uni e si i Kebangsaan Malaysia, Selango , Malaysia.
86
UKM
Medical Molecula Biology Ins i u e, Kuala Lumpu , Malaysia.
87
Uni e si i Kebangsaan Malaysia Medical Cen e, Kuala Lumpu , Malaysia.
88
In e na ional Islamic Uni e si y, Kuala
Lumpu , Malaysia.
89
Tecnologico de Mon e ey, Mon e ey, Mexico.
90
Ins i u o Nacional de Neu ologia y Neu oci ugia, Mexico Ci y, Mexico.
91
Uni e sidad Nacional Au ónoma de
México, Mexico Ci y, Mexico.
92
Mongolian Na ional Uni e si y o Medical Sciences, Ulaanbaa a , Mongolia.
93
T ibhu an Uni e si y, Ki ipu , Nepal.
94
Uni e si y o O ago, Dunedin,
New Zealand.
95
Uni e si y o Lagos, Lagos, Nige ia.
96
College o Medicine o he Uni e si y o Lagos, Lagos, Nige ia.
97
No wegian Uni e si y o Science and
Technology, T ondheim, No way.
98
Oslo Uni e si y Hospi al, Oslo, No way.
99
Uni e si y o Science and Technology Bannu, Bannu, Pakis an.
100
Uni e sidad Cien ifica del
Su , Lima, Pe u.
101
Me opoli an Medical Cen e , Manila, Philippines.
102
Uni e si y o Pue o Rico, San Juan, Pue o Rico.
103
Resea ch Cen e o Neu ology, Moscow, Russia.
104
King
Faisal Specialis Hospi al and Resea ch Cen e , Riyadh, Saudi A abia.
105
King Abdullah In e na ional Medical Resea ch Cen e , Jeddah, Saudi A abia.
106
Na ional Neu oscience
Ins i u e, Singapo e, Singapo e.
107
Nanyang Technological Uni e si y, Singapo e, Singapo e.
108
Uni e si y o KwaZulu-Na al, Du ban, Sou h A ica.
109
Uni e si y o
S ellenbosch, S ellenbosch, Sou h A ica.
110
S ellenbosch Uni e si y, S ellenbosch, Sou h A ica.
111
Seoul Na ional Uni e si y Hospi al, Seoul, Sou h Ko ea.
112
Yongin
Se e ance Hospi al, Seoul, Sou h Ko ea.
113
Hospi al Uni e si a io Bu gos, Bu gos, Spain.
114
Uni e si y Hospi al Mu ua Te assa, Ba celona, Spain.
115
Ins i u de Rece ca San Joan
de Deu, Ba celona, Spain.
116
Resea ch Ins i u e Ge mans T ias i Pujol, Ba celona, Spain.
117
Ins i u o de Biomedicina de Se illa, Se ille, Spain.
118
Uni e si y Hospi al Ge mans T ias i
Pujol, Ba celona, Spain.
119
Facul y o medicine uni e si y o Kha oum, Kha oum, Sudan.
120
Inselspi al Be n, Uni e si y o Be n, Be n, Swi ze land.
121
Na ional Taiwan Uni e si y
Hospi al, Taipei Ci y, Taiwan.
122
Chang Gung Memo ial Hospi al, Taoyuan Ci y, Taiwan.
123
Na ional Taiwan Uni e si y, Taipei Ci y, Taiwan.
124
Na ional Ins i u e Mongi Ben Hamida
o Neu ology, Tunis, Tunisia.
125
Mongi Ben Hmida Na ional Ins i u e o Neu ology, Tunis, Tunisia.
126
Koç Uni e si y, Is anbul, Tu key.
127
Sisli E al T aining and Resea ch
Hospi al, Is anbul, Tu key.
128
Queen Ma y Uni e si y o London, London, UK.
129
Uni e si y o Plymou h, Plymou h, UK.
130
Pa kinson’s UK, London, UK.
131
Uni e si y o
Glasgow, Glasgow, UK.
132
Ca di Uni e si y, Ca di , UK.
133
Royal Ve e ina y College Uni e si y o London, London, UK.
134
Uni e si y o B is ol, B is ol, UK.
135
Cu e
Pa kinson’s, London, UK.
136
Uni e si y o Cincinna i, Cincinna i, OH, USA.
137
Augus a Uni e si y / Uni e si y o Geo gia Medical Pa ne ship, Augus a, GA, USA.
138
Mid-A lan ic
Pe manen e Medical G oup, Be hesda, MD, USA.
139
Washing on Uni e si y, S . Louis, MO, USA.
140
Indiana Uni e si y, Blooming on, IN, USA.
141
Rush Uni e si y, Chicago, IL, USA.
142
Kaise Pe manen e, Oakland, CA, USA.
143
Coali ion o Aligning Science, Washing on, WA, USA.
144
Banne Sun Heal h Resea ch Ins i u e, Sun Ci y, AZ, USA.
145
Michigan S a e
Uni e si y, Eas Lansing, MI, USA.
146
Cle eland Clinic, Cle eland, OH, USA.
147
No hwes e n Uni e si y, E ans on, IL, USA.
148
Baylo College o Medicine, Hous on, TX, USA.
149
Baylo College o Medicine, Texas Child en’s Hospi al, Hous on, TX, USA.
150
Uni e si y o Miami Mille School o Medicine, Miami, FL, USA.
151
Be h Is ael Deaconess Medical
Cen e , Bos on, MA, USA.
152
No h Sho e Uni e si y Heal h Sys em, Chicago, IL, USA.
153
Ins i u e o Neu odegene a i e Diso de s, New Ha en, CT, USA.
154
Uni e si y o
Pi sbu gh, Pi sbu gh, PA, USA.
155
Uni e si y o Alabama a Bi mingham, Bi mingham, AL, USA.
156
Uni e si y o Ma yland, Bal imo e, MD, USA.
157
No hwes e n
Uni e si y, Chicago, IL, USA.
158
Uni e si o Michigan, Ann A bo , MI, USA.
159
Columbia Uni e si y, New Yo k, NY, USA.
160
James J. Pe e s Ve e ans A ai s Medical Cen e , New
Yo k, NY, USA.
161
Aligning Science Ac oss Pa kinson’s, Washing on, WA, USA.
162
Uni e si y o Chicago, Chicago, IL, USA.
163
Indiana Uni e si y School o Medicine, Indianapolis, IN,
USA.
164
Sun Heal h Resea ch Ins i u ion, Sun Ci y, AZ, USA.
165
Hue Uni e si y, Huế, Vie nam.
166
Uni e si y o Zambia, Lusaka, Zambia.
C. Towns e al.
6
npj Pa kinson’s Disease (2023) 131 Published in pa ne ship wi h he Pa kinson’s Founda ion