Impact of Demographic History and Natural Selection on Human Genomic Diversity

Sarah TishkoffDepartments of Genetics and Biology

University of Pennsylvania

•Mutation

•Genetic Drift

•Migration

•Selection

Evolutionary Forces Influencing Genetic Variation

150-195

80-105

150-160

50-80

80-125

140-150

110-130

110-12065-90

80-105

90-110(ages in Kya) 90-100

Modern Human Fossil Record

Human Molecular Genetics, 3/e (©Garland Science 2004)

Major Migration Routes

• Identical twins 0

• Unrelated humans 1/1,000

• Human vs. chimp 1/100

• Human vs. mouse 1/30

• Human vs. broccoli 2/3

• 3 billion DNA bases 3 million differences between each pair of individuals (slide courtesy of Lynn Jorde)

How much do we differ?

http://dariaburke.wordpress.com/2011/03/21/ethic-diversity-at-estee-lauder/

• There is also considerable structural variation across human genomes (insertions/deletions, gene duplications, and inversions)

• More genetic variation within (~85%) relative to between populations (~15%)

Genetic Diversity Within and Between Humans

The Genetic Structure and History of Africans and African AmericansTishkoff et al. Science May 22;324(5930):1035-44. 2009

1165 Microsatellite and In/Del Polymorphisms

Genotyped in >2,500 Africans from 121 ethnic populations

98 African Americans from four regions in the US

>1,500 comparative non-African individuals

Diversity Levels are Highest in Africa

Chart1

NtumuNtumuNtumuNtumuNtumuNtumu

BakaBakaBakaBakaBakaBaka

BurungeBurungeBurungeBurungeBurungeBurunge

BakolaBakolaBakolaBakolaBakolaBakola

SanSanSanSanSanSan

SukumaSukumaSukumaSukumaSukumaSukuma

Tikar NorthTikar NorthTikar NorthTikar NorthTikar NorthTikar North

YakomaYakomaYakomaYakomaYakomaYakoma

GogoGogoGogoGogoGogoGogo

MvaeMvaeMvaeMvaeMvaeMvae

Bantu SouthBantu SouthBantu SouthBantu SouthBantu SouthBantu South

Cape TownCape TownCape TownCape TownCape TownCape Town

SandaweSandaweSandaweSandaweSandaweSandawe

BanenBanenBanenBanenBanenBanen

IyassaIyassaIyassaIyassaIyassaIyassa

MabeaMabeaMabeaMabeaMabeaMabea

VendaVendaVendaVendaVendaVenda

BuluBuluBuluBuluBuluBulu

BedzanBedzanBedzanBedzanBedzanBedzan

LuhyaLuhyaLuhyaLuhyaLuhyaLuhya

MbugweMbugweMbugweMbugweMbugweMbugwe

ChicagoChicagoChicagoChicagoChicagoChicago

BiakaBiakaBiakaBiakaBiakaBiaka

LemandeLemandeLemandeLemandeLemandeLemande

TuruTuruTuruTuruTuruTuru

BatieBatieBatieBatieBatieBatie

IgalaIgalaIgalaIgalaIgalaIgala

BaltimoreBaltimoreBaltimoreBaltimoreBaltimoreBaltimore

ZimeZimeZimeZimeZimeZime

KanembouKanembouKanembouKanembouKanembouKanembou

IgboIgboIgboIgboIgboIgbo

YambassaYambassaYambassaYambassaYambassaYambassa

BafiaBafiaBafiaBafiaBafiaBafia

RangiRangiRangiRangiRangiRangi

DioulaDioulaDioulaDioulaDioulaDioula

DatogaDatogaDatogaDatogaDatogaDatoga

KongoKongoKongoKongoKongoKongo

BaregaBaregaBaregaBaregaBaregaBarega

LuoLuoLuoLuoLuoLuo

SamburuSamburuSamburuSamburuSamburuSamburu

GbayaGbayaGbayaGbayaGbayaGbaya

BurjiBurjiBurjiBurjiBurjiBurji

North CarolinaNorth CarolinaNorth CarolinaNorth CarolinaNorth CarolinaNorth Carolina

Tikar SouthTikar SouthTikar SouthTikar SouthTikar SouthTikar South

BamounBamounBamounBamounBamounBamoun

BalubaBalubaBalubaBalubaBalubaBaluba

HadzaHadzaHadzaHadzaHadzaHadza

TupuriTupuriTupuriTupuriTupuriTupuri

ParePareParePareParePare

KanuriKanuriKanuriKanuriKanuriKanuri

BatangaBatangaBatangaBatangaBatangaBatanga

FangFangFangFangFangFang

XhosaXhosaXhosaXhosaXhosaXhosa

LakaLakaLakaLakaLakaLaka

Yoruba CEPHYoruba CEPHYoruba CEPHYoruba CEPHYoruba CEPHYoruba CEPH

EwondoEwondoEwondoEwondoEwondoEwondo

KabaKabaKabaKabaKabaKaba

WafiomeWafiomeWafiomeWafiomeWafiomeWafiome

Bantu KenyaBantu KenyaBantu KenyaBantu KenyaBantu KenyaBantu Kenya

Maasai TanzaniaMaasai TanzaniaMaasai TanzaniaMaasai TanzaniaMaasai TanzaniaMaasai Tanzania

Samba'aSamba'aSamba'aSamba'aSamba'aSamba'a

Fulani NigeriaFulani NigeriaFulani NigeriaFulani NigeriaFulani NigeriaFulani Nigeria

MbuguMbuguMbuguMbuguMbuguMbugu

WimbumWimbumWimbumWimbumWimbumWimbum

GwariGwariGwariGwariGwariGwari

NgumbaNgumbaNgumbaNgumbaNgumbaNgumba

OuldemeOuldemeOuldemeOuldemeOuldemeOuldeme

BrongBrongBrongBrongBrongBrong

BulalaBulalaBulalaBulalaBulalaBulala

KikuyuKikuyuKikuyuKikuyuKikuyuKikuyu

PittsburghPittsburghPittsburghPittsburghPittsburghPittsburgh

TurkanaTurkanaTurkanaTurkanaTurkanaTurkana

NandiNandiNandiNandiNandiNandi

MadaMadaMadaMadaMadaMada

TugenTugenTugenTugenTugenTugen

KotokoKotokoKotokoKotokoKotokoKotoko

AshantiAshantiAshantiAshantiAshantiAshanti

YorubaYorubaYorubaYorubaYorubaYoruba

Hausa NigeriaHausa NigeriaHausa NigeriaHausa NigeriaHausa NigeriaHausa Nigeria

DoroboDoroboDoroboDoroboDoroboDorobo

YaakuYaakuYaakuYaakuYaakuYaaku

NgambayeNgambayeNgambayeNgambayeNgambayeNgambaye

SabaotSabaotSabaotSabaotSabaotSabaot

Maasai MumonyotMaasai MumonyotMaasai MumonyotMaasai MumonyotMaasai MumonyotMaasai Mumonyot

GabraGabraGabraGabraGabraGabra

MarakwetMarakwetMarakwetMarakwetMarakwetMarakwet

Maasai Il'ngwesiMaasai Il'ngwesiMaasai Il'ngwesiMaasai Il'ngwesiMaasai Il'ngwesiMaasai Il'ngwesi

BassangeBassangeBassangeBassangeBassangeBassange

GizigaGizigaGizigaGizigaGizigaGiziga

Hausa CameroonHausa CameroonHausa CameroonHausa CameroonHausa CameroonHausa Cameroon

MandaraMandaraMandaraMandaraMandaraMandara

IlchamusIlchamusIlchamusIlchamusIlchamusIlchamus

MandenkaMandenkaMandenkaMandenkaMandenkaMandenka

Sara variousSara variousSara variousSara variousSara variousSara various

!Xun/Kxoe!Xun/Kxoe!Xun/Kxoe!Xun/Kxoe!Xun/Kxoe!Xun/Kxoe

IraqwIraqwIraqwIraqwIraqwIraqw

KonsoKonsoKonsoKonsoKonsoKonso

MassaMassaMassaMassaMassaMassa

El MoloEl MoloEl MoloEl MoloEl MoloEl Molo

Fulani AdamawaFulani AdamawaFulani AdamawaFulani AdamawaFulani AdamawaFulani Adamawa

MbumMbumMbumMbumMbumMbum

ShuwaShuwaShuwaShuwaShuwaShuwa

MbutiMbutiMbutiMbutiMbutiMbuti

BoranaBoranaBoranaBoranaBoranaBorana

SengwerSengwerSengwerSengwerSengwerSengwer

RendilleRendilleRendilleRendilleRendilleRendille

PodokwoPodokwoPodokwoPodokwoPodokwoPodokwo

WataWataWataWataWataWata

KomaKomaKomaKomaKomaKoma

AkieAkieAkieAkieAkieAkie

Fulani MbororoFulani MbororoFulani MbororoFulani MbororoFulani MbororoFulani Mbororo

ShilookShilookShilookShilookShilookShilook

OgiekOgiekOgiekOgiekOgiekOgiek

NuerNuerNuerNuerNuerNuer

ZulgoZulgoZulgoZulgoZulgoZulgo

Beta IsraelBeta IsraelBeta IsraelBeta IsraelBeta IsraelBeta Israel

Beja HadandawaBeja HadandawaBeja HadandawaBeja HadandawaBeja HadandawaBeja Hadandawa

Beja BanuamirBeja BanuamirBeja BanuamirBeja BanuamirBeja BanuamirBeja Banuamir

NyiamanNyiamanNyiamanNyiamanNyiamanNyiaman

Tutsi/HutuTutsi/HutuTutsi/HutuTutsi/HutuTutsi/HutuTutsi/Hutu

PokotPokotPokotPokotPokotPokot

DinkaDinkaDinkaDinkaDinkaDinka

MozabiteMozabiteMozabiteMozabiteMozabiteMozabite

BedouinBedouinBedouinBedouinBedouinBedouin

BalochiBalochiBalochiBalochiBalochiBalochi

PalestinianPalestinianPalestinianPalestinianPalestinianPalestinian

HindiHindiHindiHindiHindiHindi

PathanPathanPathanPathanPathanPathan

KannadaKannadaKannadaKannadaKannadaKannada

PunjabiPunjabiPunjabiPunjabiPunjabiPunjabi

TemaniTemaniTemaniTemaniTemaniTemani

MakraniMakraniMakraniMakraniMakraniMakrani

UygurUygurUygurUygurUygurUygur

TeluguTeluguTeluguTeluguTeluguTelugu

SindhiSindhiSindhiSindhiSindhiSindhi

TamilTamilTamilTamilTamilTamil

HazaraHazaraHazaraHazaraHazaraHazara

KashmiriKashmiriKashmiriKashmiriKashmiriKashmiri

BrahuiBrahuiBrahuiBrahuiBrahuiBrahui

AdygeiAdygeiAdygeiAdygeiAdygeiAdygei

BengaliBengaliBengaliBengaliBengaliBengali

DruzeDruzeDruzeDruzeDruzeDruze

AssameseAssameseAssameseAssameseAssameseAssamese

ParsiParsiParsiParsiParsiParsi

OriyaOriyaOriyaOriyaOriyaOriya

KonkaniKonkaniKonkaniKonkaniKonkaniKonkani

MarathiMarathiMarathiMarathiMarathiMarathi

ItalianItalianItalianItalianItalianItalian

DaurDaurDaurDaurDaurDaur

BurushoBurushoBurushoBurushoBurushoBurusho

OrcadianOrcadianOrcadianOrcadianOrcadianOrcadian

FrenchFrenchFrenchFrenchFrenchFrench

XiboXiboXiboXiboXiboXibo

MarawariMarawariMarawariMarawariMarawariMarawari

MalayalamMalayalamMalayalamMalayalamMalayalamMalayalam

RussianRussianRussianRussianRussianRussian

GujaratGujaratGujaratGujaratGujaratGujarat

SardinianSardinianSardinianSardinianSardinianSardinian

YakutYakutYakutYakutYakutYakut

BasqueBasqueBasqueBasqueBasqueBasque

CambodianCambodianCambodianCambodianCambodianCambodian

AustralianAustralianAustralianAustralianAustralianAustralian

DogonDogonDogonDogonDogonDogon

KalashKalashKalashKalashKalashKalash

TuTuTuTuTuTu

MongolaMongolaMongolaMongolaMongolaMongola

NaxiNaxiNaxiNaxiNaxiNaxi

TujiaTujiaTujiaTujiaTujiaTujia

HanHanHanHanHanHan

JapaneseJapaneseJapaneseJapaneseJapaneseJapanese

DaiDaiDaiDaiDaiDai

HezhenHezhenHezhenHezhenHezhenHezhen

YiYiYiYiYiYi

SheSheSheSheSheShe

OroqenOroqenOroqenOroqenOroqenOroqen

LahuLahuLahuLahuLahuLahu

MiaoMiaoMiaoMiaoMiaoMiao

MayaMayaMayaMayaMayaMaya

MelanesianMelanesianMelanesianMelanesianMelanesianMelanesian

PapuanPapuanPapuanPapuanPapuanPapuan

ColumbianColumbianColumbianColumbianColumbianColumbian

PimaPimaPimaPimaPimaPima

KaritianaKaritianaKaritianaKaritianaKaritianaKaritiana

SuruiSuruiSuruiSuruiSuruiSurui

African

West, Central Eurasian

Indian

East Eurasian

Oceanian

Native American

Populations

Theta (V)

7.3229733794

7.3049552561

7.2858503558

7.2677999836

7.2566260225

7.2472133481

7.2246729245

7.2127822563

7.1999932253

7.1708918992

7.1540630261

7.1303295089

7.1278294907

7.1223120698

7.1088236368

7.107039824

7.0941356971

7.0753978638

7.0744689718

7.0540760814

7.0528720899

7.0511230207

7.0481651433

7.0404971522

7.0271009489

7.023064882

7.0230194177

7.0216842532

7.0132387562

7.0070507459

6.9923083764

6.9887745822

6.9869704415

6.978598901

6.9750255376

6.9739261764

6.9719279073

6.97112325

6.9578462181

6.9538528388

6.9530716113

6.9520686104

6.9384386432

6.9354965763

6.9336680564

6.9321312952

6.9270314448

6.9269718257

6.9215152083

6.9056284255

6.9003186565

6.8920656958

6.89123724

6.8906042715

6.8899715367

6.8830595167

6.8791792717

6.8768751534

6.8765223217

6.870591166

6.85346976

6.8531321082

6.8510771646

6.8459462857

6.8425567459

6.8421516027

6.8401883723

6.8361770647

6.8343602229

6.8321422849

6.8289994826

6.815972656

6.8108909031

6.8034792384

6.8022247624

6.8020048887

6.798987193

6.7835458159

6.781605013

6.7756360946

6.7653717853

6.7642714711

6.7630036777

6.7577278311

6.7482467861

6.7480840876

6.7453626716

6.7401382464

6.7336381372

6.7258534325

6.7246895313

6.7184183116

6.7104108654

6.7020826921

6.6985623962

6.6895289038

6.6771349661

6.6701860628

6.66877186

6.6649709818

6.6596289093

6.6570114533

6.6448859938

6.6383984688

6.6380300376

6.6269139658

6.6122868661

6.6070600358

6.6058190181

6.5959760127

6.5817371621

6.5719845085

6.558798363

6.5309984512

6.5227067678

6.5198412107

6.5151553626

6.4905117537

6.4885686146

6.4840666483

6.4751134162

6.4464037085

6.3132479966

6.1848335629

6.0767321729

6.0061304509

5.9848432752

5.9828970579

5.9810205003

5.9646373592

5.9575653865

5.9466299694

5.942984674

5.9408360514

5.927866888

5.924720491

5.9171864275

5.9115143957

5.8966732545

5.8765910253

5.8747394724

5.8736853516

5.8684838173

5.8346775983

5.8140632241

5.8133836193

5.8043319395

5.7954300423

5.7847980517

5.7708237293

5.7420939141

5.7206133402

5.7156676346

5.7123548974

5.7085427986

5.6947912258

5.6770953985

5.6525293513

5.6387274284

5.6355162274

5.6062731083

5.5760319605

5.5538282283

5.5471610609

5.4953512527

5.4751504635

5.4313916928

5.4105689045

5.386532452

5.374440497

5.3467757802

5.3197908505

5.2727982203

5.2724150328

5.2394700305

5.209570445

5.1828196319

5.1576421856

4.9600237226

4.6431244128

4.632777148

4.2854703522

3.9102788356

3.3453922844

Sheet1

PopulationAfricanWest, Central EurasianIndianEast EurasianOceanianNative American

Ntumu7.3229733794

Baka7.3049552561

Burunge7.2858503558

Bakola7.2677999836

San7.2566260225

Sukuma7.2472133481

Tikar North7.2246729245

Yakoma7.2127822563

Gogo7.1999932253

Mvae7.1708918992

Bantu South7.1540630261

Cape Town7.1303295089

Sandawe7.1278294907

Banen7.1223120698

Iyassa7.1088236368

Mabea7.107039824

Venda7.0941356971

Bulu7.0753978638

Bedzan7.0744689718

Luhya7.0540760814

Mbugwe7.0528720899

Chicago7.0511230207

Biaka7.0481651433

Lemande7.0404971522

Turu7.0271009489

Batie7.023064882

Igala7.0230194177

Baltimore7.0216842532

Zime7.0132387562

Kanembou7.0070507459

Igbo6.9923083764

Yambassa6.9887745822

Bafia6.9869704415

Rangi6.978598901

Dioula6.9750255376

Datoga6.9739261764

Kongo6.9719279073

Barega6.97112325

Luo6.9578462181

Samburu6.9538528388

Gbaya6.9530716113

Burji6.9520686104

North Carolina6.9384386432

Tikar South6.9354965763

Bamoun6.9336680564

Baluba6.9321312952

Hadza6.9270314448

Tupuri6.9269718257

Pare6.9215152083

Kanuri6.9056284255

Batanga6.9003186565

Fang6.8920656958

Xhosa6.89123724

Laka6.8906042715

Yoruba CEPH6.8899715367

Ewondo6.8830595167

Kaba6.8791792717

Wafiome6.8768751534

Bantu Kenya6.8765223217

Maasai Tanzania6.870591166

Samba'a6.85346976

Fulani Nigeria6.8531321082

Mbugu6.8510771646

Wimbum6.8459462857

Gwari6.8425567459

Ngumba6.8421516027

Ouldeme6.8401883723

Brong6.8361770647

Bulala6.8343602229

Kikuyu6.8321422849

Pittsburgh6.8289994826

Turkana6.815972656

Nandi6.8108909031

Mada6.8034792384

Tugen6.8022247624

Kotoko6.8020048887

Ashanti6.798987193

Yoruba6.7835458159

Hausa Nigeria6.781605013

Dorobo6.7756360946

Yaaku6.7653717853

Ngambaye6.7642714711

Sabaot6.7630036777

Maasai Mumonyot6.7577278311

Gabra6.7482467861

Marakwet6.7480840876

Maasai Il'ngwesi6.7453626716

Bassange6.7401382464

Giziga6.7336381372

Hausa Cameroon6.7258534325

Mandara6.7246895313

Ilchamus6.7184183116

Mandenka6.7104108654

Sara various6.7020826921

!Xun/Kxoe6.6985623962

Iraqw6.6895289038

Konso6.6771349661

Massa6.6701860628

El Molo6.66877186

Fulani Adamawa6.6649709818

Mbum6.6596289093

Shuwa6.6570114533

Mbuti6.6448859938

Borana6.6383984688

Sengwer6.6380300376

Rendille6.6269139658

Podokwo6.6122868661

Wata6.6070600358

Koma6.6058190181

Akie6.5959760127

Fulani Mbororo6.5817371621

Shilook6.5719845085

Ogiek6.558798363

Nuer6.5309984512

Zulgo6.5227067678

Beta Israel6.5198412107

Beja Hadandawa6.5151553626

Beja Banuamir6.4905117537

Nyiaman6.4885686146

Tutsi/Hutu6.4840666483

Pokot6.4751134162

Dinka6.4464037085

Mozabite6.3132479966

Bedouin6.1848335629

Balochi6.0767321729

Palestinian6.0061304509

Hindi5.9848432752

Pathan5.9828970579

Kannada5.9810205003

Punjabi5.9646373592

Temani5.9575653865

Makrani5.9466299694

Uygur5.942984674

Telugu5.9408360514

Sindhi5.927866888

Tamil5.924720491

Hazara5.9171864275

Kashmiri5.9115143957

Brahui5.8966732545

Adygei5.8765910253

Bengali5.8747394724

Druze5.8736853516

Assamese5.8684838173

Parsi5.8346775983

Oriya5.8140632241

Konkani5.8133836193

Marathi5.8043319395

Italian5.7954300423

Daur5.7847980517

Burusho5.7708237293

Orcadian5.7420939141

French5.7206133402

Xibo5.7156676346

Marawari5.7123548974

Malayalam5.7085427986

Russian5.6947912258

Gujarat5.6770953985

Sardinian5.6525293513

Yakut5.6387274284

Basque5.6355162274

Cambodian5.6062731083

Australian5.5760319605

Dogon5.5538282283

Kalash5.5471610609

Tu5.4953512527

Mongola5.4751504635

Naxi5.4313916928

Tujia5.4105689045

Han5.386532452

Japanese5.374440497

Dai5.3467757802

Hezhen5.3197908505

Yi5.2727982203

She5.2724150328

Oroqen5.2394700305

Lahu5.209570445

Miao5.1828196319

Maya5.1576421856

Melanesian4.9600237226

Papuan4.6431244128

Columbian4.632777148

Pima4.2854703522

Karitiana3.9102788356

Surui3.3453922844

Sheet1

African

West, Central Eurasian

Indian

East Eurasian

Oceanian

Native American

Populations

Theta (V)

Sheet2

Sheet3

K K Global Patterns of Substructure and Ancestry

Genetic Variation and Structure in Africa

Tishkoff, et alScience 2009

African American Ancestry

Source: David Eltis and David Richardson, Atlas of the Transatlantic Slave Trade

Volume and Direction of the Transatlantic Slave Trade from Africa to the Americas

Genetic variation correlates with geography

Bryc et al. PNAS, 2010

African Ancestry =__b__(a + b)

a and b are the chord distances from the European and African centroids

PCA-based admixture estimation

African ancestry

Shared African and European ancestry

European ancestry

Representative African American Recent admixture

High degree of European ancestryRecent European ancestry

Local Genomic Ancestry in African Americans

• Under selection– Vitamin D– Folate– Melanoma

Variation in Skin Pigmentation

Variation in Skin Pigmentation

San

N = 2200 for melanin

Anthropometric trait variation in Africa

Cardiovascular trait variation in Africa

Lipid biomarker variation in Africa

…AAGATAAT/GGTAGCTCC/GTG….

Identification of novel genetic variants associated with lactose tolerance and lactase gene expression in African pastoralist populations

136000000 136500000135500000

MCM6 36.2 kb

Intron 13

C -13907G -13915C -14010

Tishkoff et al, Nature Genetics, 2007

Ranciaro et al., AJHG, 2014

…CG/CTAAGTTACCA…….. C/T-14010 bp -13915 bp -13910 bp-13907 bp

Historically, biological classification of races has been associated with:

•hierarchical ranking of races

•biological determinism

•Eugenics

•justification for genocide (e.g. the Nazi-led holocaust)

•colonialism, slavery, and other social inequities

Problems with Racial Classifications

• Problem with using race as an identifier is the lack of a clear definition of race

• Historically, race has been classified based on both socio-cultural and biological characteristics

Race does not correlate with patterns of population structure inferred from genetic data

Problems with Racial Classifications

•Ethnic groups differ for risk to disease.

•People whose ancestors lived in distinct environments have been subject to different selection forces and have differing demographic histories.

•Some diseases may be influenced by geographically restricted susceptibility alleles.

•Information about individual ancestry could provide important medical information for diagnosis and treatment.

Implications for Medicine

•However….one must be wary of racial profiling and ignorance of the continuous nature of genetic variation and high levels of admixture in modern populations, which can result in mis-classification and mis-diagnosis.

•Cannot make generalizations based on continental region of origin because there can be tremendous variation amongst populations within a region (i.e. Africa).

•Populations/ethnic groups differ in prevalence of disease due to both genetic and environmental factors and a key issue will be to disentangle these factors and to study their interaction.

Implications for Medicine

•The goal should be personalized medicine—treating for the specific etiology in the individual patient.

•Information about “Race” could be important if trying to distinguish socio-cultural and environmental risk factors

•However, for many studies race will be an insufficient identifier and detailed information about ethnicity, geographic origin, language, religion and/or culture will be important.

•“Ancestry” should supplant “race” in discussions where the group identity of the patient is at issue.

Implications for Medicine

Tishkoff lab, University of PennsylvaniaMatt Hansen/ Nick Crawford/Joseph Lachance / Joe Jarvis / Laura Scheinfeldt / Sameer Soi / Clara Elbers / Bart Ferwerda / Bill Beggs / Simon Thompson / Alessia Ranciaro / Jibril Hirbo

Illumina 5 M GWAS studyStephen Chanock/Meredith Yeager (NCI)

African CollaboratorsSabah Omar / Muntaser Ibrahim / Thomas Nyambo / Godfrey Lema / Charles Wambebe/ Ogabara Doumbo / Mohamadu Thera/Gurja Belay/Dawit WoldeSabahOmar / Muntaser Ibrahim / Thomas Nyambo / Godfrey Lema / Charles Wambebe/ Ogabara Doumbo / Mohamadu Thera/Gurja Belay/Dawit Wolde

NIGMS NHGRI

Thanks

National Cancer Institute

National Institute of Aging

Population Studies Center at Penn

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