Cited by 1,573Forensic and Genetic ResearchGenetic diversity and population structureRace, Genetics, and Society

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The Genetic Structure and History of Africans and African Americans

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Tishkoff, Sarah A., Reed, Floyd A., Friedlaender, Françoise R., Ehret, Christopher, Ranciaro, Alessia, Froment, Alain, Hirbo, Jibril B., Awomoyi, Agnes A., Bodo, Jean-Marie, Doumbo, Ogobara, Ibrahim, Muntaser, Juma, Abdalla T., Kotze, Maritha J., Lema, Godfrey, Moore, Jason H., Mortensen, Holly, Nyambo, Thomas B., Omar, Sabah A., Powell, Kweli, Pretorius, Gideon S., Smith, Michael W., Thera, Mahamadou A., Wambebe, Charles, Weber, James L., Williams, Scott M.·2009·Science

DOI 10.1126/science.1172257PMC PMC2947357

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Abstract

African Origins: The modern human originated in Africa and subsequently spread across the globe. However, the genetic relationships among the diverse populations on the African continent have been unclear. Tishkoff et al. (p. 1035; see the cover, published online 30 April) provide a detailed genetic analysis of most major groups of African populations. The findings suggest that Africans represent 14 ancestral populations. Populations tend to be of mixed ancestry which documents historical migrations. The data mainly support but sometimes challenge proposed relationships between groups of self-identified ethnicity previously hypothesized on the basis of linguistic studies. The authors also examined populations of African Americans and individuals of mixed ancestry from Cape Town, documenting the variation and origins of admixture within these groups.

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Browne, J. L., Klipstein‐Grobusch, K., Koster, M. P. H., Ramamoorthy, D., Antwi, E., Belmouden, I., Franx, A., Grobbee, D. E., & Schielen, P. C. J. I. (2016) · PLoS ONE · doi:10.1371/journal.pone.0159592 · Cited by 18
“Ultimately, not correcting for higher levels in non-Caucasian ethnicities, especially given the known heterogeneity of people from African decent [17,18], could result in a structural underestimation of true risk for fetal aneuploidies or placental disorders [5,6].”
The SIGLEC14 null allele is associated with Mycobacterium tuberculosis- and BCG-induced clinical and immunologic outcomes
Graustein, A. D., Horne, D. J., Fong, J. J., Schwarz, F., Mefford, H. C., Peterson, G. J., Wells, R. D., Musvosvi, M., Shey, M. S., Hanekom, W. A., et al. (2017) · Tuberculosis · doi:10.1016/j.tube.2017.02.005 · Cited by 20
“The CMA ethnicity represents the genetic admixture of Khoesan, Black African, European and both east and south Asian populations that has existed for over 350 years [47,48].”
Human loci involved in drug biotransformation: worldwide genetic variation, population structure, and pharmacogenetic implications
Delser, P. M., & Fuselli, S. (2013) · Human Genetics · doi:10.1007/s00439-013-1268-5 · Cited by 19
“Exceptions to this pattern are Africa, America, and Oceania where populations are highly differentiated for reasons mainly due to the sampling scheme (few and scattered populations) and to the demographic history of these continents (Mulligan et al. 2004; Wang et al. 2007b; Tishkoff et al. 2009).”
Background
Bantu Expansion
de Luna, K. M. (2014) · doi:10.1093/obo/9780199846733-0165 · Cited by 18
Testing Spearman's hypothesis with advanced placement examination data
Warne, R. T. (2016) · Intelligence · doi:10.1016/j.intell.2016.05.002 · Cited by 19
N!aqriaxe (ǂ’Amkoe) spatial terms from a genealogical and areal perspective
Gerlach, L., & Berthold, F. (2014) · Amsterdam studies in the theory and history of linguistic science. Series 4, Current issues in linguistic theory · doi:10.1075/cilt.330.08ger · Cited by 17
BMI, physical activity, and breast cancer subtype in white, black, and Sea Island breast cancer survivors
Ford, M. E., Bauza, C. E., Findlay, V. J., Turner, D. P., Abraham-Hilaire, L., Moore, L. A., Magwood, G., Alberg, A. J., Gaymon, K., Knight, K. D., et al. (2020) · Advances in cancer research · doi:10.1016/bs.acr.2020.01.005 · Cited by 17
“Also novel is the inclusion not only of EA and AA but also Sea Islanders/Gullah (SI) women, a distinct subset of AAs that is culturally and genetically distinct (Tishkoff et al., 2009; Zakharia et al., 2009).”
Background
Pathogen-Induced Epigenetic Modifications in Cancers: Implications for Prevention, Detection and Treatment of Cancers in Africa
Zune, A. L. D., Olwal, C. O., Tapela, K., Owoicho, O., Nganyewo, N. N., Lyko, F., & Paemka, L. (2021) · Cancers · doi:10.3390/cancers13236051 · Cited by 16
“These variable factors among Africans, especially genetic, environmental and dietary are critical for understanding the genetic risk factors for disease, gene-environment interactions and epigenetic basis of human diseases [47].”
Background
The genetic and evolutionary basis of gene expression variation in East Africans
Kelly, D. E., Ramdas, S., Ma, R., Rawlings-Goss, R., Grant, G. R., Ranciaro, A., Hirbo, J. B., Beggs, W. R., Yeager, M., Chanock, S. J., et al. (2023) · Genome biology · doi:10.1186/s13059-023-02874-4 · Cited by 18
“The majority of genomic studies of sub-Saharan African individuals have focused on populations of primarily West African descent, which fails to capture much of the genetic and phenotypic diversity within sub-Saharan Africa [15].”
BackgroundSelf-citation
Comprehensive evaluation of imputation performance in African Americans
Chanda, P., Yuhki, N., Li, M., Bader, J. S., Hartz, A., Boerwinkle, E., Kao, W. L., & Arking, D. E. (2012) · Journal of Human Genetics · doi:10.1038/jhg.2012.43 · Cited by 17
The African−387 C>T TGFB1 variant is functional and associates with the ophthalmoplegic complication in juvenile myasthenia gravis
Nel, M., Buys, J.-M., Rautenbach, R. M., Mowla, S., Prince, S., & Heckmann, J. M. (2015) · Journal of Human Genetics · doi:10.1038/jhg.2015.146 · Cited by 17
Global demographic history of human populations inferred from whole mitochondrial genomes
Miller, E. F., Manica, A., & Amos, W. (2018) · Royal Society Open Science · doi:10.1098/rsos.180543 · Cited by 16
A phenome-wide association study (PheWAS) in the Population Architecture using Genomics and Epidemiology (PAGE) study reveals potential pleiotropy in African Americans
Pendergrass, S. A., Buyske, S. G., Jeff, J. M., Frase, A., DUDEK, S. M., Bradford, Y., Ambite, J. L., Avery, C. L., Bůžková, P., Deelman, E., et al. (2019) · PLoS ONE · doi:10.1371/journal.pone.0226771 · Cited by 16
“Genetic ancestry and admixture are widely recognized as useful markers of human migration [56, 58] and disease associations [86], including potential genetic interactions [87].”
Background
Signatures of Convergent Evolution and Natural Selection at the Alcohol Dehydrogenase Gene Region are Correlated with Agriculture in Ethnically Diverse Africans
McQuillan, M. A., Ranciaro, A., Hansen, M. E. B., Fan, S., Beggs, W. R., Belay, G., Meskel, D. W., & Tishkoff, S. A. (2022) · Molecular Biology and Evolution · doi:10.1093/molbev/msac183 · Cited by 18
“Modern humans arose in Africa ∼ 300,000 years ago, and African populations harbor the highest levels of genetic and phenotypic variation today (Rosenberg et al. 2002; Tishkoff et al. 2009; Hublin et al. 2017).”
Self-citation
A genome-wide association study of plasma concentrations of warfarin enantiomers and metabolites in sub-Saharan black-African patients
Asiimwe, I. G., Blockman, M., Cohen, K., Cupido, C., Hutchinson, C., Jacobson, B. F., Lamorde, M., Morgan, J., Mouton, J. P., Nakagaayi, D., et al. (2022) · Frontiers in Pharmacology · doi:10.3389/fphar.2022.967082 · Cited by 17
“Another limitation is that we only included patients from Uganda and SouthAfrica whichmeans our resultsmay not be generalizable to the rest of sub-Saharan Africa given the very high withinpopulation genetic diversity of African populations (Tishkoff et al., 2009).”
BackgroundInfluential
Lacustrine geoarchaeology in the central Kalahari: Implications for Middle Stone Age behaviour and adaptation in dryland conditions
Thomas, D. S. G., Burrough, S. L., Coulson, S. D., Mothulatshipi, S., Nash, D. J., & Staurset, S. (2022) · Quaternary Science Reviews · doi:10.1016/j.quascirev.2022.107826 · Cited by 17
Diversity and Frequencies of HLA Class I and Class II Genes of an East African Population
Peterson, T. A., Bielawny, T., Lacap, P. A., Hardie, R., Daniuk, C. A., Mendoza, L., Thavaneswaran, S., Kariri, T., Kimani, J., Wachihi, C., et al. (2014) · Open Journal of Genetics · doi:10.4236/ojgen.2014.42013 · Cited by 17
“A previous analysis of the genetic makeup of African populations and their genetic diversity has shown extensive variations in multiple genetic markers such as microsatellites, insertions/deletions and single nucleotide polymorphisms [4].”
Background
The Upper Palaeolithic Lithic Industry of Nazlet Khater 4 (Egypt): Implications for the Stone Age/Palaeolithic of Northeastern Africa
Leplongeon, A., & Pleurdeau, D. (2011) · African Archaeological Review · doi:10.1007/s10437-011-9100-x · Cited by 17
Variable Association of Reactive Intermediate Genes with Systemic Lupus Erythematosus in Populations with Different African Ancestry
Ramos, P. S., Oates, J. C., Kamen, D. L., Williams, A. H., Gaffney, P. M., Kelly, J. A., Kaufman, K. M., Kimberly, R. P., Niewold, T. B., Jacob, C. O., et al. (2013) · The Journal of Rheumatology · doi:10.3899/jrheum.120989 · Cited by 17
Research and the health of indigenous populations in low- and middle-income countries
Mohindra, K. S. (2015) · Health Promotion International · doi:10.1093/heapro/dav106 · Cited by 19
Type I Interferon Receptor Variants in Gene Regulatory Regions are Associated with Susceptibility to Cerebral Malaria in Malawi
Feintuch, C. M., Tare, A., Cusumano, L. R., Benayoun, J., Ryu, S., Sixpence, A., Seydel, K. B., Laufer, M. K., Taylor, T. E., Suh, Y., et al. (2018) · American Journal of Tropical Medicine and Hygiene · doi:10.4269/ajtmh.17-0887 · Cited by 17
“generalizability of HapMap tagSNPs usedwithin a continent of nonisolated populations.(46,47) Alternative approaches include”
Background
Population Genetics
Keats, B. J. B., & Sherman, S. L. (2013) · Elsevier eBooks · doi:10.1016/b978-0-12-383834-6.00015-x · Cited by 17
Pharmacogenomics in Africa: A Potential Catalyst for Precision Medicine in Genetically Diverse Populations
Twesigomwe, D., Mazhindu, T., Nagy, M., Agesa, G., Scholefield, J., & Masimirembwa, C. M. (2025) · Annual Review of Genomics and Human Genetics · doi:10.1146/annurev-genom-121323-104008 · Cited by 19
High resolution class I HLA-A, -B, and -C diversity in Eastern and Southern African populations
Banjoko, A. W., Nguni, T. L., Naidoo, N., Ramsuran, V., Hyrien, O., & Ndhlovu, Z. M. (2025) · Scientific Reports · doi:10.1038/s41598-025-06704-4 · Cited by 23
“Several factors such as less polymorphic nature of HLA-C compared to HLA-A and HLA-B, which result in a slower rate of allele diversification and as a result, may reflect shared deep ancestry 53,54,81 .”
Influential
Cascade Testing for Fragile X Syndrome in a Rural Setting in Cameroon (Sub-Saharan Africa)
Kamga, K. K., Nguefack, S., Minka, K., Wonkam‐Tingang, E., Esterhuizen, A. I., Munung, N. S., de Vries, J., & Wonkam, A. (2020) · Genes · doi:10.3390/genes11020136 · Cited by 17
“Also known as “Africa in miniature”, Cameroon has a diverse cultural and linguistic heritage which mimics the heterogeneity found in Africa [24].”
Background
Hearing loss and brain disorders: A review of multiple pathologies
Oluwole, O. G., James, K., Yalcouyé, A., & Wonkam, A. (2021) · Open Medicine · doi:10.1515/med-2021-0402 · Cited by 16
“Unlike the European and American populations that have been majorly investigated, research needs to focus more on the under-represented populations because they are known for genetic diversities that could play some important ramifications for diseases and treatment [70].”
Background
Polygenic risk scores improve CAD risk prediction in individuals at borderline and intermediate clinical risk
Ratman, D., Tshiaba, P., Levin, M. G., Sun, J., Tunstall, T. S., Maier, R., Shah, P., Rabinowitz, M., Rader, D. J., Kumar, A., et al. (2025) · npj Cardiovascular Health · doi:10.1038/s44325-025-00049-7 · Cited by 16
“This likely re ﬂ ects the reduced availability of GWAS summary statistics and individual-level data for PRS development, which is further compounded by the high genetic diversity of the African population 25,26 .”
Epigenetics and the Burden of Noncommunicable Disease: A Paucity of Research in Africa
Hobbs, A., & Ramsay, M. (2015) · Epigenomics · doi:10.2217/epi.15.17 · Cited by 17
Mitochondrial DNA diversity in two ethnic groups in Southeastern Kenya: Perspectives from the northeastern periphery of the bantu expansion
Batai, K., Babrowski, K. B., Arroyo, J. P., Kusimba, C. M., & Williams, S. R. (2013) · American Journal of Physical Anthropology · doi:10.1002/ajpa.22227 · Cited by 16
“…studies given the demographic, ethnic, and linguistic diversity, but the genetic studies undertaken to date (Watson et al., 1996; Semino et al., 2002; Cruciani et al., 2004; Kivisild et al., 2004; Hassan et al., 2008; Tishkoff et al., 2009), indicate that the region is genetically diverse as well.”
BackgroundSelf-citation
Increased genetic diversity of ADME genes in African Americans compared with their putative ancestral source populations and implications for Pharmacogenomics
Li, J., Lao, X., Zhang, C., Tian, L., Lu, D., & Xu, S. (2014) · BMC Genetics · doi:10.1186/1471-2156-15-52 · Cited by 18
“In this study, we investigated the genetic diversity of drug metabolism-related (ADME) genes in AfricanAmericans (ASW) compared with Europeans (CEU) and Africans (YRI), which are the representative ancestral source populations of African-Americans according to a previous study [17].”
BackgroundResultInfluential
BiDil in the Clinic: An Interdisciplinary Investigation of Physicians’ Prescription Patterns of a Race-Based Therapy
Maglo, K. N., Rubinstein, J., Huang, B., & Ittenbach, R. F. (2014) · AJOB Empirical Bioethics · doi:10.1080/23294515.2014.907371 · Cited by 16
“No subpopulation within a given continent constitutes a biologically representative subpopulation for the whole continental population or racial group (Tishkoff et al. 2009; Tishkoff and Verrelli 2003).”
Background
Gun Violence, African Ancestry, and Asthma
Rosas‐Salazar, C., Han, Y., Brehm, J. M., Forno, E., Acosta‐Pérez, E., Cloutier, M. M., Alvarez, M., Colón‐Semidey, A., CANINO, G. J., & Celedon, J. C. (2016) · CHEST Journal · doi:10.1016/j.chest.2016.02.639 · Cited by 16
Birth Prevalence of Anorectal Malformations for the Western Cape Province, South Africa, 2005 to 2012
Numanoğlu, A., & Theron, A. (2017) · European Journal of Pediatric Surgery · doi:10.1055/s-0036-1597945 · Cited by 16
Exploring Overlaps Between the Genomic and Environmental Determinants of LVH and Stroke: A Multicenter Study in West Africa
Adeoye, A. M., Ovbiagele, B. I., Kolo, P. M., Appiah, L. T., Aje, A., Adebayo, O., Sarfo, F. S., Akinyemi, J. O., Adekunle, G., Agyekum, F., et al. (2017) · Global Heart · doi:10.1016/j.gheart.2017.01.001 · Cited by 15
Homo sapiens origins and evolution in the Kalahari Basin, southern Africa
Wilkins, J. (2021) · Evolutionary Anthropology Issues News and Reviews · doi:10.1002/evan.21914 · Cited by 19
“Populations have moved significantly, most recently influenced by colonial disruptions, and before that the spread of herders and farmers across the continent.(2,7) The process of reconstructing past population dynamics based on the genetic relationships of modern populations is complex and relies on many parameters.”
Background
Conservation genetics of the black rhinoceros, Diceros bicornis bicornis, in Namibia
de Groot, P. J. V. C., Putnam, A. S., Erb, P., Scott, C., Melnick, D. J., O’Ryan, C., & Boag, P. T. (2011) · Conservation Genetics · doi:10.1007/s10592-011-0185-1 · Cited by 16
Investigating the Role of Gene-Gene Interactions in TB Susceptibility
Daya, M., van der Merwe, L., van Helden, P. D., Möller, M., & Hoal, E. G. (2015) · PLoS ONE · doi:10.1371/journal.pone.0123970 · Cited by 18
“In addition, the San has contributed a large amount of genetic material to the SAC [61, 104–108], and due to the lack of large high density reference panels for the San, this may contribute to additional inaccuracies in imputation of the SAC data set.”
Background
Maximizing genetic differentiation in core collections by PCA-based clustering of molecular marker data
van Heerwaarden, J., Odong, T. L., & van Eeuwijk, F. A. (2012) · Theoretical and Applied Genetics · doi:10.1007/s00122-012-2016-2 · Cited by 16
The contributions of admixture and genetic drift to diversity among post‐contact populations in the Americas
Koehl, A. J., & Long, J. C. (2017) · American Journal of Physical Anthropology · doi:10.1002/ajpa.23347 · Cited by 16
Ecological variation in Later Stone Age southern African biomechanical properties
Cameron, M. E., & Stock, J. T. (2017) · Journal of Archaeological Science Reports · doi:10.1016/j.jasrep.2017.10.033 · Cited by 16
Clinical and genetic analysis of spinocerebellar ataxia type 7 (SCA7) in Zambian families
Atadzhanov, M., Smith, D. C., Mwaba, M. H., Siddiqi, O. K., Bryer, A., & Greenberg, L. S. (2017) · Cerebellum & Ataxias · doi:10.1186/s40673-017-0075-5 · Cited by 16
“Although more than 70 different tribes are currently recognized, the Zambian population can be classified within the Niger-Kordofanian family, the largest of the four ethno-linguistic African macrofamilies [14].”
Background
Genetic history of the African Sahelian populations
Černý, V., Kulichová, I., Poloni, E. S., Nunes, J. M., Pereira, L., Mayor, A., & Sanchez‐Mazas, A. (2017) · HLA · doi:10.1111/tan.13189 · Cited by 15
“Although Fulani speak today a language with clear ties to west Africa (as stated above, Fulfulde is a member of the Niger-Congo west-Atlantic subfamily), they display genetic links with both east African and west African and even with Eurasian populations.(1,69,78,91,108,109) Contrary to uniparental loci the G516T cou…”
Background

Authors: Tishkoff, S. A., Reed, F. A., Friedlaender, F. R., Ehret, C., Ranciaro, A., Froment, A., Hirbo, J. B., Awomoyi, A. A., Bodo, J.-M., Doumbo, O., Ibrahim, M., Juma, A. T., Kotze, M. J., Lema, G., Moore, J. H., Mortensen, H., Nyambo, T. B., Omar, S. A., Powell, K., Pretorius, G. S., Smith, M. W., Thera, M. A., Wambebe, C., Weber, J. L., & Williams, S. M.
Affiliations: University of Maryland, College Park (US)
Niche, Nutrition, Cancer et métabolisme oxydatif (FR)
University of California, Los Angeles (US)
University of Ferrara (IT)
Muséum national d'Histoire naturelle (FR)
Ministry of Scientific Research and Innovation (CM)
University of Bamako (ML)
University of Khartoum (SD)
Stellenbosch University (ZA)
Muhimbili University of Health and Allied Sciences (TZ)
Departments of Genetics and Community and Family Medicine, Dartmouth Medical School, Lebanon, NH 03756, USA.
Kenya Medical Research Institute (KE)
National Cancer Institute (US)
International Biomedical Research in Africa, Abuja, Nigeria.
Marshfield Clinic Research Foundation (US)
Vanderbilt University (US)
Corresponding author: Tishkoff, Sarah A.
Reed, Floyd A.
Friedlaender, Françoise R.
Ranciaro, Alessia
Froment, Alain
Awomoyi, Agnes A.
Mortensen, Holly
Powell, Kweli
ORCID: Tishkoff, Sarah A.
Reed, Floyd A.
Ranciaro, Alessia
Froment, Alain
Hirbo, Jibril B.
Ibrahim, Muntaser
Kotze, Maritha J.
Moore, Jason H.
Mortensen, Holly
Nyambo, Thomas B.
Smith, Michael W.
Thera, Mahamadou A.
Williams, Scott M.
Venue: Science
Publisher: American Association for the Advancement of Science (AAAS)
Published: 2009-05-22
Volume: 324
Issue: 5930
Pages: 1035-1044
ISSN: 0036-8075 (ISSN-L)
1095-9203
DOI: 10.1126/science.1172257
PMC: PMC2947357
OpenAlex: W2129316818
Cited by: 1,573
References: 60
Citations by year: 2026: 65 · 2025: 98 · 2024: 56 · 2023: 69 · 2022: 69 · 2021: 85 · 2020: 79 · 2019: 88 · 2018: 88 · 2017: 84 · 2016: 89 · 2015: 102 · 2014: 119 · 2013: 93 · 2012: 123
FWCI: 75.46
Type: journal article
Language: en
Open access: green
www.ncbi.nlm.nih.gov
Locations: Science (publishedVersion)
PubMed (publishedVersion)
IHI Repository (acceptedVersion)
PubMed Central (acceptedVersion)
Max Planck Institute for Plasma Physics (submittedVersion)
IRIS Research product catalog (Sapienza University of Rome) (submittedVersion)
MPG.PuRe (Max Planck Society) (submittedVersion)
Funders: National Institutes of Health (n01co12400-79-0-22)
National Heart, Lung, and Blood Institute (R01 HL65234, R01HL065234)
National Human Genome Research Institute (F32HG03801, F32HG003801)
National Cancer Institute (N01-CO-012400)
National Institute of General Medical Sciences (R01GM083606, R01GM076637)
Topics: Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Forensic and Genetic Research
Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetic diversity and population structure
Social Sciences › Anthropology › Race, Genetics, and Society
Keywords: African American ancestry, admixture, microsatellite markers, InDel markers, Khoesan, Pygmies, European ancestry, migration history, self-reported ethnicity
MeSH terms: Africa, Bayes Theorem, Black or African American, Emigration and Immigration, Ethnicity, Genotype, Geography, Humans, Language, Phylogeny, Genetic Variation, Cluster Analysis, Microsatellite Repeats, Polymorphism, Single Nucleotide, Principal Component Analysis, Black People, Racial Groups, Gene Flow, INDEL Mutation

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The Genetic Structure and History of Africans and African Americans

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