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Complex History of Admixture between Modern Humans and Neandertals
Open accessAbstract
Recent analyses have found that a substantial amount of the Neandertal genome persists in the genomes of contemporary non-African individuals. East Asians have, on average, higher levels of Neandertal ancestry than do Europeans, which might be due to differences in the efficiency of purifying selection, an additional pulse of introgression into East Asians, or other unexplored scenarios. To better define the scope of plausible models of archaic admixture between Neandertals and anatomically modern humans, we analyzed patterns of introgressed sequence in whole-genome data of 379 Europeans and 286 East Asians. We found that inferences of demographic history restricted to neutrally evolving genomic regions allowed a simple one-pulse model to be robustly rejected, suggesting that differences in selection cannot explain the differences in Neandertal ancestry. We show that two additional demographic models, involving either a second pulse of Neandertal gene flow into the ancestors of East Asians or a dilution of Neandertal lineages in Europeans by admixture with an unknown ancestral population, are consistent with the data. Thus, the history of admixture between modern humans and Neandertals is most likely more complex than previously thought.
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In the citing paper
“Thus, FAs could have acted as a population that diluted NE ancestry in western Eurasia (text S4), as previously suggested (16, 17).”
From Past human expansions shaped the spatial pattern of Neanderthal ancestry (2023) · cites this paper as [17]
In the citing paper
“In Europeans, archaic ancestry is consistent with contributions by a single population of Neanderthals 3 through one or more pulses [4][5][6][7] .”
From The nature of Neanderthal introgression revealed by 27,566 Icelandic genomes (2020) · cites this paper as [6]
In the citing paper
“It has been proposed that the early FAs are partly genetically derived from the previously identified “basal Eurasian” lineage (16, 17, 51).”
From Past human expansions shaped the spatial pattern of Neanderthal ancestry (2023) · cites this paper as [17]
In the citing paper
“This lineage is thought to have diverged from other Eurasian HG populations before the latter mixed with the NE, and therefore had a lower NE ancestry (16, 17, 51).”
From Past human expansions shaped the spatial pattern of Neanderthal ancestry (2023) · cites this paper as [17]
Citations
Cites
A draft sequence of the Neandertal genome
Green, R. E., Krause, J., Briggs, A. W., Maricic, T., Stenzel, U., Kircher, M., Patterson, N., Li, H., Zhai, W., Fritz, M. H.-Y., et al. (2010) · Science · doi:10.1126/science.1188021 · Cited by 4,318
Generating samples under a Wright–Fisher neutral model of genetic variation
Hudson, R. R. (2002) · Bioinformatics · doi:10.1093/bioinformatics/18.2.337 · Cited by 2,397
The complete genome sequence of a Neanderthal from the Altai Mountains
Prüfer, K., Racimo, F., Patterson, N., Jay, F., Sankararaman, S., Sawyer, S., Heinze, A., Renaud, G., Sudmant, P. H., de Filippo, C., et al. (2014) · Nature · doi:10.1038/nature12886 · Cited by 2,254
A high-coverage genome sequence from an archaic denisovan individual
Meyer, M., Kircher, M., Gansauge, M.-T., Li, H., Racimo, F., Mallick, S., Schraiber, J. G., Jay, F., Prüfer, K., de Filippo, C., et al. (2012) · Science · doi:10.1126/science.1224344 · Cited by 1,967
Evolution and functional impact of rare coding variation from deep sequencing of human exomes
Tennessen, J. A., Bigham, A. W., O’Connor, T. D., Fu, W., Kenny, E. E., Gravel, S., McGee, S., Do, R., Liu, X., Jun, G., et al. (2012) · Science · doi:10.1126/science.1219240 · Cited by 1,613
Self-citationAncient human genomes suggest three ancestral populations for present-day Europeans
Lazaridis, I., Patterson, N., Mittnik, A., Renaud, G., Mallick, S., Kirsanow, K., Sudmant, P. H., Schraiber, J. G., Castellano, S., Lipson, M., et al. (2014) · Nature · doi:10.1038/nature13673 · Cited by 1,357
Approximate Bayesian Computation (ABC) in practice
Csilléry, K., Blum, M. G. B., Gaggiotti, O. E., & François, O. (2010) · Trends in Ecology & Evolution · doi:10.1016/j.tree.2010.04.001 · Cited by 1,302
The genomic landscape of Neanderthal ancestry in present-day humans
Sankararaman, S., Mallick, S., Dannemann, M., Prüfer, K., Kelso, J., Pääbo, S., Patterson, N., & Reich, D. (2014) · Nature · doi:10.1038/nature12961 · Cited by 1,107
abc: an R package for approximate Bayesian computation (ABC)
Csilléry, K., François, O., & Blum, M. G. B. (2012) · Methods in Ecology and Evolution · doi:10.1111/j.2041-210x.2011.00179.x · Cited by 709
Siva, N. (2008) · Nature Biotechnology · doi:10.1038/nbt0308-256b · Cited by 629
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Metadata
- Authors
- Vernot, B., & Akey, J. M.
- Affiliations
- University of Washington (US)
- Corresponding author
- Vernot, Benjamin
- Akey, Joshua M.
- Venue
- The American Journal of Human Genetics
- Publisher
- Elsevier BV
- Published
- 2015-03
- Volume
- 96
- Issue
- 3
- Pages
- 448-453
- ISSN
- 0002-9297 (ISSN-L)
- PMC
- PMC4375686
- OpenAlex
- W2056921607
- Cited by
- 149
- References
- 19
- Citations by year
- 2026: 5 · 2025: 4 · 2024: 11 · 2023: 10 · 2022: 8 · 2021: 30 · 2020: 9 · 2019: 6 · 2018: 19 · 2017: 15 · 2016: 26 · 2015: 6
- FWCI
- 92.10
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- Locations
- The American Journal of Human Genetics (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Institutes of Health
- National Institute of General Medical Sciences (R01GM110068)
- Topics
- Social Sciences › Anthropology › Pleistocene-Era Hominins and Archaeology
- 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
- Keywords
- purifying selection, demographic history, gene flow
- MeSH terms
- Animals, Genetics, Population, Humans, Models, Genetic, Genome, White People, Asian People, Gene Flow, Neanderthals
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