Genomic insights into the formation of human populations in East Asia

doi:10.1038/s41586-021-03336-2

. 2021 Mar;591(7850):413-419.

doi: 10.1038/s41586-021-03336-2. Epub 2021 Feb 22.

Genomic insights into the formation of human populations in East Asia

Chuan-Chao Wang^#^{1

2

3

4}, Hui-Yuan Yeh^#⁵, Alexander N Popov^#⁶, Hu-Qin Zhang^#⁷, Hirofumi Matsumura⁸, Kendra Sirak^{9

10}, Olivia Cheronet¹¹, Alexey Kovalev¹², Nadin Rohland⁹, Alexander M Kim^{9

13}, Swapan Mallick^{9

10

14

15}, Rebecca Bernardos⁹, Dashtseveg Tumen¹⁶, Jing Zhao⁷, Yi-Chang Liu¹⁷, Jiun-Yu Liu¹⁸, Matthew Mah^{9

14

15}, Ke Wang¹⁹, Zhao Zhang⁹, Nicole Adamski^{9

15}, Nasreen Broomandkhoshbacht^{9

15}, Kimberly Callan^{9

15}, Francesca Candilio¹¹, Kellie Sara Duffett Carlson¹¹, Brendan J Culleton²⁰, Laurie Eccles²¹, Suzanne Freilich¹¹, Denise Keating¹¹, Ann Marie Lawson^{9

15}, Kirsten Mandl¹¹, Megan Michel^{9

15}, Jonas Oppenheimer^{9

15}, Kadir Toykan Özdoğan¹¹, Kristin Stewardson^{9

15}, Shaoqing Wen²², Shi Yan²³, Fatma Zalzala^{9

15}, Richard Chuang¹⁷, Ching-Jung Huang¹⁷, Hana Looh²⁴, Chung-Ching Shiung¹⁷, Yuri G Nikitin²⁵, Andrei V Tabarev²⁶, Alexey A Tishkin²⁷, Song Lin⁷, Zhou-Yong Sun²⁸, Xiao-Ming Wu⁷, Tie-Lin Yang⁷, Xi Hu⁷, Liang Chen²⁹, Hua Du³⁰, Jamsranjav Bayarsaikhan³¹, Enkhbayar Mijiddorj³², Diimaajav Erdenebaatar³², Tumur-Ochir Iderkhangai³², Erdene Myagmar¹⁶, Hideaki Kanzawa-Kiriyama³³, Masato Nishino³⁴, Ken-Ichi Shinoda³³, Olga A Shubina³⁵, Jianxin Guo³⁶, Wangwei Cai³⁷, Qiongying Deng³⁸, Longli Kang³⁹, Dawei Li⁴⁰, Dongna Li⁴¹, Rong Lin⁴¹, Nini³⁹, Rukesh Shrestha⁴², Ling-Xiang Wang⁴², Lanhai Wei³⁶, Guangmao Xie^{43

44}, Hongbing Yao⁴⁵, Manfei Zhang⁴², Guanglin He³⁶, Xiaomin Yang³⁶, Rong Hu³⁶, Martine Robbeets⁴⁶, Stephan Schiffels¹⁹, Douglas J Kennett⁴⁷, Li Jin⁴², Hui Li⁴², Johannes Krause⁴⁸, Ron Pinhasi⁴⁹, David Reich^{50

51

52

53}

Affiliations

¹ Department of Anthropology and Ethnology, Institute of Anthropology, State Key Laboratory of Cellular Stress Biology, School of Life Sciences, Xiamen University, Xiamen, China. wang@xmu.edu.cn.
² Department of Genetics, Harvard Medical School, Boston, MA, USA. wang@xmu.edu.cn.
³ Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany. wang@xmu.edu.cn.
⁴ MOE Key Laboratory of Contemporary Anthropology, Department of Anthropology and Human Genetics, School of Life Sciences, Fudan University, Shanghai, China. wang@xmu.edu.cn.
⁵ School of Humanities, Nanyang Technological University, Nanyang, Singapore.
⁶ Scientific Museum, Far Eastern Federal University, Vladivostok, Russia.
⁷ Key Laboratory of Biomedical Information Engineering of Ministry of Education, School of Life Science and Technology, Xi'an Jiaotong University, Xi'an, China.
⁸ School of Health Science, Sapporo Medical University, Sapporo, Japan.
⁹ Department of Genetics, Harvard Medical School, Boston, MA, USA.
¹⁰ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA.
¹¹ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria.
¹² Institute of Archaeology, Russian Academy of Sciences, Moscow, Russia.
¹³ Department of Anthropology, Harvard University, Cambridge, MA, USA.
¹⁴ Broad Institute of Harvard and MIT, Cambridge, MA, USA.
¹⁵ Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA.
¹⁶ Department of Anthropology and Archaeology, National University of Mongolia, Ulaanbaatar, Mongolia.
¹⁷ Institute of Archaeology, National Cheng Kung University, Tainan, Taiwan.
¹⁸ Department of Anthropology, University of Washington, Seattle, WA, USA.
¹⁹ Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany.
²⁰ Institutes of Energy and the Environment, The Pennsylvania State University, University Park, PA, USA.
²¹ Department of Anthropology, Pennsylvania State University, University Park, PA, USA.
²² Institute of Archaeological Science, Fudan University, Shanghai, China.
²³ School of Ethnology and Sociology, Minzu University of China, Beijing, China.
²⁴ Institute of History and Philology, Institute of History and Philology, Academia Sinica, Taipei, Taiwan.
²⁵ Museum of Archaeology and Ethnology, Institute of History, Archaeology and Ethnology, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok, Russia.
²⁶ Institute of Archaeology and Ethnography, Siberian Branch of Russian Academy of Sciences, Novosibirsk, Russia.
²⁷ Department of Archeology, Ethnography and Museology, Altai State University, Barnaul, Russia.
²⁸ Shaanxi Provincial Institute of Archaeology, Xi'an, China.
²⁹ School of Cultural Heritage, Northwest University, Xi'an, China.
³⁰ Xi'an AMS Center, Institute of Earth Environment, Chinese Academy of Sciences, Xi'an, China.
³¹ Research Center at the National Museum of Mongolia, Ulaanbaatar, Mongolia.
³² Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar, Mongolia.
³³ Department of Anthropology, National Museum of Nature and Science, Tsukuba, Japan.
³⁴ Archaeological Center of Chiba City, Chiba, Japan.
³⁵ Department of Archeology, Sakhalin Regional Museum, Yuzhno-Sakhalinsk, Russia.
³⁶ Department of Anthropology and Ethnology, Institute of Anthropology, State Key Laboratory of Cellular Stress Biology, School of Life Sciences, Xiamen University, Xiamen, China.
³⁷ Department of Biochemistry and Molecular Biology, Hainan Medical University, Haikou, China.
³⁸ Department of Human Anatomy and Center for Genomics and Personalized Medicine, Guangxi Medical University, Nanning, China.
³⁹ Key Laboratory for Molecular Genetic Mechanisms and Intervention Research on High Altitude Disease of Tibet Autonomous Region, Ministry of Education, School of Medicine, Xizang Minzu University (Tibet University for Nationalities), Xianyang, China.
⁴⁰ Institute for History and Culture of Science & Technology, Guangxi University for Nationalities, Nanning, China.
⁴¹ Department of Biology, Hainan Medical University, Haikou, China.
⁴² MOE Key Laboratory of Contemporary Anthropology, Department of Anthropology and Human Genetics, School of Life Sciences, Fudan University, Shanghai, China.
⁴³ College of History, Culture and Tourism, Guangxi Normal University, Guilin, China.
⁴⁴ Guangxi Institute of Cultural Relics Protection and Archaeology, Nanning, China.
⁴⁵ Belt and Road Research Center for Forensic Molecular Anthropology, Key Laboratory of Evidence Science of Gansu Province, Gansu Institute of Political Science and Law, Lanzhou, China.
⁴⁶ Eurasia3angle Research group, Max Planck Institute for the Science of Human History, Jena, Germany.
⁴⁷ Department of Anthropology, University of California Santa Barbara, Santa Barbara, CA, USA.
⁴⁸ Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany. krause@shh.mpg.de.
⁴⁹ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria. ron.pinhasi@univie.ac.at.
⁵⁰ Department of Genetics, Harvard Medical School, Boston, MA, USA. reich@genetics.med.harvard.edu.
⁵¹ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA. reich@genetics.med.harvard.edu.
⁵² Broad Institute of Harvard and MIT, Cambridge, MA, USA. reich@genetics.med.harvard.edu.
⁵³ Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA. reich@genetics.med.harvard.edu.

^# Contributed equally.

PMID: 33618348
PMCID: PMC7993749
DOI: 10.1038/s41586-021-03336-2

Genomic insights into the formation of human populations in East Asia

Chuan-Chao Wang et al. Nature. 2021 Mar.

. 2021 Mar;591(7850):413-419.

doi: 10.1038/s41586-021-03336-2. Epub 2021 Feb 22.

Authors

Affiliations

¹ Department of Anthropology and Ethnology, Institute of Anthropology, State Key Laboratory of Cellular Stress Biology, School of Life Sciences, Xiamen University, Xiamen, China. wang@xmu.edu.cn.
² Department of Genetics, Harvard Medical School, Boston, MA, USA. wang@xmu.edu.cn.
³ Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany. wang@xmu.edu.cn.
⁴ MOE Key Laboratory of Contemporary Anthropology, Department of Anthropology and Human Genetics, School of Life Sciences, Fudan University, Shanghai, China. wang@xmu.edu.cn.
⁵ School of Humanities, Nanyang Technological University, Nanyang, Singapore.
⁶ Scientific Museum, Far Eastern Federal University, Vladivostok, Russia.
⁷ Key Laboratory of Biomedical Information Engineering of Ministry of Education, School of Life Science and Technology, Xi'an Jiaotong University, Xi'an, China.
⁸ School of Health Science, Sapporo Medical University, Sapporo, Japan.
⁹ Department of Genetics, Harvard Medical School, Boston, MA, USA.
¹⁰ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA.
¹¹ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria.
¹² Institute of Archaeology, Russian Academy of Sciences, Moscow, Russia.
¹³ Department of Anthropology, Harvard University, Cambridge, MA, USA.
¹⁴ Broad Institute of Harvard and MIT, Cambridge, MA, USA.
¹⁵ Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA.
¹⁶ Department of Anthropology and Archaeology, National University of Mongolia, Ulaanbaatar, Mongolia.
¹⁷ Institute of Archaeology, National Cheng Kung University, Tainan, Taiwan.
¹⁸ Department of Anthropology, University of Washington, Seattle, WA, USA.
¹⁹ Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany.
²⁰ Institutes of Energy and the Environment, The Pennsylvania State University, University Park, PA, USA.
²¹ Department of Anthropology, Pennsylvania State University, University Park, PA, USA.
²² Institute of Archaeological Science, Fudan University, Shanghai, China.
²³ School of Ethnology and Sociology, Minzu University of China, Beijing, China.
²⁴ Institute of History and Philology, Institute of History and Philology, Academia Sinica, Taipei, Taiwan.
²⁵ Museum of Archaeology and Ethnology, Institute of History, Archaeology and Ethnology, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok, Russia.
²⁶ Institute of Archaeology and Ethnography, Siberian Branch of Russian Academy of Sciences, Novosibirsk, Russia.
²⁷ Department of Archeology, Ethnography and Museology, Altai State University, Barnaul, Russia.
²⁸ Shaanxi Provincial Institute of Archaeology, Xi'an, China.
²⁹ School of Cultural Heritage, Northwest University, Xi'an, China.
³⁰ Xi'an AMS Center, Institute of Earth Environment, Chinese Academy of Sciences, Xi'an, China.
³¹ Research Center at the National Museum of Mongolia, Ulaanbaatar, Mongolia.
³² Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar, Mongolia.
³³ Department of Anthropology, National Museum of Nature and Science, Tsukuba, Japan.
³⁴ Archaeological Center of Chiba City, Chiba, Japan.
³⁵ Department of Archeology, Sakhalin Regional Museum, Yuzhno-Sakhalinsk, Russia.
³⁶ Department of Anthropology and Ethnology, Institute of Anthropology, State Key Laboratory of Cellular Stress Biology, School of Life Sciences, Xiamen University, Xiamen, China.
³⁷ Department of Biochemistry and Molecular Biology, Hainan Medical University, Haikou, China.
³⁸ Department of Human Anatomy and Center for Genomics and Personalized Medicine, Guangxi Medical University, Nanning, China.
³⁹ Key Laboratory for Molecular Genetic Mechanisms and Intervention Research on High Altitude Disease of Tibet Autonomous Region, Ministry of Education, School of Medicine, Xizang Minzu University (Tibet University for Nationalities), Xianyang, China.
⁴⁰ Institute for History and Culture of Science & Technology, Guangxi University for Nationalities, Nanning, China.
⁴¹ Department of Biology, Hainan Medical University, Haikou, China.
⁴² MOE Key Laboratory of Contemporary Anthropology, Department of Anthropology and Human Genetics, School of Life Sciences, Fudan University, Shanghai, China.
⁴³ College of History, Culture and Tourism, Guangxi Normal University, Guilin, China.
⁴⁴ Guangxi Institute of Cultural Relics Protection and Archaeology, Nanning, China.
⁴⁵ Belt and Road Research Center for Forensic Molecular Anthropology, Key Laboratory of Evidence Science of Gansu Province, Gansu Institute of Political Science and Law, Lanzhou, China.
⁴⁶ Eurasia3angle Research group, Max Planck Institute for the Science of Human History, Jena, Germany.
⁴⁷ Department of Anthropology, University of California Santa Barbara, Santa Barbara, CA, USA.
⁴⁸ Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany. krause@shh.mpg.de.
⁴⁹ Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria. ron.pinhasi@univie.ac.at.
⁵⁰ Department of Genetics, Harvard Medical School, Boston, MA, USA. reich@genetics.med.harvard.edu.
⁵¹ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA. reich@genetics.med.harvard.edu.
⁵² Broad Institute of Harvard and MIT, Cambridge, MA, USA. reich@genetics.med.harvard.edu.
⁵³ Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, USA. reich@genetics.med.harvard.edu.

^# Contributed equally.

PMID: 33618348
PMCID: PMC7993749
DOI: 10.1038/s41586-021-03336-2

Abstract

The deep population history of East Asia remains poorly understood owing to a lack of ancient DNA data and sparse sampling of present-day people^1,2. Here we report genome-wide data from 166 East Asian individuals dating to between 6000 BC and AD 1000 and 46 present-day groups. Hunter-gatherers from Japan, the Amur River Basin, and people of Neolithic and Iron Age Taiwan and the Tibetan Plateau are linked by a deeply splitting lineage that probably reflects a coastal migration during the Late Pleistocene epoch. We also follow expansions during the subsequent Holocene epoch from four regions. First, hunter-gatherers from Mongolia and the Amur River Basin have ancestry shared by individuals who speak Mongolic and Tungusic languages, but do not carry ancestry characteristic of farmers from the West Liao River region (around 3000 BC), which contradicts theories that the expansion of these farmers spread the Mongolic and Tungusic proto-languages. Second, farmers from the Yellow River Basin (around 3000 BC) probably spread Sino-Tibetan languages, as their ancestry dispersed both to Tibet-where it forms approximately 84% of the gene pool in some groups-and to the Central Plain, where it has contributed around 59-84% to modern Han Chinese groups. Third, people from Taiwan from around 1300 BC to AD 800 derived approximately 75% of their ancestry from a lineage that is widespread in modern individuals who speak Austronesian, Tai-Kadai and Austroasiatic languages, and that we hypothesize derives from farmers of the Yangtze River Valley. Ancient people from Taiwan also derived about 25% of their ancestry from a northern lineage that is related to, but different from, farmers of the Yellow River Basin, which suggests an additional north-to-south expansion. Fourth, ancestry from Yamnaya Steppe pastoralists arrived in western Mongolia after around 3000 BC but was displaced by previously established lineages even while it persisted in western China, as would be expected if this ancestry was associated with the spread of proto-Tocharian Indo-European languages. Two later gene flows affected western Mongolia: migrants after around 2000 BC with Yamnaya and European farmer ancestry, and episodic influences of later groups with ancestry from Turan.

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Conflict of interest statement

Competing interests

The authors declare no competing interests.

Figures

**Extended Data Figure 1|. Principal Component Analysis (PCA).**
Projection of ancient samples onto PCA dimensions 1 and 2 defined by East Asians, Europeans, Siberians and Native Americans.

**Extended Data Figure 2|. Principal Component Analysis (PCA).**
(A) PCA dimensions 1 and 2 defined by present-day East Asians, Europeans, Siberians and Native Americans. (B) PCA dimensions 1 and 2 defined by present-day East Asian groups with the little West Eurasian mixture.

**Extended Data Figure 3|. Neighbour-joining tree of present-day East Eurasians based on F_st distances using the Human Origin dataset.**
(a) The branch length is shown in F_st distance, (b) Version where internal branches are all shown as having the same length for better visualization.

**Extended Data Figure 4|. ADMIXTURE plot at K=15 using the Human Origin dataset.**
We grouped the populations roughly into six groups from A to F based on geographic and genetic affinity. (A) populations mainly from Africa (yellow), America (magenta), West Eurasia (dark green and light brown) and Oceania (light magenta); (B) populations mainly from Mongolia (blue) and Siberia (purple); (C) populations mainly from southern China and Southeast Asia (light blue); (D) populations mainly from the Tibetan Plateau (olive) and Neolithic Yellow River Basin (red); (E) mainly Han Chinese around China (light blue and red); (F) populations mainly from the Amur River Basin (blue and red) and northeast Asia.

**Extended Data Figure 5|. Estimates of population split times.**
**(A)** Cross-coalescence rates for selected population pairs. We ran MSMC for four pairs of populations: Tibetan-Ami, Tibetan-Atayal, Tibetan-Ulchi and Tibetan-Mixe. We used one individuals from each population in this analysis. The modern genomic data for those individuals are from the Simons Genome Diversity Project. The times are calculated based on the mutation rate and generation time specified on the x-axis. **(B)** Cross-coalescence rates for selected population pairs. Same analysis as in Figure SI3–1, but using MSMC2 instead of MSMC, and using two individuals per population except for the Tibetan-Atayal pair, where we used only one.

**Extended Data Figure 6|. Admixture graph model.**
(This is the same as Figure 2 except that we show the fitted genetic drifts on each lineage.) We used all available sites in the 1240K dataset, restricting to transversions only to confirm that the same model fit (Supplementary Information section 3). We started with a skeleton tree that fits the data for Denisova, Mbuti, Onge, Tianyuan and Luxembourg Loschbour and one admixture event. We grafted on Mongolia East Neolithic, Upper Yellow River Late Neolithic farmers, Liangdao2, Japan Jomon, Nepal Chokhopani, Taiwan Hanben, and West Liao River Late Neolithic farmers in turn, adding them consecutively to all possible edges in the tree and retaining only graph solutions that provided no differences of |Z|<3 between fitted and estimated statistics (maximum |Z|=2.95 here). We used the MSMC and MSMC2 relative population split time estimates to constrain models. Deep splits are not well constrained due to minimal availability of Upper Paleolithic East Asian data. (a) Locations and dates of the East Asian individuals used in model fitting, with colours indicating whether the majority ancestry is from the hypothesized coastal expansion (green), interior expansion south (red), and interior expansion north. The map is based on the “Google Map Layer” from ArcGIS Online Basemaps (Map data ©2020 Google). (b) In the model visualization, we color lineages modelled as deriving entirely from one of these expansions, and also color populations according to ancestry proportions. Dashed lines represent admixture (proportions are marked), and we show the amount of genetic drift on each lineage in units of F_ST x 1000.

**Extended Data Figure 7|. Shared genetic drift among Tibetans, measured by f₃ (X, Y; Mbuti).**
Lighter colors indicate more shared drift. Lahu groups with the Southeast Asian Cluster probably due to substantial admixture. The Tibetan_Yajiang are geographically in the Tibeto-Burman Corridor but group with Core Tibetans, presumably reflecting less genetic admixture from people of the Southeast Asian Cluster.

**Figure 1:. Overview.**
(a) Locations, sample size (in brackets) and temporal distribution of newly reported ancient individuals, plotted using the “Google Map Layer” from ArcGIS Online Basemaps (Map data ©2020 Google). (b) Plot of first and second Principal Components defined in an analysis of East Asians with minimal West Eurasian-related mixture.

**Figure 2:. Model of deep population relationships.**
We start with a skeleton tree with one admixture event that when run on all SNPs fits the data for Denisova, Mbuti, Onge, Tianyuan and Loschbour according to *qpGraph*. We grafted on Mongolia East Neolithic, Upper Yellow River Late Neolithic farmers, Liangdao2, Japan Jomon, Nepal Chokhopani, Taiwan Hanben, and West Liao River Late Neolithic farmers, adding them consecutively to all possible edges and retaining only graphs that provided no differences of |Z|<3 between fitted and estimated statistics (maximum |Z|=2.95 here). We used MSMC and MSMC2 relative population split time estimates to constrain models. (a) We colour lineages modelled as from the hypothesized coastal expansion (green), interior southern expansion (red), or interior northern expansion (blue), and populations according to ancestry proportions. Dashed lines represent admixture (proportions marked). (b) Locations and dates of East Asians used in model fitting, with colours indicating the majority ancestry source, are plotted using the “Google Map Layer” from ArcGIS Online Basemaps (Map data ©2020 Google).

Figure 3:. Estimates of mixture proportions using *qpAdm.*
(a) *qpAdm* modelling of Yellow River farmer (blue) and Liangdao-related ancestry (orange) in present-day East Asians, with numbers from Online Table 22, and plotted using the “Google Map Layer” from ArcGIS Online Basemaps (Map data ©2020 Google). (b) Mongolians and Xinjiang. As sources we explored all possible subsets of Mongolia_East_N, Afanasievo, WSHG, Sintashta_MLBA, Turkmenistan_Gonur_BA_1, and Han Chinese, adding all groups to the reference set when not used as sources, and identifying parsimonious models (fewest numbers of sources) that fit at P>0.05 based on the Hotelling T² test implemented in *qpAdm* (Online Table 25). These P-values do not incorporate any correction for multiple hypothesis testing. * indicates parsimonious models that only pass at P>0.01. ** indicates cases where multiple equally parsimonious models pass at P>0.05 so we can not determine whether the West Eurasian-related source was Afanasievo, WSHG, or Sintashta_MLBA (we plot the model with the largest p-value). Bars show ancestry proportions, and time spans are unions of all samples. We do not visualize results from singleton outliers.

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[1] Cavalli-Sforza LL The Chinese human genome diversity project. Proc. Natl. Acad. Sci. USA 95, 11501–11503 (1998). - PMC - PubMed

[2] Cavalli-Sforza LL The Chinese human genome diversity project. Proc. Natl. Acad. Sci. USA 95, 11501–11503 (1998). - PMC - PubMed

[3] HUGO Pan-Asian SNP Consortium. Mapping human genetic diversity in Asia. Science 326, 1541–1545 (2009). - PubMed

[4] HUGO Pan-Asian SNP Consortium. Mapping human genetic diversity in Asia. Science 326, 1541–1545 (2009). - PubMed

[5] Haak W, et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 522, 207–211 (2015). - PMC - PubMed

[6] Haak W, et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 522, 207–211 (2015). - PMC - PubMed

[7] Allentoft ME, et al. Population genomics of Bronze Age Eurasia. Nature 522,167–172 (2015). - PubMed

[8] Allentoft ME, et al. Population genomics of Bronze Age Eurasia. Nature 522,167–172 (2015). - PubMed

[9] de Barros Damgaard P, et al.. 137 ancient human genomes from across the Eurasian steppes. Nature 557, 369–374 (2018). - PubMed

[10] de Barros Damgaard P, et al.. 137 ancient human genomes from across the Eurasian steppes. Nature 557, 369–374 (2018). - PubMed

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Genomic insights into the formation of human populations in East Asia

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Genomic insights into the formation of human populations in East Asia

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