Reconstructing Indian population history

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We analyse 25 diverse groups in India to provide strong evidence for two ancient populations, genetically divergent, that are ancestral to most Indians today. One, the ‘Ancestral North Indians’ (ANI), is genetically close to Middle Easterners, Central Asians, and Europeans, whereas the other, the ‘Ancestral South Indians’ (ASI), is as distinct from ANI and East Asians as they are from each other. By introducing methods that can estimate ancestry without accurate ancestral populations, we show that ANI ancestry ranges from 39–71% in most Indian groups, and is higher in traditionally upper caste and Indo-European speakers. Groups with only ASI ancestry may no longer exist in mainland India. However, the indigenous Andaman Islanders are unique in being ASI-related groups without ANI ancestry. Allele frequency differences between groups in India are larger than in Europe, reflecting strong founder effects whose signatures have been maintained for thousands of years owing to endogamy. We therefore predict that there will be an excess of recessive diseases in India, which should be possible to screen and map genetically. Analysis of genetic variation in 132 individuals from 25 diverse groups in India reveals that two ancient, genetically divergent populations are ancestral to most Indians today. One lineage, termed Ancestral North Indian, is genetically close to Middle Easterners, Central Asians and Europeans. The other, Ancestral South Indian, is not close to any group outside the subcontinent. The answers to several long-standing questions emerge from this work. It seems that 'caste' has been a powerful force shaping marriage in India for thousands of years — some anthropologists argued that its current strength was a product of British colonialism. And the enigmatic 'Negritos' of the Andaman Islands are identified as an ancient isolate from the Ancestral South Indian population. Allele frequency differences between population groups are high, in part due to the custom of within-group marriages, so it is likely that there is an excess of recessive diseases in India that can be screened for and mapped genetically. Genome-wide analysis of human variation in 25 diverse groups from India reveals two ancient populations, genetically divergent, that are ancestral to most Indians today. Traditionally upper caste and Indo-European speakers tend to be descended from a group that is genetically close to Middle Easterners, Central Asians and Europeans. 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All rights reserved"/> <meta name="dc.rights" content="2009 Macmillan Publishers Limited. All rights reserved"/> <meta name="dc.rightsAgent" content="journalpermissions@springernature.com"/> <meta name="dc.description" content="India has been underrepresented in genome-wide surveys of human variation. We analyse 25 diverse groups in India to provide strong evidence for two ancient populations, genetically divergent, that are ancestral to most Indians today. One, the ‘Ancestral North Indians’ (ANI), is genetically close to Middle Easterners, Central Asians, and Europeans, whereas the other, the ‘Ancestral South Indians’ (ASI), is as distinct from ANI and East Asians as they are from each other. By introducing methods that can estimate ancestry without accurate ancestral populations, we show that ANI ancestry ranges from 39–71% in most Indian groups, and is higher in traditionally upper caste and Indo-European speakers. Groups with only ASI ancestry may no longer exist in mainland India. However, the indigenous Andaman Islanders are unique in being ASI-related groups without ANI ancestry. Allele frequency differences between groups in India are larger than in Europe, reflecting strong founder effects whose signatures have been maintained for thousands of years owing to endogamy. We therefore predict that there will be an excess of recessive diseases in India, which should be possible to screen and map genetically. Analysis of genetic variation in 132 individuals from 25 diverse groups in India reveals that two ancient, genetically divergent populations are ancestral to most Indians today. One lineage, termed Ancestral North Indian, is genetically close to Middle Easterners, Central Asians and Europeans. The other, Ancestral South Indian, is not close to any group outside the subcontinent. The answers to several long-standing questions emerge from this work. It seems that 'caste' has been a powerful force shaping marriage in India for thousands of years — some anthropologists argued that its current strength was a product of British colonialism. And the enigmatic 'Negritos' of the Andaman Islands are identified as an ancient isolate from the Ancestral South Indian population. Allele frequency differences between population groups are high, in part due to the custom of within-group marriages, so it is likely that there is an excess of recessive diseases in India that can be screened for and mapped genetically. Genome-wide analysis of human variation in 25 diverse groups from India reveals two ancient populations, genetically divergent, that are ancestral to most Indians today. Traditionally upper caste and Indo-European speakers tend to be descended from a group that is genetically close to Middle Easterners, Central Asians and Europeans. 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All rights reserved"/> <meta name="prism.rightsAgent" content="journalpermissions@springernature.com"/> <meta name="prism.url" content="https://www.nature.com/articles/nature08365"/> <meta name="prism.doi" content="doi:10.1038/nature08365"/> <meta name="citation_pdf_url" content="https://www.nature.com/articles/nature08365.pdf"/> <meta name="citation_fulltext_html_url" content="https://www.nature.com/articles/nature08365"/> <meta name="citation_journal_title" content="Nature"/> <meta name="citation_journal_abbrev" content="Nature"/> <meta name="citation_publisher" content="Nature Publishing Group"/> <meta name="citation_issn" content="1476-4687"/> <meta name="citation_title" content="Reconstructing Indian population history"/> <meta name="citation_volume" content="461"/> <meta name="citation_issue" content="7263"/> <meta name="citation_publication_date" content="2009/09"/> <meta name="citation_firstpage" content="489"/> <meta name="citation_lastpage" content="494"/> <meta name="citation_article_type" content="Article"/> <meta name="citation_language" content="en"/> <meta name="dc.identifier" content="doi:10.1038/nature08365"/> <meta name="DOI" content="10.1038/nature08365"/> <meta name="size" content="163713"/> <meta name="citation_doi" content="10.1038/nature08365"/> <meta name="citation_springer_api_url" content="http://api.springer.com/xmldata/jats?q=doi:10.1038/nature08365&api_key="/> <meta name="description" content="India has been underrepresented in genome-wide surveys of human variation. We analyse 25 diverse groups in India to provide strong evidence for two ancient populations, genetically divergent, that are ancestral to most Indians today. One, the ‘Ancestral North Indians’ (ANI), is genetically close to Middle Easterners, Central Asians, and Europeans, whereas the other, the ‘Ancestral South Indians’ (ASI), is as distinct from ANI and East Asians as they are from each other. By introducing methods that can estimate ancestry without accurate ancestral populations, we show that ANI ancestry ranges from 39–71% in most Indian groups, and is higher in traditionally upper caste and Indo-European speakers. Groups with only ASI ancestry may no longer exist in mainland India. However, the indigenous Andaman Islanders are unique in being ASI-related groups without ANI ancestry. Allele frequency differences between groups in India are larger than in Europe, reflecting strong founder effects whose signatures have been maintained for thousands of years owing to endogamy. We therefore predict that there will be an excess of recessive diseases in India, which should be possible to screen and map genetically. Analysis of genetic variation in 132 individuals from 25 diverse groups in India reveals that two ancient, genetically divergent populations are ancestral to most Indians today. One lineage, termed Ancestral North Indian, is genetically close to Middle Easterners, Central Asians and Europeans. The other, Ancestral South Indian, is not close to any group outside the subcontinent. The answers to several long-standing questions emerge from this work. It seems that 'caste' has been a powerful force shaping marriage in India for thousands of years — some anthropologists argued that its current strength was a product of British colonialism. And the enigmatic 'Negritos' of the Andaman Islands are identified as an ancient isolate from the Ancestral South Indian population. Allele frequency differences between population groups are high, in part due to the custom of within-group marriages, so it is likely that there is an excess of recessive diseases in India that can be screened for and mapped genetically. Genome-wide analysis of human variation in 25 diverse groups from India reveals two ancient populations, genetically divergent, that are ancestral to most Indians today. Traditionally upper caste and Indo-European speakers tend to be descended from a group that is genetically close to Middle Easterners, Central Asians and Europeans. 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Stat.; citation_title=On the asymptotic normality of the jackknife; citation_author=D Thorburn; citation_volume=4; citation_publication_date=1977; citation_pages=113-118; citation_id=CR53"/> <meta name="citation_author" content="Reich, David"/> <meta name="citation_author_institution" content="Department of Genetics, Harvard Medical School, Boston, Massachusetts 02115, USA, "/> <meta name="citation_author_institution" content=" Broad Institute of Harvard and MIT, Cambridge, Massachusetts 02142, USA , "/> <meta name="citation_author" content="Thangaraj, Kumarasamy"/> <meta name="citation_author_institution" content="Centre for Cellular and Molecular Biology, Hyderabad 500 007, India"/> <meta name="citation_author" content="Patterson, Nick"/> <meta name="citation_author_institution" content=" Broad Institute of Harvard and MIT, Cambridge, Massachusetts 02142, USA , "/> <meta name="citation_author" content="Price, Alkes L."/> <meta name="citation_author_institution" content=" Broad Institute of Harvard and MIT, Cambridge, Massachusetts 02142, USA , "/> <meta name="citation_author_institution" content="Departments of Epidemiology and Biostatistics, Harvard School of Public Health, Boston, Massachusetts 02115, USA, "/> <meta name="citation_author" content="Singh, Lalji"/> <meta name="citation_author_institution" content="Centre for Cellular and Molecular Biology, Hyderabad 500 007, India"/> <meta name="access_endpoint" content="https://www.nature.com/platform/readcube-access"/> <meta name="twitter:site" content="@nature"/> <meta name="twitter:card" content="summary_large_image"/> <meta name="twitter:image:alt" content="Content cover image"/> <meta name="twitter:title" content="Reconstructing Indian population history"/> <meta name="twitter:description" content="Nature - Analysis of genetic variation in 132 individuals from 25 diverse groups in India reveals that two ancient, genetically divergent populations are ancestral to most Indians today. 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class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-David-Reich-Aff1-Aff2" data-author-popup="auth-David-Reich-Aff1-Aff2" data-author-search="Reich, David" data-corresp-id="c1">David Reich<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff2">2</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Kumarasamy-Thangaraj-Aff3" data-author-popup="auth-Kumarasamy-Thangaraj-Aff3" data-author-search="Thangaraj, Kumarasamy">Kumarasamy Thangaraj</a><sup class="u-js-hide"><a href="#Aff3">3</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Nick-Patterson-Aff2" data-author-popup="auth-Nick-Patterson-Aff2" data-author-search="Patterson, Nick">Nick Patterson</a><sup class="u-js-hide"><a href="#Aff2">2</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Alkes_L_-Price-Aff2-Aff4" data-author-popup="auth-Alkes_L_-Price-Aff2-Aff4" data-author-search="Price, Alkes L.">Alkes L. Price</a><sup class="u-js-hide"><a href="#Aff2">2</a>,<a href="#Aff4">4</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup> & </li><li class="c-article-author-list__show-more" aria-label="Show all 5 authors for this article" title="Show all 5 authors for this article">…</li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Lalji-Singh-Aff3" data-author-popup="auth-Lalji-Singh-Aff3" data-author-search="Singh, Lalji" data-corresp-id="c2">Lalji Singh<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><sup class="u-js-hide"><a href="#Aff3">3</a></sup> </li></ul><button aria-expanded="false" class="c-article-author-list__button"><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-down-medium"></use></svg><span>Show authors</span></button> <p class="c-article-info-details" data-container-section="info"> <a data-test="journal-link" href="/" data-track="click" data-track-action="journal homepage" data-track-category="article body" data-track-label="link"><i data-test="journal-title">Nature</i></a> <b data-test="journal-volume"><span class="u-visually-hidden">volume</span> 461</b>, <span class="u-visually-hidden">pages </span>489–494 (<span data-test="article-publication-year">2009</span>)<a href="#citeas" class="c-article-info-details__cite-as u-hide-print" data-track="click" data-track-action="cite this article" data-track-label="link">Cite this article</a> </p> <div class="c-article-metrics-bar__wrapper u-clear-both"> <ul class="c-article-metrics-bar u-list-reset"> <li class=" c-article-metrics-bar__item" data-test="access-count"> <p class="c-article-metrics-bar__count">28k <span class="c-article-metrics-bar__label">Accesses</span></p> </li> <li class="c-article-metrics-bar__item" data-test="citation-count"> <p class="c-article-metrics-bar__count">1145 <span class="c-article-metrics-bar__label">Citations</span></p> </li> <li class="c-article-metrics-bar__item" data-test="altmetric-score"> <p class="c-article-metrics-bar__count">391 <span class="c-article-metrics-bar__label">Altmetric</span></p> </li> <li class="c-article-metrics-bar__item"> <p class="c-article-metrics-bar__details"><a href="/articles/nature08365/metrics" data-track="click" data-track-action="view metrics" data-track-label="link" rel="nofollow">Metrics <span class="u-visually-hidden">details</span></a></p> </li> </ul> </div> </header> </div> <div class="c-article-body"> <section aria-labelledby="Abs1" data-title="Abstract" lang="en"><div class="c-article-section" id="Abs1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Abs1">Abstract</h2><div class="c-article-section__content" id="Abs1-content"><p>India has been underrepresented in genome-wide surveys of human variation. We analyse 25 diverse groups in India to provide strong evidence for two ancient populations, genetically divergent, that are ancestral to most Indians today. One, the ‘Ancestral North Indians’ (ANI), is genetically close to Middle Easterners, Central Asians, and Europeans, whereas the other, the ‘Ancestral South Indians’ (ASI), is as distinct from ANI and East Asians as they are from each other. By introducing methods that can estimate ancestry without accurate ancestral populations, we show that ANI ancestry ranges from 39–71% in most Indian groups, and is higher in traditionally upper caste and Indo-European speakers. Groups with only ASI ancestry may no longer exist in mainland India. However, the indigenous Andaman Islanders are unique in being ASI-related groups without ANI ancestry. Allele frequency differences between groups in India are larger than in Europe, reflecting strong founder effects whose signatures have been maintained for thousands of years owing to endogamy. 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G. Reddy, A. Shah and R. Tamang for generating the Y chromosome and mtDNA data; J. Neubauer for sample preparation; and A. Tandon for data curation. We thank B. N. Sarkar and A. G. Roy for helping with group census size estimates, and D. Falush, J. Novembre, A. Ruiz-Linares and S. Watkins for comments on the manuscript. D.R., N.P. and A.L.P. were supported by NIH grant HG004168, and D.R. was supported by a Burroughs Wellcome Career Development Award in the Biomedical Sciences. K.T. and L.S. were supported by grants from the Council of Scientific and Industrial Research of the Government of India, and K.T. was supported by a UKIERI Major Award (RG-4772).</p><p><b>Author Contributions</b> K.T. and L.S. collected the DNA samples, D.R., K.T. and L.S. collected the genetic data, N.P. developed the mathematical theory for <i>f</i>-statistics, and D.R., K.T., N.P. and A.L.P. analysed the data. D.R. wrote the manuscript and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/nature08365#MOESM280">Supplementary Information</a> with input from all authors.</p></div></div></section><section aria-labelledby="author-information" data-title="Author information"><div class="c-article-section" id="author-information-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="author-information">Author information</h2><div class="c-article-section__content" id="author-information-content"><span class="c-article-author-information__subtitle u-visually-hidden" id="author-notes">Author notes</span><ol class="c-article-author-information__list"><li class="c-article-author-information__item" id="na1"><p>David Reich, Kumarasamy Thangaraj, Nick Patterson and Alkes L. Price: These authors contributed equally to this work.</p></li></ol><h3 class="c-article__sub-heading" id="affiliations">Authors and Affiliations</h3><ol class="c-article-author-affiliation__list"><li id="Aff1"><p class="c-article-author-affiliation__address">Department of Genetics, Harvard Medical School, Boston, Massachusetts 02115, USA, </p><p class="c-article-author-affiliation__authors-list">David Reich</p></li><li id="Aff2"><p class="c-article-author-affiliation__address"> Broad Institute of Harvard and MIT, Cambridge, Massachusetts 02142, USA , </p><p class="c-article-author-affiliation__authors-list">David Reich, Nick Patterson & Alkes L. Price</p></li><li id="Aff3"><p class="c-article-author-affiliation__address">Centre for Cellular and Molecular Biology, Hyderabad 500 007, India</p><p class="c-article-author-affiliation__authors-list">Kumarasamy Thangaraj & Lalji Singh</p></li><li id="Aff4"><p class="c-article-author-affiliation__address">Departments of Epidemiology and Biostatistics, Harvard School of Public Health, Boston, Massachusetts 02115, USA, </p><p class="c-article-author-affiliation__authors-list">Alkes L. Price</p></li></ol><div class="u-js-hide u-hide-print" data-test="author-info"><span class="c-article__sub-heading">Authors</span><ol class="c-article-authors-search u-list-reset"><li id="auth-David-Reich-Aff1-Aff2"><span class="c-article-authors-search__title u-h3 js-search-name">David Reich</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?author=David%20Reich" class="c-article-button" data-track="click" data-track-action="author link - publication" data-track-label="link" rel="nofollow">View author publications</a></div><div class="c-article-authors-search__item c-article-authors-search__list-item--right"><p class="search-in-title-js c-article-authors-search__text">You can also search for this author in <span class="c-article-identifiers"><a class="c-article-identifiers__item" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=search&term=David%20Reich" data-track="click" data-track-action="author link - pubmed" data-track-label="link" rel="nofollow">PubMed</a><span class="u-hide"> </span><a class="c-article-identifiers__item" href="http://scholar.google.co.uk/scholar?as_q=&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_sauthors=%22David%20Reich%22&as_publication=&as_ylo=&as_yhi=&as_allsubj=all&hl=en" data-track="click" data-track-action="author link - scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li><li id="auth-Kumarasamy-Thangaraj-Aff3"><span class="c-article-authors-search__title u-h3 js-search-name">Kumarasamy Thangaraj</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?author=Kumarasamy%20Thangaraj" class="c-article-button" data-track="click" data-track-action="author link - publication" data-track-label="link" rel="nofollow">View author publications</a></div><div class="c-article-authors-search__item c-article-authors-search__list-item--right"><p class="search-in-title-js c-article-authors-search__text">You can also search for this author in <span class="c-article-identifiers"><a class="c-article-identifiers__item" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=search&term=Kumarasamy%20Thangaraj" data-track="click" data-track-action="author link - pubmed" data-track-label="link" rel="nofollow">PubMed</a><span class="u-hide"> </span><a class="c-article-identifiers__item" href="http://scholar.google.co.uk/scholar?as_q=&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_sauthors=%22Kumarasamy%20Thangaraj%22&as_publication=&as_ylo=&as_yhi=&as_allsubj=all&hl=en" data-track="click" data-track-action="author link - scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li><li id="auth-Nick-Patterson-Aff2"><span class="c-article-authors-search__title u-h3 js-search-name">Nick Patterson</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?author=Nick%20Patterson" class="c-article-button" data-track="click" data-track-action="author link - publication" data-track-label="link" rel="nofollow">View author publications</a></div><div class="c-article-authors-search__item c-article-authors-search__list-item--right"><p class="search-in-title-js c-article-authors-search__text">You can also search for this author in <span class="c-article-identifiers"><a class="c-article-identifiers__item" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=search&term=Nick%20Patterson" data-track="click" data-track-action="author link - pubmed" data-track-label="link" rel="nofollow">PubMed</a><span class="u-hide"> </span><a class="c-article-identifiers__item" href="http://scholar.google.co.uk/scholar?as_q=&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_sauthors=%22Nick%20Patterson%22&as_publication=&as_ylo=&as_yhi=&as_allsubj=all&hl=en" data-track="click" data-track-action="author link - scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li><li id="auth-Alkes_L_-Price-Aff2-Aff4"><span class="c-article-authors-search__title u-h3 js-search-name">Alkes L. Price</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?author=Alkes%20L.%20Price" class="c-article-button" data-track="click" data-track-action="author link - publication" data-track-label="link" rel="nofollow">View author publications</a></div><div class="c-article-authors-search__item c-article-authors-search__list-item--right"><p class="search-in-title-js c-article-authors-search__text">You can also search for this author in <span class="c-article-identifiers"><a class="c-article-identifiers__item" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=search&term=Alkes%20L.%20Price" data-track="click" data-track-action="author link - pubmed" data-track-label="link" rel="nofollow">PubMed</a><span class="u-hide"> </span><a class="c-article-identifiers__item" href="http://scholar.google.co.uk/scholar?as_q=&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_sauthors=%22Alkes%20L.%20Price%22&as_publication=&as_ylo=&as_yhi=&as_allsubj=all&hl=en" data-track="click" data-track-action="author link - scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li><li id="auth-Lalji-Singh-Aff3"><span class="c-article-authors-search__title u-h3 js-search-name">Lalji Singh</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?author=Lalji%20Singh" class="c-article-button" data-track="click" data-track-action="author link - publication" data-track-label="link" rel="nofollow">View author publications</a></div><div class="c-article-authors-search__item c-article-authors-search__list-item--right"><p class="search-in-title-js c-article-authors-search__text">You can also search for this author in <span class="c-article-identifiers"><a class="c-article-identifiers__item" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=search&term=Lalji%20Singh" data-track="click" data-track-action="author link - pubmed" data-track-label="link" rel="nofollow">PubMed</a><span class="u-hide"> </span><a class="c-article-identifiers__item" href="http://scholar.google.co.uk/scholar?as_q=&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_sauthors=%22Lalji%20Singh%22&as_publication=&as_ylo=&as_yhi=&as_allsubj=all&hl=en" data-track="click" data-track-action="author link - scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li></ol></div><h3 class="c-article__sub-heading" id="corresponding-author">Corresponding authors</h3><p id="corresponding-author-list">Correspondence to <a id="corresp-c1" href="mailto:reich@genetics.med.harvard.edu">David Reich</a> or <a id="corresp-c2" href="mailto:lalji@ccmb.res.in">Lalji Singh</a>.</p></div></div></section><section data-title="Supplementary information"><div class="c-article-section" id="Sec18-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec18">Supplementary information</h2><div class="c-article-section__content" id="Sec18-content"><div data-test="supplementary-info"><div id="figshareContainer" class="c-article-figshare-container" data-test="figshare-container"></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM280"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="supplementary information" href="https://static-content.springer.com/esm/art%3A10.1038%2Fnature08365/MediaObjects/41586_2009_BFnature08365_MOESM280_ESM.pdf" data-supp-info-image="">Supplementary Information</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>This file contains Supplementary Tables S1-S6, Supplementary Figures S1-S7, Supplementary Notes S1-S5 and Supplementary References. (PDF 1777 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM281"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="supplementary appendix" href="https://static-content.springer.com/esm/art%3A10.1038%2Fnature08365/MediaObjects/41586_2009_BFnature08365_MOESM281_ESM.pdf" data-supp-info-image="">Supplementary Appendix</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>This file contains Supplementary Data, Supplementary Statistics and Supplementary References. (PDF 162 kb)</p></div></div></div></div></div></section><section data-title="PowerPoint slides"><div class="c-article-section" id="Sec19-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec19">PowerPoint slides</h2><div class="c-article-section__content" id="Sec19-content"><div data-test="supplementary-info"><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM276"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="powerpoint slide for fig. 1" href="https://static-content.springer.com/esm/art%3A10.1038%2Fnature08365/MediaObjects/41586_2009_BFnature08365_MOESM276_ESM.ppt" data-supp-info-image="">PowerPoint slide for Fig. 1</a></h3></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM277"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" 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Summary</h2><h3 class="c-article-editorial-summary__article-title u-mb-8">Meet the ancestors: Indian population history from gene screening</h3><div class="c-article-editorial-summary__content c-article-editorial-summary__content--less"><p>Analysis of genetic variation in 132 individuals from 25 diverse groups in India reveals that two ancient, genetically divergent populations are ancestral to most Indians today. 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