Convergent adaptation of human lactase persistence in Africa and Europe

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Identification of the Palaeocene/Eocene thermal maximum in a marine sedimentary sequence. A SNP in the gene encoding lactase (LCT) (C/T-13910) is associated with the ability to digest milk as adults (lactase persistence) in Europeans, but the genetic basis of lactase persistence in Africans was previously unknown. We conducted a genotype-phenotype association study in 470 Tanzanians, Kenyans and Sudanese and identified three SNPs (G/C-14010, T/G-13915 and C/G-13907) that are associated with lactase persistence and that have derived alleles that significantly enhance transcription from the LCT promoter in vitro. These SNPs originated on different haplotype backgrounds from the European C/T-13910 SNP and from each other. Genotyping across a 3-Mb region demonstrated haplotype homozygosity extending >2.0 Mb on chromosomes carrying C-14010, consistent with a selective sweep over the past ∼7,000 years. These data provide a marked example of convergent evolution due to strong selective pressure resulting from shared cultural traits—animal domestication and adult milk consumption.","datePublished":"2006-12-10T00:00:00Z","dateModified":"2006-12-10T00:00:00Z","pageStart":"31","pageEnd":"40","sameAs":"https://doi.org/10.1038/ng1946","keywords":["Biomedicine","general","Human Genetics","Cancer Research","Agriculture","Gene Function","Animal Genetics and 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data-test="authors-list" data-component-authors-activator="authors-list"><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-Sarah_A-Tishkoff-Aff1" data-author-popup="auth-Sarah_A-Tishkoff-Aff1" data-author-search="Tishkoff, Sarah A" data-corresp-id="c1">Sarah A Tishkoff<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></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-Floyd_A-Reed-Aff1" data-author-popup="auth-Floyd_A-Reed-Aff1" data-author-search="Reed, Floyd A">Floyd A Reed</a><sup class="u-js-hide"><a href="#Aff1">1</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-Alessia-Ranciaro-Aff1-Aff2" data-author-popup="auth-Alessia-Ranciaro-Aff1-Aff2" data-author-search="Ranciaro, Alessia">Alessia Ranciaro</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff2">2</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-Benjamin_F-Voight-Aff3" data-author-popup="auth-Benjamin_F-Voight-Aff3" data-author-search="Voight, Benjamin F">Benjamin F Voight</a><sup class="u-js-hide"><a href="#Aff3">3</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-Courtney_C-Babbitt-Aff4" data-author-popup="auth-Courtney_C-Babbitt-Aff4" data-author-search="Babbitt, Courtney C">Courtney C Babbitt</a><sup class="u-js-hide"><a href="#Aff4">4</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-Jesse_S-Silverman-Aff4" data-author-popup="auth-Jesse_S-Silverman-Aff4" data-author-search="Silverman, Jesse S">Jesse S Silverman</a><sup class="u-js-hide"><a href="#Aff4">4</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-Kweli-Powell-Aff1" data-author-popup="auth-Kweli-Powell-Aff1" data-author-search="Powell, Kweli">Kweli Powell</a><sup class="u-js-hide"><a href="#Aff1">1</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-Holly_M-Mortensen-Aff1" data-author-popup="auth-Holly_M-Mortensen-Aff1" data-author-search="Mortensen, Holly M">Holly M Mortensen</a><sup class="u-js-hide"><a href="#Aff1">1</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-Jibril_B-Hirbo-Aff1" data-author-popup="auth-Jibril_B-Hirbo-Aff1" data-author-search="Hirbo, Jibril B">Jibril B Hirbo</a><sup class="u-js-hide"><a href="#Aff1">1</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-Maha-Osman-Aff5" data-author-popup="auth-Maha-Osman-Aff5" data-author-search="Osman, Maha">Maha Osman</a><sup class="u-js-hide"><a href="#Aff5">5</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-Muntaser-Ibrahim-Aff5" data-author-popup="auth-Muntaser-Ibrahim-Aff5" data-author-search="Ibrahim, Muntaser">Muntaser Ibrahim</a><sup class="u-js-hide"><a href="#Aff5">5</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-Sabah_A-Omar-Aff6" data-author-popup="auth-Sabah_A-Omar-Aff6" data-author-search="Omar, Sabah A">Sabah A Omar</a><sup class="u-js-hide"><a href="#Aff6">6</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-Godfrey-Lema-Aff7" data-author-popup="auth-Godfrey-Lema-Aff7" data-author-search="Lema, Godfrey">Godfrey Lema</a><sup class="u-js-hide"><a href="#Aff7">7</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-Thomas_B-Nyambo-Aff7" data-author-popup="auth-Thomas_B-Nyambo-Aff7" data-author-search="Nyambo, Thomas B">Thomas B Nyambo</a><sup class="u-js-hide"><a href="#Aff7">7</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-Jilur-Ghori-Aff8" data-author-popup="auth-Jilur-Ghori-Aff8" data-author-search="Ghori, Jilur">Jilur Ghori</a><sup class="u-js-hide"><a href="#Aff8">8</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-Suzannah-Bumpstead-Aff8" data-author-popup="auth-Suzannah-Bumpstead-Aff8" data-author-search="Bumpstead, Suzannah">Suzannah Bumpstead</a><sup class="u-js-hide"><a href="#Aff8">8</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-Jonathan_K-Pritchard-Aff3" data-author-popup="auth-Jonathan_K-Pritchard-Aff3" data-author-search="Pritchard, Jonathan K">Jonathan K Pritchard</a><sup class="u-js-hide"><a href="#Aff3">3</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-Gregory_A-Wray-Aff4" data-author-popup="auth-Gregory_A-Wray-Aff4" data-author-search="Wray, Gregory A">Gregory A Wray</a><sup class="u-js-hide"><a href="#Aff4">4</a></sup> & </li><li class="c-article-author-list__show-more" aria-label="Show all 19 authors for this article" title="Show all 19 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-Panos-Deloukas-Aff8" data-author-popup="auth-Panos-Deloukas-Aff8" data-author-search="Deloukas, Panos">Panos Deloukas</a><sup class="u-js-hide"><a href="#Aff8">8</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="/ng" data-track="click" data-track-action="journal homepage" data-track-category="article body" data-track-label="link"><i data-test="journal-title">Nature Genetics</i></a> <b data-test="journal-volume"><span class="u-visually-hidden">volume</span> 39</b>, <span class="u-visually-hidden">pages </span>31–40 (<span data-test="article-publication-year">2007</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">29k <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">969 <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">376 <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/ng1946/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="Abs2" data-title="Abstract" lang="en"><div class="c-article-section" id="Abs2-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Abs2">Abstract</h2><div class="c-article-section__content" id="Abs2-content"><p>A SNP in the gene encoding lactase (<i>LCT</i>) (C/T-13910) is associated with the ability to digest milk as adults (lactase persistence) in Europeans, but the genetic basis of lactase persistence in Africans was previously unknown. We conducted a genotype-phenotype association study in 470 Tanzanians, Kenyans and Sudanese and identified three SNPs (G/C-14010, T/G-13915 and C/G-13907) that are associated with lactase persistence and that have derived alleles that significantly enhance transcription from the <i>LCT</i> promoter <i>in vitro</i>. These SNPs originated on different haplotype backgrounds from the European C/T-13910 SNP and from each other. Genotyping across a 3-Mb region demonstrated haplotype homozygosity extending >2.0 Mb on chromosomes carrying C-14010, consistent with a selective sweep over the past <span class="stix">∼</span>7,000 years. These data provide a marked example of convergent evolution due to strong selective pressure resulting from shared cultural traits—animal domestication and adult milk consumption.</p></div></div></section> <noscript> <div class="c-nature-box c-nature-box--side " data-component="entitlement-box"> <div class="js-access-button"> <a href="https://wayf.springernature.com?redirect_uri=https%3A%2F%2Fwww.nature.com%2Farticles%2Fng1946" class="c-article__button" data-test="ra21"> <svg class="u-icon" width="18" height="18" aria-hidden="true" focusable="false"><use href="#icon-institution"></use></svg> <span class="c-article__button-text">Access through your institution</span> </a> </div> <div class="js-buy-button"> <a href="#access-options" class="c-article__button c-article__button--inverted" data-test="ra21"> <span>Buy or subscribe</span> </a> </div> </div> </noscript> <div class="js-context-bar-sticky-point-mobile" data-track-context="article body"> </div> <div 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data-track-label="link">Contact customer support</a> </li> </ul> </nav> <div class="u-display-none"> <div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-1"><figure><figcaption><b id="Fig1" class="c-article-section__figure-caption" data-test="figure-caption-text">Figure 1: Map of the <i>LCT</i> and <i>MCM6</i> gene region and location of genotyped SNPs.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><picture><source type="image/webp" srcset="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig1_HTML.jpg?as=webp"><img src="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig1_HTML.jpg" alt="" loading="lazy" width="312" height="206"></picture></div></div></figure></div><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-2"><figure><figcaption><b id="Fig2" class="c-article-section__figure-caption" data-test="figure-caption-text">Figure 2: Map of phenotype and genotype proportions for each population group considered in this study.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><picture><source type="image/webp" srcset="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig2_HTML.jpg?as=webp"><img src="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig2_HTML.jpg" alt="" loading="lazy" width="312" height="181"></picture></div></div></figure></div><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-3"><figure><figcaption><b id="Fig3" class="c-article-section__figure-caption" data-test="figure-caption-text">Figure 3: Genotype-phenotype association for G/C-14010, T/G-13915 and C/G-13907.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><picture><source type="image/webp" srcset="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig3_HTML.jpg?as=webp"><img src="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig3_HTML.jpg" alt="" loading="lazy" width="312" height="274"></picture></div></div></figure></div><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-4"><figure><figcaption><b id="Fig4" class="c-article-section__figure-caption" data-test="figure-caption-text">Figure 4: Haplotype networks consisting of 55 SNPs spanning a 98-kb region encompassing <i>LCT</i> and <i>MCM6</i>.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><picture><source type="image/webp" srcset="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig4_HTML.jpg?as=webp"><img src="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig4_HTML.jpg" alt="" loading="lazy" width="287" height="312"></picture></div></div></figure></div><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-5"><figure><figcaption><b id="Fig5" class="c-article-section__figure-caption" data-test="figure-caption-text">Figure 5: Dual-luciferase reporter assay of <i>LCT</i> promoter and <i>MCM6</i> introns.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><picture><source type="image/webp" srcset="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig5_HTML.jpg?as=webp"><img src="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig5_HTML.jpg" alt="" loading="lazy" width="312" height="177"></picture></div></div></figure></div><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-6"><figure><figcaption><b id="Fig6" class="c-article-section__figure-caption" data-test="figure-caption-text">Figure 6: Comparison of tracts of homozygous genotypes flanking the lactase persistence–associated SNPs.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><picture><source type="image/webp" srcset="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig6_HTML.jpg?as=webp"><img src="//media.springernature.com/m312/springer-static/image/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_Article_BFng1946_Fig6_HTML.jpg" alt="" loading="lazy" width="219" height="312"></picture></div></div></figure></div><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-7"><figure><figcaption><b id="Fig7" class="c-article-section__figure-caption" data-test="figure-caption-text">Figure 7: Plots of the extent and decay of haplotype homozygosity in the region surrounding the C-14010 allele.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><picture><source type="image/webp" 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Panchapakesan, E. King, S. Morrow and T. Severson for technical assistance. We thank E. Sibley and L.C. Olds for sharing advice and materials and T. Bersaglieri and J. Hirschhorn for sharing data. We thank S.J. Deo, P. Lufungulo, W. Ntandu, A. Mabulla, J.L. Mountain, J. Hanby, D. Bygott, A. Tibwitta, D. Kariuki, L. Alando, E. Aluvala, F. Mohammed, A. Teia and A.A. Mohamed for their assistance with sample collection. We thank A. Clark for critical review of the manuscript and for helpful suggestions and we thank L. Peltonen, N. Enattah and C. Ehret for discussion. We thank the African participants who generously donated DNA and phenotype information so that we might learn more about their population history and the genetic basis of lactase persistence in Africa. This study was funded by L.S.B. Leakey and Wenner Gren Foundation grants, US National Science Foundation (NSF) grants BSC-0196183 and BSC-0552486, US National Institutes of Health (NIH) grant R01GM076637 and David and Lucile Packard and Burroughs Wellcome Foundation Career Awards to S.A.T. K.P. and H.M.M. were funded by NSF grant IGERT-9987590 to S.A.T. F.A.R. was supported by US National Institutes of Health (NIH) grant F32HG03801. B.F.V. and J.K.P. were supported by NIH grant HG002772-1. The Institute for Genome Sciences and Policy of Duke University supported the work of C.C.B., J.S.S. and G.A.W. The Wellcome Trust supported the work of J.G., S.B. and P.D.</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>Sarah A Tishkoff and Floyd A Reed: 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 Biology, University of Maryland, College Park, 20742, Maryland, USA</p><p class="c-article-author-affiliation__authors-list">Sarah A Tishkoff, Floyd A Reed, Alessia Ranciaro, Kweli Powell, Holly M Mortensen & Jibril B Hirbo</p></li><li id="Aff2"><p class="c-article-author-affiliation__address">Department of Biology, University of Ferrara, Ferrara, 44100, Italy</p><p class="c-article-author-affiliation__authors-list">Alessia Ranciaro</p></li><li id="Aff3"><p class="c-article-author-affiliation__address">Department of Human Genetics, University of Chicago, Chicago, 60637, Illinois, USA</p><p class="c-article-author-affiliation__authors-list">Benjamin F Voight & Jonathan K Pritchard</p></li><li id="Aff4"><p class="c-article-author-affiliation__address">Institute for Genome Sciences & Policy and Department of Biology, Duke University, Durham, 27708, North Carolina, USA</p><p class="c-article-author-affiliation__authors-list">Courtney C Babbitt, Jesse S Silverman & Gregory A Wray</p></li><li id="Aff5"><p class="c-article-author-affiliation__address">Department of Molecular Biology, Institute of Endemic Diseases, University of Khartoum, Khartoum, 15-13, Sudan</p><p class="c-article-author-affiliation__authors-list">Maha Osman & Muntaser Ibrahim</p></li><li id="Aff6"><p class="c-article-author-affiliation__address">Kenya Medical Research Institute, Centre for Biotechnology Research and Development, Nairobi, 54840-00200, Kenya</p><p class="c-article-author-affiliation__authors-list">Sabah A Omar</p></li><li id="Aff7"><p class="c-article-author-affiliation__address">Department of Biochemistry, Muhimbili University College of Health Sciences, Dar es Salaam, Tanzania</p><p class="c-article-author-affiliation__authors-list">Godfrey Lema & Thomas B Nyambo</p></li><li id="Aff8"><p class="c-article-author-affiliation__address">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, CB10 1SA, Cambridge, UK</p><p class="c-article-author-affiliation__authors-list">Jilur Ghori, Suzannah Bumpstead & Panos Deloukas</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-Sarah_A-Tishkoff-Aff1"><span class="c-article-authors-search__title u-h3 js-search-name">Sarah A Tishkoff</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=Sarah%20A%20Tishkoff" class="c-article-button" data-track="click" data-track-action="author link - 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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=Panos%20Deloukas" 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=%22Panos%20Deloukas%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="contributions">Contributions</h3><p>S.A.T. conceived and supervised the study. S.A.T., K.P., H.M.M., A.R., J.B.H., M.O., M.I., S.A.O., G.L. and T.B.N. were involved in DNA collection and phenotype testing. A.R. performed the resequencing and initial identification of association of candidate SNPs with the phenotype. S.A.T. and F.A.R. selected the SNPs to be genotyped and samples to test for gene expression. P.D., J.G. and S.B. performed the SNP design and genotyping. F.A.R. processed and phased the raw data and performed the genotype-phenotype association analyses, plots of haplotype homozygosity from unphased data, dominance estimates and pairwise plot of LD. B.F.V. performed, and J.K.P. co-supervised, the iHS test to detect positive selection and plots of haplotype homozygosity from phased data as well as rejection-sampling analyses to estimate age of alleles and selection parameters. H.M.M. constructed the haplotype networks. C.C.B., J.S.S. and G.A.W. built the expression constructs, carried out transcription assays and analyzed the results of expression assays. The paper was written primarily by S.A.T., with contributions from F.A.R., B.F.V., J.K.P., C.C.B., G.A.W. and P.D. The supplementary information was written by S.A.T. and F.A.R. with contributions from B.F.V., J.K.P., C.C.B., G.A.W. and P.D.</p><h3 class="c-article__sub-heading" id="corresponding-author">Corresponding author</h3><p id="corresponding-author-list">Correspondence to <a id="corresp-c1" href="mailto:Tishkoff@umd.edu">Sarah A Tishkoff</a>.</p></div></div></section><section data-title="Ethics declarations"><div class="c-article-section" id="ethics-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="ethics">Ethics declarations</h2><div class="c-article-section__content" id="ethics-content"> <h3 class="c-article__sub-heading">Competing interests</h3> <p>The authors declare no competing financial interests.</p> </div></div></section><section data-title="Supplementary information"><div class="c-article-section" id="Sec28-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec28">Supplementary information</h2><div class="c-article-section__content" id="Sec28-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="MOESM1"><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 fig. 1" href="https://static-content.springer.com/esm/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_BFng1946_MOESM1_ESM.pdf" data-supp-info-image="">Supplementary Fig. 1</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>Distribution of phenotype values for the pooled African data set. (PDF 611 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM2"><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 fig. 2" href="https://static-content.springer.com/esm/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_BFng1946_MOESM2_ESM.pdf" data-supp-info-image="">Supplementary Fig. 2</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>Plots of the extent and decay of haplotype homozygosity in the region surrounding the G-rs2322813, G-13907 and G-13915 alleles. (PDF 400 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM3"><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 fig. 3" href="https://static-content.springer.com/esm/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_BFng1946_MOESM3_ESM.pdf" data-supp-info-image="">Supplementary Fig. 3</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>Plot of the degree of LD between each pair of genotyped SNPs. (PDF 2978 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM4"><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 table 1" href="https://static-content.springer.com/esm/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_BFng1946_MOESM4_ESM.pdf" data-supp-info-image="">Supplementary Table 1</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>Language, subsistence and lactase persistence classifications of sampled populations and LP-associated allele frequencies. (PDF 15 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM5"><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 table 2" href="https://static-content.springer.com/esm/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_BFng1946_MOESM5_ESM.pdf" data-supp-info-image="">Supplementary Table 2</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>Genotyped SNP identifications and locations. (PDF 27 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM6"><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 table 3" href="https://static-content.springer.com/esm/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_BFng1946_MOESM6_ESM.pdf" data-supp-info-image="">Supplementary Table 3</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>Significance of iHS under assorted demographic models (PDF 22 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM7"><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 methods (pdf 161 kb)" href="https://static-content.springer.com/esm/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_BFng1946_MOESM7_ESM.pdf" data-supp-info-image="">Supplementary Methods (PDF 161 kb)</a></h3></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM8"><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 discussion (pdf 115 kb)" href="https://static-content.springer.com/esm/art%3A10.1038%2Fng1946/MediaObjects/41588_2007_BFng1946_MOESM8_ESM.pdf" data-supp-info-image="">Supplementary Discussion (PDF 115 kb)</a></h3></div></div></div></div></section><section data-title="Rights and permissions"><div class="c-article-section" id="rightslink-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="rightslink">Rights and permissions</h2><div class="c-article-section__content" 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