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Here, we investigate the relationship between host genetic variation, parasitism and natural selection in a population of red colobus (<jats:italic>Procolobus rufomitratus tephrosceles<\/jats:italic>) in Kibale National Park, Uganda. We collected parasitological, cellular and genomic data to test the following hypotheses: (i) <jats:italic><jats:styled-content style=\"fixed-case\">MHC<\/jats:styled-content>\u2010<jats:styled-content style=\"fixed-case\">DQA<\/jats:styled-content>1<\/jats:italic> regulatory genetic variation is associated with control of whipworm (<jats:italic>Trichuris<\/jats:italic>) infection in a natural population of red colobus; (ii) infection\u2010associated <jats:styled-content style=\"fixed-case\">SNP<\/jats:styled-content>s are functional in driving differential gene expression in\u00a0vitro; and (iii) balancing selection has shaped patterns of variation in the <jats:italic><jats:styled-content style=\"fixed-case\">MHC<\/jats:styled-content>\u2010<jats:styled-content style=\"fixed-case\">DQA<\/jats:styled-content>1<\/jats:italic> promoter. We identified two <jats:styled-content style=\"fixed-case\">SNP<\/jats:styled-content>s in the <jats:italic><jats:styled-content style=\"fixed-case\">MHC<\/jats:styled-content>\u2010<jats:styled-content style=\"fixed-case\">DQA<\/jats:styled-content>1<\/jats:italic> promoter, both in transcription factor binding sites, and both of which are associated with decreased control of <jats:italic>Trichuris<\/jats:italic> infection. We characterized the function of both <jats:styled-content style=\"fixed-case\">SNP<\/jats:styled-content>s by testing differences in gene expression between the two alleles of each <jats:styled-content style=\"fixed-case\">SNP<\/jats:styled-content> in two mammalian cell lines. Alleles of one of the <jats:styled-content style=\"fixed-case\">SNP<\/jats:styled-content>s drove differential gene expression in both cell lines, while the other <jats:styled-content style=\"fixed-case\">SNP<\/jats:styled-content> drove differences in expression in one of the cell lines. Additionally, we found evidence of balancing selection acting on the <jats:italic><jats:styled-content style=\"fixed-case\">MHC<\/jats:styled-content>\u2010<jats:styled-content style=\"fixed-case\">DQA<\/jats:styled-content>1<\/jats:italic> promoter, including extensive trans\u2010species polymorphisms between red colobus and other primates, and an excess of intermediate\u2010frequency alleles relative to genome\u2010wide, coding and noncoding <jats:styled-content style=\"fixed-case\">RAD<\/jats:styled-content>seq data. Our data suggest that balancing selection provides adaptive regulatory flexibility that outweighs the consequences of increased parasite infection intensity in heterozygotes.<\/jats:p>","DOI":"10.1111\/mec.14221","type":"journal-article","created":{"date-parts":[[2017,6,30]],"date-time":"2017-06-30T11:32:57Z","timestamp":1498822377000},"page":"4523-4535","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":8,"title":["<i>Cis<\/i>\u2010regulatory evolution in a wild primate: Infection\u2010associated genetic variation drives differential expression of <i><scp>MHC<\/scp>\u2010<scp>DQA<\/scp>1<\/i> in\u00a0vitro"],"prefix":"10.1111","volume":"26","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9703-5802","authenticated-orcid":false,"given":"Noah D.","family":"Simons","sequence":"first","affiliation":[{"name":"Department of Anthropology University of Oregon Eugene OR USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Geeta N.","family":"Eick","sequence":"additional","affiliation":[{"name":"Department of Anthropology University of Oregon Eugene OR USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Maria J.","family":"Ruiz\u2010Lopez","sequence":"additional","affiliation":[{"name":"Institute of Ecology and Evolution University of Oregon Eugene OR USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Patrick A.","family":"Omeja","sequence":"additional","affiliation":[{"name":"Makerere University Biological Field Station Fort Portal Uganda"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Colin A.","family":"Chapman","sequence":"additional","affiliation":[{"name":"Makerere University Biological Field Station Fort Portal Uganda"},{"name":"Department of Anthropology and McGill School of Environment McGill University Montreal QC Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tony L.","family":"Goldberg","sequence":"additional","affiliation":[{"name":"Department of Pathobiological Sciences University of Wisconsin\u2010Madison Madison WI USA"},{"name":"Global Health Institute University of Wisconsin\u2010Madison Madison WI USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Nelson","family":"Ting","sequence":"additional","affiliation":[{"name":"Department of Anthropology University of Oregon Eugene OR USA"},{"name":"Institute of Ecology and Evolution University of Oregon Eugene OR USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kirstin N.","family":"Sterner","sequence":"additional","affiliation":[{"name":"Department of Anthropology University of Oregon Eugene OR USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2017,7,24]]},"reference":[{"key":"e_1_2_8_2_1","doi-asserted-by":"publisher","DOI":"10.1038\/sj.gene.6363765"},{"key":"e_1_2_8_3_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg3891"},{"key":"e_1_2_8_4_1","doi-asserted-by":"publisher","DOI":"10.1002\/ece3.395"},{"key":"e_1_2_8_5_1","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/25.17.3389"},{"key":"e_1_2_8_6_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msp190"},{"key":"e_1_2_8_7_1","doi-asserted-by":"publisher","DOI":"10.1002\/evan.20251"},{"key":"e_1_2_8_8_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.meegid.2011.09.020"},{"key":"e_1_2_8_9_1","doi-asserted-by":"publisher","DOI":"10.1002\/ece3.484"},{"key":"e_1_2_8_10_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0090714"},{"key":"e_1_2_8_11_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.162046399"},{"key":"e_1_2_8_12_1","doi-asserted-by":"publisher","DOI":"10.1016\/S1286-4579(99)00225-7"},{"key":"e_1_2_8_13_1","first-page":"684","article-title":"Identification of SNP markers for the endangered Ugandan red colobus (Procolobus rufomitratus tephrosceles) using RAD sequencing","volume":"15","author":"Blanchet S.","year":"2015","journal-title":"Molecular Ecology Resources"},{"key":"e_1_2_8_14_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00251-003-0630-5"},{"key":"e_1_2_8_15_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1744-313X.2008.00823.x"},{"key":"e_1_2_8_16_1","doi-asserted-by":"publisher","DOI":"10.1051\/parasite\/1998052137"},{"key":"e_1_2_8_17_1","doi-asserted-by":"publisher","DOI":"10.1186\/gb-2008-9-9-r143"},{"key":"e_1_2_8_18_1","doi-asserted-by":"publisher","DOI":"10.1111\/mec.12354"},{"key":"e_1_2_8_19_1","first-page":"1","article-title":"Protozoan parasites in group\u2010living primates: Testing the biological island hypothesis","volume":"73","author":"Chapman C. 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