{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,12]],"date-time":"2026-09-12T19:31:26Z","timestamp":1789241486926,"version":"build-2803163510"},"reference-count":142,"publisher":"National Academy of Sciences","issue":"22","license":[{"start":{"date-parts":[[2025,5,30]],"date-time":"2025-05-30T00:00:00Z","timestamp":1748563200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000051","name":"HHS | NIH | National Human Genome Research Institute","doi-asserted-by":"publisher","award":["Intramural Program"],"award-info":[{"award-number":["Intramural Program"]}],"id":[{"id":"10.13039\/100000051","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2025,6,3]]},"abstract":"<jats:p>\n                    Gastric cancer ranks as the fifth most common human cancer worldwide and has a poor survival rate and limited treatment options. Despite the high prevalence and mortality rate, the genetic etiology is largely unknown. In dogs, a clinically and histologically similar disease disproportionately affects two breeds, the Belgian Tervuren and Belgian Sheepdog, which develop the intestinal and diffuse tumor subtypes observed in humans. We performed a Bayesian genome-wide association study and selection analyses in this naturally occurring canine model to elucidate underlying genetic risk factors for gastric cancer and identified 15 associated loci. Variant filtering revealed germline putative regulatory variants for the\n                    <jats:italic toggle=\"yes\">EPAS1<\/jats:italic>\n                    (\n                    <jats:italic toggle=\"yes\">HIF2A<\/jats:italic>\n                    ) and\n                    <jats:italic toggle=\"yes\">PTEN<\/jats:italic>\n                    genes and a coding variant in\n                    <jats:italic toggle=\"yes\">CD101<\/jats:italic>\n                    . Two loci are overrepresented among cases of intestinal tumor subtype. Although closely related to Tervuren and Sheepdogs, Belgian Malinois rarely develop gastric cancer. Across-breed analyses uncovered haplotypes enriched in Malinois at\n                    <jats:italic toggle=\"yes\">SOX2-OT<\/jats:italic>\n                    and\n                    <jats:italic toggle=\"yes\">IGF2BP2<\/jats:italic>\n                    that are at significantly higher frequency among genome-wide association study controls. Among Tervuren and Sheepdogs,\n                    <jats:italic toggle=\"yes\">HDAC2<\/jats:italic>\n                    putative regulatory variants were present at comparatively high frequency and were associated with risk of gastric cancer. Here, we describe a complex genetic architecture governing gastric cancer in a dog model, including genes such as\n                    <jats:italic toggle=\"yes\">PDZRN3<\/jats:italic>\n                    and\n                    <jats:italic toggle=\"yes\">KLHL29<\/jats:italic>\n                    , that have not been associated with human gastric cancer.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2416723122","type":"journal-article","created":{"date-parts":[[2025,5,30]],"date-time":"2025-05-30T12:16:44Z","timestamp":1748607404000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":4,"title":["Genomic analyses identify 15 risk loci and reveal\n                    <i>HDAC2<\/i>\n                    ,\n                    <i>SOX2-OT<\/i>\n                    , and\n                    <i>IGF2BP2<\/i>\n                    in a naturally occurring canine model of gastric cancer"],"prefix":"10.1073","volume":"122","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7967-9607","authenticated-orcid":false,"given":"Shawna R.","family":"Cook","sequence":"first","affiliation":[{"name":"Baker Institute for Animal Health, College of Veterinary Medicine, Cornell University","place":["Ithaca"]},{"name":"Department of Biomedical Sciences, College of Veterinary Medicine, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sanne","family":"Hugen","sequence":"additional","affiliation":[{"name":"Expertisecentre of Genetics, Department of Clinical Sciences, Utrecht University","place":["Utrecht, The Netherlands"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6213-5013","authenticated-orcid":false,"given":"Jessica J.","family":"Hayward","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences, College of Veterinary Medicine, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3139-9849","authenticated-orcid":false,"given":"Thomas R.","family":"Famula","sequence":"additional","affiliation":[{"name":"Department of Animal Science, University of California","place":["Davis"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Janelle M.","family":"Belanger","sequence":"additional","affiliation":[{"name":"Department of Animal Science, University of California","place":["Davis"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elizabeth","family":"McNiel","sequence":"additional","affiliation":[{"name":"Cummings School of Veterinary Medicine, Tufts University","place":["Grafton"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0562-6964","authenticated-orcid":false,"given":"Hille","family":"Fieten","sequence":"additional","affiliation":[{"name":"Expertisecentre of Genetics, Department of Clinical Sciences, Utrecht University","place":["Utrecht, The Netherlands"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3945-2851","authenticated-orcid":false,"given":"Anita M.","family":"Oberbauer","sequence":"additional","affiliation":[{"name":"Department of Animal Science, University of California","place":["Davis"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter A. J.","family":"Leegwater","sequence":"additional","affiliation":[{"name":"Expertisecentre of Genetics, Department of Clinical Sciences, Utrecht University","place":["Utrecht, The Netherlands"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6075-9738","authenticated-orcid":false,"given":"Elaine A.","family":"Ostrander","sequence":"additional","affiliation":[{"name":"Cancer Genetics and Comparative Genomics Branch, National Human Genome Research Institute, National Institutes of Health","place":["Bethesda"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paul J. J.","family":"Mandigers","sequence":"additional","affiliation":[{"name":"Expertisecentre of Genetics, Department of Clinical Sciences, Utrecht University","place":["Utrecht, The Netherlands"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1272-0059","authenticated-orcid":false,"given":"Jacquelyn M.","family":"Evans","sequence":"additional","affiliation":[{"name":"Baker Institute for Animal Health, College of Veterinary Medicine, Cornell University","place":["Ithaca"]},{"name":"Department of Biomedical Sciences, College of Veterinary Medicine, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2025,5,30]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"publisher","DOI":"10.3322\/caac.21834"},{"key":"e_1_3_4_2_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0140-6736(17)33326-3"},{"key":"e_1_3_4_3_2","doi-asserted-by":"publisher","DOI":"10.1016\/S1470-2045(19)30456-5"},{"key":"e_1_3_4_4_2","first-page":"1","article-title":"Recent estimates and predictions of 5-year survival in patients with gastric cancer: A model-based period analysis","volume":"29","author":"Li Y.","year":"2022","unstructured":"Y. Li, A. Feng, S. Zheng, C. Chen, J. Lyu, Recent estimates and predictions of 5-year survival in patients with gastric cancer: A model-based period analysis. Cancer Control J. Moffitt Cancer Cent. 29, 1\u20139 (2022).","journal-title":"Cancer Control J. Moffitt Cancer Cent."},{"key":"e_1_3_4_5_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.annonc.2022.07.004"},{"key":"e_1_3_4_6_2","doi-asserted-by":"publisher","DOI":"10.1136\/gutjnl-2020-321600"},{"key":"e_1_3_4_7_2","doi-asserted-by":"publisher","DOI":"10.1111\/apm.1965.64.1.31"},{"key":"e_1_3_4_8_2","first-page":"3554","article-title":"A human model of gastric carcinogenesis","volume":"48","author":"Correa P.","year":"1988","unstructured":"P. Correa, A human model of gastric carcinogenesis. 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