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Adv."],"published-print":{"date-parts":[[2026,8,14]]},"abstract":"<jats:p>\n                    How did\n                    <jats:italic toggle=\"yes\">Homo sapiens<\/jats:italic>\n                    thrive in extreme environments during their dispersal across Eurasia? Unraveling adaptations that enabled human survival during the Last Glacial Maximum (LGM) remains critical. Here, we analyze 42 genomes from Holocene hunter-gatherers (Jomon) in the Japanese archipelago, revealing their deep ancestry from an isolated Upper Paleolithic (UP) East Eurasian lineage. We detect gene flow from northern UP Siberians into Jomon, reconciling long-standing debates between bioanthropology and archaeology regarding East Asian peopling. We find no Jomon-specific Denisovan ancestry, suggesting shared introgression before their LGM divergence from mainlanders. Furthermore, we find evidence consistent with cold adaptation in Jomon UP ancestry, highlighting diverse physiological pathways. Our study provides one of the first paleogenomic frameworks for investigating ancient adaptations in UP East Eurasians, revealing an evolutionary legacy from a colder past in present-day humans.\n                  <\/jats:p>","DOI":"10.1126\/sciadv.aea7469","type":"journal-article","created":{"date-parts":[[2026,8,12]],"date-time":"2026-08-12T18:01:07Z","timestamp":1786557667000},"update-policy":"https:\/\/doi.org\/10.34133\/aaas_crossmark","source":"Crossref","is-referenced-by-count":2,"title":["Jomon genomics reveal cold adaptation in Upper Paleolithic hunter-gatherers of eastern Eurasia"],"prefix":"10.1126","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4490-340X","authenticated-orcid":true,"given":"Yusuke","family":"Watanabe","sequence":"first","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yoshiki","family":"Wakiyama","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3949-9235","authenticated-orcid":true,"given":"Daisuke","family":"Waku","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."},{"name":"Department of International Agricultural Development, Faculty of International Agriculture and Food Studies, Tokyo University of Agriculture, Setagaya-ku, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6201-6030","authenticated-orcid":true,"given":"Guido","family":"Valverde","sequence":"additional","affiliation":[{"name":"Department of Anatomy, Kitasato University School of Medicine, Sagamihara, Kanagawa, Japan."},{"name":"Institute for Mummy Studies, Eurac Research, Bolzano, Italy."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Akio","family":"Tanino","sequence":"additional","affiliation":[{"name":"Department of Anatomy, Kitasato University School of Medicine, Sagamihara, Kanagawa, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-1540-2385","authenticated-orcid":true,"given":"Mizuki","family":"Hayashi","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."},{"name":"Department of Anatomy, Kitasato University School of Medicine, Sagamihara, Kanagawa, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Izumi","family":"Naka","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuka","family":"Nakamura","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tsubasa","family":"Suzuki","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."},{"name":"Department of Biosciences, Kitasato University School of Science, Sagamihara, Kanagawa, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6972-2218","authenticated-orcid":true,"given":"Kae","family":"Koganebuchi","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1855-8371","authenticated-orcid":true,"given":"Takashi","family":"Gakuhari","sequence":"additional","affiliation":[{"name":"Center for the Study of Ancient Civilizations and Cultural Resources, College of Human and Social Sciences, Kanazawa University, Kanazawa, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7192-150X","authenticated-orcid":true,"given":"Takafumi","family":"Katsumura","sequence":"additional","affiliation":[{"name":"Department of Anatomy, Kitasato University School of Medicine, Sagamihara, Kanagawa, Japan."},{"name":"Department of Human Life Design and Science, Faculty of Design, Kyushu University, Minami-Ku, Fukuoka, 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Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5791-7512","authenticated-orcid":true,"given":"Takayuki","family":"Nishimura","sequence":"additional","affiliation":[{"name":"Department of Human Life Design and Science, Faculty of Design, Kyushu University, Minami-Ku, Fukuoka, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tadayuki","family":"Masuyama","sequence":"additional","affiliation":[{"name":"Tahara City Board of Education, Tahara, Aichi, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4686-7243","authenticated-orcid":true,"given":"Yasuhiro","family":"Yamada","sequence":"additional","affiliation":[{"name":"Department of History and Archaeology, Tokyo Metropolitan University, Hachioji, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryuzaburo","family":"Takahashi","sequence":"additional","affiliation":[{"name":"Waseda University, Shinjuku-ku, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6598-1562","authenticated-orcid":true,"given":"Masami","family":"Izuho","sequence":"additional","affiliation":[{"name":"Department of History and Archaeology, Tokyo Metropolitan University, Hachioji, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jun","family":"Ohashi","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"name":"NCBN Controls WGS Consortium","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0746-2729","authenticated-orcid":true,"given":"Hiroki","family":"Oota","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Bunkyo-ku, Tokyo, Japan."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"221","reference":[{"key":"e_1_3_2_2_2","doi-asserted-by":"publisher","DOI":"10.3378\/027.085.0302"},{"key":"e_1_3_2_3_2","doi-asserted-by":"publisher","DOI":"10.1038\/nature23452"},{"key":"e_1_3_2_4_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.1137564"},{"key":"e_1_3_2_5_2","unstructured":"Y. Kaifu M. Izuho T. Goebel \u201cModern human dispersal and behavior in Paleolithic Asia: Summary and discussion \u201d in Emergence and Diversity of Modern Human Behavior in Paleolithic Asia Y. Kaifu M. Izuho T. Goebel H. Sato A. Ono Eds. (Texas A&M Univ. Press 2015) pp. 535\u2013566."},{"key":"e_1_3_2_6_2","doi-asserted-by":"crossref","unstructured":"R. Dennell From Arabia to the Pacific: How Our Species Colonised Asia (Routledge 2020).","DOI":"10.4324\/9781003038788"},{"key":"e_1_3_2_7_2","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-020-01162-2"},{"key":"e_1_3_2_8_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.aat3628"},{"key":"e_1_3_2_9_2","doi-asserted-by":"publisher","DOI":"10.2486\/indhealth.47.221"},{"key":"e_1_3_2_10_2","first-page":"51","article-title":"Periodization and classification of the Upper Paleolithic of Sakhalin and Hokkaido in the light of the research conducted at the Ogonki-5 site","volume":"3","author":"Vasilevsky A. A.","year":"2003","unstructured":"A. A. Vasilevsky, Periodization and classification of the Upper Paleolithic of Sakhalin and Hokkaido in the light of the research conducted at the Ogonki-5 site. Archaeol. Ethnol. Anthropol. Eurasia 3, 51\u201369 (2003).","journal-title":"Archaeol. Ethnol. Anthropol. Eurasia"},{"key":"e_1_3_2_11_2","unstructured":"M. Izuho \u201cHuman technological and behavioral adaptation to landscape changes around the Last Glacial Maximum in Japan: A focus on Hokkaido \u201d in Paleoamerican Odyssey K. E. Graf C. V. Ketron M. R. Waters Eds. (Texas A&M Univ. Press 2014) pp. 45\u201364."},{"key":"e_1_3_2_12_2","unstructured":"M. Izuho Y. Kaifu \u201cThe appearance and characteristics of the early Upper Palaeolithic in the Japanese Archipelago \u201d in Emergence and Diversity of Modern Human Behavior in Paleolithic Asia Y. Kaifu M. Izuho T. Goebel H. Sato A. Ono Eds. (Texas A&M Univ. Press 2015) pp. 289\u2013313."},{"key":"e_1_3_2_13_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.ara.2018.03.001"},{"key":"e_1_3_2_14_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.1172873"},{"key":"e_1_3_2_15_2","doi-asserted-by":"publisher","DOI":"10.1038\/jhg.2016.110"},{"key":"e_1_3_2_16_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-019-0484-x"},{"key":"e_1_3_2_17_2","doi-asserted-by":"publisher","DOI":"10.1080\/20555563.2018.1457392"},{"key":"e_1_3_2_18_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.quaint.2021.01.009"},{"key":"e_1_3_2_19_2","doi-asserted-by":"publisher","DOI":"10.15184\/aqy.2015.23"},{"key":"e_1_3_2_20_2","doi-asserted-by":"publisher","DOI":"10.1007\/s41982-024-00175-1"},{"key":"e_1_3_2_21_2","first-page":"122","article-title":"The nature of the transition from Late Upper Paleolithic to Incipient Jomon in the Japanese Archipelago: An archaeological perspective","volume":"7","author":"Izuho M.","year":"2023","unstructured":"M. Izuho, The nature of the transition from Late Upper Paleolithic to Incipient Jomon in the Japanese Archipelago: An archaeological perspective. Anthropol. Pap. Univ. Alaska 7, 122\u2013135 (2023).","journal-title":"Anthropol. Pap. Univ. Alaska"},{"key":"e_1_3_2_22_2","doi-asserted-by":"publisher","DOI":"10.1080\/20555563.2018.1563406"},{"key":"e_1_3_2_23_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.quascirev.2020.106605"},{"key":"e_1_3_2_24_2","doi-asserted-by":"crossref","unstructured":"I. Buvit K. Terry M. Izuho \u201cPathways along the Pacific: Using early stone tools to. Reconstruct coastal migration between Japan and the Americas \u201d in Palaeolandscapes in Archaeology M. T. Carson Ed. (Routledge 2021) pp. 39\u201381.","DOI":"10.4324\/9781003139553-4"},{"key":"e_1_3_2_25_2","doi-asserted-by":"publisher","DOI":"10.1002\/gea.21477"},{"key":"e_1_3_2_26_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.quaint.2016.02.050"},{"key":"e_1_3_2_27_2","unstructured":"K. Morisaki \u201cAppearance of Hakuhen-Sentoki (HS points) and second modern human migration into Kyushu Japan \u201d in Emergence and Diversity of Modern Human Behavior in Paleolithic Asia Y. Kaifu M. Izuho T. Gobel H. Sato Eds. (Texas A&M Univ. 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