{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T08:07:13Z","timestamp":1788941233596,"version":"build-2803163510"},"reference-count":61,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T00:00:00Z","timestamp":1788912000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell Dev. Biol."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>Extraembryonic lineages, specifically the trophectoderm (TE) and primitive endoderm (PrE), are indispensable for early embryogenesis and successful pregnancy. Expanded potential stem cells (EPSCs) can contribute to both embryonic and extraembryonic fates, providing a platform to model early lineage specification.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>This study investigated the effects of fibroblast growth factor 2 (FGF2) and bone morphogenetic protein 4 (BMP4) (2F condition) on lineage-associated priming in porcine EPSCs (pEPSCs).<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Following 2F treatment, cells maintained a normal karyotype and underwent morphological changes accompanied by downregulation of pluripotency markers (\n                      <jats:italic>OCT4<\/jats:italic>\n                      ,\n                      <jats:italic>SOX2<\/jats:italic>\n                      ) and upregulation of TE (\n                      <jats:italic>CDX2<\/jats:italic>\n                      and\n                      <jats:italic>GATA3<\/jats:italic>\n                      ) and PrE markers (\n                      <jats:italic>GATA4<\/jats:italic>\n                      ,\n                      <jats:italic>GATA6<\/jats:italic>\n                      ,\n                      <jats:italic>SOX17<\/jats:italic>\n                      , and\n                      <jats:italic>HNF4A<\/jats:italic>\n                      ). Transcriptomic and immunofluorescence analyses consistently demonstrated activation of TE- and PrE-associated molecular programs, although markers of additional developmental lineages were also detected. These findings suggest that combined FGF2 and BMP4 treatment promotes extraembryonic lineage-associated priming in pEPSCs, and generates a heterogeneous population exhibiting features of both TE- and PrE-associated states.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion<\/jats:title>\n                    <jats:p>\n                      Collectively, this study establishes a simple and defined\n                      <jats:italic>in vitro<\/jats:italic>\n                      platform for investigating signaling-dependent lineage priming and cell state remodeling in porcine EPSCs.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fcell.2026.1909585","type":"journal-article","created":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T07:45:01Z","timestamp":1788939901000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["BMP4 and FGF2 promote heterogeneous priming including extraembryonic lineages in porcine expanded potential stem cells"],"prefix":"10.3389","volume":"14","author":[{"given":"Haneul","family":"Kim","sequence":"first","affiliation":[{"name":"Laboratory of Molecular Diagnostics and Cell Biology, College of Veterinary Medicine, Gyeongsang National University","place":["Jinju, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lian","family":"Cai","sequence":"additional","affiliation":[{"name":"Laboratory of Veterinary Embryology and Biotechnology, Veterinary Medical Center and College of Veterinary Medicine, Chungbuk National University","place":["Cheongju, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sang-Hwan","family":"Hyun","sequence":"additional","affiliation":[{"name":"Laboratory of Veterinary Embryology and Biotechnology, Veterinary Medical Center and College of Veterinary Medicine, Chungbuk National University","place":["Cheongju, Republic of Korea"]},{"name":"Vet-ICT Convergence Education and Research Center (VICERC), Chungbuk National University","place":["Cheongju, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eunhye","family":"Kim","sequence":"additional","affiliation":[{"name":"Laboratory of Molecular Diagnostics and Cell Biology, College of Veterinary Medicine, Gyeongsang National University","place":["Jinju, Republic of Korea"]},{"name":"Vet-ICT Convergence Education and Research Center (VICERC), Chungbuk National University","place":["Cheongju, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,9]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1093\/bioinformatics\/btu638","article-title":"HTSeq\u2014a python framework to work with high-throughput sequencing data","volume":"31","author":"Anders","year":"2015","journal-title":"Bioinformatics"},{"key":"B2","doi-asserted-by":"publisher","first-page":"1341","DOI":"10.1038\/s41556-022-00984-y","article-title":"Stem cell-derived synthetic embryos self-assemble by exploiting cadherin codes and cortical tension","volume":"24","author":"Bao","year":"2022","journal-title":"Nat. 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