{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T23:06:03Z","timestamp":1791846363843,"version":"4.3.4"},"reference-count":107,"publisher":"Oxford University Press (OUP)","issue":"5","license":[{"start":{"date-parts":[[2016,11,1]],"date-time":"2016-11-01T00:00:00Z","timestamp":1477958400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"content-domain":{"domain":["rep.bioscientifica.com"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2016,11,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Human life begins with sperm and oocyte fusion. After fertilization, various fusion events occur during human embryogenesis and morphogenesis. For example, the fusion of trophoblastic cells constitutes a key process for normal placental development. Fusion in the placenta is facilitated by syncytin 1 and syncytin 2. These syncytins arose from retroviral sequences that entered the primate genome 25 million and more than 40 million years ago respectively. About 8% of the human genome consists of similar human endogenous retroviral (HERVs) sequences. Many are inactive because of mutations or deletions. However, the role of the few that remain transcriptionally active has not been fully elucidated. Syncytin proteins maintain cell\u2013cell fusogenic activity based on env gene-mediated viral cell entry. In this review, we summarize how syncytins and their receptors are involved in fusion events during human reproduction. The significance of syncytins in tumorigenesis is also discussed.<\/jats:p>","DOI":"10.1530\/rep-16-0031","type":"journal-article","created":{"date-parts":[[2016,8,2]],"date-time":"2016-08-02T21:33:07Z","timestamp":1470173587000},"page":"R167-R178","update-policy":"https:\/\/doi.org\/10.1530\/crossmarkpolicy-1","source":"Crossref","is-referenced-by-count":60,"title":["The role of syncytins in human reproduction and reproductive organ cancers"],"prefix":"10.1093","volume":"152","author":[{"given":"Bikem","family":"Soygur","sequence":"first","affiliation":[{"name":"Department of Histology and Embryology Akdeniz University School of Medicine, Antalya, Turkey"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leyla","family":"Sati","sequence":"additional","affiliation":[{"name":"Department of Histology and Embryology Akdeniz University School of Medicine, Antalya, Turkey"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2016,11,1]]},"reference":[{"key":"2026010712452523600_bib1","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1016\/0020-7292(90)90350-T","article-title":"Perinatal morbidity and mortality in pregnancy hypertensive disorders: prognostic value of the clinical and laboratory findings","volume":"32","author":"Acien","year":"1990","journal-title":"International Journal of Gynecology and Obstetrics"},{"key":"2026010712452523600_bib2","doi-asserted-by":"crossref","DOI":"10.1007\/978-0-387-88441-7","volume-title":"AJCC Cancer Staging Manual","author":"American Joint Committee on Cancer","year":"2010"},{"key":"2026010712452523600_bib3","doi-asserted-by":"crossref","first-page":"826","DOI":"10.1096\/fasebj.7.10.8344482","article-title":"Tumor suppressor genes studied by cell hybridization and chromosome transfer","volume":"7","author":"Anderson","year":"1993","journal-title":"FASEB Journal"},{"key":"2026010712452523600_bib4","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1038\/77076","article-title":"The glial cells missing-1 protein is essential for branching morphogenesis in the chorioallantoic placenta","volume":"25","author":"Anson-Cartwright","year":"2000","journal-title":"Nature Genetics"},{"key":"2026010712452523600_bib5","doi-asserted-by":"crossref","DOI":"10.1007\/978-1-4757-4199-5","author":"Benirschke","year":"2000","journal-title":"Pathology of the Human Placenta"},{"key":"2026010712452523600_bib6","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1111\/j.1447-0756.2008.00719.x","article-title":"Surface area of chorionic villi of placentas: an index of intrauterine growth restriction of fetuses","volume":"34","author":"Biswas","year":"2008","journal-title":"Journal of Obstetrics and Gynaecology Research"},{"key":"2026010712452523600_bib7","doi-asserted-by":"crossref","first-page":"1906","DOI":"10.1007\/s00018-006-6201-9","article-title":"Syncytin is involved in breast cancer-endothelial cell fusions","volume":"63","author":"Bjerregaard","year":"2006","journal-title":"Cellular and Molecular Life Sciences"},{"key":"2026010712452523600_bib8","doi-asserted-by":"crossref","first-page":"533","DOI":"10.1007\/s10815-014-0224-1","article-title":"Syncytin-1 and its receptor is present in human gametes","volume":"31","author":"Bjerregaard","year":"2014","journal-title":"Journal of Assisted Reproduction and Genetics"},{"key":"2026010712452523600_bib9","doi-asserted-by":"crossref","first-page":"13013","DOI":"10.1073\/pnas.2132646100","article-title":"Genomewide screening for fusogenic human endogenous retrovirus envelopes identifies syncytin 2, a gene conserved on primate evolution","volume":"100","author":"Blaise","year":"2003","journal-title":"PNAS"},{"key":"2026010712452523600_bib10","doi-asserted-by":"crossref","first-page":"3321","DOI":"10.1128\/JVI.74.7.3321-3329.2000","article-title":"An envelope glycoprotein of the human endogenous retrovirus HERV-W is expressed in the human placenta and fuses cells expressing the type D mammalian retrovirus receptor","volume":"74","author":"Blond","year":"2000","journal-title":"Journal of Virology"},{"key":"2026010712452523600_bib11","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1530\/REP-10-0294","article-title":"Remodelling at the maternal-fetal interface: relevance to human pregnancy disorders","volume":"140","author":"Cartwright","year":"2010","journal-title":"Reproduction"},{"key":"2026010712452523600_bib12","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1111\/j.1471-0528.2005.00843.x","article-title":"Altered placental syncytin and its receptor ASCT2 expression in placental development and pre-eclampsia","volume":"113","author":"Chen","year":"2006","journal-title":"BJOG"},{"key":"2026010712452523600_bib13","doi-asserted-by":"crossref","first-page":"815","DOI":"10.1095\/biolreprod.108.069765","article-title":"Functional characterization of the human placental fusogenic membrane protein syncytin 2","volume":"79","author":"Chen","year":"2008","journal-title":"Biology of Reproduction"},{"key":"2026010712452523600_bib14","doi-asserted-by":"crossref","first-page":"17411","DOI":"10.1074\/jbc.M109.016170","article-title":"Mechanism of hypoxia-induced GCM1 degradation: implications for the pathogenesis of preeclampsia","volume":"284","author":"Chiang","year":"2009","journal-title":"Journal of Biological Chemistry"},{"key":"2026010712452523600_bib15","doi-asserted-by":"crossref","first-page":"1857","DOI":"10.1126\/science.281.5384.1857","article-title":"Fertilization defects in sperm from mice lacking fertilin beta","volume":"281","author":"Cho","year":"1998","journal-title":"Science"},{"key":"2026010712452523600_bib16","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1080\/00016340801908783","article-title":"Placental pathology in women with gestational diabetes","volume":"87","author":"Daskalakis","year":"2008","journal-title":"Acta Obstetricia et Gynecologica Scandinavica"},{"key":"2026010712452523600_bib17","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1159\/000084964","article-title":"Human endogenous retroviruses: from infectious elements to human genes","volume":"110","author":"de Parseval","year":"2005","journal-title":"Cytogenetic and Genome Research"},{"key":"2026010712452523600_bib18","doi-asserted-by":"crossref","first-page":"10414","DOI":"10.1128\/JVI.77.19.10414-10422.2003","article-title":"Survey of human genes of retroviral origin: identification and transcriptome of the genes with coding capacity for complete envelope proteins","volume":"77","author":"de Parseval","year":"2003","journal-title":"Journal of Virology"},{"key":"2026010712452523600_bib19","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1186\/1471-2164-6-117","article-title":"Comprehensive search for intra- and inter-specific sequence polymorphisms among coding envelope genes of retroviral origin found in the human genome: genes and pseudogenes","volume":"6","author":"de Parseval","year":"2005","journal-title":"BMC Genomics"},{"key":"2026010712452523600_bib20","article-title":"Endogenous retroviruses","volume-title":"Retroviruses Molecular Biology, Genomics and Pathogenesis","author":"Denner","year":"2010"},{"key":"2026010712452523600_bib21","doi-asserted-by":"crossref","first-page":"646","DOI":"10.1016\/j.coviro.2013.08.005","article-title":"Endogenous retroviruses: acquisition, amplification and taming of genome invaders","volume":"3","author":"Dewannieux","year":"2013","journal-title":"Current Opinion in Virology"},{"key":"2026010712452523600_bib22","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1016\/S1535-6108(03)00114-4","article-title":"Cell fusion: a hidden enemy?","volume":"3","author":"Duelli","year":"2003","journal-title":"Cancer Cell"},{"key":"2026010712452523600_bib23","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1073\/pnas.0406509102","article-title":"Syncytin-A and syncytin-B, two fusogenic placenta-specific murine envelope genes of retroviral origin conserved in Muridae","volume":"102","author":"Dupressoir","year":"2005","journal-title":"PNAS"},{"key":"2026010712452523600_bib24","doi-asserted-by":"crossref","first-page":"12127","DOI":"10.1073\/pnas.0902925106","article-title":"Syncytin-A knockout mice demonstrate the critical role in placentation of a fusogenic, endogenous retrovirus-derived, envelope gene","volume":"106","author":"Dupressoir","year":"2009","journal-title":"PNAS"},{"key":"2026010712452523600_bib25","doi-asserted-by":"crossref","first-page":"E1164","DOI":"10.1073\/pnas.1112304108","article-title":"A pair of co-opted retroviral envelope syncytin genes is required for formation of the two-layered murine placental syncytiotrophoblast","volume":"108","author":"Dupressoir","year":"2011","journal-title":"PNAS"},{"key":"2026010712452523600_bib26","doi-asserted-by":"crossref","first-page":"17532","DOI":"10.1073\/pnas.0807413105","article-title":"A placenta-specific receptor for the fusogenic, endogenous retrovirus-derived, human syncytin-2","volume":"105","author":"Esnault","year":"2008","journal-title":"PNAS"},{"key":"2026010712452523600_bib27","doi-asserted-by":"crossref","first-page":"e1003400","DOI":"10.1371\/journal.pgen.1003400","article-title":"Differential evolutionary fate of an ancestral primate endogenous retrovirus envelope gene, the EnvV syncytin, captured for a function in placentation","volume":"9","author":"Esnault","year":"2013","journal-title":"PLoS Genetics"},{"key":"2026010712452523600_bib28","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1146\/annurev-physiol-020911-153339","article-title":"Sperm-egg interaction","volume":"74","author":"Evans","year":"2012","journal-title":"Annual Review of Physiology"},{"key":"2026010712452523600_bib29","doi-asserted-by":"crossref","first-page":"3566","DOI":"10.1128\/MCB.23.10.3566-3574.2003","article-title":"Direct involvement of HERV-W Env glycoprotein in human trophoblast cell fusion and differentiation","volume":"23","author":"Frendo","year":"2003","journal-title":"Molecular and Cellular Biology"},{"key":"2026010712452523600_bib30","doi-asserted-by":"crossref","first-page":"2229","DOI":"10.1093\/nar\/gkp1214","article-title":"Custom human endogenous retroviruses dedicated microarray identifies self-induced HERV-W family elements reactivated in testicular cancer upon methylation control","volume":"38","author":"Gimenez","year":"2010","journal-title":"Nucleic Acids Research"},{"key":"2026010712452523600_bib31","doi-asserted-by":"crossref","DOI":"10.1186\/gb-2001-2-6-reviews1017","article-title":"Endogenous retroviruses in the human genome sequence","volume":"2","author":"Griffiths","year":"2001","journal-title":"Genome Biology"},{"key":"2026010712452523600_bib32","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1002\/(SICI)1521-1878(199804)20:4<307::AID-BIES7>3.0.CO;2-M","article-title":"Placental endogenous retrovirus (ERV): structural, functional, and evolutionary significance","volume":"20","author":"Harris","year":"1998","journal-title":"BioEssays"},{"key":"2026010712452523600_bib33","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1038\/223363a0","article-title":"Suppression of malignancy by cell fusion","volume":"223","author":"Harris","year":"1969","journal-title":"Nature"},{"key":"2026010712452523600_bib34","doi-asserted-by":"crossref","first-page":"460","DOI":"10.1016\/j.placenta.2012.02.012","article-title":"Syncytin 1 in the human placenta","volume":"33","author":"Holder","year":"2012","journal-title":"Placenta"},{"key":"2026010712452523600_bib35","doi-asserted-by":"crossref","first-page":"1027","DOI":"10.1016\/j.cellsig.2013.01.008","article-title":"Syncytin-1 modulates placental trophoblast cell proliferation by promoting G1\/S transition","volume":"25","author":"Huang","year":"2013","journal-title":"Cellular Signalling"},{"key":"2026010712452523600_bib36","doi-asserted-by":"crossref","first-page":"648","DOI":"10.1016\/j.cellsig.2013.11.002","article-title":"Epigenetic and non-epigenetic regulation of syncytin-1 expression in human placenta and cancer tissues","volume":"26","author":"Huang","year":"2014","journal-title":"Cellular Signalling"},{"key":"2026010712452523600_bib37","doi-asserted-by":"crossref","first-page":"3151","DOI":"10.1007\/s00018-013-1533-8","article-title":"Reduced syncytin-1 expression in choriocarcinoma BeWo cells activates the calpain1-AIF-mediated apoptosis, implication for preeclampsia","volume":"71","author":"Huang","year":"2014","journal-title":"Cellular and Molecular Life Sciences"},{"key":"2026010712452523600_bib38","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1111\/gtc.12278","article-title":"Baton pass hypothesis: successive incorporation of unconserved endogenous retroviral genes for placentation during mammalian evolution","volume":"20","author":"Imakawa","year":"2015","journal-title":"Genes to Cells"},{"key":"2026010712452523600_bib39","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1038\/nature03362","article-title":"The immunoglobulin superfamily protein Izumo is required for sperm to fuse with eggs","volume":"434","author":"Inoue","year":"2005","journal-title":"Nature"},{"key":"2026010712452523600_bib40","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1067\/mob.2002.119176","article-title":"Increased apoptosis in the syncytiotrophoblast in human term placentas complicated by either preeclampsia or intrauterine growth retardation","volume":"186","author":"Ishihara","year":"2002","journal-title":"American Journal of Obstetrics and Gynecology"},{"key":"2026010712452523600_bib41","first-page":"1552","article-title":"Fragments of target cells are internalized into retroviral envelope protein-expressing cells during cell-cell fusion by endocytosis","volume":"6","author":"Izumida","year":"2015","journal-title":"Frontiers in Microbiology"},{"key":"2026010712452523600_bib42","volume-title":"Histology and Cell Biology: An Introduction to Pathology","author":"Kierszenbaum","year":"2002"},{"key":"2026010712452523600_bib43","article-title":"Retroviruses and cell fusion: overview","volume-title":"Cell Fusions Regulation and Control","author":"Kjeldbjerg","year":"2010"},{"key":"2026010712452523600_bib44","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1067\/mob.2002.119636","article-title":"Syncytin, a novel human endogenous retroviral gene in human placenta: evidence for its dysregulation in preeclampsia and HELLP syndrome","volume":"186","author":"Knerr","year":"2002","journal-title":"American Journal of Obstetrics and Gynecology"},{"key":"2026010712452523600_bib45","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1007\/s10495-006-0329-9","article-title":"Fusiogenic endogenous-retroviral syncytin-1 exerts anti-apoptotic functions in staurosporine-challenged CHO cells","volume":"12","author":"Knerr","year":"2007","journal-title":"Apoptosis"},{"key":"2026010712452523600_bib46","doi-asserted-by":"crossref","first-page":"766","DOI":"10.1002\/jcb.21874","article-title":"Response of HEK293 and CHO cells overexpressing fusiogenic syncytin-1 to mitochondrion-mediated apoptosis induced by antimycin A","volume":"105","author":"Knerr","year":"2008","journal-title":"Journal of Cellular Biochemistry"},{"key":"2026010712452523600_bib47","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1038\/256495a0","article-title":"Continuous cultures of fused cells secreting antibody of predefined specificity","volume":"256","author":"Kohler","year":"1975","journal-title":"Nature"},{"key":"2026010712452523600_bib48","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1016\/j.placenta.2007.11.009","article-title":"Cellular localization of placenta-specific human endogenous retrovirus (HERV) transcripts and their possible implication in pregnancy-induced hypertension","volume":"29","author":"Kudaka","year":"2008","journal-title":"Placenta"},{"key":"2026010712452523600_bib49","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/S0925-4439(03)00043-7","article-title":"Hypoxia alters expression and function of syncytin and its receptor during trophoblast cell fusion of human placental BeWo cells: implications for impaired trophoblast syncytialisation in pre-eclampsia","volume":"1638","author":"Kudo","year":"2003","journal-title":"Biochimica et Biophysica Acta"},{"key":"2026010712452523600_bib50","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1002\/mrd.20729","article-title":"Impaired cytotrophoblast cell-cell fusion is associated with reduced Syncytin and increased apoptosis in patients with placental dysfunction","volume":"75","author":"Langbein","year":"2008","journal-title":"Molecular Reproduction and Development"},{"key":"2026010712452523600_bib51","doi-asserted-by":"crossref","first-page":"1193","DOI":"10.1100\/tsw.2007.212","article-title":"Syncytin and cancer cell fusions","volume":"7","author":"Larsson","year":"2007","journal-title":"Scientific World Journal"},{"key":"2026010712452523600_bib52","doi-asserted-by":"crossref","first-page":"726","DOI":"10.1016\/j.humpath.2006.10.018","article-title":"Prognostic role of syncytin expression in breast cancer","volume":"38","author":"Larsson","year":"2007","journal-title":"Human Pathology"},{"key":"2026010712452523600_bib53","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1007\/s00418-008-0411-1","article-title":"Cell fusions in mammals","volume":"129","author":"Larsson","year":"2008","journal-title":"Histochemistry and Cell Biology"},{"key":"2026010712452523600_bib54","doi-asserted-by":"crossref","first-page":"20120507","DOI":"10.1098\/rstb.2012.0507","article-title":"Paleovirology of \u2018syncytins\u2019, retroviral env genes exapted for a role in placentation","volume":"368","author":"Lavialle","year":"2013","journal-title":"Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences"},{"key":"2026010712452523600_bib55","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1126\/science.287.5451.319","article-title":"Severely reduced female fertility in CD9-deficient mice","volume":"287","author":"Le Naour","year":"2000","journal-title":"Science"},{"key":"2026010712452523600_bib56","doi-asserted-by":"crossref","first-page":"808","DOI":"10.1053\/plac.2001.0722","article-title":"Downregulation of placental syncytin expression and abnormal protein localization in pre-eclampsia","volume":"22","author":"Lee","year":"2001","journal-title":"Placenta"},{"key":"2026010712452523600_bib57","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1095\/biolreprod.110.083915","article-title":"GCM1 regulation of the expression of syncytin 2 and its cognate receptor MFSD2A in human placenta","volume":"83","author":"Liang","year":"2010","journal-title":"Biology of Reproduction"},{"key":"2026010712452523600_bib58","doi-asserted-by":"crossref","first-page":"5177","DOI":"10.1073\/pnas.93.11.5177","article-title":"The viruses in all of us: characteristics and biological significance of human endogenous retrovirus sequences","volume":"93","author":"Lower","year":"1996","journal-title":"PNAS"},{"key":"2026010712452523600_bib59","doi-asserted-by":"crossref","DOI":"10.1128\/microbiolspec.MDNA3-0009-2014","article-title":"Mammalian endogenous retroviruses","volume":"3","author":"Mager","year":"2015","journal-title":"Microbiology Spectrum"},{"key":"2026010712452523600_bib60","doi-asserted-by":"crossref","first-page":"556","DOI":"10.1016\/j.placenta.2004.09.002","article-title":"Expression of HERV-W Env glycoprotein (syncytin) in the extravillous trophoblast of first trimester human placenta","volume":"26","author":"Malassine","year":"2005","journal-title":"Placenta"},{"key":"2026010712452523600_bib61","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1016\/j.placenta.2006.03.001","article-title":"Expression of the fusogenic HERV-FRD Env glycoprotein (syncytin 2) in human placenta is restricted to villous cytotrophoblastic cells","volume":"28","author":"Malassine","year":"2007","journal-title":"Placenta"},{"key":"2026010712452523600_bib62","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1186\/1742-4690-5-6","article-title":"Human endogenous retrovirus-FRD envelope protein (syncytin 2) expression in normal and trisomy 21-affected placenta","volume":"5","author":"Malassine","year":"2008","journal-title":"Retrovirology"},{"key":"2026010712452523600_bib63","doi-asserted-by":"crossref","first-page":"e76281","DOI":"10.1371\/journal.pone.0076281","article-title":"Expression of human endogenous retrovirus-w including syncytin-1 in cutaneous T-cell lymphoma","volume":"8","author":"Maliniemi","year":"2013","journal-title":"PLoS ONE"},{"key":"2026010712452523600_bib64","doi-asserted-by":"crossref","first-page":"20534","DOI":"10.1073\/pnas.0707873105","article-title":"Placental syncytins: genetic disjunction between the fusogenic and immunosuppressive activity of retroviral envelope proteins","volume":"104","author":"Mangeney","year":"2007","journal-title":"PNAS"},{"key":"2026010712452523600_bib65","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1186\/1476-4598-3-12","article-title":"L1 and HERV-W retrotransposons are hypomethylated in human ovarian carcinomas","volume":"3","author":"Menendez","year":"2004","journal-title":"Molecular Cancer"},{"key":"2026010712452523600_bib66","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1038\/35001608","article-title":"Syncytin is a captive retroviral envelope protein involved in human placental morphogenesis","volume":"403","author":"Mi","year":"2000","journal-title":"Nature"},{"key":"2026010712452523600_bib67","first-page":"556","article-title":"Human endogenous retroviral syncytin exerts inhibitory effect on invasive phenotype of B16F10 melanoma cells","volume":"25","author":"Mo","year":"2013","journal-title":"Chinese Journal of Cancer Research"},{"key":"2026010712452523600_bib68","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1067\/mob.2002.127458","article-title":"Birth weight and fetal death in the United States: the effect of maternal diabetes during pregnancy","volume":"187","author":"Mondestin","year":"2002","journal-title":"American Journal of Obstetrics and Gynecology"},{"key":"2026010712452523600_bib69","doi-asserted-by":"crossref","first-page":"2125","DOI":"10.1007\/s00018-004-4200-2","article-title":"Spontaneous fusion between cancer cells and endothelial cells","volume":"61","author":"Mortensen","year":"2004","journal-title":"Cellular and Molecular Life Sciences"},{"key":"2026010712452523600_bib70","doi-asserted-by":"crossref","first-page":"2067","DOI":"10.1099\/vir.0.81412-0","article-title":"Human endogenous retrovirus-W envelope (syncytin) is expressed in both villous and extravillous trophoblast populations","volume":"87","author":"Muir","year":"2006","journal-title":"Journal of General Virology"},{"key":"2026010712452523600_bib71","doi-asserted-by":"crossref","first-page":"945","DOI":"10.1530\/REP-09-0082","article-title":"CD9 regulates transcription factor GCM1 and ERVWE1 expression through the cAMP\/protein kinase A signaling pathway","volume":"138","author":"Muroi","year":"2009","journal-title":"Reproduction"},{"key":"2026010712452523600_bib72","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1093\/humupd\/dmq052","article-title":"Epigenetics and the placenta","volume":"17","author":"Nelissen","year":"2011","journal-title":"Human Reproduction Update"},{"key":"2026010712452523600_bib73","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1023\/A:1008004332513","article-title":"Expression of envelope proteins of endogeneous C-type retrovirus on the surface of mouse and human oocytes at fertilization","volume":"18","author":"Nilsson","year":"1999","journal-title":"Virus Genes"},{"key":"2026010712452523600_bib74","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1006\/dbio.2001.0166","article-title":"Analysis of loss of adhesive function in sperm lacking cyritestin or fertilin beta","volume":"233","author":"Nishimura","year":"2001","journal-title":"Developmental Biology"},{"key":"2026010712452523600_bib75","doi-asserted-by":"crossref","first-page":"1805","DOI":"10.1007\/s00018-007-6552-x","article-title":"New insights into the molecular mechanisms of sperm-egg interaction","volume":"64","author":"Nixon","year":"2007","journal-title":"Cellular and Molecular Life Sciences"},{"key":"2026010712452523600_bib76","first-page":"1","article-title":"Maternal diabetes in pregnancy","volume-title":"Diabetes in America","author":"O\u2019Sullivan","year":"1985"},{"key":"2026010712452523600_bib77","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1159\/000084967","article-title":"Endogenous retroviruses and animal reproduction","volume":"110","author":"Prudhomme","year":"2005","journal-title":"Cytogenetic and Genome Research"},{"key":"2026010712452523600_bib78","doi-asserted-by":"crossref","first-page":"478","DOI":"10.1016\/S0167-5699(97)01127-4","article-title":"Th1-type immunity is incompatible with successful pregnancy","volume":"18","author":"Raghupathy","year":"1997","journal-title":"Immunology Today"},{"key":"2026010712452523600_bib79","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1146\/annurev.cellbio.24.110707.175418","article-title":"The evolution, regulation, and function of placenta-specific genes","volume":"24","author":"Rawn","year":"2008","journal-title":"Annual Review of Cell and Developmental Biology"},{"key":"2026010712452523600_bib80","doi-asserted-by":"crossref","first-page":"673","DOI":"10.1016\/j.placenta.2004.02.008","article-title":"The role of human endogenous retroviruses in trophoblast differentiation and placental development","volume":"25","author":"Rote","year":"2004","journal-title":"Placenta"},{"key":"2026010712452523600_bib81","doi-asserted-by":"crossref","first-page":"935","DOI":"10.4161\/epi.25578","article-title":"Gestational diabetes mellitus epigenetically affects genes predominantly involved in metabolic diseases","volume":"8","author":"Ruchat","year":"2013","journal-title":"Epigenetics"},{"key":"2026010712452523600_bib82","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1007\/s00109-010-0656-8","article-title":"Impaired cell fusion and differentiation in placentae from patients with intrauterine growth restriction correlate with reduced levels of HERV envelope genes","volume":"88","author":"Ruebner","year":"2010","journal-title":"Journal of Molecular Medicine"},{"key":"2026010712452523600_bib83","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1095\/biolreprod.102.013078","article-title":"Temporal regulation of the expression of syncytin (HERV-W), maternally imprinted PEG10, and SGCE in human placenta","volume":"69","author":"Smallwood","year":"2003","journal-title":"Biology of Reproduction"},{"key":"2026010712452523600_bib84","first-page":"1037","article-title":"Altered expression of human endogenous retroviruses syncytin-1, syncytin-2 and their receptors in human normal and gestational diabetic placenta","volume":"31","author":"Soygur","year":"2016","journal-title":"Histol Histopathol"},{"key":"2026010712452523600_bib85","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1038\/nrmicro2783","article-title":"Studies of endogenous retroviruses reveal a continuing evolutionary saga","volume":"10","author":"Stoye","year":"2012","journal-title":"Nature Reviews Microbiology"},{"key":"2026010712452523600_bib86","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1007\/s00109-006-0104-y","article-title":"Proliferation and cell-cell fusion of endometrial carcinoma are induced by the human endogenous retroviral Syncytin-1 and regulated by TGF-beta","volume":"85","author":"Strick","year":"2007","journal-title":"Journal of Molecular Medicine"},{"key":"2026010712452523600_bib87","doi-asserted-by":"crossref","first-page":"1204","DOI":"10.18632\/oncotarget.679","article-title":"Reactivation of codogenic endogenous retroviral (ERV) envelope genes in human endometrial carcinoma and prestages: emergence of new molecular targets","volume":"3","author":"Strissel","year":"2012","journal-title":"Oncotarget"},{"key":"2026010712452523600_bib88","doi-asserted-by":"crossref","first-page":"1462","DOI":"10.1038\/srep01462","article-title":"A novel human endogenous retroviral protein inhibits cell-cell fusion","volume":"3","author":"Sugimoto","year":"2013","journal-title":"Scientific Reports"},{"key":"2026010712452523600_bib89","doi-asserted-by":"crossref","first-page":"933","DOI":"10.1016\/j.placenta.2012.08.004","article-title":"The endogenous retroviral envelope protein syncytin-1 inhibits LPS\/PHA-stimulated cytokine responses in human blood and is sorted into placental exosomes","volume":"33","author":"Tolosa","year":"2012","journal-title":"Placenta"},{"key":"2026010712452523600_bib90","doi-asserted-by":"crossref","first-page":"e0121468","DOI":"10.1371\/journal.pone.0121468","article-title":"A CRE\/AP-1-like motif is essential for induced syncytin-2 expression and fusion in human trophoblast-like model","volume":"10","author":"Toufaily","year":"2015","journal-title":"PLoS ONE"},{"key":"2026010712452523600_bib91","doi-asserted-by":"crossref","first-page":"8728","DOI":"10.1093\/nar\/gkr562","article-title":"Epigenetic regulation of transcription and splicing of syncytins, fusogenic glycoproteins of retroviral origin","volume":"39","author":"Trejbalova","year":"2011","journal-title":"Nucleic Acids Research"},{"key":"2026010712452523600_bib92","doi-asserted-by":"crossref","first-page":"654","DOI":"10.1038\/ncb1596","article-title":"Exosome-mediated transfer of mRNAs and microRNAs is a novel mechanism of genetic exchange between cells","volume":"9","author":"Valadi","year":"2007","journal-title":"Nature Cell Biology"},{"key":"2026010712452523600_bib93","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1016\/j.jmb.2009.07.025","article-title":"Syncytin-2 plays an important role in the fusion of human trophoblast cells","volume":"392","author":"Vargas","year":"2009","journal-title":"Journal of Molecular Biology"},{"key":"2026010712452523600_bib94","doi-asserted-by":"crossref","first-page":"1085","DOI":"10.1177\/1933719111404608","article-title":"Reduced expression of both syncytin 1 and syncytin 2 correlates with severity of preeclampsia","volume":"18","author":"Vargas","year":"2011","journal-title":"Reproductive Sciences"},{"key":"2026010712452523600_bib95","doi-asserted-by":"crossref","first-page":"3703","DOI":"10.1096\/fj.13-239053","article-title":"Syncytin proteins incorporated in placenta exosomes are important for cell uptake and show variation in abundance in serum exosomes from patients with preeclampsia","volume":"28","author":"Vargas","year":"2014","journal-title":"FASEB Journal"},{"key":"2026010712452523600_bib96","doi-asserted-by":"crossref","first-page":"901","DOI":"10.1038\/nature01539","article-title":"Transplanted bone marrow regenerates liver by cell fusion","volume":"422","author":"Vassilopoulos","year":"2003","journal-title":"Nature"},{"key":"2026010712452523600_bib97","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1186\/1742-4690-1-32","article-title":"Identification of endogenous retroviral reading frames in the human genome","volume":"1","author":"Villesen","year":"2004","journal-title":"Retrovirology"},{"key":"2026010712452523600_bib98","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1016\/S0378-1119(97)00376-4","article-title":"Positioning of 72 potentially full size LTRs of human endogenous retroviruses HERV-K on the human chromosome 19 map. Occurrences of the LTRs in human gene sites","volume":"199","author":"Vinogradova","year":"1997","journal-title":"Gene"},{"key":"2026010712452523600_bib99","doi-asserted-by":"crossref","first-page":"1529","DOI":"10.1089\/088922299309810","article-title":"Phylogeny of a novel family of human endogenous retrovirus sequences, HERV-W, in humans and other primates","volume":"15","author":"Voisset","year":"1999","journal-title":"AIDS Research and Human Retroviruses"},{"key":"2026010712452523600_bib100","doi-asserted-by":"crossref","first-page":"3707","DOI":"10.1128\/MCB.00223-12","article-title":"High-temperature requirement protein A4 (HtrA4) suppresses the fusogenic activity of syncytin-1 and promotes trophoblast invasion","volume":"32","author":"Wang","year":"2012","journal-title":"Molecular and Cellular Biology"},{"key":"2026010712452523600_bib101","doi-asserted-by":"crossref","first-page":"1","DOI":"10.2478\/immun-2013-0001","article-title":"Exposure to maternal diabetes in utero and DNA methylation patterns in the offspring","volume":"1","author":"West","year":"2013","journal-title":"Immunometabolism"},{"key":"2026010712452523600_bib102","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1242\/jcs.15.1.177","article-title":"The analysis of malignancy by cell fusion. V. Further evidence of the ability of normal diploid cells to suppress malignancy","volume":"15","author":"Wiener","year":"1974","journal-title":"Journal of Cell Science"},{"key":"2026010712452523600_bib103","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1097\/00006254-200301000-00022","article-title":"Molecular epidemiology of preeclampsia","volume":"58","author":"Wilson","year":"2003","journal-title":"Obstetrical and Gynecological Survey"},{"key":"2026010712452523600_bib104","doi-asserted-by":"crossref","first-page":"50062","DOI":"10.1074\/jbc.M209316200","article-title":"GCMa regulates the syncytin-mediated trophoblastic fusion","volume":"277","author":"Yu","year":"2002","journal-title":"Journal of Biological Chemistry"},{"key":"2026010712452523600_bib105","doi-asserted-by":"crossref","first-page":"9429","DOI":"10.1093\/nar\/15.22.9429","article-title":"Hypermethylation of human DNA sequences in embryonal carcinoma cells and somatic tissues but not in sperm","volume":"15","author":"Zhang","year":"1987","journal-title":"Nucleic Acids Research"},{"key":"2026010712452523600_bib106","doi-asserted-by":"crossref","first-page":"1786","DOI":"10.2174\/13816128113199990540","article-title":"Hypomethylation and activation of syncytin-1 gene in endometriotic tissue","volume":"20","author":"Zhou","year":"2014","journal-title":"Current Pharmaceutical Design"},{"key":"2026010712452523600_bib107","doi-asserted-by":"crossref","first-page":"1796","DOI":"10.2174\/13816128113199990541","article-title":"Decreased expression and altered methylation of syncytin-1 gene in human placentas associated with preeclampsia","volume":"20","author":"Zhuang","year":"2014","journal-title":"Current Pharmaceutical Design"}],"container-title":["Reproduction"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/rep.bioscientifica.com\/view\/journals\/rep\/152\/5\/R167.xml","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/academic.oup.com\/reproduction\/article-pdf\/152\/5\/R167\/64135017\/r167.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/reproduction\/article-pdf\/152\/5\/R167\/64135017\/r167.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,7]],"date-time":"2026-01-07T17:46:32Z","timestamp":1767807992000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/reproduction\/article\/152\/5\/R167\/8236231"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2016,11,1]]},"references-count":107,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2016,11,1]]}},"URL":"https:\/\/doi.org\/10.1530\/rep-16-0031","relation":{},"ISSN":["1470-1626","1741-7899"],"issn-type":[{"value":"1470-1626","type":"print"},{"value":"1741-7899","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2016,11,1]]},"published":{"date-parts":[[2016,11,1]]}}}