{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T16:17:29Z","timestamp":1779207449384,"version":"3.51.4"},"reference-count":56,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,4,7]],"date-time":"2026-04-07T00:00:00Z","timestamp":1775520000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T00:00:00Z","timestamp":1779148800000},"content-version":"vor","delay-in-days":42,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/501100004281","name":"Narodowe Centrum Nauki","doi-asserted-by":"publisher","award":["NCN 2020\/36\/C\/NZ9\/00192"],"award-info":[{"award-number":["NCN 2020\/36\/C\/NZ9\/00192"]}],"id":[{"id":"10.13039\/501100004281","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Genomics"],"DOI":"10.1186\/s12864-026-12822-8","type":"journal-article","created":{"date-parts":[[2026,4,7]],"date-time":"2026-04-07T00:24:10Z","timestamp":1775521450000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Comparative analysis of the transcriptome and methylome in distinct trophoblast populations at the time of embryo implantation in mares"],"prefix":"10.1186","volume":"27","author":[{"given":"Joanna","family":"Jaworska","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Dawid","family":"Tobolski","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shebl E","family":"Salem","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Katarzyna","family":"Ropka-Molik","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tomasz","family":"Z\u0105bek","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tomasz","family":"Szmato\u0142a","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Anne","family":"Kahler","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Izabela","family":"Woc\u0142awek-Potocka","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Katarzyna","family":"Pi\u00f3rkowska","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Amanda M","family":"de Mestre","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2026,4,7]]},"reference":[{"key":"12822_CR1","doi-asserted-by":"publisher","first-page":"1486608","DOI":"10.3389\/fendo.2024.1486608","volume":"15","author":"L Liu","year":"2024","unstructured":"Liu L, Tang L, Chen S, Zheng L, Ma X. Decoding the molecular pathways governing trophoblast migration and placental development; a literature review. Front Endocrinol. 2024;15:1486608. https:\/\/doi.org\/10.3389\/fendo.2024.1486608.","journal-title":"Front Endocrinol"},{"key":"12822_CR2","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/j.ando.2016.04.006","volume":"77","author":"P Chavatte-Palmer","year":"2016","unstructured":"Chavatte-Palmer P, Tarrade A. Placentation in different mammalian species. Ann Endocrinol. 2016;77:67\u201374. https:\/\/doi.org\/10.1016\/j.ando.2016.04.006.","journal-title":"Ann Endocrinol"},{"key":"12822_CR3","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1016\/j.bpobgyn.2010.10.010","volume":"25","author":"AM Carter","year":"2011","unstructured":"Carter AM, Pijnenborg R. Evolution of invasive placentation with special reference to non-human primates. Best Pract Res Clin Obstet Gynaecol. 2011;25:249\u201357. https:\/\/doi.org\/10.1016\/j.bpobgyn.2010.10.010.","journal-title":"Best Pract Res Clin Obstet Gynaecol"},{"key":"12822_CR4","doi-asserted-by":"publisher","first-page":"R39","DOI":"10.1530\/REP-21-0115","volume":"163","author":"S Loux","year":"2022","unstructured":"Loux S, Robles M, Chavatte-Palmer P, de Mestre A. Markers of equine placental differentiation: insights from gene expression studies. Reprod Camb Engl. 2022;163:R39\u201354. https:\/\/doi.org\/10.1530\/REP-21-0115.","journal-title":"Reprod Camb Engl"},{"key":"12822_CR5","doi-asserted-by":"publisher","first-page":"3054","DOI":"10.1210\/en.2013-2116","volume":"155","author":"V Cabrera-Sharp","year":"2014","unstructured":"Cabrera-Sharp V, Read JE, Richardson S, Kowalski AA, Antczak DF, Cartwright JE, et al. SMAD1\/5 signaling in the early equine placenta regulates trophoblast differentiation and chorionic gonadotropin secretion. Endocrinology. 2014;155:3054\u201364. https:\/\/doi.org\/10.1210\/en.2013-2116.","journal-title":"Endocrinology"},{"key":"12822_CR6","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1530\/REP-18-0270","volume":"156","author":"JE Read","year":"2018","unstructured":"Read JE, Cabrera-Sharp V, Offord V, Mirczuk SM, Allen SP, Fowkes RC, et al. Dynamic changes in gene expression and signalling during trophoblast development in the horse. Reprod Camb Engl. 2018;156:313\u201330. https:\/\/doi.org\/10.1530\/REP-18-0270.","journal-title":"Reprod Camb Engl"},{"key":"12822_CR7","doi-asserted-by":"publisher","first-page":"1612","DOI":"10.1095\/biolreprod66.6.1612","volume":"66","author":"SJ Bacon","year":"2002","unstructured":"Bacon SJ, Ellis SA, Antczak DF. Control of expression of major histocompatibility complex genes in horse trophoblast. Biol Reprod. 2002;66:1612\u201320. https:\/\/doi.org\/10.1095\/biolreprod66.6.1612.","journal-title":"Biol Reprod"},{"key":"12822_CR8","doi-asserted-by":"publisher","unstructured":"Allen WR. Maternal recognition of pregnancy and immunological implications of trophoblast-endometrium interactions in equids. Ciba Found Symp. 1978;:323\u201352. https:\/\/doi.org\/10.1002\/9780470720479.ch14.","DOI":"10.1002\/9780470720479.ch14"},{"key":"12822_CR9","doi-asserted-by":"publisher","first-page":"3564","DOI":"10.3390\/nu15163564","volume":"15","author":"L Lawless","year":"2023","unstructured":"Lawless L, Qin Y, Xie L, Zhang K. Trophoblast Differentiation: Mechanisms and Implications for Pregnancy Complications. Nutrients. 2023;15:3564. https:\/\/doi.org\/10.3390\/nu15163564.","journal-title":"Nutrients"},{"key":"12822_CR10","doi-asserted-by":"publisher","first-page":"484","DOI":"10.1038\/nrg3230","volume":"13","author":"PA Jones","year":"2012","unstructured":"Jones PA. Functions of DNA methylation: islands, start sites, gene bodies and beyond. Nat Rev Genet. 2012;13:484\u201392. https:\/\/doi.org\/10.1038\/nrg3230.","journal-title":"Nat Rev Genet"},{"key":"12822_CR11","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1016\/j.placenta.2017.07.007","volume":"57","author":"C Gibson","year":"2017","unstructured":"Gibson C, De Ruijter-Villani M, Stout TAE. Negative uterine asynchrony retards early equine conceptus development and upregulation of placental imprinted genes. Placenta. 2017;57:175\u201382. https:\/\/doi.org\/10.1016\/j.placenta.2017.07.007.","journal-title":"Placenta"},{"key":"12822_CR12","doi-asserted-by":"publisher","first-page":"7084","DOI":"10.3390\/ijms24087084","volume":"24","author":"D Orellana-Guerrero","year":"2023","unstructured":"Orellana-Guerrero D, Uribe-Salazar JM, El-Sheikh Ali H, Scoggin KE, Ball B, Daels P, et al. Dynamics of the Equine Placental DNA Methylome and Transcriptome from Mid- to Late Gestation. Int J Mol Sci. 2023;24:7084. https:\/\/doi.org\/10.3390\/ijms24087084.","journal-title":"Int J Mol Sci"},{"key":"12822_CR13","doi-asserted-by":"publisher","first-page":"458","DOI":"10.1093\/biolre\/ioaf004","volume":"112","author":"J Jaworska","year":"2025","unstructured":"Jaworska J, Tobolski D, Salem SE, Kahler A, Woc\u0142awek-Potocka I, de Mestre AM. Single-cell atlas of the pregnant equine endometrium before and after implantation\u2020. Biol Reprod. 2025;112:458\u201373. https:\/\/doi.org\/10.1093\/biolre\/ioaf004.","journal-title":"Biol Reprod"},{"key":"12822_CR14","doi-asserted-by":"publisher","first-page":"675","DOI":"10.3390\/ani11030675","volume":"11","author":"J Jaworska","year":"2021","unstructured":"Jaworska J, Ropka-Molik K, Pi\u00f3rkowska K, Szmato\u0142a T, Kowalczyk-Zi\u0119ba I, Woc\u0142awek-Potocka I, et al. Transcriptome Profiling of the Retained Fetal Membranes-An Insight in the Possible Pathogenesis of the Disease. Anim Open Access J MDPI. 2021;11:675. https:\/\/doi.org\/10.3390\/ani11030675.","journal-title":"Anim Open Access J MDPI"},{"key":"12822_CR15","doi-asserted-by":"publisher","first-page":"895","DOI":"10.3390\/biology1030895","volume":"1","author":"M Dodt","year":"2012","unstructured":"Dodt M, Roehr JT, Ahmed R, Dieterich C. FLEXBAR-Flexible Barcode and Adapter Processing for Next-Generation Sequencing Platforms. Biology. 2012;1:895\u2013905. https:\/\/doi.org\/10.3390\/biology1030895.","journal-title":"Biology"},{"key":"12822_CR16","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","volume":"29","author":"A Dobin","year":"2013","unstructured":"Dobin A, Davis CA, Schlesinger F, Drenkow J, Zaleski C, Jha S, et al. STAR: ultrafast universal RNA-seq aligner. Bioinforma Oxf Engl. 2013;29:15\u201321. https:\/\/doi.org\/10.1093\/bioinformatics\/bts635.","journal-title":"Bioinforma Oxf Engl"},{"key":"12822_CR17","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1093\/bioinformatics\/btu638","volume":"31","author":"S Anders","year":"2015","unstructured":"Anders S, Pyl PT, Huber W. HTSeq\u2013a Python framework to work with high-throughput sequencing data. Bioinforma Oxf Engl. 2015;31:166\u20139. https:\/\/doi.org\/10.1093\/bioinformatics\/btu638.","journal-title":"Bioinforma Oxf Engl"},{"key":"12822_CR18","doi-asserted-by":"publisher","unstructured":"Mi L, W H. Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2. Genome Biol. 2014;15. https:\/\/doi.org\/10.1186\/s13059-014-0550-8.","DOI":"10.1186\/s13059-014-0550-8"},{"key":"12822_CR19","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","volume":"57","author":"Y Benjamini","year":"1995","unstructured":"Benjamini Y, Hochberg Y. Controlling the False Discovery Rate: A Practical and Powerful Approach to Multiple Testing. J R Stat Soc Ser B Methodol. 1995;57:289\u2013300. https:\/\/doi.org\/10.1111\/j.2517-6161.1995.tb02031.x.","journal-title":"J R Stat Soc Ser B Methodol"},{"key":"12822_CR20","doi-asserted-by":"publisher","first-page":"811","DOI":"10.1186\/1471-2164-15-811","volume":"15","author":"G Li","year":"2014","unstructured":"Li G, Jia Q, Zhao J, Li X, Yu M, Samuel MS, et al. Dysregulation of genome-wide gene expression and DNA methylation in abnormal cloned piglets. BMC Genomics. 2014;15:811. https:\/\/doi.org\/10.1186\/1471-2164-15-811.","journal-title":"BMC Genomics"},{"key":"12822_CR21","doi-asserted-by":"publisher","first-page":"916","DOI":"10.1038\/s41467-020-14566-9","volume":"11","author":"A Liakos","year":"2020","unstructured":"Liakos A, Konstantopoulos D, Lavigne MD, Fousteri M. Continuous transcription initiation guarantees robust repair of all transcribed genes and regulatory regions. Nat Commun. 2020;11:916. https:\/\/doi.org\/10.1038\/s41467-020-14566-9.","journal-title":"Nat Commun"},{"key":"12822_CR22","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1016\/j.tig.2020.04.006","volume":"36","author":"R van der Lee","year":"2020","unstructured":"van der Lee R, Correard S, Wasserman WW. Deregulated Regulators: Disease-Causing cis Variants in Transcription Factor Genes. Trends Genet TIG. 2020;36:523\u201339. https:\/\/doi.org\/10.1016\/j.tig.2020.04.006.","journal-title":"Trends Genet TIG"},{"key":"12822_CR23","doi-asserted-by":"publisher","first-page":"13314","DOI":"10.1038\/s41598-020-69967-z","volume":"10","author":"CA Shilton","year":"2020","unstructured":"Shilton CA, Kahler A, Davis BW, Crabtree JR, Crowhurst J, McGladdery AJ, et al. Whole genome analysis reveals aneuploidies in early pregnancy loss in the horse. Sci Rep. 2020;10:13314. https:\/\/doi.org\/10.1038\/s41598-020-69967-z.","journal-title":"Sci Rep"},{"key":"12822_CR24","doi-asserted-by":"publisher","first-page":"P3","DOI":"10.1186\/gb-2003-4-5-p3","volume":"4","author":"G Dennis","year":"2003","unstructured":"Dennis G, Sherman BT, Hosack DA, Yang J, Gao W, Lane HC, et al. DAVID: Database for Annotation, Visualization, and Integrated Discovery. Genome Biol. 2003;4:P3. https:\/\/doi.org\/10.1186\/gb-2003-4-5-p3.","journal-title":"Genome Biol"},{"key":"12822_CR25","doi-asserted-by":"publisher","unstructured":"Noronha LE, Antczak DF. Maternal immune responses to trophoblast: the contribution of the horse to pregnancy immunology. Am J Reprod Immunol N Y N. 1989. 2010;64:231\u201344. https:\/\/doi.org\/10.1111\/j.1600-0897.2010.00895.x.","DOI":"10.1111\/j.1600-0897.2010.00895.x"},{"key":"12822_CR26","doi-asserted-by":"publisher","first-page":"753","DOI":"10.1095\/biolreprod64.3.753","volume":"64","author":"AP Adams","year":"2001","unstructured":"Adams AP, Antczak DF. Ectopic transplantation of equine invasive trophoblast. Biol Reprod. 2001;64:753\u201363. https:\/\/doi.org\/10.1095\/biolreprod64.3.753.","journal-title":"Biol Reprod"},{"key":"12822_CR27","doi-asserted-by":"publisher","first-page":"e0224497","DOI":"10.1371\/journal.pone.0224497","volume":"14","author":"SC Loux","year":"2019","unstructured":"Loux SC, Dini P, El-Sheikh Ali H, Kalbfleisch T, Ball BA. Characterization of the placental transcriptome through mid to late gestation in the mare. PLoS ONE. 2019;14:e0224497. https:\/\/doi.org\/10.1371\/journal.pone.0224497.","journal-title":"PLoS ONE"},{"key":"12822_CR28","doi-asserted-by":"publisher","first-page":"438","DOI":"10.4049\/jimmunol.1401344","volume":"194","author":"X Gao","year":"2015","unstructured":"Gao X, Wang X, Yang Q, Zhao X, Wen W, Li G, et al. Tumoral expression of IL-33 inhibits tumor growth and modifies the tumor microenvironment through CD8\u2009+\u2009T and NK cells. J Immunol Baltim Md 1950. 2015;194:438\u201345. https:\/\/doi.org\/10.4049\/jimmunol.1401344.","journal-title":"J Immunol Baltim Md 1950"},{"key":"12822_CR29","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1038\/s44321-023-00012-y","volume":"16","author":"K Che","year":"2024","unstructured":"Che K, Luo Y, Song X, Yang Z, Wang H, Shi T, et al. Macrophages reprogramming improves immunotherapy of IL-33 in peritoneal metastasis of gastric cancer. EMBO Mol Med. 2024;16:251\u201366. https:\/\/doi.org\/10.1038\/s44321-023-00012-y.","journal-title":"EMBO Mol Med"},{"key":"12822_CR30","doi-asserted-by":"publisher","unstructured":"Valero-Pacheco N, Tang EK, Massri N, Loia R, Chemerinski A, Wu T, et al. Maternal IL-33 critically regulates tissue remodeling and type 2 immune responses in the uterus during early pregnancy in mice. Proc Natl Acad Sci. 2022;119. https:\/\/doi.org\/10.1073\/pnas.2123267119.","DOI":"10.1073\/pnas.2123267119"},{"key":"12822_CR31","doi-asserted-by":"publisher","first-page":"16882","DOI":"10.3390\/ijms242316882","volume":"24","author":"B Wen","year":"2023","unstructured":"Wen B, Liao H, Lin W, Li Z, Ma X, Xu Q, et al. The Role of TGF-\u03b2 during Pregnancy and Pregnancy Complications. Int J Mol Sci. 2023;24:16882. https:\/\/doi.org\/10.3390\/ijms242316882.","journal-title":"Int J Mol Sci"},{"key":"12822_CR32","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1016\/j.tem.2020.12.005","volume":"32","author":"Y Li","year":"2021","unstructured":"Li Y, Yan J, Chang H-M, Chen Z-J, Leung PCK. Roles of TGF-\u03b2 Superfamily Proteins in Extravillous Trophoblast Invasion. Trends Endocrinol Metab TEM. 2021;32:170\u201389. https:\/\/doi.org\/10.1016\/j.tem.2020.12.005.","journal-title":"Trends Endocrinol Metab TEM"},{"key":"12822_CR33","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1111\/j.1439-0531.2011.01843.x","volume":"46","author":"AJ Ziecik","year":"2011","unstructured":"Ziecik AJ, Waclawik A, Kaczmarek MM, Blitek A, Jalali BM, Andronowska A. Mechanisms for the establishment of pregnancy in the pig. Reprod Domest Anim Zuchthyg. 2011;46:31\u201341. https:\/\/doi.org\/10.1111\/j.1439-0531.2011.01843.x.","journal-title":"Reprod Domest Anim Zuchthyg"},{"key":"12822_CR34","doi-asserted-by":"publisher","first-page":"465","DOI":"10.1530\/REP-09-0447","volume":"139","author":"DA Massuto","year":"2010","unstructured":"Massuto DA, Kneese EC, Johnson GA, Burghardt RC, Hooper RN, Ing NH, et al. Transforming growth factor beta (TGFB) signaling is activated during porcine implantation: proposed role for latency-associated peptide interactions with integrins at the conceptus-maternal interface. Reprod Camb Engl. 2010;139:465\u201378. https:\/\/doi.org\/10.1530\/REP-09-0447.","journal-title":"Reprod Camb Engl"},{"key":"12822_CR35","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1038\/nrneph.2016.48","volume":"12","author":"X-M Meng","year":"2016","unstructured":"Meng X-M, Nikolic-Paterson DJ, Lan HY. TGF-\u03b2: the master regulator of fibrosis. Nat Rev Nephrol. 2016;12:325\u201338. https:\/\/doi.org\/10.1038\/nrneph.2016.48.","journal-title":"Nat Rev Nephrol"},{"key":"12822_CR36","doi-asserted-by":"publisher","first-page":"1458","DOI":"10.3389\/fimmu.2019.01458","volume":"10","author":"JP Vago","year":"2019","unstructured":"Vago JP, Sugimoto MA, Lima KM, Negreiros-Lima GL, Baik N, Teixeira MM, et al. Plasminogen and the Plasminogen Receptor, Plg-RKT, Regulate Macrophage Phenotypic, and Functional Changes. Front Immunol. 2019;10:1458. https:\/\/doi.org\/10.3389\/fimmu.2019.01458.","journal-title":"Front Immunol"},{"key":"12822_CR37","doi-asserted-by":"publisher","first-page":"876","DOI":"10.1038\/cdd.2014.7","volume":"21","author":"CH Bird","year":"2014","unstructured":"Bird CH, Christensen ME, Mangan MSJ, Prakash MD, Sedelies KA, Smyth MJ, et al. The granzyme B-Serpinb9 axis controls the fate of lymphocytes after lysosomal stress. Cell Death Differ. 2014;21:876\u201387. https:\/\/doi.org\/10.1038\/cdd.2014.7.","journal-title":"Cell Death Differ"},{"key":"12822_CR38","doi-asserted-by":"publisher","first-page":"121723","DOI":"10.1016\/j.biomaterials.2022.121723","volume":"288","author":"R Han","year":"2022","unstructured":"Han R, Yu L, Zhao C, Li Y, Ma Y, Zhai Y, et al. Inhibition of SerpinB9 to enhance granzyme B-based tumor therapy by using a modified biomimetic nanoplatform with a cascade strategy. Biomaterials. 2022;288:121723. https:\/\/doi.org\/10.1016\/j.biomaterials.2022.121723.","journal-title":"Biomaterials"},{"key":"12822_CR39","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1146\/annurev-animal-031412-103703","volume":"1","author":"DF Antczak","year":"2013","unstructured":"Antczak DF, de Mestre AM, Wilsher S, Allen WR. The equine endometrial cup reaction: a fetomaternal signal of significance. Annu Rev Anim Biosci. 2013;1:419\u201342. https:\/\/doi.org\/10.1146\/annurev-animal-031412-103703.","journal-title":"Annu Rev Anim Biosci"},{"key":"12822_CR40","doi-asserted-by":"publisher","first-page":"445","DOI":"10.1387\/ijdb.082795ad","volume":"54","author":"A de Mestre","year":"2010","unstructured":"de Mestre A, Noronha L, Wagner B, Antczak DF. Split immunological tolerance to trophoblast. Int J Dev Biol. 2010;54:445\u201355. https:\/\/doi.org\/10.1387\/ijdb.082795ad.","journal-title":"Int J Dev Biol"},{"key":"12822_CR41","doi-asserted-by":"publisher","first-page":"813","DOI":"10.1038\/nri3339","volume":"12","author":"KS Kobayashi","year":"2012","unstructured":"Kobayashi KS, van den Elsen PJ. NLRC5: a key regulator of MHC class I-dependent immune responses. Nat Rev Immunol. 2012;12:813\u201320. https:\/\/doi.org\/10.1038\/nri3339.","journal-title":"Nat Rev Immunol"},{"key":"12822_CR42","doi-asserted-by":"publisher","first-page":"831","DOI":"10.1093\/biolre\/iox009","volume":"96","author":"T Tilburgs","year":"2017","unstructured":"Tilburgs T, Meissner TB, Ferreira LMR, Mulder A, Musunuru K, Ye J, et al. NLRP2 is a suppressor of NF-\u0199B signaling and HLA-C expression in human trophoblasts\u2020\u2021. Biol Reprod. 2017;96:831\u201342. https:\/\/doi.org\/10.1093\/biolre\/iox009.","journal-title":"Biol Reprod"},{"key":"12822_CR43","doi-asserted-by":"publisher","first-page":"1964","DOI":"10.3390\/ijms22041964","volume":"22","author":"A Shukla","year":"2021","unstructured":"Shukla A, Cloutier M, Appiya Santharam M, Ramanathan S, Ilangumaran S. The MHC Class-I Transactivator NLRC5: Implications to Cancer Immunology and Potential Applications to Cancer Immunotherapy. Int J Mol Sci. 2021;22:1964. https:\/\/doi.org\/10.3390\/ijms22041964.","journal-title":"Int J Mol Sci"},{"key":"12822_CR44","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1007\/s00251-002-0514-0","volume":"54","author":"RL Tallmadge","year":"2003","unstructured":"Tallmadge RL, Lear TL, Johnson AK, Gu\u00e9rin G, Millon LV, Carpenter SL, et al. Characterization of the \u03b22-microglobulin gene of the horse. Immunogenetics. 2003;54:725\u201333. https:\/\/doi.org\/10.1007\/s00251-002-0514-0.","journal-title":"Immunogenetics"},{"key":"12822_CR45","doi-asserted-by":"publisher","first-page":"2837","DOI":"10.3390\/ijms20112837","volume":"20","author":"C Apicella","year":"2019","unstructured":"Apicella C, Ruano CSM, M\u00e9hats C, Miralles F, Vaiman D. The Role of Epigenetics in Placental Development and the Etiology of Preeclampsia. Int J Mol Sci. 2019;20:2837. https:\/\/doi.org\/10.3390\/ijms20112837.","journal-title":"Int J Mol Sci"},{"key":"12822_CR46","doi-asserted-by":"publisher","first-page":"547","DOI":"10.1093\/molehr\/gan046","volume":"14","author":"B Novakovic","year":"2008","unstructured":"Novakovic B, Rakyan V, Ng HK, Manuelpillai U, Dewi C, Wong NC, et al. Specific tumour-associated methylation in normal human term placenta and first-trimester cytotrophoblasts. Mol Hum Reprod. 2008;14:547\u201354. https:\/\/doi.org\/10.1093\/molehr\/gan046.","journal-title":"Mol Hum Reprod"},{"key":"12822_CR47","doi-asserted-by":"publisher","first-page":"1461","DOI":"10.4161\/15592294.2014.971608","volume":"9","author":"H Cardenas","year":"2014","unstructured":"Cardenas H, Vieth E, Lee J, Segar M, Liu Y, Nephew KP, et al. TGF-\u03b2 induces global changes in DNA methylation during the epithelial-to-mesenchymal transition in ovarian cancer cells. Epigenetics. 2014;9:1461\u201372. https:\/\/doi.org\/10.4161\/15592294.2014.971608.","journal-title":"Epigenetics"},{"key":"12822_CR48","doi-asserted-by":"publisher","first-page":"e100148","DOI":"10.1371\/journal.pone.0100148","volume":"9","author":"L Anton","year":"2014","unstructured":"Anton L, Brown AG, Bartolomei MS, Elovitz MA. Differential methylation of genes associated with cell adhesion in preeclamptic placentas. PLoS ONE. 2014;9:e100148. https:\/\/doi.org\/10.1371\/journal.pone.0100148.","journal-title":"PLoS ONE"},{"key":"12822_CR49","doi-asserted-by":"publisher","first-page":"848","DOI":"10.1095\/biolreprod.108.073353","volume":"80","author":"SP Murphy","year":"2009","unstructured":"Murphy SP, Tayade C, Ashkar AA, Hatta K, Zhang J, Croy BA. Interferon gamma in successful pregnancies. Biol Reprod. 2009;80:848\u201359. https:\/\/doi.org\/10.1095\/biolreprod.108.073353.","journal-title":"Biol Reprod"},{"key":"12822_CR50","doi-asserted-by":"publisher","first-page":"2092","DOI":"10.1182\/blood-2017-03-768507","volume":"130","author":"M Bai","year":"2017","unstructured":"Bai M, Grieshaber-Bouyer R, Wang J, Schmider AB, Wilson ZS, Zeng L, et al. CD177 modulates human neutrophil migration through activation-mediated integrin and chemoreceptor regulation. Blood. 2017;130:2092\u2013100. https:\/\/doi.org\/10.1182\/blood-2017-03-768507.","journal-title":"Blood"},{"key":"12822_CR51","doi-asserted-by":"publisher","first-page":"10705","DOI":"10.1073\/pnas.1308998110","volume":"110","author":"X Wang","year":"2013","unstructured":"Wang X, Miller DC, Harman R, Antczak DF, Clark AG. Paternally expressed genes predominate in the placenta. Proc Natl Acad Sci. 2013;110:10705\u201310. https:\/\/doi.org\/10.1073\/pnas.1308998110.","journal-title":"Proc Natl Acad Sci"},{"key":"12822_CR52","doi-asserted-by":"publisher","first-page":"e2006474118","DOI":"10.1073\/pnas.2006474118","volume":"118","author":"P Dini","year":"2021","unstructured":"Dini P, Kalbfleisch T, Uribe-Salazar JM, Carossino M, Ali HE-S, Loux SC, et al. Parental bias in expression and interaction of genes in the equine placenta. Proc Natl Acad Sci. 2021;118:e2006474118. https:\/\/doi.org\/10.1073\/pnas.2006474118.","journal-title":"Proc Natl Acad Sci"},{"key":"12822_CR53","doi-asserted-by":"publisher","first-page":"1439","DOI":"10.1038\/s41418-024-01337-y","volume":"31","author":"M Cesana","year":"2024","unstructured":"Cesana M, Tufano G, Panariello F, Zampelli N, Soldati C, Mutarelli M, et al. TFEB controls syncytiotrophoblast formation and hormone production in placenta. Cell Death Differ. 2024;31:1439\u201351. https:\/\/doi.org\/10.1038\/s41418-024-01337-y.","journal-title":"Cell Death Differ"},{"key":"12822_CR54","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1007\/s00441-021-03502-y","volume":"386","author":"C Desterke","year":"2021","unstructured":"Desterke C, Dang J, Lorenzo H-K, Candelier J-J. Roles of tetraspanins during trophoblast development: bioinformatics and new perspectives. Cell Tissue Res. 2021;386:157\u201371. https:\/\/doi.org\/10.1007\/s00441-021-03502-y.","journal-title":"Cell Tissue Res"},{"key":"12822_CR55","doi-asserted-by":"publisher","first-page":"11649","DOI":"10.1074\/jbc.M112.445304","volume":"288","author":"D Perez-Hernandez","year":"2013","unstructured":"Perez-Hernandez D, Guti\u00e9rrez-V\u00e1zquez C, Jorge I, L\u00f3pez-Mart\u00edn S, Ursa A, S\u00e1nchez-Madrid F, et al. The intracellular interactome of tetraspanin-enriched microdomains reveals their function as sorting machineries toward exosomes. J Biol Chem. 2013;288:11649\u201361. https:\/\/doi.org\/10.1074\/jbc.M112.445304.","journal-title":"J Biol Chem"},{"key":"12822_CR56","doi-asserted-by":"publisher","first-page":"e3","DOI":"10.1017\/S1462399409001355","volume":"12","author":"CS Stipp","year":"2010","unstructured":"Stipp CS. Laminin-binding integrins and their tetraspanin partners as potential antimetastatic targets. Expert Rev Mol Med. 2010;12:e3. https:\/\/doi.org\/10.1017\/S1462399409001355.","journal-title":"Expert Rev Mol Med"}],"container-title":["BMC Genomics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12864-026-12822-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12864-026-12822-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12864-026-12822-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T16:05:12Z","timestamp":1779206712000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s12864-026-12822-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,7]]},"references-count":56,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["12822"],"URL":"https:\/\/doi.org\/10.1186\/s12864-026-12822-8","relation":{},"ISSN":["1471-2164"],"issn-type":[{"value":"1471-2164","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,7]]},"assertion":[{"value":"8 August 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"31 March 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"7 April 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"All animal procedures were conducted in accordance with the UK Animals (Scientific Procedures) Act 1986 (as revised) and the EU Directive 2010\/63\/EU for animal experiments. The study was approved by the Royal Veterinary College Ethics and Welfare Committee, specifically its Animal Welfare Ethical Review Body (AWERB) under UK Home Office project license PL70\/8577. For the non-surgical uterine lavage procedure, mares were restrained in stocks and sedated with detomidine hydrochloride (Dormosedan, 10\u00a0mg\/mL; Pfizer, UK) at a dose of 0.01\u00a0mg\/kg body weight, administered intravenously. The mares were not euthanised, anaesthetised, or rendered unconscious; detomidine hydrochloride is a potent sedative and analgesic that induces a state of reduced awareness and provides pain relief.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"492"}}