{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,9]],"date-time":"2026-06-09T18:44:46Z","timestamp":1781030686817,"version":"3.54.1"},"reference-count":65,"publisher":"Oxford University Press (OUP)","issue":"3","funder":[{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"crossref","award":["2019R1A2C1004670"],"award-info":[{"award-number":["2019R1A2C1004670"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"crossref","award":["2020R1A6A3A01098213"],"award-info":[{"award-number":["2020R1A6A3A01098213"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["rep.bioscientifica.com"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2025,1,29]]},"abstract":"<jats:sec>\n                    <jats:title>In brief<\/jats:title>\n                    <jats:p>The CD40 ligand\u2013CD40 signaling system expressed at the maternal-conceptus interface in pigs is involved in regulating maternal immunity by modifying endometrial endothelial cell function during early pregnancy. This paper reveals the role of CD40 ligand\u2013CD40 signaling at the maternal-conceptus interface in pigs.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      To successfully establish and maintain pregnancy in pigs, a variety of factors must work together at the maternal-conceptus interface to form an immune environment appropriate for both the mother and the conceptus. Our transcriptomics study has shown that cluster of differentiation ligand 40 (CD40L) and its receptor CD40, which are known to play important roles in regulating cell- and antibody-mediated immunity, are expressed in the endometrium during early pregnancy. However, the roles of the CD40L and CD40 signaling system are not well understood. Therefore, we determined the expression, regulation and function of CD40L and CD40 at the maternal-conceptus interface in pigs. The endometrium expressed\n                      <jats:italic>CD40L<\/jats:italic>\n                      and\n                      <jats:italic>CD40<\/jats:italic>\n                      mRNAs at the greatest levels on day 15 of pregnancy. The CD40L protein was localized predominantly to luminal epithelial cells on day 15 of pregnancy, and the CD40 protein was found in luminal epithelial, stromal and vascular endothelial cells in the endometrium during pregnancy. During early pregnancy, the conceptus expressed\n                      <jats:italic>CD40<\/jats:italic>\n                      but not\n                      <jats:italic>CD40L<\/jats:italic>\n                      and chorioallantoic tissues during mid- to late pregnancy expressed both\n                      <jats:italic>CD40L<\/jats:italic>\n                      and\n                      <jats:italic>CD40<\/jats:italic>\n                      . Interferon-\u03b3 increased the expression of\n                      <jats:italic>CD40L<\/jats:italic>\n                      and\n                      <jats:italic>CD40<\/jats:italic>\n                      in endometrial explants. CD40L increased the migration but not the proliferation of cultured porcine uterine endothelial (pENDO) cells\n                      <jats:italic>in vitro<\/jats:italic>\n                      . In addition, CD40L affected the expression of genes related to angiogenesis, cell adhesion, chemokines and immunity in pENDO cells. These results suggest that the CD40L\u2013CD40 system might play an important role in the establishment of pregnancy in pigs by regulating endometrial endothelial cell function during the implantation period.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1530\/rep-24-0396","type":"journal-article","created":{"date-parts":[[2025,1,28]],"date-time":"2025-01-28T12:09:22Z","timestamp":1738066162000},"update-policy":"https:\/\/doi.org\/10.1530\/crossmarkpolicy-11","source":"Crossref","is-referenced-by-count":1,"title":["CD40 ligand and CD40: expression, regulation and function at the maternal-conceptus interface in pigs"],"prefix":"10.1093","volume":"169","author":[{"given":"Inkyu","family":"Yoo","sequence":"first","affiliation":[{"name":"Division of Biological Science and Technology, Yonsei University, Wonju, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Soohyung","family":"Lee","sequence":"additional","affiliation":[{"name":"Division of Biological Science and Technology, Yonsei University, Wonju, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yugyeong","family":"Cheon","sequence":"additional","affiliation":[{"name":"Division of Biological Science and Technology, Yonsei University, Wonju, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tae Sub","family":"Park","sequence":"additional","affiliation":[{"name":"Graduate School of International Agricultural Technology and Institute of Green-Bio Science and Technology, Seoul National University, Pyeongchang, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0213-5884","authenticated-orcid":true,"given":"Hakhyun","family":"Ka","sequence":"additional","affiliation":[{"name":"Division of Biological Science and Technology, Yonsei University, Wonju, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","reference":[{"key":"bib1","series-title":"Blood Coagul Fibrinolysis","first-page":"71","article-title":"The expression and concentration of CD40 ligand in normal pregnancy, preeclampsia, and hemolytic anemia, elevated liver enzymes and low platelet count (HELLP) syndrome","volume":"24","author":"Azzam HA","year":"2013","unstructured":"Azzam HA\n, \nAbousamra NK\n, \nGoda H\n, et al.\n 2013 The expression and concentration of CD40 ligand in normal pregnancy, preeclampsia, and hemolytic anemia, elevated liver enzymes and low platelet count (HELLP) syndrome. Blood Coagul Fibrinolysis 24 71\u201375. (https:\/\/doi.org\/10.1097\/MBC.0b013e32835a8aca)"},{"key":"bib2","series-title":"Differentiation","first-page":"52","article-title":"Pig blastocyst-uterine interactions","volume":"87","author":"Bazer FW","year":"2014","unstructured":"Bazer FW\n & \nJohnson GA\n\n 2014 Pig blastocyst-uterine interactions. Differentiation 87 52\u201365. (https:\/\/doi.org\/10.1016\/j.diff.2013.11.005)"},{"key":"bib3","series-title":"Biol Reprod","first-page":"613","article-title":"Characterization of a polarized porcine uterine epithelial model system","volume":"55","author":"Bowen JA","year":"1996","unstructured":"Bowen JA\n, \nNewton GR\n, \nWeise DW\n, et al.\n 1996 Characterization of a polarized porcine uterine epithelial model system. Biol Reprod 55 613\u2013619. (https:\/\/doi.org\/10.1095\/biolreprod55.3.613)"},{"key":"bib4","series-title":"Am J Reprod Immunol","first-page":"487","article-title":"Interleukin-10: a pleiotropic regulator in pregnancy","volume":"73","author":"Cheng SB","year":"2015","unstructured":"Cheng SB\n & \nSharma S\n\n 2015 Interleukin-10: a pleiotropic regulator in pregnancy. Am J Reprod Immunol 73 487\u2013500. (https:\/\/doi.org\/10.1111\/aji.12329)"},{"key":"bib5","series-title":"J Thromb Haemost","first-page":"1364","article-title":"CD40 ligand shedding is regulated by interaction between matrix metalloproteinase\u20102 and platelet integrin \u03b1IIb\u03b23","volume":"8","author":"Choi WS","year":"2010","unstructured":"Choi WS\n, \nJeon OH\n & \nKim DS\n\n 2010 CD40 ligand shedding is regulated by interaction between matrix metalloproteinase\u20102 and platelet integrin \u03b1IIb\u03b23. J Thromb Haemost 8 1364\u20131371. (https:\/\/doi.org\/10.1111\/j.1538-7836.2010.03837.x)"},{"key":"bib6","series-title":"Anat Rec","first-page":"177","article-title":"Initial vascularisation in the pig placenta: I. demonstration of nonglandular areas by histology and corrosion casts","volume":"238","author":"Dantzer V","year":"1994","unstructured":"Dantzer V\n & \nLeiser R\n\n 1994 Initial vascularisation in the pig placenta: I. demonstration of nonglandular areas by histology and corrosion casts. Anat Rec 238 177\u2013190. (https:\/\/doi.org\/10.1002\/ar.1092380204)"},{"key":"bib7","series-title":"Biol Reprod","first-page":"62","article-title":"KLF10 mediated epigenetic dysregulation of epithelial CD40\/CD154 promotes endometriosis","volume":"95","author":"Delaney AA","year":"2016","unstructured":"Delaney AA\n, \nKhan Z\n, \nZheng Y\n, et al.\n 2016 KLF10 mediated epigenetic dysregulation of epithelial CD40\/CD154 promotes endometriosis. Biol Reprod 95 62. (https:\/\/doi.org\/10.1095\/biolreprod.116.140764)"},{"key":"bib8","series-title":"Semin Immunol","article-title":"CD40-CD154: a perspective from type 2 immunity","volume":"53","author":"Diaz A","year":"2021","unstructured":"Diaz A\n, \nGonzalez-Alayon I\n, \nPerez-Torrado V\n, et al.\n 2021 CD40-CD154: a perspective from type 2 immunity. Semin Immunol 53 101528. (https:\/\/doi.org\/10.1016\/j.smim.2021.101528)"},{"key":"bib9","series-title":"J Immunol","first-page":"8088","article-title":"CD40-induced signaling in human endothelial cells results in mTORC2- and Akt-dependent expression of vascular endothelial growth factor in vitro and in vivo","volume":"181","author":"Dormond O","year":"2008","unstructured":"Dormond O\n, \nContreras AG\n, \nMeijer E\n, et al.\n 2008 CD40-induced signaling in human endothelial cells results in mTORC2- and Akt-dependent expression of vascular endothelial growth factor in vitro and in vivo. J Immunol 181 8088\u20138095. (https:\/\/doi.org\/10.4049\/jimmunol.181.11.8088)"},{"key":"bib10","series-title":"Am J Respir Cell Mol Biol","first-page":"442","article-title":"Human lung tissue macrophages, but not alveolar macrophages, express matrix metalloproteinases after direct contact with activated T lymphocytes","volume":"24","author":"Ferrari-Lacraz S","year":"2001","unstructured":"Ferrari-Lacraz S\n, \nNicod LP\n, \nChicheportiche R\n, et al.\n 2001 Human lung tissue macrophages, but not alveolar macrophages, express matrix metalloproteinases after direct contact with activated T lymphocytes. Am J Respir Cell Mol Biol 24 442\u2013451. (https:\/\/doi.org\/10.1165\/ajrcmb.24.4.4008)"},{"key":"bib11","series-title":"Biol Reprod","first-page":"1438","article-title":"Dendritic cells in the human decidua","volume":"69","author":"Gardner L","year":"2003","unstructured":"Gardner L\n & \nMoffett A\n\n 2003 Dendritic cells in the human decidua. Biol Reprod 69 1438\u20131446. (https:\/\/doi.org\/10.1095\/biolreprod.103.017574)"},{"key":"bib12","series-title":"Trends Immunol","first-page":"242","article-title":"Tolerance, DCs and tryptophan: much ado about Ido","volume":"24","author":"Grohmann U","year":"2003","unstructured":"Grohmann U\n, \nFallarino F\n & \nPuccetti P\n\n 2003 Tolerance, DCs and tryptophan: much ado about Ido. Trends Immunol 24 242\u2013248. (https:\/\/doi.org\/10.1016\/s1471-4906(03)00072-3)"},{"key":"bib13","series-title":"Biol Reprod","first-page":"1527","article-title":"Immunolocalization of acidic and basic fibroblast growth factors in porcine uterine and conceptus tissues","volume":"56","author":"Gupta A","year":"1997","unstructured":"Gupta A\n, \nBazer FW\n & \nJaeger LA\n\n 1997 Immunolocalization of acidic and basic fibroblast growth factors in porcine uterine and conceptus tissues. Biol Reprod 56 1527\u20131536. (https:\/\/doi.org\/10.1095\/biolreprod56.6.1527)"},{"key":"bib14","series-title":"Biol Reprod","first-page":"69","article-title":"Analysis of cysteine-X-cysteine motif chemokine ligands 9, 10, and 11, their receptor CXCR3, and their possible role on the recruitment of immune cells at the maternal\u2013conceptus interface in pigs","volume":"97","author":"Han J","year":"2017","unstructured":"Han J\n, \nGu MJ\n, \nYoo I\n, et al.\n 2017 Analysis of cysteine-X-cysteine motif chemokine ligands 9, 10, and 11, their receptor CXCR3, and their possible role on the recruitment of immune cells at the maternal\u2013conceptus interface in pigs. Biol Reprod 97 69\u201380. (https:\/\/doi.org\/10.1093\/biolre\/iox074)"},{"key":"bib15","series-title":"Biol Reprod","first-page":"1137","article-title":"Cysteine-X-cysteine motif chemokine ligand 12 and its receptor CXCR4: expression, regulation, and possible function at the maternal\u2013conceptus interface during early pregnancy in pigs","volume":"99","author":"Han J","year":"2018","unstructured":"Han J\n, \nJeong W\n, \nGu MJ\n, et al.\n 2018 Cysteine-X-cysteine motif chemokine ligand 12 and its receptor CXCR4: expression, regulation, and possible function at the maternal\u2013conceptus interface during early pregnancy in pigs. Biol Reprod 99 1137\u20131148. (https:\/\/doi.org\/10.1093\/biolre\/ioy147)"},{"key":"bib16","series-title":"Biol Reprod","first-page":"1159","article-title":"Interleukin-10 and its receptors at the maternal\u2013conceptus interface: expression, regulation, and implication for T helper 2 cytokine predominance and maternal immune tolerance in the pig, a true epitheliochorial placentation species","volume":"106","author":"Han J","year":"2022","unstructured":"Han J\n, \nYoo I\n, \nLee S\n, et al.\n 2022 Interleukin-10 and its receptors at the maternal\u2013conceptus interface: expression, regulation, and implication for T helper 2 cytokine predominance and maternal immune tolerance in the pig, a true epitheliochorial placentation species. Biol Reprod 106 1159\u20131174. (https:\/\/doi.org\/10.1093\/biolre\/ioac058)"},{"key":"bib17","series-title":"J Exp Med","first-page":"33","article-title":"Expression of functional CD40 by vascular endothelial cells","volume":"182","author":"Hollenbaugh D","year":"1995","unstructured":"Hollenbaugh D\n, \nMischel-Petty N\n, \nEdwards CP\n, et al.\n 1995 Expression of functional CD40 by vascular endothelial cells. J Exp Med 182 33\u201340. (https:\/\/doi.org\/10.1084\/jem.182.1.33)"},{"key":"bib18","series-title":"Theriogenology","first-page":"166","article-title":"Characterization of interferon \u03b1 and \u03b2 receptor IFNAR1 and IFNAR2 expression and regulation in the uterine endometrium during the estrous cycle and pregnancy in pigs","volume":"88","author":"Jang H","year":"2017","unstructured":"Jang H\n, \nChoi Y\n, \nYoo I\n, et al.\n 2017 Characterization of interferon \u03b1 and \u03b2 receptor IFNAR1 and IFNAR2 expression and regulation in the uterine endometrium during the estrous cycle and pregnancy in pigs. Theriogenology 88 166\u2013173. (https:\/\/doi.org\/10.1016\/j.theriogenology.2016.09.025)"},{"key":"bib19","series-title":"Biol Reprod","first-page":"104","article-title":"Lysophosphatidic acid (LPA) receptor 3-mediated LPA signal transduction pathways: a possible relationship with early development of peri-implantation porcine conceptus","volume":"94","author":"Jeong W","year":"2016","unstructured":"Jeong W\n, \nSeo H\n, \nSung Y\n, et al.\n 2016 Lysophosphatidic acid (LPA) receptor 3-mediated LPA signal transduction pathways: a possible relationship with early development of peri-implantation porcine conceptus. Biol Reprod 94 104. (https:\/\/doi.org\/10.1095\/biolreprod.115.137174)"},{"key":"bib20","series-title":"Adv Anat Embryol Cell Biol","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1007\/978-3-030-77360-1_5","article-title":"The early stages of implantation and placentation in the pig","volume":"234","author":"Johnson GA","year":"2021","unstructured":"Johnson GA\n, \nBazer FW\n & \nSeo H\n\n 2021 The early stages of implantation and placentation in the pig. Adv Anat Embryol Cell Biol 234 61\u201389. (https:\/\/doi.org\/10.1007\/978-3-030-77360-1_5)"},{"key":"bib21","series-title":"Biol Reprod","first-page":"292","article-title":"Pig conceptuses increase uterine interferon-regulatory factor 1 (IRF1), but restrict expression to stroma through estrogen-induced IRF2 in luminal epithelium","volume":"77","author":"Joyce MM","year":"2007a","unstructured":"Joyce MM\n, \nBurghardt JR\n, \nBurghardt RC\n, et al.\n 2007a Pig conceptuses increase uterine interferon-regulatory factor 1 (IRF1), but restrict expression to stroma through estrogen-induced IRF2 in luminal epithelium. Biol Reprod 77 292\u2013302. (https:\/\/doi.org\/10.1095\/biolreprod.107.060939)"},{"key":"bib22","series-title":"Endocrinology","first-page":"4420","article-title":"Pig conceptuses secrete estrogen and interferons to differentially regulate uterine STAT1 in a temporal and cell type-specific manner","volume":"148","author":"Joyce MM","year":"2007b","unstructured":"Joyce MM\n, \nBurghardt RC\n, \nGeisert RD\n, et al.\n 2007b Pig conceptuses secrete estrogen and interferons to differentially regulate uterine STAT1 in a temporal and cell type-specific manner. Endocrinology 148 4420\u20134431. (https:\/\/doi.org\/10.1210\/en.2007-0505)"},{"key":"bib23","series-title":"J Immunol","first-page":"2494","article-title":"Uterine MHC class I molecules and \u03b22-microglobulin are regulated by progesterone and conceptus interferons during pig pregnancy","volume":"181","author":"Joyce MM","year":"2008","unstructured":"Joyce MM\n, \nBurghardt JR\n, \nBurghardt RC\n, et al.\n 2008 Uterine MHC class I molecules and \u03b22-microglobulin are regulated by progesterone and conceptus interferons during pig pregnancy. J Immunol 181 2494\u20132505. (https:\/\/doi.org\/10.4049\/jimmunol.181.4.2494)"},{"key":"bib24","series-title":"Endocrinology","first-page":"2303","article-title":"Keratinocyte growth factor is up-regulated by estrogen in the porcine uterine endometrium and functions in trophectoderm cell proliferation and differentiation","volume":"142","author":"Ka H","year":"2001","unstructured":"Ka H\n, \nJaeger LA\n, \nJohnson GA\n, et al.\n 2001 Keratinocyte growth factor is up-regulated by estrogen in the porcine uterine endometrium and functions in trophectoderm cell proliferation and differentiation. Endocrinology 142 2303\u20132310. (https:\/\/doi.org\/10.1210\/endo.142.6.8194)"},{"key":"bib25","series-title":"J Anim Sci Biotechnol","first-page":"44","article-title":"Endometrial response to conceptus-derived estrogen and interleukin-1\u03b2 at the time of implantation in pigs","volume":"9","author":"Ka H","year":"2018","unstructured":"Ka H\n, \nSeo H\n, \nChoi Y\n, et al.\n 2018 Endometrial response to conceptus-derived estrogen and interleukin-1\u03b2 at the time of implantation in pigs. J Anim Sci Biotechnol 9 44. (https:\/\/doi.org\/10.1186\/s40104-018-0259-8)"},{"key":"bib26","series-title":"Mol Reprod Dev","first-page":"362","article-title":"Expression of the vascular endothelial growth factor-receptor system in the porcine endometrium throughout the estrous cycle and early pregnancy","volume":"75","author":"Kaczmarek MM","year":"2008","unstructured":"Kaczmarek MM\n, \nWaclawik A\n, \nBlitek A\n, et al.\n 2008 Expression of the vascular endothelial growth factor-receptor system in the porcine endometrium throughout the estrous cycle and early pregnancy. Mol Reprod Dev 75 362\u2013372. (https:\/\/doi.org\/10.1002\/mrd.20815)"},{"key":"bib27","series-title":"Theriogenology","first-page":"169","article-title":"Prostaglandin F2\u03b1 stimulates angiogenesis at the embryo\u2013maternal interface during early pregnancy in the pig","volume":"142","author":"Kaczynski P","year":"2020","unstructured":"Kaczynski P\n, \nGoryszewska E\n, \nBaryla M\n, et al.\n 2020 Prostaglandin F2\u03b1 stimulates angiogenesis at the embryo\u2013maternal interface during early pregnancy in the pig. Theriogenology 142 169\u2013176. (https:\/\/doi.org\/10.1016\/j.theriogenology.2019.09.046)"},{"key":"bib28","series-title":"Mol Reprod Dev","first-page":"658","article-title":"Novel insights into conceptus-maternal signaling during pregnancy establishment in pigs","volume":"90","author":"Kaczynski P","year":"2023","unstructured":"Kaczynski P\n, \nGoryszewska-Szczurek E\n, \nBaryla M\n, et al.\n 2023 Novel insights into conceptus-maternal signaling during pregnancy establishment in pigs. Mol Reprod Dev 90 658\u2013672. (https:\/\/doi.org\/10.1002\/mrd.23567)"},{"key":"bib29","series-title":"Proc Natl Acad Sci U S A","first-page":"4342","article-title":"CD40 on human endothelial cells: inducibility by cytokines and functional regulation of adhesion molecule expression","volume":"92","author":"Karmann K","year":"1995","unstructured":"Karmann K\n, \nHughes CC\n, \nSchechner J\n, et al.\n 1995 CD40 on human endothelial cells: inducibility by cytokines and functional regulation of adhesion molecule expression. Proc Natl Acad Sci U S A 92 4342\u20134346. (https:\/\/doi.org\/10.1073\/pnas.92.10.4342)"},{"key":"bib30","series-title":"J Immunol","first-page":"4917","article-title":"Stimulation of CD40 with purified soluble gp39 induces proinflammatory responses in human monocytes","volume":"155","author":"Kiener PA","year":"1995","unstructured":"Kiener PA\n, \nMoran-Davis P\n, \nRankin BM\n, et al.\n 1995 Stimulation of CD40 with purified soluble gp39 induces proinflammatory responses in human monocytes. J Immunol 155 4917\u20134925. (https:\/\/doi.org\/10.4049\/jimmunol.155.10.4917)"},{"key":"bib31","series-title":"Biol Reprod","first-page":"43","article-title":"Swine leukocyte antigen-DQ expression and its regulation by interferon-gamma at the maternal\u2013fetal interface in pigs","volume":"86","author":"Kim M","year":"2012","unstructured":"Kim M\n, \nSeo H\n, \nChoi Y\n, et al.\n 2012 Swine leukocyte antigen-DQ expression and its regulation by interferon-gamma at the maternal\u2013fetal interface in pigs. Biol Reprod 86 43. (https:\/\/doi.org\/10.1095\/biolreprod.111.094011)"},{"key":"bib32","series-title":"J Clin Endocrinol Metab","first-page":"405","article-title":"Cd40 expression in uterine tissues: a key regulator of cytokine expression by fibroblasts","volume":"86","author":"King AE","year":"2001","unstructured":"King AE\n, \nKelly RW\n, \nCritchley HO\n, et al.\n 2001 Cd40 expression in uterine tissues: a key regulator of cytokine expression by fibroblasts. J Clin Endocrinol Metab 86 405\u2013412. (https:\/\/doi.org\/10.1210\/jcem.86.1.7133)"},{"key":"bib33","series-title":"Proc Natl Acad Sci U S A","first-page":"5205","article-title":"CD40 ligand (CD154) stimulation of macrophages to produce HIV-1-suppressive \u03b2-chemokines","volume":"95","author":"Kornbluth RS","year":"1998","unstructured":"Kornbluth RS\n, \nKee K\n & \nRichman DD\n\n 1998 CD40 ligand (CD154) stimulation of macrophages to produce HIV-1-suppressive \u03b2-chemokines. Proc Natl Acad Sci U S A 95 5205\u20135210. (https:\/\/doi.org\/10.1073\/pnas.95.9.5205)"},{"key":"bib34","series-title":"Immunology","first-page":"441","article-title":"Biological function of CD40 on human endothelial cells: costimulation with CD40 ligand and interleukin-4 selectively induces expression of vascular cell adhesion molecule-1 and P-selectin resulting in preferential adhesion of lymphocytes","volume":"100","author":"Kotowicz K","year":"2000","unstructured":"Kotowicz K\n, \nDixon GL\n, \nKlein NJ\n, et al.\n 2000 Biological function of CD40 on human endothelial cells: costimulation with CD40 ligand and interleukin-4 selectively induces expression of vascular cell adhesion molecule-1 and P-selectin resulting in preferential adhesion of lymphocytes. Immunology 100 441\u2013448. (https:\/\/doi.org\/10.1046\/j.1365-2567.2000.00061.x)"},{"key":"bib35","series-title":"J Reprod Fertil","first-page":"469","article-title":"Production of two species of interferon by large white and Meishan pig conceptuses during the peri-attachment period","volume":"91","author":"La Bonnardiere C","year":"1991","unstructured":"La Bonnardiere C\n, \nMartinat-Botte F\n, \nTerqui M\n, et al.\n 1991 Production of two species of interferon by large white and Meishan pig conceptuses during the peri-attachment period. J Reprod Fertil 91 469\u2013478. (https:\/\/doi.org\/10.1530\/jrf.0.0910469)"},{"key":"bib36","series-title":"Biochimie","first-page":"779","article-title":"Interferon-delta: the first member of a novel type I interferon family","volume":"80","author":"Lefevre F","year":"1998","unstructured":"Lefevre F\n, \nGuillomot M\n, \nD'Andrea S\n, et al.\n 1998 Interferon-delta: the first member of a novel type I interferon family. Biochimie 80 779\u2013788. (https:\/\/doi.org\/10.1016\/s0300-9084(99)80030-3)"},{"key":"bib37","series-title":"J Immunol","first-page":"4681","article-title":"Depletion of invariant NKT cells reduces inflammation-induced preterm delivery in mice","volume":"188","author":"Li LP","year":"2012","unstructured":"Li LP\n, \nFang YC\n, \nDong GF\n, et al.\n 2012 Depletion of invariant NKT cells reduces inflammation-induced preterm delivery in mice. J Immunol 188 4681\u20134689. (https:\/\/doi.org\/10.4049\/jimmunol.1102628)"},{"key":"bib38","series-title":"Methods","first-page":"402","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2-\u0394\u0394CT method","volume":"25","author":"Livak KJ","year":"2001","unstructured":"Livak KJ\n & \nSchmittgen TD\n\n 2001 Analysis of relative gene expression data using real-time quantitative PCR and the 2-\u0394\u0394CT method. Methods 25 402\u2013408. (https:\/\/doi.org\/10.1006\/meth.2001.1262)"},{"key":"bib39","series-title":"Int J Gynaecol Obstet","first-page":"257","article-title":"Upregulation of the CD40-CD40 ligand system in pre-eclampsia","volume":"105","author":"Lukanov TH","year":"2009","unstructured":"Lukanov TH\n, \nBojinova SI\n, \nEmin AL\n, et al.\n 2009 Upregulation of the CD40-CD40 ligand system in pre-eclampsia. Int J Gynaecol Obstet 105 257\u2013260. (https:\/\/doi.org\/10.1016\/j.ijgo.2009.01.034)"},{"key":"bib40","series-title":"Semin Immunol","first-page":"265","article-title":"The role of CD40 and CD154\/CD40L in dendritic cells","volume":"21","author":"Ma DY","year":"2009","unstructured":"Ma DY\n & \nClark EA\n\n 2009 The role of CD40 and CD154\/CD40L in dendritic cells. Semin Immunol 21 265\u2013272. (https:\/\/doi.org\/10.1016\/j.smim.2009.05.010)"},{"key":"bib41","series-title":"J Immunol","first-page":"3952","article-title":"Activation of human monocytes through CD40 induces matrix metalloproteinases","volume":"156","author":"Malik N","year":"1996","unstructured":"Malik N\n, \nGreenfield BW\n, \nWahl AF\n, et al.\n 1996 Activation of human monocytes through CD40 induces matrix metalloproteinases. J Immunol 156 3952\u20133960. (https:\/\/doi.org\/10.4049\/jimmunol.156.10.3952)"},{"key":"bib42","series-title":"Nat Med","first-page":"540","article-title":"Reciprocal CD40 signals through p38MAPK and ERK-1\/2 induce counteracting immune responses","volume":"10","author":"Mathur RK","year":"2004","unstructured":"Mathur RK\n, \nAwasthi A\n, \nWadhone P\n, et al.\n 2004 Reciprocal CD40 signals through p38MAPK and ERK-1\/2 induce counteracting immune responses. Nat Med 10 540\u2013544. (https:\/\/doi.org\/10.1038\/nm1045)"},{"key":"bib43","series-title":"Hypertens Res","first-page":"407","article-title":"Immune activation during the implantation phase causes preeclampsia-like symptoms via the CD40-CD40 ligand pathway in pregnant mice","volume":"39","author":"Matsubara K","year":"2016","unstructured":"Matsubara K\n, \nMatsubara Y\n, \nMori M\n, et al.\n 2016 Immune activation during the implantation phase causes preeclampsia-like symptoms via the CD40-CD40 ligand pathway in pregnant mice. Hypertens Res 39 407\u2013414. (https:\/\/doi.org\/10.1038\/hr.2015.160)"},{"key":"bib44","series-title":"Biol Reprod","first-page":"1018","article-title":"Pig conceptuses secrete interferon gamma to recruit T cells to the endometrium during the peri-implantation period","volume":"103","author":"McLendon BA","year":"2020","unstructured":"McLendon BA\n, \nSeo H\n, \nKramer AC\n, et al.\n 2020 Pig conceptuses secrete interferon gamma to recruit T cells to the endometrium during the peri-implantation period. Biol Reprod 103 1018\u20131029. (https:\/\/doi.org\/10.1093\/biolre\/ioaa132)"},{"key":"bib45","series-title":"Blood","first-page":"3801","article-title":"Ligation of CD40 induces the expression of vascular endothelial growth factor by endothelial cells and monocytes and promotes angiogenesis in vivo","volume":"96","author":"Melter M","year":"2000","unstructured":"Melter M\n, \nReinders ME\n, \nSho M\n, et al.\n 2000 Ligation of CD40 induces the expression of vascular endothelial growth factor by endothelial cells and monocytes and promotes angiogenesis in vivo. Blood 96 3801\u20133808. (https:\/\/doi.org\/10.1182\/blood.v96.12.3801.h8003801_3801_3808)"},{"key":"bib46","series-title":"Reproduction","first-page":"503","article-title":"IFNT-independent effects of intrauterine extracellular vesicles (EVs) in cattle","volume":"159","author":"Nakamura K","year":"2020","unstructured":"Nakamura K\n, \nKusama K\n, \nIdeta A\n, et al.\n 2020 IFNT-independent effects of intrauterine extracellular vesicles (EVs) in cattle. Reproduction 159 503\u2013511. (https:\/\/doi.org\/10.1530\/REP-19-0314)"},{"key":"bib47","series-title":"Immunity","first-page":"415","article-title":"CD40 and its ligand in host defense","volume":"4","author":"Noelle RJ","year":"1996","unstructured":"Noelle RJ\n\n 1996 CD40 and its ligand in host defense. Immunity 4 415\u2013419. (https:\/\/doi.org\/10.1016\/s1074-7613(00)80408-2)"},{"key":"bib48","series-title":"Annu Rev Immunol","first-page":"307","article-title":"CD40\/CD154 interactions at the interface of tolerance and immunity","volume":"22","author":"Quezada SA","year":"2004","unstructured":"Quezada SA\n, \nJarvinen LZ\n, \nLind EF\n, et al.\n 2004 CD40\/CD154 interactions at the interface of tolerance and immunity. Annu Rev Immunol 22 307\u2013328. (https:\/\/doi.org\/10.1146\/annurev.immunol.22.012703.104533)"},{"key":"bib49","series-title":"Platelets","first-page":"441","article-title":"Preliminary evidence for a matrix metalloproteinase-2 (MMP-2)-dependent shedding of soluble CD40 ligand (sCD40L) from activated platelets","volume":"20","author":"Reinboldt S","year":"2009","unstructured":"Reinboldt S\n, \nWenzel F\n, \nRauch BH\n, et al.\n 2009 Preliminary evidence for a matrix metalloproteinase-2 (MMP-2)-dependent shedding of soluble CD40 ligand (sCD40L) from activated platelets. Platelets 20 441\u2013444. (https:\/\/doi.org\/10.1080\/09537100903096684)"},{"key":"bib50","series-title":"Biol Reprod","first-page":"1251","article-title":"Characterization of the interleukin-1beta system during porcine trophoblastic elongation and early placental attachment","volume":"69","author":"Ross JW","year":"2003","unstructured":"Ross JW\n, \nMalayer JR\n, \nRitchey JW\n, et al.\n 2003 Characterization of the interleukin-1beta system during porcine trophoblastic elongation and early placental attachment. Biol Reprod 69 1251\u20131259. (https:\/\/doi.org\/10.1095\/biolreprod.103.015842)"},{"key":"bib51","series-title":"Transplantation","first-page":"127","article-title":"Human CD154 induces activation of porcine endothelial cells and up-regulation of MHC class II expression","volume":"72","author":"Rushworth SA","year":"2001","unstructured":"Rushworth SA\n, \nBravery CA\n & \nThompson S\n\n 2001 Human CD154 induces activation of porcine endothelial cells and up-regulation of MHC class II expression. Transplantation 72 127\u2013132. (https:\/\/doi.org\/10.1097\/00007890-200107150-00025)"},{"key":"bib52","series-title":"Biol Reprod","first-page":"84","article-title":"Deep sequencing of the porcine endometrial transcriptome on day 14 of pregnancy","volume":"88","author":"Samborski A","year":"2013","unstructured":"Samborski A\n, \nGraf A\n, \nKrebs S\n, et al.\n 2013 Deep sequencing of the porcine endometrial transcriptome on day 14 of pregnancy. Biol Reprod 88 84. (https:\/\/doi.org\/10.1095\/biolreprod.113.107870)"},{"key":"bib53","series-title":"Cell Mol Life Sci","first-page":"4","article-title":"The CD40\/CD154 receptor\/ligand dyad","volume":"58","author":"Schonbeck U","year":"2001","unstructured":"Schonbeck U\n & \nLibby P\n\n 2001 The CD40\/CD154 receptor\/ligand dyad. Cell Mol Life Sci 58 4\u201343. (https:\/\/doi.org\/10.1007\/pl00000776)"},{"key":"bib54","series-title":"Endocrinology","first-page":"6166","article-title":"Analysis of lysophosphatidic acid (LPA) receptor and LPA-induced endometrial prostaglandin-endoperoxide synthase 2 expression in the porcine uterus","volume":"149","author":"Seo H","year":"2008","unstructured":"Seo H\n, \nKim M\n, \nChoi Y\n, et al.\n 2008 Analysis of lysophosphatidic acid (LPA) receptor and LPA-induced endometrial prostaglandin-endoperoxide synthase 2 expression in the porcine uterus. Endocrinology 149 6166\u20136175. (https:\/\/doi.org\/10.1210\/en.2008-0354)"},{"key":"bib55","series-title":"J Reprod Immunol","article-title":"Tryptophan metabolism and immune regulation in the human placenta","volume":"147","author":"Silvano A","year":"2021","unstructured":"Silvano A\n, \nSeravalli V\n, \nStrambi N\n, et al.\n 2021 Tryptophan metabolism and immune regulation in the human placenta. J Reprod Immunol 147 103361. (https:\/\/doi.org\/10.1016\/j.jri.2021.103361)"},{"key":"bib56","series-title":"J Immunol","first-page":"8","article-title":"Impaired T cell-mediated macrophage activation in CD40 ligand-deficient mice","volume":"156","author":"Stout RD","year":"1996","unstructured":"Stout RD\n, \nSuttles J\n, \nXu J\n, et al.\n 1996 Impaired T cell-mediated macrophage activation in CD40 ligand-deficient mice. J Immunol 156 8\u201311. (https:\/\/doi.org\/10.4049\/jimmunol.156.1.8)"},{"key":"bib57","series-title":"J Leukoc Biol","first-page":"877","article-title":"Lymphocyte contributions to altered endometrial angiogenesis during early and midgestation fetal loss","volume":"82","author":"Tayade C","year":"2007","unstructured":"Tayade C\n, \nFang Y\n, \nHilchie D\n, et al.\n 2007 Lymphocyte contributions to altered endometrial angiogenesis during early and midgestation fetal loss. J Leukoc Biol 82 877\u2013886. (https:\/\/doi.org\/10.1189\/jlb.0507330)"},{"key":"bib58","series-title":"J Leukoc Biol","first-page":"2","article-title":"CD40-CD40 ligand","volume":"67","author":"van Kooten C","year":"2000","unstructured":"van Kooten C\n & \nBanchereau J\n\n 2000 CD40-CD40 ligand. J Leukoc Biol 67 2\u201317. (https:\/\/doi.org\/10.1002\/jlb.67.1.2)"},{"key":"bib59","series-title":"Reproduction","first-page":"389","article-title":"Novel insights into the mechanisms of pregnancy establishment: regulation of prostaglandin synthesis and signaling in the pig","volume":"142","author":"Waclawik A","year":"2011","unstructured":"Waclawik A\n\n 2011 Novel insights into the mechanisms of pregnancy establishment: regulation of prostaglandin synthesis and signaling in the pig. Reproduction 142 389\u2013399. (https:\/\/doi.org\/10.1530\/rep-11-0033)"},{"key":"bib60","series-title":"Eur J Immunol","first-page":"3148","article-title":"Role of the CD40-CD40 ligand interaction in CD4+ T cell contact-dependent activation of monocyte interleukin-1 synthesis","volume":"24","author":"Wagner DH Jr","year":"1994","unstructured":"Wagner DH Jr\n, \nStout RD\n & \nSuttles J\n\n 1994 Role of the CD40-CD40 ligand interaction in CD4+ T cell contact-dependent activation of monocyte interleukin-1 synthesis. Eur J Immunol 24 3148\u20133154. (https:\/\/doi.org\/10.1002\/eji.1830241235)"},{"key":"bib61","series-title":"Biochem Biophys Res Commun","first-page":"808","article-title":"Induction and regulation of matrix metalloproteinase-12 by cytokines and CD40 signaling in monocyte\/macrophages","volume":"269","author":"Wu L","year":"2000","unstructured":"Wu L\n, \nFan J\n, \nMatsumoto S\n, et al.\n 2000 Induction and regulation of matrix metalloproteinase-12 by cytokines and CD40 signaling in monocyte\/macrophages. Biochem Biophys Res Commun 269 808\u2013815. (https:\/\/doi.org\/10.1006\/bbrc.2000.2368)"},{"key":"bib62","series-title":"Mol Reprod Dev","first-page":"1993","article-title":"Analysis of interferon-gamma receptor IFNGR1 and IFNGR2 expression and regulation at the maternal\u2013conceptus interface and the role of interferon-gamma on endometrial expression of interferon signaling molecules during early pregnancy in pigs","volume":"86","author":"Yoo I","year":"2019","unstructured":"Yoo I\n, \nSeo H\n, \nChoi Y\n, et al.\n 2019 Analysis of interferon-gamma receptor IFNGR1 and IFNGR2 expression and regulation at the maternal\u2013conceptus interface and the role of interferon-gamma on endometrial expression of interferon signaling molecules during early pregnancy in pigs. Mol Reprod Dev 86 1993\u20132004. (https:\/\/doi.org\/10.1002\/mrd.23287)"},{"key":"bib63","series-title":"Theriogenology","first-page":"114","article-title":"Transcriptomic analysis of interferon-gamma-regulated genes in endometrial explants and their possible role in regulating maternal endometrial immunity during the implantation period in pigs, a true epitheliochorial placentation species","volume":"155","author":"Yoo I","year":"2020a","unstructured":"Yoo I\n, \nKim D\n, \nHan J\n, et al.\n 2020a Transcriptomic analysis of interferon-gamma-regulated genes in endometrial explants and their possible role in regulating maternal endometrial immunity during the implantation period in pigs, a true epitheliochorial placentation species. Theriogenology 155 114\u2013124. (https:\/\/doi.org\/10.1016\/j.theriogenology.2020.05.045)"},{"key":"bib64","series-title":"Biol Reprod","first-page":"828","article-title":"Uterine epithelial expression of the tumor necrosis factor superfamily: a strategy for immune privilege during pregnancy in a true epitheliochorial placentation species","volume":"102","author":"Yoo I","year":"2020b","unstructured":"Yoo I\n, \nKye YC\n, \nHan J\n, et al.\n 2020b Uterine epithelial expression of the tumor necrosis factor superfamily: a strategy for immune privilege during pregnancy in a true epitheliochorial placentation species. Biol Reprod 102 828\u2013842. (https:\/\/doi.org\/10.1093\/biolre\/ioz233)"},{"key":"bib65","series-title":"Anim Biosci","first-page":"1167","article-title":"Matrix metalloproteinases: expression and regulation in the endometrium during the estrous cycle and at the maternal\u2013conceptus interface during pregnancy in pigs","volume":"36","author":"Yoo I","year":"2023","unstructured":"Yoo I\n, \nLee S\n, \nCheon Y\n, et al.\n 2023 Matrix metalloproteinases: expression and regulation in the endometrium during the estrous cycle and at the maternal\u2013conceptus interface during pregnancy in pigs. Anim Biosci 36 1167\u20131179. 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