{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T07:39:55Z","timestamp":1790840395795,"version":"4.1.0"},"reference-count":55,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2021,9,23]],"date-time":"2021-09-23T00:00:00Z","timestamp":1632355200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Med."],"abstract":"<jats:p>The prevalence of end-stage liver diseases has reached very high levels globally. The election treatment for affected patients is orthotopic liver transplantation, which is a very complex procedure, and due to the limited number of suitable organ donors, considerable research is being done on alternative therapeutic options. For instance, the use of cell therapy, such as the transplantation of hepatocytes to promote liver repair\/regeneration, has been explored, but standardized protocols to produce suitable human hepatocytes are still limited. On the other hand, liver progenitor and multipotent stem cells offer potential cell sources that could be used clinically. Different studies have reported regarding the therapeutic effects of transplanted mesenchymal stromal\/stem cells (MSCs) on end-stage liver diseases. Moreover, it has been shown that delivery of MSC-derived conditioned medium (MSC-CM) can reduce cell death and enhance liver proliferation in fulminant hepatic failure. Therefore, it is believed that MSC-CM contains many factors that probably support liver regeneration. In our work, we used an <jats:italic>in vitro<\/jats:italic> model of human liver organoids to study if the paracrine components secreted by human amnion-derived MSCs (hAMSCs) affected liver stem\/progenitor cell differentiation. In particular, we differentiated liver organoids derived from bipotent EpCAM<jats:sup>+<\/jats:sup> human liver cells and tested the effects of hAMSC secretome, derived from both two-dimensional (2D) and three-dimensional (3D) hAMSC cultures, on that model. Our analysis showed that conditioned medium (CM) produced by 3D hAMSCs was able to induce an over-expression of mature hepatocyte markers, such as ALB, NTCP, and CYP3A4, compared with both 2D hAMSC cultures and the conventional differentiation medium (DM). These data were confirmed by the over-production of ALB protein and over-activity of CYP3A4 observed in organoids grown in 3D hAMSC-CM. Liver repair dysfunction plays a role in the development of liver diseases, and effective repair likely requires the normal functioning of liver stem\/progenitor cells. Herein, we showed that hAMSC-CM produced mainly by 3D cultures had the potential to increase hepatic stem\/progenitor cell differentiation, demonstrating that soluble factors secreted by those cells are potentially responsible for the reaction. This work shows a potential approach to improve liver repair\/regeneration also in a transplantation setting.<\/jats:p>","DOI":"10.3389\/fmed.2021.746298","type":"journal-article","created":{"date-parts":[[2021,9,23]],"date-time":"2021-09-23T05:48:33Z","timestamp":1632376113000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":23,"title":["Amnion-Derived Mesenchymal Stromal\/Stem Cell Paracrine Signals Potentiate Human Liver Organoid Differentiation: Translational Implications for Liver Regeneration"],"prefix":"10.3389","volume":"8","author":[{"given":"Antonio","family":"Lo Nigro","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alessia","family":"Gallo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matteo","family":"Bulati","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Giampiero","family":"Vitale","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Diego Sebastian","family":"Paini","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mariangela","family":"Pampalone","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniele","family":"Galvagno","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pier Giulio","family":"Conaldi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vitale","family":"Miceli","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2021,9,23]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1016\/S2468-1253(19)30349-8","article-title":"The global, regional, and national burden of cirrhosis by cause in 195 countries and territories, 1990\u20132017: a systematic analysis for the Global Burden of Disease Study 2017","volume":"5","author":"Collaborators","year":"2020","journal-title":"Lancet Gastroenterol Hepatol"},{"key":"B2","doi-asserted-by":"publisher","first-page":"584","DOI":"10.1002\/hep.31047","article-title":"The von Willebrand factor facilitates model for end-stage liver disease-independent risk stratification on the waiting list for liver transplantation","volume":"72","author":"Gyori","year":"2020","journal-title":"Hepatology"},{"key":"B3","doi-asserted-by":"publisher","first-page":"1307","DOI":"10.1007\/s00018-017-2713-8","article-title":"Liver cell therapy: is this the end of the beginning?","volume":"75","author":"Alwahsh","year":"2018","journal-title":"Cell Mol Life Sci"},{"key":"B4","doi-asserted-by":"publisher","first-page":"824","DOI":"10.1097\/TP.0000000000003318","article-title":"Live donor liver transplantation in the united states: impact of share 35 on live donor utilization","volume":"105","author":"Braun","year":"2021","journal-title":"Transplantation"},{"key":"B5","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1097\/SLA.0000000000002383","article-title":"Expanding the margins: high volume utilization of marginal liver grafts among >2000 liver transplants at a single institution","volume":"266","author":"Halazun","year":"2017","journal-title":"Ann Surg"},{"key":"B6","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1007\/s13770-019-00178-y","article-title":"Stem cell-based therapies for liver diseases: an overview and update","volume":"16","author":"Wang","year":"2019","journal-title":"Tissue Engineer Regenerat Med"},{"key":"B7","doi-asserted-by":"publisher","first-page":"2593745","DOI":"10.1155\/2018\/2593745","article-title":"Domino hepatocyte transplantation: a therapeutic alternative for the treatment of acute liver failure","volume":"2018","author":"Cardoso","year":"2018","journal-title":"Cana J Gastroenterol Hepatol"},{"key":"B8","doi-asserted-by":"publisher","first-page":"232","DOI":"10.1038\/pr.2017.284","article-title":"Human hepatocyte transplantation for liver disease: current status and future perspectives","volume":"83","author":"Iansante","year":"2018","journal-title":"Pediatr Res"},{"key":"B9","doi-asserted-by":"publisher","first-page":"987","DOI":"10.1016\/j.jhep.2016.12.017","article-title":"Host conditioning and rejection monitoring in hepatocyte transplantation in humans","volume":"66","author":"Soltys","year":"2017","journal-title":"J Hepatol"},{"key":"B10","doi-asserted-by":"publisher","first-page":"1068","DOI":"10.1016\/j.jhep.2016.01.004","article-title":"Transplantation of a human iPSC-derived hepatocyte sheet increases survival in mice with acute liver failure","volume":"64","author":"Nagamoto","year":"2016","journal-title":"J Hepatol"},{"key":"B11","doi-asserted-by":"publisher","first-page":"2","DOI":"10.3390\/biomedicines4010002","article-title":"Stem cell therapies for treatment of liver disease","volume":"4","author":"Nicolas","year":"2016","journal-title":"Biomedicines"},{"key":"B12","doi-asserted-by":"publisher","first-page":"987","DOI":"10.15252\/embj.201490756","article-title":"Application of biomaterials to advance induced pluripotent stem cell research and therapy","volume":"34","author":"Tong","year":"2015","journal-title":"EMBO J"},{"key":"B13","doi-asserted-by":"publisher","first-page":"30","DOI":"10.1186\/s40164-018-0122-9","article-title":"Organoid technology in disease modelling, drug development, personalized treatment and regeneration medicine","volume":"7","author":"Xu","year":"2018","journal-title":"Exp Hematol Oncol"},{"key":"B14","doi-asserted-by":"publisher","first-page":"1586","DOI":"10.1016\/j.cell.2016.05.082","article-title":"Modeling development and disease with organoids","volume":"165","author":"Clevers","year":"2016","journal-title":"Cell"},{"key":"B15","doi-asserted-by":"publisher","first-page":"299","DOI":"10.1016\/j.cell.2014.11.050","article-title":"Long-term culture of genome-stable bipotent stem cells from adult human liver","volume":"160","author":"Huch","year":"2015","journal-title":"Cell"},{"key":"B16","doi-asserted-by":"publisher","first-page":"329","DOI":"10.1038\/nrg2318","article-title":"Genetic programming of liver and pancreas progenitors: lessons for stem-cell differentiation","volume":"9","author":"Zaret","year":"2008","journal-title":"Nat Rev Genet"},{"key":"B17","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1016\/j.scr.2010.02.002","article-title":"Generating hepatic cell lineages from pluripotent stem cells for drug toxicity screening","volume":"5","author":"Baxter","year":"2010","journal-title":"Stem Cell Res"},{"key":"B18","doi-asserted-by":"publisher","first-page":"S157","DOI":"10.1016\/j.jhep.2015.02.040","article-title":"Cell therapy for liver disease: from liver transplantation to cell factory","volume":"62","author":"Forbes","year":"2015","journal-title":"J Hepatol"},{"key":"B19","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1007\/978-1-4939-7471-9_2","article-title":"induced pluripotent stem cells in disease modeling and gene identification","volume":"1706","author":"Kumar","year":"2018","journal-title":"Methods Mol Biol"},{"key":"B20","doi-asserted-by":"publisher","first-page":"559","DOI":"10.1126\/science.1063889","article-title":"Liver organogenesis promoted by endothelial cells prior to vascular function","volume":"294","author":"Matsumoto","year":"2001","journal-title":"Science"},{"key":"B21","doi-asserted-by":"publisher","first-page":"262","DOI":"10.1196\/annals.1386.028","article-title":"Metabolic profiles of androgens in malignant human liver cell lines","volume":"1089","author":"Granata","year":"2006","journal-title":"Ann N Y Acad Sci"},{"key":"B22","doi-asserted-by":"publisher","first-page":"511","DOI":"10.1111\/jcmm.12482","article-title":"The multiple functional roles of mesenchymal stem cells in participating in treating liver diseases","volume":"19","author":"Liu","year":"2015","journal-title":"J Cell Mol Med"},{"key":"B23","doi-asserted-by":"publisher","first-page":"1199","DOI":"10.1097\/MEG.0b013e32832a1f6c","article-title":"Improvement of liver function in liver cirrhosis patients after autologous mesenchymal stem cell injection: a phase I-II clinical trial","volume":"21","author":"Kharaziha","year":"2009","journal-title":"Eur J Gastroenterol Hepatol"},{"key":"B24","doi-asserted-by":"publisher","first-page":"9156560","DOI":"10.1155\/2018\/9156560","article-title":"The effects of conditioned medium derived from mesenchymal stem cells cocultured with hepatocytes on damaged hepatocytes and acute liver failure in rats","volume":"2018","author":"Chen","year":"2018","journal-title":"Stem Cells Int"},{"key":"B25","doi-asserted-by":"publisher","first-page":"e941","DOI":"10.1371\/journal.pone.0000941","article-title":"Mesenchymal stem cell-derived molecules reverse fulminant hepatic failure","volume":"2","author":"Parekkadan","year":"2007","journal-title":"PLoS ONE"},{"key":"B26","doi-asserted-by":"publisher","first-page":"907","DOI":"10.1016\/j.jss.2013.02.009","article-title":"Mesenchymal stem cell-conditioned medium reduces liver injury and enhances regeneration in reduced-size rat liver transplantation","volume":"183","author":"Du","year":"2013","journal-title":"J Surg Res"},{"key":"B27","doi-asserted-by":"publisher","first-page":"763","DOI":"10.1007\/s00535-019-01599-1","article-title":"The mesenchymal stem cell secretome as an acellular regenerative therapy for liver disease","volume":"54","author":"Driscoll","year":"2019","journal-title":"J Gastroenterol"},{"key":"B28","doi-asserted-by":"publisher","first-page":"54","DOI":"10.3389\/fimmu.2020.00054","article-title":"The immunomodulatory properties of the human amnion-derived mesenchymal stromal\/stem cells are induced by INF-gamma produced by activated lymphomonocytes and are mediated by cell-to-cell contact and soluble factors","volume":"11","author":"Bulati","year":"2020","journal-title":"Front Immunol"},{"key":"B29","doi-asserted-by":"publisher","first-page":"853","DOI":"10.1007\/s12015-020-10006-9","article-title":"Specificity of 3D MSC spheroids microenvironment: impact on MSC behavior and properties","volume":"16","author":"Jaukovic","year":"2020","journal-title":"Stem Cell Rev Rep"},{"key":"B30","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1016\/j.bbrc.2019.11.069","article-title":"Comparative study of the production of soluble factors in human placenta-derived mesenchymal stromal\/stem cells grown in adherent conditions or as aggregates in a catheter-like device","volume":"522","author":"Miceli","year":"2020","journal-title":"Biochem Biophys Res Commun"},{"key":"B31","doi-asserted-by":"publisher","first-page":"763","DOI":"10.3390\/ijms22020763","article-title":"Therapeutic properties of mesenchymal stromal\/stem cells: the need of cell priming for cell-free therapies in regenerative medicine","volume":"22","author":"Miceli","year":"2021","journal-title":"Int J Mol Sci"},{"key":"B32","doi-asserted-by":"publisher","first-page":"510","DOI":"10.3390\/ijms22020510","article-title":"Conditioned medium from human amnion-derived mesenchymal stromal\/stem cells attenuating the effects of cold ischemia-reperfusion injury in an in vitro model using human alveolar epithelial cells","volume":"22","author":"Miceli","year":"2021","journal-title":"Int J Mol Sci"},{"key":"B33","doi-asserted-by":"publisher","first-page":"7486279","DOI":"10.1155\/2019\/7486279","article-title":"Comparison of immunosuppressive and angiogenic properties of human amnion-derived mesenchymal stem cells between 2D and 3D culture systems","volume":"2019","author":"Miceli","year":"2019","journal-title":"Stem Cells Int"},{"key":"B34","doi-asserted-by":"publisher","first-page":"1098","DOI":"10.1053\/j.gastro.2015.12.042","article-title":"Organoid models of human gastrointestinal development and disease","volume":"150","author":"Dedhia","year":"2016","journal-title":"Gastroenterology"},{"key":"B35","doi-asserted-by":"publisher","first-page":"958","DOI":"10.1242\/dev.140731","article-title":"Pluripotent stem cell-derived organoids: using principles of developmental biology to grow human tissues in a dish","volume":"144","author":"McCauley","year":"2017","journal-title":"Development"},{"key":"B36","doi-asserted-by":"publisher","first-page":"530674","DOI":"10.1155\/2012\/530674","article-title":"Amniotic stem cells: potential in regenerative medicine","volume":"2012","author":"Bani-Yaghoub","year":"2012","journal-title":"Stem Cells Int"},{"key":"B37","doi-asserted-by":"publisher","first-page":"275","DOI":"10.2217\/17460751.4.2.275","article-title":"Amniotic membrane and amniotic fluid-derived cells: potential tools for regenerative medicine?","volume":"4","author":"Parolini","year":"2009","journal-title":"Regenerat Med"},{"key":"B38","doi-asserted-by":"publisher","first-page":"879","DOI":"10.1080\/22221751.2019.1625728","article-title":"Down-regulation of cell membrane localized NTCP expression in proliferating hepatocytes prevents hepatitis B virus infection","volume":"8","author":"Yan","year":"2019","journal-title":"Emerg Microbes Infect"},{"key":"B39","doi-asserted-by":"publisher","first-page":"11452","DOI":"10.1073\/pnas.1106083108","article-title":"R-spondins function as ligands of the orphan receptors LGR4 and LGR5 to regulate Wnt\/beta-catenin signaling","volume":"108","author":"Carmon","year":"2011","journal-title":"Proc Natl Acad Sci USA"},{"key":"B40","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1016\/j.stem.2009.11.013","article-title":"Lgr5(+ve) stem cells drive self-renewal in the stomach and build long-lived gastric units in vitro","volume":"6","author":"Barker","year":"2010","journal-title":"Cell Stem Cell"},{"key":"B41","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1038\/nature11826","article-title":"In vitro expansion of single Lgr5+ liver stem cells induced by Wnt-driven regeneration","volume":"494","author":"Huch","year":"2013","journal-title":"Nature"},{"key":"B42","doi-asserted-by":"publisher","first-page":"1445","DOI":"10.1002\/hep.24457","article-title":"CXC chemokine signaling in the liver: impact on repair and regeneration","volume":"54","author":"Van Sweringen","year":"2011","journal-title":"Hepatology"},{"key":"B43","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1016\/j.stem.2019.04.005","article-title":"YAP, but not RSPO-LGR4\/5, signaling in biliary epithelial cells promotes a ductular reaction in response to liver injury","volume":"25","author":"Planas-Paz","year":"2019","journal-title":"Cell Stem Cell"},{"key":"B44","doi-asserted-by":"publisher","first-page":"1565","DOI":"10.1096\/fasebj.13.12.1565","article-title":"Novel CXCR2-dependent liver regenerative qualities of ELR-containing CXC chemokines","volume":"13","author":"Hogaboam","year":"1999","journal-title":"FASEB J"},{"key":"B45","doi-asserted-by":"publisher","first-page":"G203","DOI":"10.1152\/ajpgi.00193.2016","article-title":"Stromal cell-derived factor-1 (SDF-1) as a target in liver diseases","volume":"311","author":"Liepelt","year":"2016","journal-title":"Am J Physiol Gastrointest Liver Physiol"},{"key":"B46","doi-asserted-by":"publisher","first-page":"e96053","DOI":"10.1371\/journal.pone.0096053","article-title":"Synthesis of IL-6 by hepatocytes is a normal response to common hepatic stimuli","volume":"9","author":"Norris","year":"2014","journal-title":"PLoS ONE"},{"key":"B47","doi-asserted-by":"publisher","first-page":"836","DOI":"10.1038\/nrm1489","article-title":"Liver regeneration: from myth to mechanism","volume":"5","author":"Taub","year":"2004","journal-title":"Nat Rev Mol Cell Biol"},{"key":"B48","doi-asserted-by":"publisher","first-page":"1379","DOI":"10.1126\/science.274.5291.1379","article-title":"Liver failure and defective hepatocyte regeneration in interleukin-6-deficient mice","volume":"274","author":"Cressman","year":"1996","journal-title":"Science"},{"key":"B49","first-page":"15","article-title":"Expression of leukemia inhibitory factor and its receptor during liver regeneration in the adult rat","volume":"75","author":"Omori","year":"1996","journal-title":"Lab Investig"},{"key":"B50","doi-asserted-by":"publisher","first-page":"68","DOI":"10.3978\/j.issn.2304-3881.2012.10.11","article-title":"Expression of vascular endothelial growth factors and their receptors by hepatic progenitor cells in human liver diseases","volume":"2","author":"Franchitto","year":"2013","journal-title":"Hepatobiliary Surg Nutr"},{"key":"B51","doi-asserted-by":"publisher","first-page":"385","DOI":"10.3748\/wjg.v10.i3.385","article-title":"Expression of platelet-derived growth factor-BB in liver tissues of patients with chronic hepatitis B","volume":"10","author":"Lou","year":"2004","journal-title":"World J Gastroenterol"},{"key":"B52","doi-asserted-by":"publisher","first-page":"7879","DOI":"10.3892\/mmr.2017.7641","article-title":"PDGF signaling pathway in hepatic fibrosis pathogenesis and therapeutics (Review)","volume":"16","author":"Ying","year":"2017","journal-title":"Mol Med Rep"},{"key":"B53","doi-asserted-by":"publisher","first-page":"851","DOI":"10.1002\/stem.2575","article-title":"Concise review: MSC-derived exosomes for cell-free therapy","volume":"35","author":"Phinney","year":"2017","journal-title":"Stem Cells"},{"key":"B54","doi-asserted-by":"publisher","first-page":"1451","DOI":"10.1089\/scd.2019.0085","article-title":"Human mesenchymal stromal cell-derived extracellular vesicles improve liver regeneration after ischemia reperfusion injury in mice","volume":"28","author":"Anger","year":"2019","journal-title":"Stem Cells Dev"},{"key":"B55","doi-asserted-by":"publisher","first-page":"3715","DOI":"10.3390\/ijms19123715","article-title":"Liver-derived exosomes and their implications in liver pathobiology","volume":"19","author":"Sung","year":"2018","journal-title":"Int J Mol Sci"}],"container-title":["Frontiers in Medicine"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmed.2021.746298\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,9,23]],"date-time":"2021-09-23T05:48:40Z","timestamp":1632376120000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmed.2021.746298\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,9,23]]},"references-count":55,"alternative-id":["10.3389\/fmed.2021.746298"],"URL":"https:\/\/doi.org\/10.3389\/fmed.2021.746298","relation":{},"ISSN":["2296-858X"],"issn-type":[{"value":"2296-858X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,9,23]]},"article-number":"746298"}}