{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T15:41:46Z","timestamp":1788968506055,"version":"build-2803163510"},"reference-count":40,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2019,7,5]],"date-time":"2019-07-05T00:00:00Z","timestamp":1562284800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2019,7,5]],"date-time":"2019-07-05T00:00:00Z","timestamp":1562284800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004686","name":"Deanship of Scientific Research, King Faisal University","doi-asserted-by":"publisher","award":["186063"],"award-info":[{"award-number":["186063"]}],"id":[{"id":"10.13039\/501100004686","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["J Biomed Sci"],"published-print":{"date-parts":[[2019,12]]},"DOI":"10.1186\/s12929-019-0544-7","type":"journal-article","created":{"date-parts":[[2019,7,5]],"date-time":"2019-07-05T07:25:13Z","timestamp":1562311513000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":32,"title":["5\u2032-hydroxy Auraptene stimulates osteoblast differentiation of bone marrow-derived mesenchymal stem cells via a BMP-dependent mechanism"],"prefix":"10.1186","volume":"26","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0324-8055","authenticated-orcid":false,"given":"Basem M.","family":"Abdallah","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Enas M.","family":"Ali","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2019,7,5]]},"reference":[{"issue":"3","key":"544_CR1","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1016\/j.ecl.2015.05.002","volume":"44","author":"BC Lupsa","year":"2015","unstructured":"Lupsa BC, Insogna K. Bone health and osteoporosis. Endocrinol Metab Clin N Am. 2015;44(3):517\u201330.","journal-title":"Endocrinol Metab Clin N Am"},{"issue":"8","key":"544_CR2","doi-asserted-by":"publisher","first-page":"756","DOI":"10.1056\/NEJMoa0809493","volume":"361","author":"SR Cummings","year":"2009","unstructured":"Cummings SR, Martin JS, McClung MR, Siris ES, Eastell R, Reid IR, Delmas P, Zoog HB, Austin M, Wang A, et al. Denosumab for prevention of fractures in postmenopausal women with osteoporosis. N Engl J Med. 2009;361(8):756\u201365.","journal-title":"N Engl J Med"},{"issue":"3","key":"544_CR3","first-page":"284","volume":"15","author":"P Valverde","year":"2008","unstructured":"Valverde P. Pharmacotherapies to manage bone loss-associated diseases: a quest for the perfect benefit-to-risk ratio. CurrMedChem. 2008;15(3):284\u2013304.","journal-title":"CurrMedChem"},{"issue":"4","key":"544_CR4","doi-asserted-by":"publisher","first-page":"555","DOI":"10.1016\/j.coms.2015.07.001","volume":"27","author":"S Weinerman","year":"2015","unstructured":"Weinerman S, Usera GL. Antiresorptive therapies for osteoporosis. Oral Maxillofac Surg Clin North Am. 2015;27(4):555\u201360.","journal-title":"Oral Maxillofac Surg Clin North Am"},{"key":"544_CR5","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1007\/978-1-59745-104-8_6","volume":"455","author":"BM Abdallah","year":"2008","unstructured":"Abdallah BM, Ditzel N, Kassem M. Assessment of bone formation capacity using in vivo transplantation assays: procedure and tissue analysis. Methods Mol Biol. 2008;455:89\u2013100.","journal-title":"Methods Mol Biol"},{"key":"544_CR6","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1016\/j.bone.2014.07.028","volume":"70","author":"BM Abdallah","year":"2015","unstructured":"Abdallah BM, Jafari A, Zaher W, Qiu W, Kassem M. Skeletal (stromal) stem cells: an update on intracellular signaling pathways controlling osteoblast differentiation. Bone. 2015;70:28\u201336.","journal-title":"Bone"},{"issue":"1","key":"544_CR7","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1002\/jcp.21572","volume":"218","author":"BM Abdallah","year":"2009","unstructured":"Abdallah BM, Kassem M. The use of mesenchymal (skeletal) stem cells for treatment of degenerative diseases: current status and future perspectives. JCell Physiol. 2009;218(1):9\u201312.","journal-title":"JCell Physiol"},{"issue":"6","key":"544_CR8","doi-asserted-by":"publisher","first-page":"1023","DOI":"10.1242\/dev.102210","volume":"142","author":"P Bianco","year":"2015","unstructured":"Bianco P, Robey PG. Skeletal stem cells. Development. 2015;142(6):1023\u20137.","journal-title":"Development"},{"issue":"4","key":"544_CR9","doi-asserted-by":"publisher","first-page":"718","DOI":"10.1111\/j.1582-4934.2010.01224.x","volume":"15","author":"K Arvidson","year":"2011","unstructured":"Arvidson K, Abdallah BM, Applegate LA, Baldini N, Cenni E, Gomez-Barrena E, Granchi D, Kassem M, Konttinen YT, Mustafa K, et al. Bone regeneration and stem cells. J Cell Mol Med. 2011;15(4):718\u201346.","journal-title":"J Cell Mol Med"},{"key":"544_CR10","doi-asserted-by":"crossref","unstructured":"Zhu JJ, Jiang JG. Pharmacological and Nutritional Effects of Natural Coumarins and Their Structure-Activity Relationships. Mol Nutr Food Res. 2018:e1701073. [Epub ahead of print].","DOI":"10.1002\/mnfr.201701073"},{"issue":"9\u201310","key":"544_CR11","doi-asserted-by":"publisher","first-page":"693","DOI":"10.1515\/znc-2005-9-1006","volume":"60","author":"SM de Souza","year":"2005","unstructured":"de Souza SM, Delle Monache F, Smania A Jr. Antibacterial activity of coumarins. Zeitschrift fur Naturforschung C, Journal of biosciences. 2005;60(9\u201310):693\u2013700.","journal-title":"Zeitschrift fur Naturforschung C, Journal of biosciences"},{"issue":"11","key":"544_CR12","doi-asserted-by":"publisher","first-page":"1201","DOI":"10.1080\/13543776.2017.1360284","volume":"27","author":"A Detsi","year":"2017","unstructured":"Detsi A, Kontogiorgis C, Hadjipavlou-Litina D. Coumarin derivatives: an updated patent review (2015-2016). Expert opinion on therapeutic patents. 2017;27(11):1201\u201326.","journal-title":"Expert opinion on therapeutic patents"},{"issue":"4","key":"544_CR13","doi-asserted-by":"publisher","first-page":"437","DOI":"10.1517\/13543776.2012.678835","volume":"22","author":"C Kontogiorgis","year":"2012","unstructured":"Kontogiorgis C, Detsi A, Hadjipavlou-Litina D. Coumarin-based drugs: a patent review (2008 -- present). Expert opinion on therapeutic patents. 2012;22(4):437\u201354.","journal-title":"Expert opinion on therapeutic patents"},{"issue":"1\u20133","key":"544_CR14","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1016\/j.ejphar.2007.10.013","volume":"579","author":"CH Tang","year":"2008","unstructured":"Tang CH, Yang RS, Chien MY, Chen CC, Fu WM. Enhancement of bone morphogenetic protein-2 expression and bone formation by coumarin derivatives via p38 and ERK-dependent pathway in osteoblasts. Eur J Pharmacol. 2008;579(1\u20133):40\u20139.","journal-title":"Eur J Pharmacol"},{"issue":"2","key":"544_CR15","doi-asserted-by":"publisher","first-page":"2304","DOI":"10.1002\/jcb.27555","volume":"120","author":"Xifang Liu","year":"2018","unstructured":"Liu X, Gao X, Liu Y, Liang D, Fu T, Song Y, Zhao C, Dong B, Han W. Daphnetin inhibits RANKL-induced osteoclastogenesis in vitro. J Cell Biochem. 2018;120:2304-12.","journal-title":"Journal of Cellular Biochemistry"},{"issue":"6","key":"544_CR16","doi-asserted-by":"publisher","first-page":"588","DOI":"10.3390\/nu9060588","volume":"9","author":"Zhong-Rong Zhang","year":"2017","unstructured":"Zhang ZR, Leung WN, Li G, Kong SK, Lu X, Wong YM, Chan CW. Osthole enhances osteogenesis in osteoblasts by elevating transcription factor Osterix via cAMP\/CREB signaling in vitro and in vivo. Nutrients. 2017;9(6):588-601.","journal-title":"Nutrients"},{"issue":"2","key":"544_CR17","doi-asserted-by":"publisher","first-page":"158","DOI":"10.1002\/jor.21124","volume":"29","author":"RW Wong","year":"2011","unstructured":"Wong RW, Rabie AB. Effect of psoralen on bone formation. J Orthop Res. 2011;29(2):158\u201364.","journal-title":"J Orthop Res"},{"issue":"5","key":"544_CR18","doi-asserted-by":"publisher","first-page":"1662","DOI":"10.1182\/blood-2003-09-3070","volume":"103","author":"A Peister","year":"2004","unstructured":"Peister A, Mellad JA, Larson BL, Hall BM, Gibson LF, Prockop DJ. Adult stem cells from bone marrow (MSCs) isolated from different strains of inbred mice vary in surface epitopes, rates of proliferation, and differentiation potential. Blood. 2004;103(5):1662\u20138.","journal-title":"Blood"},{"key":"544_CR19","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1186\/s12929-017-0321-4","volume":"24","author":"BM Abdallah","year":"2017","unstructured":"Abdallah BM. Marrow adipocytes inhibit the differentiation of mesenchymal stem cells into osteoblasts via suppressing BMP-signaling. J Biomed Sci. 2017;24:11.","journal-title":"J Biomed Sci"},{"issue":"8","key":"544_CR20","first-page":"643","volume":"59","author":"F Meng","year":"2004","unstructured":"Meng F, Xiong Z, Sun Y, Li F. Coumarins from Cnidium monnieri (L.) and their proliferation stimulating activity on osteoblast-like UMR106 cells. Die Pharmazie. 2004;59(8):643\u20135.","journal-title":"Die Pharmazie"},{"issue":"1","key":"544_CR21","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1055\/s-2006-951724","volume":"73","author":"Q Zhang","year":"2007","unstructured":"Zhang Q, Qin L, He W, Van Puyvelde L, Maes D, Adams A, Zheng H, De Kimpe N. Coumarins from Cnidium monnieri and their antiosteoporotic activity. Planta Med. 2007;73(1):13\u20139.","journal-title":"Planta Med"},{"issue":"2","key":"544_CR22","doi-asserted-by":"publisher","first-page":"256","DOI":"10.1016\/j.bbrc.2011.01.021","volume":"405","author":"DZ Tang","year":"2011","unstructured":"Tang DZ, Yang F, Yang Z, Huang J, Shi Q, Chen D, Wang YJ. Psoralen stimulates osteoblast differentiation through activation of BMP signaling. Biochem Biophys Res Commun. 2011;405(2):256\u201361.","journal-title":"Biochem Biophys Res Commun"},{"issue":"3","key":"544_CR23","doi-asserted-by":"publisher","first-page":"527","DOI":"10.3892\/ijmm.2017.2880","volume":"39","author":"J Wang","year":"2017","unstructured":"Wang J, Li SF, Wang T, Sun CH, Wang L, Huang MJ, Chen J, Zheng SW, Wang N, Zhang YJ, et al. Isopsoralen-mediated suppression of bone marrow adiposity and attenuation of the adipogenic commitment of bone marrow-derived mesenchymal stem cells. Int J Mol Med. 2017;39(3):527\u201338.","journal-title":"Int J Mol Med"},{"issue":"7","key":"544_CR24","doi-asserted-by":"publisher","first-page":"2124","DOI":"10.3390\/ijms19072124","volume":"19","author":"Patricia Corrales","year":"2018","unstructured":"Corrales P, Vidal-Puig A, Medina-Gomez G. PPARs and metabolic disorders associated with challenged adipose tissue plasticity. Int J Mol Sci. 2018;19(7):2124-36.","journal-title":"International Journal of Molecular Sciences"},{"issue":"1","key":"544_CR25","doi-asserted-by":"publisher","first-page":"219","DOI":"10.1016\/j.bbrc.2007.11.119","volume":"366","author":"K Kuroyanagi","year":"2008","unstructured":"Kuroyanagi K, Kang MS, Goto T, Hirai S, Ohyama K, Kusudo T, Yu R, Yano M, Sasaki T, Takahashi N, et al. Citrus auraptene acts as an agonist for PPARs and enhances adiponectin production and MCP-1 reduction in 3T3-L1 adipocytes. Biochem Biophys Res Commun. 2008;366(1):219\u201325.","journal-title":"Biochem Biophys Res Commun"},{"issue":"3","key":"544_CR26","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1016\/j.cbi.2009.08.003","volume":"181","author":"HJ Liang","year":"2009","unstructured":"Liang HJ, Suk FM, Wang CK, Hung LF, Liu DZ, Chen NQ, Chen YC, Chang CC, Liang YC. Osthole, a potential antidiabetic agent, alleviates hyperglycemia in db\/db mice. Chem Biol Interact. 2009;181(3):309\u201315.","journal-title":"Chem Biol Interact"},{"issue":"4","key":"544_CR27","doi-asserted-by":"publisher","first-page":"787","DOI":"10.1016\/j.bbrc.2014.10.027","volume":"453","author":"A Watanabe","year":"2014","unstructured":"Watanabe A, Kato T, Ito Y, Yoshida I, Harada T, Mishima T, Fujita K, Watai M, Nakagawa K, Miyazawa T. Aculeatin, a coumarin derived from Toddalia asiatica (L.) lam., enhances differentiation and lipolysis of 3T3-L1 adipocytes. Biochem Biophys Res Commun. 2014;453(4):787\u201392.","journal-title":"Biochem Biophys Res Commun"},{"issue":"2","key":"544_CR28","doi-asserted-by":"publisher","first-page":"723","DOI":"10.1016\/j.foodchem.2013.04.036","volume":"141","author":"JR Noh","year":"2013","unstructured":"Noh JR, Kim YH, Hwang JH, Gang GT, Yeo SH, Kim KS, Oh WK, Ly SY, Lee IK, Lee CH. Scoparone inhibits adipocyte differentiation through down-regulation of peroxisome proliferators-activated receptor gamma in 3T3-L1 preadipocytes. Food Chem. 2013;141(2):723\u201330.","journal-title":"Food Chem"},{"issue":"10","key":"544_CR29","doi-asserted-by":"publisher","first-page":"1177","DOI":"10.1016\/j.nut.2014.01.015","volume":"30","author":"RN Nugara","year":"2014","unstructured":"Nugara RN, Inafuku M, Takara K, Iwasaki H, Oku H. Pteryxin: a coumarin in Peucedanum japonicum Thunb leaves exerts antiobesity activity through modulation of adipogenic gene network. Nutrition (Burbank, Los Angeles County, Calif). 2014;30(10):1177\u201384.","journal-title":"Nutrition (Burbank, Los Angeles County, Calif)"},{"issue":"9","key":"544_CR30","doi-asserted-by":"publisher","first-page":"1610","DOI":"10.1248\/bpb.33.1610","volume":"33","author":"E Shin","year":"2010","unstructured":"Shin E, Choi KM, Yoo HS, Lee CK, Hwang BY, Lee MK. Inhibitory effects of coumarins from the stem barks of Fraxinus rhynchophylla on adipocyte differentiation in 3T3-L1 cells. Biol Pharm Bull. 2010;33(9):1610\u20134.","journal-title":"Biol Pharm Bull"},{"issue":"3","key":"544_CR31","doi-asserted-by":"publisher","first-page":"601","DOI":"10.1142\/S0192415X18500313","volume":"46","author":"JH Ko","year":"2018","unstructured":"Ko JH, Nam D, Um JY, Jung SH, Ahn KS. Bergamottin inhibits Adipogenesis in 3T3-L1 cells and weight regulation in diet-induced obese mice. The American journal of Chinese medicine. 2018;46(3):601\u201315.","journal-title":"The American journal of Chinese medicine"},{"issue":"1","key":"544_CR32","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1002\/jcp.20258","volume":"204","author":"F Afzal","year":"2005","unstructured":"Afzal F, Pratap J, Ito K, Ito Y, Stein JL, van Wijnen AJ, Stein GS, Lian JB, Javed A. Smad function and intranuclear targeting share a Runx2 motif required for osteogenic lineage induction and BMP2 responsive transcription. J Cell Physiol. 2005;204(1):63\u201372.","journal-title":"J Cell Physiol"},{"issue":"3","key":"544_CR33","doi-asserted-by":"publisher","first-page":"446","DOI":"10.1002\/jcb.10369","volume":"88","author":"RT Franceschi","year":"2003","unstructured":"Franceschi RT, Xiao G. Regulation of the osteoblast-specific transcription factor, Runx2: responsiveness to multiple signal transduction pathways. J Cell Biochem. 2003;88(3):446\u201354.","journal-title":"J Cell Biochem"},{"key":"544_CR34","doi-asserted-by":"publisher","first-page":"16009","DOI":"10.1038\/boneres.2016.9","volume":"4","author":"M Wu","year":"2016","unstructured":"Wu M, Chen G, Li YP. TGF-beta and BMP signaling in osteoblast, skeletal development, and bone formation, homeostasis and disease. Bone research. 2016;4:16009.","journal-title":"Bone research"},{"issue":"20","key":"544_CR35","doi-asserted-by":"publisher","first-page":"15577","DOI":"10.1074\/jbc.M109.028019","volume":"285","author":"J Nojima","year":"2010","unstructured":"Nojima J, Kanomata K, Takada Y, Fukuda T, Kokabu S, Ohte S, Takada T, Tsukui T, Yamamoto TS, Sasanuma H, et al. Dual roles of smad proteins in the conversion from myoblasts to osteoblastic cells by bone morphogenetic proteins. J Biol Chem. 2010;285(20):15577\u201386.","journal-title":"J Biol Chem"},{"issue":"Pt 21","key":"544_CR36","doi-asserted-by":"publisher","first-page":"4974","DOI":"10.1242\/jcs.131953","volume":"126","author":"VS Salazar","year":"2013","unstructured":"Salazar VS, Zarkadis N, Huang L, Norris J, Grimston SK, Mbalaviele G, Civitelli R. Embryonic ablation of osteoblast Smad4 interrupts matrix synthesis in response to canonical Wnt signaling and causes an osteogenesis-imperfecta-like phenotype. J Cell Sci. 2013;126(Pt 21):4974\u201384.","journal-title":"J Cell Sci"},{"key":"544_CR37","doi-asserted-by":"publisher","first-page":"2162","DOI":"10.1242\/jcs.03466","volume":"120","author":"X Tan","year":"2007","unstructured":"Tan X, Weng T, Zhang J, Wang J, Li W, Wan H, Lan Y, Cheng X, Hou N, Liu H, et al. Smad4 is required for maintaining normal murine postnatal bone homeostasis. J Cell Sci. 2007;120(Pt 13:2162\u201370.","journal-title":"J Cell Sci"},{"key":"544_CR38","doi-asserted-by":"publisher","first-page":"388","DOI":"10.1016\/j.ijcard.2016.11.202","volume":"228","author":"JC Liu","year":"2017","unstructured":"Liu JC, Wang F, Xie ML, Cheng ZQ, Qin Q, Chen L, Chen R. Osthole inhibits the expressions of collagen I and III through Smad signaling pathway after treatment with TGF-beta1 in mouse cardiac fibroblasts. Int J Cardiol. 2017;228:388\u201393.","journal-title":"Int J Cardiol"},{"issue":"2","key":"544_CR39","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1007\/s10549-012-2282-3","volume":"136","author":"ML Panno","year":"2012","unstructured":"Panno ML, Giordano F, Rizza P, Pellegrino M, Zito D, Giordano C, Mauro L, Catalano S, Aquila S, Sisci D, et al. Bergapten induces ER depletion in breast cancer cells through SMAD4-mediated ubiquitination. Breast Cancer Res Treat. 2012;136(2):443\u201355.","journal-title":"Breast Cancer Res Treat"},{"issue":"6","key":"544_CR40","doi-asserted-by":"publisher","first-page":"1234","DOI":"10.1002\/jbmr.21","volume":"25","author":"DZ Tang","year":"2010","unstructured":"Tang DZ, Hou W, Zhou Q, Zhang M, Holz J, Sheu TJ, Li TF, Cheng SD, Shi Q, Harris SE, et al. Osthole stimulates osteoblast differentiation and bone formation by activation of beta-catenin-BMP signaling. J Bone Miner Res. 2010;25(6):1234\u201345.","journal-title":"J Bone Miner Res"}],"container-title":["Journal of Biomedical Science"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1186\/s12929-019-0544-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/article\/10.1186\/s12929-019-0544-7\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1186\/s12929-019-0544-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,7,3]],"date-time":"2020-07-03T23:14:59Z","timestamp":1593818099000},"score":1,"resource":{"primary":{"URL":"https:\/\/jbiomedsci.biomedcentral.com\/articles\/10.1186\/s12929-019-0544-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,7,5]]},"references-count":40,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2019,12]]}},"alternative-id":["544"],"URL":"https:\/\/doi.org\/10.1186\/s12929-019-0544-7","relation":{},"ISSN":["1423-0127"],"issn-type":[{"value":"1423-0127","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,7,5]]},"assertion":[{"value":"30 January 2019","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"26 June 2019","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"5 July 2019","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare that they have no competing interests.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"51"}}