{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,29]],"date-time":"2026-10-29T23:04:51Z","timestamp":1793315091432,"version":"4.5.0"},"reference-count":172,"publisher":"MDPI AG","issue":"24","license":[{"start":{"date-parts":[[2020,12,18]],"date-time":"2020-12-18T00:00:00Z","timestamp":1608249600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["AR071115"],"award-info":[{"award-number":["AR071115"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Scars are generated in mature skin as a result of the normal repair process, but the replacement of normal tissue with scar tissue can lead to biomechanical and functional deficiencies in the skin as well as psychological and social issues for patients that negatively affect quality of life. Abnormal scars, such as hypertrophic scars and keloids, and cutaneous fibrosis that develops in diseases such as systemic sclerosis and graft-versus-host disease can be even more challenging for patients. There is a large body of literature suggesting that inflammation promotes the deposition of scar tissue by fibroblasts. Mast cells represent one inflammatory cell type in particular that has been implicated in skin scarring and fibrosis. Most published studies in this area support a pro-fibrotic role for mast cells in the skin, as many mast cell-derived mediators stimulate fibroblast activity and studies generally indicate higher numbers of mast cells and\/or mast cell activation in scars and fibrotic skin. However, some studies in mast cell-deficient mice have suggested that these cells may not play a critical role in cutaneous scarring\/fibrosis. Here, we will review the data for and against mast cells as key regulators of skin fibrosis and discuss scientific gaps in the field.<\/jats:p>","DOI":"10.3390\/ijms21249673","type":"journal-article","created":{"date-parts":[[2020,12,18]],"date-time":"2020-12-18T11:42:09Z","timestamp":1608291729000},"page":"9673","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":52,"title":["A Review of the Evidence for and against a Role for Mast Cells in Cutaneous Scarring and Fibrosis"],"prefix":"10.3390","volume":"21","author":[{"given":"Traci A.","family":"Wilgus","sequence":"first","affiliation":[{"name":"Department of Pathology, Ohio State University, Columbus, OH 43210, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sara","family":"Ud-Din","sequence":"additional","affiliation":[{"name":"Centre for Dermatology Research, NIHR Manchester Biomedical Research Centre, Plastic and Reconstructive Surgery Research, University of Manchester, Manchester M13 9PT, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4116-6491","authenticated-orcid":false,"given":"Ardeshir","family":"Bayat","sequence":"additional","affiliation":[{"name":"Centre for Dermatology Research, NIHR Manchester Biomedical Research Centre, Plastic and Reconstructive Surgery Research, University of Manchester, Manchester M13 9PT, UK"},{"name":"MRC-SA Wound Healing Unit, Division of Dermatology, University of Cape Town, Observatory, Cape Town 7945, South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2020,12,18]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"265sr6","DOI":"10.1126\/scitranslmed.3009337","article-title":"Wound repair and regeneration: Mechanisms, signaling, and translation","volume":"6","author":"Eming","year":"2014","journal-title":"Sci. Transl. Med."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"314","DOI":"10.1038\/nature07039","article-title":"Wound repair and regeneration","volume":"453","author":"Gurtner","year":"2008","journal-title":"Nature"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1126\/science.276.5309.75","article-title":"Wound healing\u2014Aiming for perfect skin regeneration","volume":"276","author":"Martin","year":"1997","journal-title":"Science"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1111\/j.1524-475X.2009.00463.x","article-title":"Biomechanical behavior of scar tissue and uninjured skin in a porcine model","volume":"17","author":"Corr","year":"2009","journal-title":"Wound Repair Regen."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1111\/1523-1747.ep12274528","article-title":"Mechanical analysis of hypertrophic scar tissue: Structural basis for apparent increased rigidity","volume":"84","author":"Dunn","year":"1985","journal-title":"J. Investig. Dermatol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1049","DOI":"10.1016\/j.bjps.2008.03.020","article-title":"The hidden cost of skin scars: Quality of life after skin scarring","volume":"61","author":"Brown","year":"2008","journal-title":"J. Plast. Reconstr. Aesthet. Surg."},{"key":"ref_7","first-page":"54","article-title":"Inflammation as an orchestrator of cutaneous scar formation: A review of the literature","volume":"7","author":"Wilgus","year":"2020","journal-title":"Plast. Aesthet. Res."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"4519","DOI":"10.1007\/s12325-020-01499-4","article-title":"Mast Cells in Diabetes and Diabetic Wound Healing","volume":"37","author":"Dong","year":"2020","journal-title":"Adv. Ther."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1007\/s40629-020-00118-6","article-title":"Mast cells and IgE in defense against lethality of venoms: Possible \u201cbenefit\u201d of allergy","volume":"29","author":"Galli","year":"2020","journal-title":"Allergo J. Int."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"S4","DOI":"10.1016\/j.jaci.2018.10.054","article-title":"Mast cells as protectors of health","volume":"144","author":"Dudeck","year":"2019","journal-title":"J. Allergy Clin. Immunol."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Varricchi, G., de Paulis, A., Marone, G., and Galli, S.J. (2019). Future Needs in Mast Cell Biology. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20184397"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1002\/jlb.49.2.197","article-title":"Granule changes of human skin mast cells characteristic of piecemeal degranulation and associated with recovery during wound healing in situ","volume":"49","author":"Dvorak","year":"1991","journal-title":"J. Leukoc. Biol."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Lateef, Z., Stuart, G., Jones, N., Mercer, A., Fleming, S., and Wise, L. (2019). The Cutaneous Inflammatory Response to Thermal Burn Injury in a Murine Model. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20030538"},{"key":"ref_14","first-page":"38","article-title":"Migration of stromal cells supporting mast-cell differentiation into open wound produced in the skin of mice","volume":"9","author":"Matsuda","year":"1981","journal-title":"Exp. Hematol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1002\/(SICI)1096-9896(200001)190:1<100::AID-PATH496>3.0.CO;2-Q","article-title":"Mast cell involvement in normal human skin wound healing: Expression of monocyte chemoattractant protein-1 is correlated with recruitment of mast cells which synthesize interleukin-4 in vivo","volume":"190","author":"Trautmann","year":"2000","journal-title":"J. Pathol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1007\/s004030050351","article-title":"Skin mast cell promotion of matrix remodeling in burn wound healing in mice: Relevance of chymase","volume":"290","author":"Nishikori","year":"1998","journal-title":"Arch. Dermatol. Res."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1680","DOI":"10.1016\/j.jid.2019.01.030","article-title":"A Double-Blind, Randomized Trial Shows the Role of Zonal Priming and Direct Topical Application of Epigallocatechin-3-Gallate in the Modulation of Cutaneous Scarring in Human Skin","volume":"139","author":"Foden","year":"2019","journal-title":"J. Investig. Dermatol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1007\/s12016-019-08729-w","article-title":"A Review of the Contribution of Mast Cells in Wound Healing: Involved Molecular and Cellular Mechanisms","volume":"58","author":"Komi","year":"2020","journal-title":"Clin. Rev. Allergy Immunol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"552205","DOI":"10.3389\/fimmu.2020.552205","article-title":"Mast Cells in Skin Scarring: A Review of Animal and Human Research","volume":"11","author":"Wilgus","year":"2020","journal-title":"Front. Immunol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"20500","DOI":"10.1073\/pnas.1908816116","article-title":"Mast cells are critical for controlling the bacterial burden and the healing of infected wounds","volume":"116","author":"Zimmermann","year":"2019","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1111\/imr.12634","article-title":"Mast cells as sources of cytokines, chemokines, and growth factors","volume":"282","author":"Mukai","year":"2018","journal-title":"Immunol. Rev."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"e309","DOI":"10.1002\/wdev.309","article-title":"Scarless wound healing: Transitioning from fetal research to regenerative healing","volume":"7","author":"Moore","year":"2018","journal-title":"Wiley Interdiscip. Rev. Dev. Biol."},{"key":"ref_23","first-page":"16","article-title":"Regenerative healing in fetal skin: A review of the literature","volume":"53","author":"Wilgus","year":"2007","journal-title":"Ostomy Wound Manag."},{"key":"ref_24","unstructured":"Bagchi, D., Das, A., and Roy, S. (2020). Fetal wound healing. Wound Healing, Tissue Repair, and Regeneration in Diabetes, Academic Press."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1006\/dbio.1995.1141","article-title":"Ontogeny of the skin and the transition from scar-free to scarring phenotype during wound healing in the pouch young of a marsupial, Monodelphis domestica","volume":"169","author":"Armstrong","year":"1995","journal-title":"Dev. Biol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1002\/(SICI)1097-0177(199807)212:3<385::AID-AJA6>3.0.CO;2-D","article-title":"Endogenous inflammatory response to dermal wound healing in the fetal and adult mouse","volume":"212","author":"Cowin","year":"1998","journal-title":"Dev. Dyn."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"360","DOI":"10.1016\/0007-1226(94)90096-5","article-title":"Acute inflammation in foetal and adult sheep: The response to subcutaneous injection of turpentine and carrageenan","volume":"47","author":"Kumta","year":"1994","journal-title":"Br. J. Plast. Surg."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1016\/S0002-9440(10)63338-X","article-title":"The impact of cyclooxygenase-2 mediated inflammation on scarless fetal wound healing","volume":"165","author":"Wilgus","year":"2004","journal-title":"Am. J. Pathol."},{"key":"ref_29","first-page":"345","article-title":"Mast cells in the skin of rats during development","volume":"9","author":"Savall","year":"1981","journal-title":"Med. Cutanea Ibero Lat. Am."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"458","DOI":"10.1038\/jid.2011.324","article-title":"Mast cells contribute to scar formation during fetal wound healing","volume":"132","author":"Wulff","year":"2012","journal-title":"J. Investig. Dermatol."},{"key":"ref_31","first-page":"1337","article-title":"Deficient differentiation of mast cells in the skin of mi\/mi mice. Usefulness of in situ hybridization for evaluation of mast cell phenotype","volume":"143","author":"Kasugai","year":"1993","journal-title":"Am. J. Pathol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"533","DOI":"10.1111\/wrr.12421","article-title":"Healthy human second-trimester fetal skin is deficient in leukocytes and associated homing chemokines","volume":"24","author":"Walraven","year":"2016","journal-title":"Wound Repair Regen."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1083\/jcb.25.3.577","article-title":"Differentiation and proliferation of embryonic mast cells of the rat","volume":"25","author":"Combs","year":"1965","journal-title":"J. Cell Biol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1007\/978-1-62703-505-7_29","article-title":"Examining the role of mast cells in fetal wound healing using cultured cells in vitro","volume":"1037","author":"Wulff","year":"2013","journal-title":"Methods Mol. Biol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"1053","DOI":"10.1111\/j.1365-2133.2012.11190.x","article-title":"Site-specific immunophenotyping of keloid disease demonstrates immune upregulation and the presence of lymphoid aggregates","volume":"167","author":"Bagabir","year":"2012","journal-title":"Br. J. Dermatol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1046\/j.1523-1747.2000.00837.x","article-title":"Altered expression of mast cell chymase and tryptase and of c-Kit in human cutaneous scar tissue","volume":"114","author":"Hermes","year":"2000","journal-title":"J. Investig. Dermatol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1046\/j.1523-1747.2001.01284.x","article-title":"Expression of mast cell growth modulating and chemotactic factors and their receptors in human cutaneous scars","volume":"116","author":"Hermes","year":"2001","journal-title":"J. Investig. Dermatol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1136\/bjo.82.6.691","article-title":"Mast cells in pathological and surgical scars","volume":"82","author":"Beer","year":"1998","journal-title":"Br. J. Ophthalmol."},{"key":"ref_39","first-page":"195","article-title":"The effect of disodium cromoglycate on the skin wound healing and collagen content in the wounds of rats","volume":"41","author":"Dabrowski","year":"1990","journal-title":"Acta Physiol. Pol."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Chen, L., Schrementi, M.E., Ranzer, M.J., Wilgus, T.A., and DiPietro, L.A. (2014). Blockade of mast cell activation reduces cutaneous scar formation. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0085226"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1159\/000232573","article-title":"Pathophysiological role of skin mast cells in wound healing after scald injury: Study with mast cell-deficient W\/W(V) mice","volume":"151","author":"Shiota","year":"2010","journal-title":"Int. Arch. Allergy Immunol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"7681","DOI":"10.4049\/jimmunol.1002803","article-title":"The inflammatory response after an epidermal burn depends on the activities of mouse mast cell proteases 4 and 5","volume":"185","author":"Younan","year":"2010","journal-title":"J. Immunol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1873","DOI":"10.1016\/j.intimp.2004.08.009","article-title":"Possible involvement of mast cells in collagen remodeling in the late phase of cutaneous wound healing in mice","volume":"4","author":"Iba","year":"2004","journal-title":"Int. Immunopharmacol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"649e","DOI":"10.1097\/PRS.0b013e318230c55d","article-title":"Mast cells are required in the proliferation and remodeling phases of microdeformational wound therapy","volume":"128","author":"Younan","year":"2011","journal-title":"Plast. Reconstr. Surg."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Nauta, A.C., Grova, M., Montoro, D.T., Zimmermann, A., Tsai, M., Gurtner, G.C., Galli, S.J., and Longaker, M.T. (2013). Evidence that mast cells are not required for healing of splinted cutaneous excisional wounds in mice. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0059167"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"6930","DOI":"10.1182\/blood-2011-03-343962","article-title":"Reduced mast cell and basophil numbers and function in Cpa3-Cre; Mcl-1fl\/fl mice","volume":"118","author":"Lilla","year":"2011","journal-title":"Blood"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"832","DOI":"10.1016\/j.immuni.2011.09.015","article-title":"Cre-mediated cell ablation contests mast cell contribution in models of antibody- and T cell-mediated autoimmunity","volume":"35","author":"Feyerabend","year":"2011","journal-title":"Immunity"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"6147","DOI":"10.4049\/jimmunol.1301350","article-title":"Mast cells are dispensable for normal and activin-promoted wound healing and skin carcinogenesis","volume":"191","author":"Antsiferova","year":"2013","journal-title":"J. Immunol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"973","DOI":"10.1016\/j.immuni.2011.03.028","article-title":"Mast cells are key promoters of contact allergy that mediate the adjuvant effects of haptens","volume":"34","author":"Dudeck","year":"2011","journal-title":"Immunity"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"2005","DOI":"10.1038\/jid.2014.12","article-title":"Genetic ablation of mast cells redefines the role of mast cells in skin wound healing and bleomycin-induced fibrosis","volume":"134","author":"Willenborg","year":"2014","journal-title":"J. Investig. Dermatol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1802","DOI":"10.1038\/jid.2014.142","article-title":"Rethinking the potential roles of mast cells in skin wound healing and bleomycin-induced skin fibrosis","volume":"134","author":"Galli","year":"2014","journal-title":"J. Investig. Dermatol."},{"key":"ref_52","first-page":"509","article-title":"Histamine and collagen synthesis in keloid and hypertrophic scar","volume":"23","author":"Cohen","year":"1972","journal-title":"Surg. Forum"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1097\/00004630-198703000-00008","article-title":"The possible role of mast cells (allergy) in the production of keloid and hypertrophic scarring","volume":"8","author":"Smith","year":"1987","journal-title":"J. Burn Care Rehabil."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1097\/BCR.0000000000000392","article-title":"Clinical and Histopathological Features of Post Burn Pruritus","volume":"37","author":"Kwak","year":"2016","journal-title":"J. Burn Care Res."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"669","DOI":"10.1016\/j.burns.2006.03.015","article-title":"Histology of the thick scar on the female, red Duroc pig: Final similarities to human hypertrophic scar","volume":"32","author":"Harunari","year":"2006","journal-title":"Burns"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1317","DOI":"10.1097\/00006534-199810000-00001","article-title":"Transforming growth factor-beta in thermally injured patients with hypertrophic scars: Effects of interferon alpha-2b","volume":"102","author":"Tredget","year":"1998","journal-title":"Plast. Reconstr. Surg."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"428","DOI":"10.5021\/ad.2013.25.4.428","article-title":"Clinical and histological correlation in post-burn hypertrophic scar for pain and itching sensation","volume":"25","author":"Choi","year":"2013","journal-title":"Ann. Dermatol."},{"key":"ref_58","first-page":"4438","article-title":"Mast cell chymase promotes hypertrophic scar fibroblast proliferation and collagen synthesis by activating TGF-beta1\/Smads signaling pathway","volume":"14","author":"Chen","year":"2017","journal-title":"Exp. Ther. Med."},{"key":"ref_59","first-page":"327","article-title":"The mast cell in hypertrophic scars","volume":"30","author":"Kischer","year":"1972","journal-title":"Tex. Rep. Biol. Med."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1111\/1523-1747.ep12543553","article-title":"Mast cell analyses in hypertrophic scars, hypertrophic scars treated with pressure and mature scars","volume":"70","author":"Kischer","year":"1978","journal-title":"J. Investig. Dermatol."},{"key":"ref_61","first-page":"32","article-title":"Mast cells and angiogenesis in wound healing","volume":"36","author":"Gaber","year":"2014","journal-title":"Anal. Quant. Cytopathol. Histopathol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1272","DOI":"10.1111\/j.1524-4725.1989.tb03146.x","article-title":"Exogenous electric current can reduce the formation of hypertrophic scars","volume":"15","author":"Weiss","year":"1989","journal-title":"J. Dermatol. Surg. Oncol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1002\/path.1502","article-title":"Hypertrophic scar formation is associated with an increased number of epidermal Langerhans cells","volume":"202","author":"Niessen","year":"2004","journal-title":"J. Pathol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1111\/j.1600-0625.1999.tb00370.x","article-title":"Mast cell chymase and tryptase during tissue turnover: Analysis on in vitro mitogenesis of fibroblasts and keratinocytes and alterations in cutaneous scars","volume":"8","author":"Algermissen","year":"1999","journal-title":"Exp. Dermatol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"1564","DOI":"10.1016\/j.bjps.2014.06.014","article-title":"The structure of the basement membrane zone differs between keloids, hypertrophic scars and normal skin: A possible background to an impaired function","volume":"67","author":"Hellstrom","year":"2014","journal-title":"J. Plast. Reconstr. Aesthetic Surg."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"226","DOI":"10.1111\/j.1524-475X.2008.00363.x","article-title":"The mast cell stabilizer ketotifen prevents development of excessive skin wound contraction and fibrosis in red Duroc pigs","volume":"16","author":"Hildebrand","year":"2008","journal-title":"Wound Repair Regen."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"3250","DOI":"10.1096\/fj.07-8218com","article-title":"Mechanical load initiates hypertrophic scar formation through decreased cellular apoptosis","volume":"21","author":"Aarabi","year":"2007","journal-title":"FASEB J."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1111\/wrr.12238","article-title":"A novel immune competent murine hypertrophic scar contracture model: A tool to elucidate disease mechanism and develop new therapies","volume":"22","author":"Ibrahim","year":"2014","journal-title":"Wound Repair Regen."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1111\/j.1524-475X.2011.00672.x","article-title":"Human hypertrophic scar-like nude mouse model: Characterization of the molecular and cellular biology of the scar process","volume":"19","author":"Wang","year":"2011","journal-title":"Wound Repair Regen."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"854","DOI":"10.1007\/BF01897918","article-title":"Direct histochemical demonstration of histamine in cutaneous mast cells: Urticaria pigmentosa and keloids","volume":"25","author":"Hakanson","year":"1969","journal-title":"Experientia"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1111\/exd.12854","article-title":"The role of aldo-keto reductase 1C3 (AKR1C3)-mediated prostaglandin D2 (PGD2) metabolism in keloids","volume":"25","author":"Mantel","year":"2016","journal-title":"Exp. Dermatol."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1007\/s13304-012-0183-y","article-title":"Tryptase-positive mast cells and angiogenesis in keloids: A new possible post-surgical target for prevention","volume":"65","author":"Ammendola","year":"2013","journal-title":"Updates Surg."},{"key":"ref_73","first-page":"3596","article-title":"Mast cell chymase in keloid induces profibrotic response via transforming growth factor-beta1\/Smad activation in keloid fibroblasts","volume":"7","author":"Dong","year":"2014","journal-title":"Int. J. Clin. Exp. Pathol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1080\/09546630310016808","article-title":"Silicone gel sheets relieve pain and pruritus with clinical improvement of keloid: Possible target of mast cells","volume":"14","author":"Eishi","year":"2003","journal-title":"J. Dermatol. Treat."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"2429","DOI":"10.1038\/jid.2008.103","article-title":"Green tea polyphenol epigallocatechin-3-gallate suppresses collagen production and proliferation in keloid fibroblasts via inhibition of the STAT3-signaling pathway","volume":"128","author":"Park","year":"2008","journal-title":"J. Investig. Dermatol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"2607","DOI":"10.1038\/sj.jid.5700472","article-title":"Green tea extract and (-)-epigallocatechin-3-gallate inhibit mast cell-stimulated type I collagen expression in keloid fibroblasts via blocking PI-3K\/AkT signaling pathways","volume":"126","author":"Zhang","year":"2006","journal-title":"J. Investig. Dermatol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1038\/labinvest.2013.82","article-title":"Ex vivo evaluation of antifibrotic compounds in skin scarring: EGCG and silencing of PAI-1 independently inhibit growth and induce keloid shrinkage","volume":"93","author":"Syed","year":"2013","journal-title":"Lab. Investig."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"783","DOI":"10.1111\/j.1532-950X.2013.12055.x","article-title":"Equine exuberant granulation tissue and human keloids: A comparative histopathologic study","volume":"42","author":"Theoret","year":"2013","journal-title":"Vet. Surg."},{"key":"ref_79","first-page":"427","article-title":"Mast cells in human keloid, small intestine, and lung by an immunoperoxidase technique using a murine monoclonal antibody against tryptase","volume":"124","author":"Craig","year":"1986","journal-title":"Am. J. Pathol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"15002","DOI":"10.1038\/nrdp.2015.2","article-title":"Systemic sclerosis","volume":"1","author":"Allanore","year":"2015","journal-title":"Nat. Rev. Dis. Primers"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1007\/s10067-003-0706-5","article-title":"The roles of transforming growth factor type beta3 (TGF-beta3) and mast cells in the pathogenesis of scleroderma","volume":"22","author":"Ozbilgin","year":"2003","journal-title":"Clin. Rheumatol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"288","DOI":"10.1007\/BF00431065","article-title":"Increased number of mast cells accompany enhanced collagen synthesis in linear localized scleroderma","volume":"281","author":"Takeda","year":"1989","journal-title":"Arch. Dermatol. Res."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1046\/j.1365-2133.1998.02114.x","article-title":"Dermal mast cells in scleroderma: Their skin density, tryptase\/chymase phenotypes and degranulation","volume":"138","author":"Akimoto","year":"1998","journal-title":"Br. J. Dermatol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1002\/path.1711660307","article-title":"Sequential dermal microvascular and perivascular changes in the development of scleroderma","volume":"166","author":"Prescott","year":"1992","journal-title":"J. Pathol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1702","DOI":"10.1002\/art.1780331114","article-title":"Dermal mast cell degranulation in systemic sclerosis","volume":"33","author":"Seibold","year":"1990","journal-title":"Arthritis Rheumatol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1001\/archderm.1987.01660260075017","article-title":"Mast cell numbers in diffuse scleroderma","volume":"123","author":"Nishioka","year":"1987","journal-title":"Arch. Dermatol."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"182","DOI":"10.7326\/0003-4819-102-2-182","article-title":"Increased dermal mast cell populations in progressive systemic sclerosis: A link in chronic fibrosis?","volume":"102","author":"Hawkins","year":"1985","journal-title":"Ann. Intern. Med."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"795","DOI":"10.1002\/art.30190","article-title":"Mast cells are a source of transforming growth factor beta in systemic sclerosis","volume":"63","author":"Hugle","year":"2011","journal-title":"Arthritis Rheumatol."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1016\/S0923-1811(98)00077-2","article-title":"Increased cutaneous immunoreactive stem cell factor expression and serum stem cell factor level in systemic scleroderma","volume":"20","author":"Kihira","year":"1998","journal-title":"J. Dermatol. Sci."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"933","DOI":"10.1002\/art.1780350813","article-title":"Mast cell changes in scleroderma. Presence of MCT cells in the skin and evidence of mast cell activation","volume":"35","author":"Irani","year":"1992","journal-title":"Arthritis Rheumatol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"1347","DOI":"10.1002\/art.37860","article-title":"Interleukin-17A+ cell counts are increased in systemic sclerosis skin and their number is inversely correlated with the extent of skin involvement","volume":"65","author":"Truchetet","year":"2013","journal-title":"Arthritis Rheumatol."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1111\/1523-1747.ep12276876","article-title":"Mast cell changes in a case of rapidly progressive scleroderma-ultrastructural analysis","volume":"92","author":"Claman","year":"1989","journal-title":"J. Investig. Dermatol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"336","DOI":"10.1001\/archderm.1990.01670270068011","article-title":"Elevated plasma histamine levels in systemic sclerosis (scleroderma)","volume":"126","author":"Falanga","year":"1990","journal-title":"Arch. Dermatol."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1002\/art.1780310514","article-title":"Immunologic dysfunction in scleroderma: Evidence for increased mast cell releasability and HLA-DR positivity in the dermis","volume":"31","author":"Pearson","year":"1988","journal-title":"Arthritis Rheumatol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"561","DOI":"10.1097\/BOR.0000000000000331","article-title":"Animal models of scleroderma: Recent progress","volume":"28","author":"Marangoni","year":"2016","journal-title":"Curr. Opin. Rheumatol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"456","DOI":"10.1046\/j.1523-1747.1999.00528.x","article-title":"Animal model of sclerotic skin. I: Local injections of bleomycin induce sclerotic skin mimicking scleroderma","volume":"112","author":"Yamamoto","year":"1999","journal-title":"J. Investig. Dermatol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1016\/j.ajpath.2011.09.013","article-title":"Blockade of interleukin-6 receptor alleviates disease in mouse model of scleroderma","volume":"180","author":"Kitaba","year":"2012","journal-title":"Am. J. Pathol."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1016\/j.biopha.2005.04.004","article-title":"Inhibition of skin sclerosis by 15deoxy delta12,14-prostaglandin J2 and retrovirally transfected prostaglandin D synthase in a mouse model of bleomycin-induced scleroderma","volume":"60","author":"Kohno","year":"2006","journal-title":"Biomed. Pharmacother."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"3562","DOI":"10.1002\/art.23994","article-title":"A CD40-CD154 interaction in tissue fibrosis","volume":"58","author":"Kawai","year":"2008","journal-title":"Arthritis Rheumatol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"843","DOI":"10.1046\/j.1523-1747.1999.00758.x","article-title":"Effect of superoxide dismutase on bleomycin-induced dermal sclerosis: Implications for the treatment of systemic sclerosis","volume":"113","author":"Yamamoto","year":"1999","journal-title":"J. Investig. Dermatol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1006\/clim.1999.4720","article-title":"Anti-sclerotic effect of transforming growth factor-beta antibody in a mouse model of bleomycin-induced scleroderma","volume":"92","author":"Yamamoto","year":"1999","journal-title":"Clin. Immunol."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1159\/000098208","article-title":"Increased bleomycin-induced skin fibrosis in mice lacking the Th1-specific transcription factor T-bet","volume":"73","author":"Lakos","year":"2006","journal-title":"Pathobiology"},{"key":"ref_103","doi-asserted-by":"crossref","unstructured":"Ishikawa, M., and Yamamoto, T. (2020). Antifibrogenic effects of C-C chemokine receptor type 2 antagonist in a bleomycin-induced scleroderma model. Exp. Dermatol.","DOI":"10.1111\/exd.14088"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"3813","DOI":"10.1111\/bph.14450","article-title":"VCE-004.3, a cannabidiol aminoquinone derivative, prevents bleomycin-induced skin fibrosis and inflammation through PPARgamma- and CB2 receptor-dependent pathways","volume":"175","author":"Cantarero","year":"2018","journal-title":"Br. J. Pharmacol."},{"key":"ref_105","first-page":"S15","article-title":"Spleen tyrosine kinase (Syk) inhibitor fostamatinib limits tissue damage and fibrosis in a bleomycin-induced scleroderma mouse model","volume":"33","author":"Pamuk","year":"2015","journal-title":"Clin. Exp. Rheumatol."},{"key":"ref_106","first-page":"2628","article-title":"Animal model of sclerotic skin. II. Bleomycin induced scleroderma in genetically mast cell deficient WBB6F1-W\/W(V) mice","volume":"26","author":"Yamamoto","year":"1999","journal-title":"J. Rheumatol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"300","DOI":"10.1101\/gr.6.4.300","article-title":"A tandem duplication within the fibrillin 1 gene is associated with the mouse tight skin mutation","volume":"6","author":"Siracusa","year":"1996","journal-title":"Genome Res."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"323","DOI":"10.3181\/00379727-180-42183","article-title":"Mast cells and their degranulation in the Tsk mouse model of scleroderma","volume":"180","author":"Walker","year":"1985","journal-title":"Proc. Soc. Exp. Biol. Med."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1111\/j.0906-6705.2005.00315.x","article-title":"Mast cell accumulation and cytokine expression in the tight skin mouse model of scleroderma","volume":"14","author":"Wang","year":"2005","journal-title":"Exp. Dermatol."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1046\/j.1523-1747.2001.00165.x","article-title":"Isoform-selective upregulation of mast cell chymase in the development of skin fibrosis in scleroderma model mice","volume":"116","author":"Kakizoe","year":"2001","journal-title":"J. Investig. Dermatol."},{"key":"ref_111","first-page":"57","article-title":"Ketotifen prevents skin fibrosis in the tight skin mouse","volume":"17","author":"Walker","year":"1990","journal-title":"J. Rheumatol."},{"key":"ref_112","first-page":"299","article-title":"Inhibition of fibrosis in TSK mice by blocking mast cell degranulation","volume":"14","author":"Walker","year":"1987","journal-title":"J. Rheumatol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"424","DOI":"10.1038\/sj.bjp.0706209","article-title":"Effect of mast cell chymase inhibitor on the development of scleroderma in tight-skin mice","volume":"145","author":"Shiota","year":"2005","journal-title":"Br. J. Pharmacol."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"196","DOI":"10.3181\/00379727-192-42979","article-title":"Adoptive transfer of tsk skin fibrosis to +\/+ recipients by tsk bone marrow and spleen cells","volume":"192","author":"Walker","year":"1989","journal-title":"Proc. Soc. Exp. Biol. Med."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1016\/0300-9629(94)00127-F","article-title":"The role of mast cells in the development of skin fibrosis in tight-skin mutant mice","volume":"110","author":"Everett","year":"1995","journal-title":"Comp. Biochem. Physiol. A Physiol."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"1807","DOI":"10.1172\/JCI99088","article-title":"PAI1 mediates fibroblast-mast cell interactions in skin fibrosis","volume":"128","author":"Pincha","year":"2018","journal-title":"J. Clin. Investig."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"1424","DOI":"10.1001\/archderm.1977.01640100102020","article-title":"Scleroderma-like changes in chronic graft vs host disease","volume":"113","author":"Spielvogel","year":"1977","journal-title":"Arch. Dermatol."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"2470","DOI":"10.3389\/fimmu.2019.02470","article-title":"Mast Cells Are Mediators of Fibrosis and Effector Cell Recruitment in Dermal Chronic Graft-vs.-Host Disease","volume":"10","author":"Strattan","year":"2019","journal-title":"Front. Immunol."},{"key":"ref_119","first-page":"296","article-title":"Immunohistologic and ultrastructural study of the sclerotic skin in chronic graft-versus-host disease in man","volume":"115","author":"Devergie","year":"1984","journal-title":"Am. J. Pathol."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"412","DOI":"10.1111\/1523-1747.ep12522551","article-title":"Murine graft-versus-host skin disease: A chronologic and quantitative analysis of two histologic patterns","volume":"81","author":"Charley","year":"1983","journal-title":"J. Investig. Dermatol."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1111\/1523-1747.ep12265302","article-title":"Mast cell depletion in murine chronic graft-versus-host disease","volume":"84","author":"Claman","year":"1985","journal-title":"J. Investig. Dermatol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/0008-8749(85)90086-3","article-title":"Chronic graft-versus-host disease as a model for scleroderma. II. Mast cell depletion with deposition of immunoglobulins in the skin and fibrosis","volume":"94","author":"Claman","year":"1985","journal-title":"Cell. Immunol."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"2009","DOI":"10.4049\/jimmunol.137.6.2009","article-title":"Mast cell \u201cdisappearance\u201d in chronic murine graft-vs-host disease (GVHD)-ultrastructural demonstration of \u201cphantom mast cells\u201d","volume":"137","author":"Claman","year":"1986","journal-title":"J. Immunol."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"4093","DOI":"10.4049\/jimmunol.138.12.4093","article-title":"Cutaneous mast cell depletion and recovery in murine graft-vs-host disease","volume":"138","author":"Choi","year":"1987","journal-title":"J. Immunol."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/0008-8749(83)90001-1","article-title":"Chronic graft-versus-host disease (GVHD) as a model for scleroderma. I. Description of model systems","volume":"77","author":"Jaffee","year":"1983","journal-title":"Cell. Immunol."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"823","DOI":"10.1038\/sj.bmt.1700734","article-title":"Nedocromil sodium ameliorates skin manifestations in a murine model of chronic graft-versus-host disease","volume":"19","author":"Goldenhersh","year":"1997","journal-title":"Bone Marrow Transpl."},{"key":"ref_127","first-page":"340","article-title":"Effect of activated human mast cells and mast cell-derived mediators on proliferation, type I collagen production and glycosaminoglycans synthesis by human dermal fibroblasts","volume":"12","author":"Abe","year":"2002","journal-title":"Eur. J. Dermatol."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1016\/0014-4827(90)90275-F","article-title":"Mast cells enhance migration and proliferation of fibroblasts into an in vitro wound","volume":"188","author":"Kupietzky","year":"1990","journal-title":"Exp. Cell Res."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"2027","DOI":"10.1084\/jem.180.6.2027","article-title":"Promotion of mouse fibroblast collagen gene expression by mast cells stimulated via the Fc epsilon RI. Role for mast cell-derived transforming growth factor beta and tumor necrosis factor alpha","volume":"180","author":"Gordon","year":"1994","journal-title":"J. Exp. Med."},{"key":"ref_130","first-page":"113","article-title":"Promotion of mouse fibroblast proliferation by IgE-dependent activation of mouse mast cells: Role for mast cell tumor necrosis factor-alpha and transforming growth factor-beta 1","volume":"99","author":"Kendall","year":"1997","journal-title":"J. Allergy Clin. Immunol."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"2867","DOI":"10.1007\/s00018-005-5289-7","article-title":"Divergent effects of the major mast cell products histamine, tryptase and TNF-alpha on human fibroblast behaviour","volume":"62","author":"Albrecht","year":"2005","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"1509","DOI":"10.1046\/j.1365-2222.1998.00360.x","article-title":"Mast cell tryptase stimulates both human dermal fibroblast proliferation and type I collagen production","volume":"28","author":"Abe","year":"1998","journal-title":"Clin. Exp. Allergy"},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1046\/j.1365-2222.2002.01293.x","article-title":"Human mast cells stimulate fibroblast proliferation, collagen synthesis and lattice contraction: A direct role for mast cells in skin fibrosis","volume":"32","author":"Garbuzenko","year":"2002","journal-title":"Clin. Exp. Allergy"},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"2310","DOI":"10.4049\/jimmunol.158.5.2310","article-title":"Human mast cells activate fibroblasts: Tryptase is a fibrogenic factor stimulating collagen messenger ribonucleic acid synthesis and fibroblast chemotaxis","volume":"158","author":"Gruber","year":"1997","journal-title":"J. Immunol."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"1113","DOI":"10.1046\/j.1523-1747.2001.15211.x","article-title":"The differentiation and function of myofibroblasts is regulated by mast cell mediators","volume":"117","author":"Gailit","year":"2001","journal-title":"J. Investig. Dermatol."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"S78","DOI":"10.1067\/mai.2000.106058","article-title":"Effect of mast cell-derived mediators and mast cell-related neutral proteases on human dermal fibroblast proliferation and type I collagen production","volume":"106","author":"Abe","year":"2000","journal-title":"J. Allergy Clin. Immunol."},{"key":"ref_137","first-page":"1487","article-title":"Mast cell chymase promotes cell proliferation and expression of certain cytokines in a dose-dependent manner","volume":"5","author":"Dong","year":"2012","journal-title":"Mol. Med. Rep."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"955","DOI":"10.3892\/etm.2015.2216","article-title":"High concentrations of mast cell chymase facilitate the transduction of the transforming growth factor-beta1\/Smads signaling pathway in skin fibroblasts","volume":"9","author":"Dong","year":"2015","journal-title":"Exp. Ther. Med."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"7127","DOI":"10.1074\/jbc.272.11.7127","article-title":"Cleavage of type I procollagen by human mast cell chymase initiates collagen fibril formation and generates a unique carboxyl-terminal propeptide","volume":"272","author":"Kofford","year":"1997","journal-title":"J. Biol. Chem."},{"key":"ref_140","first-page":"245","article-title":"Emedastine difumarate inhibits histamine-induced collagen synthesis in dermal fibroblasts","volume":"18","author":"Murota","year":"2008","journal-title":"J. Investig. Allergol. Clin. Immunol."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"S72","DOI":"10.1067\/mai.2000.106059","article-title":"Influence of IgE-mediated activation of cultured human mast cells on proliferation and type I collagen production by human dermal fibroblasts","volume":"106","author":"Abe","year":"2000","journal-title":"J. Allergy Clin. Immunol."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1007\/BF02285158","article-title":"The role of mast cell-derived histamine in the closure of an in vitro wound","volume":"45","author":"Kupietzky","year":"1996","journal-title":"Inflamm. Res."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"5053","DOI":"10.4049\/jimmunol.160.10.5053","article-title":"Human mast cells augment fibroblast proliferation by heterotypic cell-cell adhesion and action of IL-4","volume":"160","author":"Trautmann","year":"1998","journal-title":"J. Immunol."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"2147","DOI":"10.1172\/JCI115247","article-title":"Fibroblast chemotaxis induction by human recombinant interleukin-4. Identification by synthetic peptide analysis of two chemotactic domains residing in amino acid sequences 70\u201388 and 89\u2013122","volume":"87","author":"Postlethwaite","year":"1991","journal-title":"J. Clin. Investig."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"1479","DOI":"10.1172\/JCI116015","article-title":"Human fibroblasts synthesize elevated levels of extracellular matrix proteins in response to interleukin 4","volume":"90","author":"Postlethwaite","year":"1992","journal-title":"J. Clin. Investig."},{"key":"ref_146","first-page":"823","article-title":"Interleukin-4 stimulates collagen synthesis by normal and scleroderma fibroblasts in dermal equivalents","volume":"37","author":"Fertin","year":"1991","journal-title":"Cell. Mol. Biol."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1016\/j.jdermsci.2011.02.008","article-title":"Plasminogen activator inhibitor-2 (PAI-2) secreted from activated mast cells induces alpha-smooth muscle actin (alpha-SMA) expression in dermal fibroblasts","volume":"62","author":"Kim","year":"2011","journal-title":"J. Dermatol. Sci."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1067\/mai.2000.107036","article-title":"Monocyte chemoattractant peptide-1 expression during cutaneous allergic reactions in mice is mast cell dependent and largely mediates the monocyte recruitment response","volume":"106","author":"Gordon","year":"2000","journal-title":"J. Allergy Clin. Immunol."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1016\/S0923-1811(00)00164-X","article-title":"Role of stem cell factor and monocyte chemoattractant protein-1 in the interaction between fibroblasts and mast cells in fibrosis","volume":"26","author":"Yamamoto","year":"2001","journal-title":"J. Dermatol. Sci."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"6653","DOI":"10.1016\/j.febslet.2005.09.104","article-title":"IL-4 modulates the histamine content of mast cells in a mast cell\/fibroblast co-culture through a Stat6 signaling pathway in fibroblasts","volume":"579","author":"Nabeshima","year":"2005","journal-title":"FEBS Lett."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1006\/cimm.2000.1631","article-title":"TGFbeta1 and TNFalpha secreted by mast cells stimulated via the FcepsilonRI activate fibroblasts for high-level production of monocyte chemoattractant protein-1 (MCP-1)","volume":"201","author":"Gordon","year":"2000","journal-title":"Cell. Immunol."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"1900","DOI":"10.1038\/sj.jid.5700302","article-title":"Diminished induction of skin fibrosis in mice with MCP-1 deficiency","volume":"126","author":"Ferreira","year":"2006","journal-title":"J. Investig. Dermatol."},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1038\/nm.2574","article-title":"Focal adhesion kinase links mechanical force to skin fibrosis via inflammatory signaling","volume":"18","author":"Wong","year":"2011","journal-title":"Nat. Med."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"1036","DOI":"10.1097\/PRS.0b013e3182865c3f","article-title":"Through gap junction communications, co-cultured mast cells and fibroblasts generate fibroblast activities allied with hypertrophic scarring","volume":"131","author":"Foley","year":"2013","journal-title":"Plast. Reconstr. Surg."},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"1170","DOI":"10.1002\/jcb.21107","article-title":"Rat mast cells communicate with fibroblasts via gap junction intercellular communications","volume":"100","author":"Au","year":"2007","journal-title":"J. Cell. Biochem."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1046\/j.1365-2567.2000.00973.x","article-title":"Mast cells enhance contraction of three-dimensional collagen lattices by fibroblasts by cell-cell interaction: Role of stem cell factor\/c-kit","volume":"99","author":"Yamamoto","year":"2000","journal-title":"Immunology"},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"638e","DOI":"10.1097\/PRS.0000000000000103","article-title":"Mast cells prevent dexamethasone-induced cell death of cultured fibroblasts: Relationship to gap junctional intercellular communications","volume":"133","author":"Foley","year":"2014","journal-title":"Plast. Reconstr. Surg."},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"1441","DOI":"10.1002\/jcb.23061","article-title":"Modulatory effects of connexin-43 expression on gap junction intercellular communications with mast cells and fibroblasts","volume":"112","author":"Pistorio","year":"2011","journal-title":"J. Cell. Biochem."},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"95","DOI":"10.3109\/01913123.2014.981326","article-title":"Premature collagen fibril formation, fibroblast-mast cell interactions and mast cell-mediated phagocytosis of collagen in keloids","volume":"39","author":"Arbi","year":"2015","journal-title":"Ultrastruct. Pathol."},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1097\/PAI.0b013e3181efa2ef","article-title":"Cell talk: A phenomenon observed in the keloid scar by immunohistochemical study","volume":"19","author":"Shaker","year":"2011","journal-title":"Appl. Immunohistochem. Mol. Morphol."},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"901","DOI":"10.1016\/j.jid.2019.08.449","article-title":"Topical Application of a Mast Cell Stabilizer Improves Impaired Diabetic Wound Healing","volume":"140","author":"Tellechea","year":"2020","journal-title":"J. Investig. Dermatol."},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1016\/j.bmcl.2016.12.062","article-title":"Structure-activity relationship study and discovery of indazole 3-carboxamides as calcium-release activated calcium channel blockers","volume":"27","author":"Bai","year":"2017","journal-title":"Bioorg. Med. Chem. Lett."},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1016\/j.it.2012.09.008","article-title":"New models for analyzing mast cell functions in vivo","volume":"33","author":"Reber","year":"2012","journal-title":"Trends Immunol."},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1016\/S0002-9440(10)62055-X","article-title":"Mast cell-deficient W-sash c-kit mutant Kit W-sh\/W-sh mice as a model for investigating mast cell biology in vivo","volume":"167","author":"Grimbaldeston","year":"2005","journal-title":"Am. J. Pathol."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"668","DOI":"10.1016\/j.immuni.2011.11.004","article-title":"Mast cell deficiency, a game of kit and mouse","volume":"35","author":"Katz","year":"2011","journal-title":"Immunity"},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1146\/annurev-immunol-071719-094903","article-title":"Mast Cells in Inflammation and Disease: Recent Progress and Ongoing Concerns","volume":"38","author":"Galli","year":"2020","journal-title":"Annu. Rev. Immunol."},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1111\/wrr.12687","article-title":"Interleukin-33 encourages scar formation in murine fetal skin wounds","volume":"27","author":"Wulff","year":"2019","journal-title":"Wound Repair Regen."},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"1275","DOI":"10.3389\/fimmu.2018.01275","article-title":"Genetic and Imaging Approaches Reveal Pro-Inflammatory and Immunoregulatory Roles of Mast Cells in Contact Hypersensitivity","volume":"9","author":"Gaudenzio","year":"2018","journal-title":"Front. Immunol."},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1007\/978-1-4419-9533-9_11","article-title":"Mast cells and immunoregulation\/immunomodulation","volume":"716","author":"Tsai","year":"2011","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.cyto.2019.02.002","article-title":"The anti-fibrotic role of mast cells in the liver is mediated by HLA-G and interaction with hepatic stellate cells","volume":"117","author":"Amiot","year":"2019","journal-title":"Cytokine"},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1038\/ki.2013.98","article-title":"Mast cell chymase protects against renal fibrosis in murine unilateral ureteral obstruction","volume":"84","author":"Beghdadi","year":"2013","journal-title":"Kidney Int."},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"580","DOI":"10.3389\/fimmu.2019.00580","article-title":"Mast Cells in Cardiac Fibrosis: New Insights Suggest Opportunities for Intervention","volume":"10","author":"Legere","year":"2019","journal-title":"Front. 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