RAGE and TGF-β1 cross-talk regulate extracellular matrix turnover and cytokine synthesis in AGes exposed fibroblast cells

PLoS ONE

Volumn 11, Issue 3, 2016, Pages

  Serban, Andreea Iren ^a   Stanca, Loredana ^a   Geicu, Ovidiu Ionut ^a,b   Munteanu, Maria Cristina ^c   Dinischiotu, Anca ^b  

^a UNIVERSITY OF AGRONOMIC SCIENCES AND VETERINARY MEDICINE OF BUCHAREST   (Romania)

^b UNIVERSITY OF BUCHAREST   (Romania)

^c OKLAHOMA STATE UNIVERSITY   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

ADVANCED GLYCATION END PRODUCT; ADVANCED GLYCATION END PRODUCT RECEPTOR; ADVANCED GLYCATION END PRODUCTS-BOVINE SERUM ALBUMIN; BOVINE SERUM ALBUMIN; COLLAGEN; CONDITIONED MEDIUM; CYTOKINE; GRANULOCYTE MACROPHAGE COLONY STIMULATING FACTOR; INTERLEUKIN DERIVATIVE; MEMBRANE PROTEIN; TGFB1 PROTEIN, HUMAN; TRANSFORMING GROWTH FACTOR BETA1;

CELL CULTURE; CHEMISTRY; CONDITIONED MEDIUM; EXTRACELLULAR MATRIX; FIBROBLAST; HUMAN; INFLAMMATION; METABOLISM; SIGNAL TRANSDUCTION; UPREGULATION;

ADVANCED GLYCOSYLATION END PRODUCT-SPECIFIC RECEPTOR; CELLS, CULTURED; COLLAGEN; CULTURE MEDIA, CONDITIONED; CYTOKINES; EXTRACELLULAR MATRIX; FIBROBLASTS; GLYCOSYLATION END PRODUCTS, ADVANCED; GRANULOCYTE-MACROPHAGE COLONY-STIMULATING FACTOR; HUMANS; INFLAMMATION; INTERLEUKINS; MEMBRANE PROTEINS; SERUM ALBUMIN, BOVINE; SIGNAL TRANSDUCTION; TRANSFORMING GROWTH FACTOR BETA1; UP-REGULATION;

EID: 85006229888     PISSN: None     EISSN: 19326203     Source Type: Journal    
DOI: 10.1371/journal.pone.0152376     Document Type: Article

References (93)

1. Maillard LC (1912) Action des acides aminés sur les sucres: formation des mélanoidines par voie méthodique. C R Acad Sci Paris 154: 66–68.
2. Bookchin RM, Gallop PM (1968) Structure of hemoglobin A1c: nature of the N-terminal ß-chain blocking group. Biochem Biophys Res Commun 32: 86–93. PMID: 4874776
3. Gkogkolou P, Böhm M (2012) Advanced glycation end products: Key players in skin aging? Dermato-Endocrinol 4: 259–270.
4. Dyer DG, Dunn JA, Thorpe SR, Bailie KE, Lyons TJ, McCance DR, et al. (1993) Accumulation of Maillard reaction products in skin collagen in diabetes and aging. J Clin Invest 91: 2463–2469. PMID: 8514858
5. Sell DR, Biemel KM, Reihl O, Lederer MO, Strauch CM, Monnier VM (2005) Glucosepane is a major protein cross-link of the senescent human extracellular matrix. Relationship with diabetes. J Biol Chem 280: 12310–12315. PMID: 15677467
6. Sell DR, Lapolla A, Odetti P, Fogarty J, Monnier VM (1992) Pentosidine formation in skin correlates with severity of complication in individuals with long-standing IDDM. Diabetes 41: 1286–1292. PMID: 1397702
7. Fowlkes V, Clark J, Fix C, Law BA, Morales MO, Qiao X, et al. (2013) Type II diabetes promotes a myofibroblast phenotype in cardiac fibroblasts. Life Sci 92: 669–676. doi: 10.1016/j.lfs.2013.01.003 PMID: 23333820
8. Meng J, Sakata N, Takebayashi S, Asano T, Futata T, Nagai R, et al. (1998) Glycoxidation in aortic collagen from STZ-induced diabetic rats and its relevance to vascular damage. Atherosclerosis 136: 355–365. PMID: 9543107
9. Brownlee M (1995) Advanced protein glycosylation in diabetes and aging. Annu Rev Med 46: 223–234. PMID: 7598459
10. Vlassara H, Bucala R, Striker L (1994) Pathogenic effects of advanced glycosylation: biochemical, biologic, and clinical implications for diabetes and aging. Lab Invest 70: 138–151. PMID: 8139257
11. Liao H, Zakhaleva J, Chen W (2009) Cells and tissue interactions with glycated collagen and their relevance to delayed diabetic wound healing. Biomaterials 30: 1689–1696. doi: 10.1016/j.biomaterials. 2008.11.038 PMID: 19157537
12. Ott C, Jacobs K, Haucke E, Santos AN, Grune T, Simm A (2014) Role of advanced glycation end products in cellular signaling. Redox Biology 2: 411–429. doi: 10.1016/j.redox.2013.12.016 PMID: 24624331
13. Okano Y, Masaki H, Sakurai H (2002) Dysfunction of dermal fibroblasts induced by advanced glycation end-products (AGEs) and the contribution of a nonspecific interaction with cell membrane and AGEs. J Dermatol Sci 29: 171–180. PMID: 12234706
14. Candido R, Forbes JM, Thomas MC, Thallas V, Dean RG, Burns WC, et al. (2003) A breaker of advanced glycation end products attenuates diabetes-induced myocardial structural changes. Circ Res 92: 785–792. PMID: 12623881
15. Hobizal KB, Wukich DK (2012) Diabetic foot infections: current concept review. Diabet Foot Ankle 3.
16. Frykberg RG (1998) Diabetic foot ulcers: current concepts. J Foot Ankle Surg 37: 440–446. PMID: 9798178
17. Jordana M, Särnstrand B, Sime PJ, Ramis I (1994) Immune-inflammatory functions of fibroblasts. Eur Respir J 7: 2212–2222. PMID: 7713206
18. Gabbiani G (2003) The myofibroblast in wound healing and fibrocontractive diseases. J Pathol 200: 500–503. PMID: 12845617
19. Sappino A, Schürch W, Gabbiani G (1990) Differentiation repertoire of fibroblastic cells: expression of cytoskeletal proteins as marker of phenotypic modulations. Lab Invest 63: 144–161. PMID: 2116562
20. Darby IA, Hewitson TD (2007) Fibroblast differentiation in wound healing and fibrosis. Int Rev Cytol 257: 143–179. PMID: 17280897
21. Ban CR, Twigg SM (2008) Fibrosis in diabetes complications: pathogenic mechanisms and circulating and urinary markers. Vasc Health Risk Manag 4: 575. PMID: 18827908
22. Yang J, Xue Q, Miao L, Cai L (2011) Pulmonary fibrosis: a possible diabetic complication. Diabetes Metab Res Rev 27: 311–317. doi: 10.1002/dmrr.1175 PMID: 21309056
23. Lanng S, Thorsteinsson B, Lund-Andersen C, Nerup J, Schiøtz P, Koch C (1994) Diabetes mellitus in Danish cystic fibrosis patients: prevalence and late diabetic complications. Acta Paediatrica 83: 72–77. PMID: 8193478
24. Leask A, Abraham DJ (2004) TGF-β signaling and the fibrotic response. FASEB J 18: 816–827. PMID: 15117886
25. Howard EW, Benton R, Ahern-Moore J, Tomasek JJ (1996) Cellular contraction of collagen lattices is inhibited by nonenzymatic glycation. Exp Cell Res 228: 132–137. PMID: 8892980
26. Ramasamy R, Yan SF, Herold K, Clynes R, Schmidt AM (2008) Receptor for advanced glycation end products: fundamental roles in the inflammatory response: winding the way to the pathogenesis of endothelial dysfunction and atherosclerosis. Ann N Y Acad Sci 1126: 7–13. doi: 10.1196/annals.1433. 056 PMID: 18448789
27. Wendt TM, Tanji N, Guo J, Kislinger TR, Qu W, Lu Y, et al. (2003) RAGE drives the development of glomerulosclerosis and implicates podocyte activation in the pathogenesis of diabetic nephropathy. Am J Pathol 162: 1123–1137. PMID: 12651605
28. Giacomoni PU, Rein G (2001) Factors of skin ageing share common mechanisms. Biogerontology 4: 219–229.
29. Yan SF, Ramasamy R, Schmidt AM (2008) Mechanisms of Disease: advanced glycation end-products and their receptor in inflammation and diabetes complications. Nat Clin Pract Endocrinol Metab 4: 285–293. doi: 10.1038/ncpendmet0786 PMID: 18332897
30. Li J, Schmidt AM (1997) Characterization and functional analysis of the promoter of RAGE, the receptor for advancedglycation end products. J Biol Chem 272: 16498–16506. PMID: 9195959
31. Sanchez AP, Sharma K (2009) Transcription factors in the pathogenesis of diabetic nephropathy. Expert Rev Mol Med 11: e13. doi: 10.1017/S1462399409001057 PMID: 19397838
32. Ishihara K, Tsutsumi K, Kawane S, Nakajima M, Kasaoka T (2003) The receptor for advanced glycation end-products (RAGE) directly binds to ERK by a D-domain-like docking site. FEBS Lett 550: 107–113. PMID: 12935895
33. Lee MK, Pardoux C, Hall MC, Lee PS, Warburton D, Qing J, et al. (2007) TGF-β activates Erk MAP kinase signalling through direct phosphorylation of ShcA. EMBO J 26: 3957–3967. PMID: 17673906
34. Xie L, Law BK, Chytil AM, Brown KA, Aakre ME, Moses HL (2004) Activation of the Erk pathway is required for TGF-β1-induced EMT in vitro. Neoplasia 6: 603–610. PMID: 15548370
35. Zhao J, Randive R, Stewart JA (2014) Molecular mechanisms of AGE/RAGE-mediated fibrosis in the diabetic heart. World J Diabetes 5: 860. doi: 10.4239/wjd.v5.i6.860 PMID: 25512788
36. Chuah YK, Basir R, Talib H, Tie TH, Nordin N (2013) Receptor for advanced glycation end products and its involvement in inflammatory diseases. Int J Inflam 2013.
37. Smith WB, Noack L, Khew-Goodall Y, Isenmann S, Vadas MA, Gamble JR (1996) Transforming growth factor-beta 1 inhibits the production of IL-8 and the transmigration of neutrophils through activated endothelium. J Immunol 157: 360–368. PMID: 8683138
38. Ling E, Robinson DS (2002) Transforming growth factor-bβ1: its anti-inflammatory and pro-fibrotic effects. Clin Exp Allergy 32: 175–178. PMID: 11929477
39. Wang M, Zhao D, Spinetti G, Zhang J, Jiang LQ, Pintus G, et al. (2006) Matrix metalloproteinase 2 activation of transforming growth factor-beta1 (TGF-beta1) and TGF-beta1-type II receptor signaling within the aged arterial wall. Arterioscler Thromb Vasc Biol 26: 1503–1509. PMID: 16690877
40. Varani J, Dame MK, Rittie L, Fligiel SEG, Sewon Kang S, Fisher GJ, et al. (2006) Decreased collagen production in chronologically aged skin: Roles of age-dependent alteration in fibroblast function and defective mechanical stimulation. Am J Pathol 168: 1861–1868. PMID: 16723701
41. Serban AI, Stanca L, Geicu OI, Munteanu MC, Costache M, Dinischiotu A (2015) Extracellular matrix is modulated in advanced glycation end products milieu via a RAGE receptor dependent pathway boosted by transforming growth factor-β1. J Diabetes 7: 114–124. doi: 10.1111/1753-0407.12154 PMID: 24666836
42. Serban AI, Stanca L, Geicu OI, Dinischiotu A (2015) AGEs-induced IL-6 synthesis precedes RAGE upregulation in HEK 293 cells: An alternative inflammatory mechanism? Int J Mol Sci 16: 20100–20117. doi: 10.3390/ijms160920100 PMID: 26307981
43. Bradford M (1976) A rapid and sensitive methode for quantitation of microgram quantities of protein utilizing the principle pf protein-dye binding. Anal Biochem 72: 248–254. PMID: 942051
44. Ngo P, Ramalingam P, Phillips JA, Furuta G (2006) Collagen gel contraction assay. Methods Mol Biol 341: 103–109. PMID: 16799192
45. Romero-Calvo I, Ocón B, Martínez-Moya P, Suárez MD, Zarzuelo A, Martínez-Augustin O, et al. (2010) Reversible Ponceau staining as a loading control alternative to actin in Western blots. Anal Biochem 401: 318–320. doi: 10.1016/j.ab.2010.02.036 PMID: 20206115
46. Chomezynski P (1993) A reagent for the single-step simultaneous isolation of RNA, DNA and proteins from cell and tissue samples. Biotechniques 15: 532–537. PMID: 7692896
47. Pfaffl MW (2001) A new mathematical model for relative quantification in real-time RT-PCR. Nucleic Acids Res 29: 2002–2007.
48. Lohwasser C, Neureiter D, Weigle B, Kirchner T, Schuppan D (2006) The receptor for advanced glycation end products is highly expressed in the skin and upregulated by advanced glycation end products and tumor necrosis factor-alpha. J Invest Dermatol 126: 291–299. PMID: 16374460
49. Chen Q, Lee CE, Denard B, Ye J (2014) Sustained induction of collagen synthesis by TGF-β requires regulated intramembrane proteolysis of CREB3L1. PLoS One 9: e108528. doi: 10.1371/journal.pone. 0108528 PMID: 25310401
50. Denard B, Seemann J, Chen Q, Gay A, Huang H, Chen Y, et al. (2011) The membrane-bound transcription factor CREB3L1 is activated in response to virus infection to inhibit proliferation of virus-infected cells. Cell Host Microbe 10: 65–74. doi: 10.1016/j.chom.2011.06.006 PMID: 21767813
51. Uhlén M, Fagerberg L, Hallström BM, Lindskog C, Oksvold P, Mardinoglu A, et al. (2015) Proteomics. Tissue-based map of the human proteome. Science 347: 1260419. doi: 10.1126/science.1260419 PMID: 25613900
52. Goel SA, Guo LW, Shi XD, Kundi R, Sovinski G, Seedial S, et al. (2013) Preferential secretion of collagen type 3 versus type 1 from adventitial fibroblasts stimulated by TGF-β/Smad3-treated medial smooth muscle cells. Cell Signal 25: 955–960. doi: 10.1016/j.cellsig.2012.12.021 PMID: 23280188
53. Stallmach A, Schuppan D, Riese HH, Matthes H, Riecken EO (1992) Increased collagen type III synthesis by fibroblasts isolated from strictures of patients with Crohn's disease. Gastroenterology 102: 1920–1929. PMID: 1587410
54. Bermudez DM, Herdrich BJ, Xu J, Lind R, Beason DP, Mitchell ME, et al. (2011) Impaired biomechanical properties of diabetic skin implications in pathogenesis of diabetic wound complications. Am J Pathol 178: 2215–2223. doi: 10.1016/j.ajpath.2011.01.015 PMID: 21514435
55. Karagiannis ED, Popel AS (2004) A theoretical model of type I collagen proteolysis by matrix metalloproteinase (MMP) 2 and membrane type 1 MMP in the presence of tissue inhibitor of metalloproteinase 2. J Biol Chem 279: 39105–39114. PMID: 15252025
56. Nagase H, Visse R (2011) Triple helicase activity and structural basis of collagenolysis. In: Parks WC, Mecham R, editors. Extracellular Matrix Degradation. Berlin Heidelberg: Springer-Verlag pp. 95–122.
57. Parks WC, Wilson CL, López-Boado YS (2004) Matrix metalloproteinases as modulators of inflammation and innate immunity. Nat Rev Immunol 4: 617–629. PMID: 15286728
58. Brown CN, Finch JG (2010) Which mesh for hernia repair? Ann R Coll Surg Engl 92: 272–278. doi: 10. 1308/003588410X12664192076296 PMID: 20501011
59. Peng Y, Kim J-M, Park H-S, Yang A, Islam C, Lakatta EG, et al. (2016) AGE-RAGE signal generates a specific NF-κB RelA “barcode” that directs collagen I expression. Sci Rep 6: 18822. doi: 10.1038/ srep18822 PMID: 26729520
60. Bierhaus A, Humpert PM, Morcos M, Wendt T, Chavakis T, Arnold B, et al. (2005) Understanding RAGE, the receptor for advanced glycation end products. J Mol Med 83: 876–886. PMID: 16133426
61. Xu X, Chen H, Zhu X, Ma Y, Liu Q, Xue Y, et al. (2013) S100A9 promotes human lung fibroblast cells activation through receptor for advanced glycation end-product-mediated extracellular-regulated kinase 1/2, mitogen-activated protein-kinase and nuclear factor-κB-dependent pathways. Clin Exp Immunol 173: 523–535. doi: 10.1111/cei.12139 PMID: 23682982
62. Hough C, Radu M, Doré J (2012) TGF-beta induced Erk phosphorylation of smad linker region regulates smad signaling. PLoS One 7: e42513. doi: 10.1371/journal.pone.0042513 PMID: 22880011
63. Roomi MW, Monterrey JC, Kalinovsky T, Rath M, Niedzwiecki A (2009) Distinct patterns of matrix metalloproteinase-2 and -9 expression in normal human cell lines. Oncol Rep 21: 821–826. PMID: 19212645
64. Duivenvoorden WC, Hirte HW, Singh G (1999) Transforming growth factor beta1 acts as an inducer of matrix metalloproteinase expression and activity in human bone-metastasizing cancer cells. Clin Exp Metastasis 17: 27–34. PMID: 10390144
65. Qin H, Moellinger JD, Wells A, Windsor LJ, Sun Y, Benveniste EN (1998) Transcriptional suppression of matrix metalloproteinase-2 gene expression in human astroglioma cells by TNF-alpha and IFN-gamma. J Immunol 161: 6664–6673. PMID: 9862695
66. Fukami K, Yamagishi S-I, Coughlan MT, Harcourt BE, Kantharidis P, Thallas-Bonke V, et al. (2014) Ramipril inhibits AGE-RAGE-induced matrix metalloproteinase-2 activation in experimental diabetic nephropathy. Diabetol Metab Syndr 6: 86. doi: 10.1186/1758-5996-6-86 PMID: 25143788
67. Lalla E, Lamster IB, Feit M, Huang L, Spessot A, Qu W, et al. (2000) Blockade of RAGE suppresses periodontitis-associated bone loss in diabetic mice. J Clin Invest 105: 1117–1124. PMID: 10772656
68. Karsdal MA, Larsen L, Engsig MT, Lou H, Ferreras M, Lochter A, et al. (2002) Matrix metalloproteinase-dependent activation of latent transforming growth factor-beta controls the conversion of osteoblasts into osteocytes by blocking osteoblast apoptosis. J Biol Chem 277: 44061–44067. PMID: 12226090
69. George SJ, Johnson JL, Smith MA, Angelini GD, Jackson CL (2005) Transforming growth factor-beta is activated by plasmin and inhibits smooth muscle cell death in human saphenous vein. J Vasc Res 42: 247–254. PMID: 15870504
70. Banchereau J, Pascual V, O'Garra A (2012) From IL-2 to IL-37: the expanding spectrum of anti-inflammatory cytokines. Nat Immunol 13: 925–931. doi: 10.1038/ni.2406 PMID: 22990890
71. Zhou Y, Hagood JS, Murphy-Ullrich JE (2004) Thy-1 expression regulates the ability of rat lung fibroblasts to activate transforming growth factor-beta in response to fibrogenic stimuli. Am J Pathol 165: 659–669. PMID: 15277239
72. Aoudjehane L, Pissaia AJ, Scatton O, Podevin P, Massault PP, Chouzenoux S, et al. (2008) Interleukin-4 induces the activation and collagen production of cultured human intrahepatic fibroblasts via the STAT-6 pathway. Lab Invest 88: 973–985. doi: 10.1038/labinvest.2008.61 PMID: 18626468
73. Kanellakis P, Ditiatkovski M, Kostolias G, Bobik A (2012) A pro-fibrotic role for interleukin-4 in cardiac pressure overload. Cardiovasc Res 95: 77–85. doi: 10.1093/cvr/cvs142 PMID: 22492684
74. Rasheed Z, Akhtar N, Haqqi TM (2011) Advanced glycation end products induce the expression of interleukin-6 and interleukin-8 by receptor for advanced glycation end product-mediated activation of mitogen-activated protein kinases and nuclear factor-κB in human osteoarthritis chondrocytes. Rheumatology 50: 838–851. doi: 10.1093/rheumatology/keq380 PMID: 21172926
75. Lonnemann G, Engler-Blum G, Müller GA, Koch KM, Dinarello CA (1995) Cytokines in human renal interstitial fibrosis. II. Intrinsic interleukin (IL)-1 synthesis and IL-1-dependent production of IL-6 and IL-8 by cultured kidney fibroblasts. Kidney Int 47: 845–854. PMID: 7752583
76. Sullivan DE, Ferris M, Nguyen H, Abboud E, Brody AR (2009) TNF-alpha induces TGF-beta1 expression in lung fibroblasts at the transcriptional level via AP-1 activation. J Cell Mol Med 13: 1866–1876. doi: 10.1111/j.1582-4934.2009.00647.x PMID: 20141610
77. Nah SS, Choi IY, Yoo B, Kim YG, Moon HB, Lee CK (2007) Advanced glycation end products increases matrix metalloproteinase-1, -3, and -13, and TNF-alpha in human osteoarthritic chondrocytes. FEBS Lett 581: 1928–1932. PMID: 17434489
78. Mohan MJ, Seaton T, Mitchell J, Howe A, Blackburn K, Burkhart W, et al. (2002) The tumor necrosis factor-alpha converting enzyme (TACE): a unique metalloproteinase with highly defined substrate selectivity. Biochemistry 41: 9462–9469. PMID: 12135369
79. Gross TJ, Hunninghake GW (2001) Idiopathic pulmonary fibrosis. N Engl J Med 345: 517–525. PMID: 11519507
80. King TEJ, Pardo A, Selman M (2011) Idiopathic pulmonary fibrosis. Lancet 378: 1949–1961. doi: 10. 1016/S0140-6736(11)60052-4 PMID: 21719092
81. Crosby LM, Waters CM (2010) Epithelial repair mechanisms in the lung. Am J Physiol Lung Cell Mol Physiol 298: L715–L731. doi: 10.1152/ajplung.00361.2009 PMID: 20363851
82. Shiomi A, Usui T (2015) Pivotal roles of GM-CSF in autoimmunity and inflammation. Mediators Inflamm 2015: 568543. doi: 10.1155/2015/568543 PMID: 25838639
83. Leivonen SK, Lazaridis K, Decock J, Chantry A, Edwards DR, Kähäri VM (2013) TGF-β-elicited induction of tissue inhibitor of metalloproteinases (TIMP)-3 expression in fibroblasts involves complex interplay between Smad3, p38α, and ERK1/2. PLoS One 8: e57474. doi: 10.1371/journal.pone.0057474 PMID: 23468994
84. Amour A, Slocombe PM, Webster A, Butler M, Knight CG, Smith BJ, et al. (1998) TNF-alpha converting enzyme (TACE) is inhibited by TIMP-3. FEBS Lett 435: 39–44. PMID: 9755855
85. Oppenheim JJ, Neta R (1994) Pathophysiological roles of cytokines in development, immunity, and inflammation. FASEB J 8: 158–162. PMID: 8119486
86. Jiang K, Cao S, Cui JZ, Matsubara JA (2013) Immuno-modulatory Effect of IFN-gamma in AMD and its Role as a Possible Target for Therapy. J Clinic Experiment Ophthalmol S2: 007.
87. Kahan A, Amor B, Menkes CJ, Strauch G (1989) Recombinant interferon-gamma in the treatment of systemic sclerosis. Am J Med 87: 273–277. PMID: 2505614
88. Hein R, Behr J, Hündgen M, Hunzelmann N, Meurer M, Braun-Falco O, et al. (1992) Treatment of systemic sclerosis with gamma-interferon. Br J Dermatol 126: 496–501. PMID: 1610690
89. Sáez-Royuela F, Porres JC, Moreno A, Castillo I, Martinez G, Galiana F, et al. (1991) High doses of recombinant alpha-interferon or gamma-interferon for chronic hepatitis C: a randomized, controlled trial. Hepatology 13: 327–331. PMID: 1899852
90. Tsoutsou PG, Gourgoulianis KI, Petinaki E, Germenis A, Tsoutsou AG, Mpaka M, et al. (2006) Cytokine levels in the sera of patients with idiopathic pulmonary fibrosis. Respir Med 100: 938–945. PMID: 16236490
91. Raghu G, Brown KK, Bradford WZ, Starko K, Noble PW, Schwartz DA, et al. (2004) A placebo-controlled trial of interferon gamma-1b in patients with idiopathic pulmonary fibrosis. N Engl J Med 350: 125–133. PMID: 14711911
92. Balaji S, King A, Marsh E, LeSaint M, Bhattacharya SS, Han N, et al. (2015) The role of interleukin-10 and hyaluronan in murine fetal fibroblast function in vitro: implications for recapitulating fetal regenerative wound healing. PLoS One 10: e0124302. doi: 10.1371/journal.pone.0124302 PMID: 25951109
93. Englert JM, Hanford LE, Kaminski N, Tobolewski JM, Tan RJ, Fattman CL, et al. (2008) A role for the receptor for advanced glycation end products in idiopathic pulmonary fibrosis. Am J Pathol 172: 583–591. doi: 10.2353/ajpath.2008.070569 PMID: 18245812