Influence of age on wound healing and fibrosis

Journal of Pathology

Volumn 229, Issue 2, 2013, Pages 310-322

  Kapetanaki, Maria G ^a,b   Mora, Ana L ^b   Rojas, Mauricio ^a,b  

^a UNIVERSITY OF PITTSBURGH MEDICAL CENTER   (United States)

^b UNIVERSITY OF PITTSBURGH SCHOOL OF MEDICINE   (United States)

Author keywords

ageing; chronic obstructive pulmonary disease; extracellular matrix; lung fibrosis; telomeres

Indexed keywords

DNA;

AGING; APOPTOSIS; CELL REGENERATION; CELL STRESS; EPIGENETICS; EPITHELIUM CELL; EXTRACELLULAR MATRIX; FIBROBLAST; GENE EXPRESSION; HUMAN; LUNG FIBROSIS; LUNG PARENCHYMA; MESENCHYMAL STEM CELL; MUTATION; NONHUMAN; OXIDATIVE STRESS; PATHOLOGY; PRIORITY JOURNAL; REVIEW; TELOMERE; WOUND HEALING;

AGE FACTORS; AGING; ANIMALS; APOPTOSIS; EPIGENESIS, GENETIC; EPITHELIAL CELLS; EXTRACELLULAR MATRIX; FIBROBLASTS; FIBROSIS; HUMANS; LUNG; MESENCHYMAL STROMAL CELLS; PULMONARY FIBROSIS; TELOMERE; WOUND HEALING;

EID: 84871049210     PISSN: 00223417     EISSN: 10969896     Source Type: Journal    
DOI: 10.1002/path.4122     Document Type: Review

References (156)

1. Kirkwood TB,. Understanding the odd science of aging. Cell 2005; 120: 437-447. (Pubitemid 40269759)
2. Marshman E, Booth C, Potten CS,. The intestinal epithelial stem cell. Bioessays 2002; 24: 91-98. (Pubitemid 34108424)
3. Martin K, Potten CS, Roberts SA, et al,. Altered stem cell regeneration in irradiated intestinal crypts of senescent mice. J Cell Sci 1998; 111 (16): 2297-2303. (Pubitemid 28434917)
4. Kasahara Y, Tuder RM, Cool CD, et al,. Endothelial cell death and decreased expression of vascular endothelial growth factor and vascular endothelial growth factor receptor 2 in emphysema. Am J Respir Crit Care Med 2001; 163: 737-744. (Pubitemid 32230838)
5. Tsuji T, Aoshiba K, Nagai A,. Cigarette smoke induces senescence in alveolar epithelial cells. Am J Respir Cell Mol Biol 2004; 31: 643-649. (Pubitemid 39587494)
6. Yokohori N, Aoshiba K, Nagai A,. Increased levels of cell death and proliferation in alveolar wall cells in patients with pulmonary emphysema. Chest 2004; 125: 626-632. (Pubitemid 38223645)
7. Warburton D, Bellusci S, De Langhe S, et al,. Molecular mechanisms of early lung specification and branching morphogenesis. Pediatr Res 2005; 57: 26-37R.
8. Warburton D, Bellusci S, Del Moral PM, et al,. Growth factor signaling in lung morphogenetic centers: automaticity, stereotypy and symmetry. Respir Res 2003; 4: 5.
9. Warburton D, Schwarz M, Tefft D, et al,. The molecular basis of lung morphogenesis. Mech Dev 2000; 92: 55-81. (Pubitemid 30125962)
10. Janssens JP, Pache JC, Nicod LP,. Physiological changes in respiratory function associated with ageing. Eur Respir J 1999; 13: 197-205. (Pubitemid 29079244)
11. Ito K, Barnes PJ,. COPD as a disease of accelerated lung aging. Chest 2009; 135: 173-180.
12. Miravitlles M, Soriano JB, Garcia-Rio F, et al,. Prevalence of COPD in Spain: impact of undiagnosed COPD on quality of life and daily life activities. Thorax 2009; 64: 863-868.
13. Navaratnam V, Fleming KM, West J, et al,. The rising incidence of idiopathic pulmonary fibrosis in the UK. Thorax 2011; 66: 462-467.
14. Faner R, Rojas M, Macnee W, et al,. Abnormal lung aging in chronic obstructive pulmonary disease and idiopathic pulmonary fibrosis. Am J Respir Crit Care Med 2012; 186: 306-313.
15. Crosby LM, Waters CM,. Epithelial repair mechanisms in the lung. Am J Physiol Lung Cell Mol Physiol 2010; 298: L715-731.
16. Crystal RG, Randell SH, Engelhardt JF, et al,. Airway epithelial cells: current concepts and challenges. Proc Am Thorac Soc 2008; 5: 772-777.
17. Strieter RM,. What differentiates normal lung repair and fibrosis? Inflammation, resolution of repair, and fibrosis. Proc Am Thorac Soc 2008; 5: 305-310. (Pubitemid 351520676)
18. Kim DS, Collard HR, King TE, Jr,. Classification and natural history of the idiopathic interstitial pneumonias. Proc Am Thorac Soc 2006; 3: 285-292. (Pubitemid 43845404)
19. Xu J, Gonzalez ET, Iyer SS, et al,. Use of senescence-accelerated mouse model in bleomycin-induced lung injury suggests that bone marrow-derived cells can alter the outcome of lung injury in aged mice. J Gerontol A Biol Sci Med Sci 2009; 64: 731-739.
20. Torres-Gonzalez E, Bueno M, Tanaka A, et al,. Role of endoplasmic reticulum stress in age-related susceptibility to lung fibrosis. Am J Respir Cell Mol Biol 2012; 46: 748-756.
21. Silver FH, DeVore D, Siperko LM,. Invited review: role of mechanophysiology in aging of ECM: effects of changes in mechanochemical transduction. J Appl Physiol 2003; 95: 2134-2141. (Pubitemid 37310401)
22. Limper AH, Roman J. Fibronectin. A versatile matrix protein with roles in thoracic development, repair and infection. Chest 1992; 101: 1663-1673.
23. Mays PK, Bishop JE, Laurent GJ,. Age-related changes in the proportion of types I and III collagen. Mech Ageing Dev 1988; 45: 203-212. (Pubitemid 19004062)
24. Huang K, Mitzner W, Rabold R, et al,. Variation in senescent-dependent lung changes in inbred mouse strains. J Appl Physiol 2007; 102: 1632-1639. (Pubitemid 46571061)
25. Shuckett R, Malemud CJ,. Distinct cartilage proteoglycan chromatographic elution patterns in advanced human hip osteoarthritis: correlations with histologic analysis. J Rheumatol 1990; 17: 357-363. (Pubitemid 20106615)
26. Roman J, Ritzenthaler JD, Bechara R, et al,. Ethanol stimulates the expression of fibronectin in lung fibroblasts via kinase-dependent signals that activate CREB. Am J Physiol Lung Cell Mol Physiol 2005; 288: L975-987. (Pubitemid 40547926)
27. Muro AF, Moretti FA, Moore BB, et al,. An essential role for fibronectin extra type III domain A in pulmonary fibrosis. Am J Respir Crit Care Med 2008; 177: 638-645. (Pubitemid 351442171)
28. McDonald JA, Kelley DG, Broekelmann TJ,. Role of fibronectin in collagen deposition: Fab′ to the gelatin-binding domain of fibronectin inhibits both fibronectin and collagen organization in fibroblast extracellular matrix. J Cell Biol 1982; 92: 485-492. (Pubitemid 12157421)
29. Bitterman PB, Rennard SI, Adelberg S, et al,. Role of fibronectin as a growth factor for fibroblasts. J Cell Biol 1983; 97: 1925-1932. (Pubitemid 14208917)
30. Magnuson VL, Young M, Schattenberg DG, et al,. The alternative splicing of fibronectin pre-mRNA is altered during aging and in response to growth factors. J Biol Chem 1991; 266: 14654-14662. (Pubitemid 21907552)
31. Sueblinvong V, Neujahr DC, Mills ST, et al,. Predisposition for disrepair in the aged lung. Am J Med Sci 2012; 344: 41-51.
32. White ES, Sagana RL, Booth AJ, et al,. Control of fibroblast fibronectin expression and alternative splicing via the PI3K/Akt/mTOR pathway. Exp Cell Res 2010; 316: 2644-2653.
33. Singer NG, Caplan AI,. Mesenchymal stem cells: mechanisms of inflammation. Annu Rev Pathol 2011; 6: 457-478.
34. Dominici M, Le Blanc K, Mueller I, et al,. Minimal criteria for defining multipotent mesenchymal stromal cells. The International Society for Cellular Therapy position statement. Cytotherapy 2006; 8: 315-317.
35. Wang G, Bunnell BA, Painter RG, et al,. Adult stem cells from bone marrow stroma differentiate into airway epithelial cells: potential therapy for cystic fibrosis. Proc Natl Acad Sci USA 2005; 102: 186-191. (Pubitemid 40094450)
36. Spees JL, Olson SD, Ylostalo J, et al,. Differentiation, cell fusion, and nuclear fusion during ex vivo repair of epithelium by human adult stem cells from bone marrow stroma. Proc Natl Acad Sci USA 2003; 100: 2397-2402. (Pubitemid 36297510)
37. Xu J, Woods CR, Mora AL, et al,. Prevention of endotoxin-induced systemic response by bone marrow-derived mesenchymal stem cells in mice. Am J Physiol Lung Cell Mol Physiol 2007; 293: L131-141. (Pubitemid 47080721)
38. Kotton DN, Ma BY, Cardoso WV, et al,. Bone marrow-derived cells as progenitors of lung alveolar epithelium. Development 2001; 128: 5181-5188. (Pubitemid 34083043)
39. Hashimoto N, Jin H, Liu T, et al,. Bone marrow-derived progenitor cells in pulmonary fibrosis. J Clin Invest 2004; 113: 243-252. (Pubitemid 38544209)
40. Phillips RJ, Burdick MD, Hong K, et al,. Circulating fibrocytes traffic to the lungs in response to CXCL12 and mediate fibrosis. J Clin Invest 2004; 114: 438-446. (Pubitemid 39071616)
41. Moeller A, Gilpin SE, Ask K, et al,. Circulating fibrocytes are an indicator of poor prognosis in idiopathic pulmonary fibrosis. Am J Respir Crit Care Med 2009; 179: 588-594.
42. Jun D, Garat C, West J, et al,. The pathology of bleomycin-induced fibrosis is associated with loss of resident lung mesenchymal stem cells that regulate effector T-cell proliferation. Stem Cells 2011; 29: 725-735.
43. Stolzing A, Jones E, McGonagle D, et al,. Age-related changes in human bone marrow-derived mesenchymal stem cells: consequences for cell therapies. Mech Ageing Dev 2008; 129: 163-173.
44. Bork S, Pfister S, Witt H, et al,. DNA methylation pattern changes upon long-term culture and aging of human mesenchymal stromal cells. Aging Cell 2010; 9: 54-63.
45. Roobrouck VD, Ulloa-Montoya F, Verfaillie CM,. Self-renewal and differentiation capacity of young and aged stem cells. Exp Cell Res 2008; 314: 1937-1944.
46. Choumerianou DM, Martimianaki G, Stiakaki E, et al,. Comparative study of stemness characteristics of mesenchymal cells from bone marrow of children and adults. Cytotherapy 2010; 12: 881-887.
47. Zhou S, Greenberger JS, Epperly MW, et al,. Age-related intrinsic changes in human bone-marrow-derived mesenchymal stem cells and their differentiation to osteoblasts. Aging Cell 2008; 7: 335-343. (Pubitemid 351663902)
48. Lavasani M, Robinson AR, Lu A, et al,. Muscle-derived stem/progenitor cell dysfunction limits healthspan and lifespan in a murine progeria model. Nat Commun 2012; 3: 608.
49. Conboy IM, Conboy MJ, Wagers AJ, et al,. Rejuvenation of aged progenitor cells by exposure to a young systemic environment. Nature 2005; 433: 760-764. (Pubitemid 40292986)
50. Fan M, Chen W, Liu W, et al,. The effect of age on the efficacy of human mesenchymal stem cell transplantation after a myocardial infarction. Rejuvenat Res 2010; 13: 429-438.
51. Hermann A, List C, Habisch HJ, et al,. Age-dependent neuroectodermal differentiation capacity of human mesenchymal stromal cells: limitations for autologous cell replacement strategies. Cytotherapy 2010; 12: 17-30.
52. Walenda T, Bork S, Horn P, et al,. Co-culture with mesenchymal stromal cells increases proliferation and maintenance of haematopoietic progenitor cells. J Cell Mol Med 2010; 14: 337-350.
53. Khan M, Mohsin S, Khan SN, et al,. Repair of senescent myocardium by mesenchymal stem cells is dependent on the age of donor mice. J Cell Mol Med 2011; 15: 1515-1527.
54. Guillot PV, Gotherstrom C, Chan J, et al,. Human first-trimester fetal MSC express pluripotency markers and grow faster and have longer telomeres than adult MSC. Stem Cells 2007; 25: 646-654. (Pubitemid 46354223)
55. Baxter MA, Wynn RF, Jowitt SN, et al,. Study of telomere length reveals rapid aging of human marrow stromal cells following in vitro expansion. Stem Cells 2004; 22: 675-682. (Pubitemid 39172200)
56. Blackburn EH,. Structure and function of telomeres. Nature 1991; 350: 569-573. (Pubitemid 21912269)
57. Greider CW, Blackburn EH,. A telomeric sequence in the RNA of Tetrahymena telomerase required for telomere repeat synthesis. Nature 1989; 337: 331-337.
58. McClintock B,. The behavior in successive nuclear divisions of a chromosome broken at meiosis. Proc Natl Acad Sci USA 1939; 25: 405-416.
59. Oeseburg H, de Boer RA, van Gilst WH, et al,. Telomere biology in healthy aging and disease. Pflugers Arch 2009; 459: 259-268.
60. Allsopp RC, Vaziri H, Patterson C, et al,. Telomere length predicts replicative capacity of human fibroblasts. Proc Natl Acad Sci USA 1992; 89: 10114-10118.
61. Olovnikov AM,. A theory of marginotomy. The incomplete copying of template margin in enzymic synthesis of polynucleotides and biological significance of the phenomenon. J Theor Biol 1973; 41: 181-190.
62. Harman D,. Role of free radicals in aging and disease. Ann NY Acad Sci 1992; 673: 126-141. (Pubitemid 23057567)
63. Hastie ND, Dempster M, Dunlop MG, et al,. Telomere reduction in human colorectal carcinoma and with ageing. Nature 1990; 346: 866-868.
64. Loeb LA, Wallace DC, Martin GM,. The mitochondrial theory of aging and its relationship to reactive oxygen species damage and somatic mtDNA mutations. Proc Natl Acad Sci USA 2005; 102: 18769-18770. (Pubitemid 43049521)
65. Mooi WJ, Peeper DS,. Oncogene-induced cell senescence-halting on the road to cancer. N Engl J Med 2006; 355: 1037-1046. (Pubitemid 44343547)
66. Zhu J, Woods D, McMahon M, et al,. Senescence of human fibroblasts induced by oncogenic Raf. Genes Dev 1998; 12: 2997-3007. (Pubitemid 28469302)
67. d'Adda di, Fagagna F, Reaper PM, Clay-Farrace L, et al,. A DNA damage checkpoint response in telomere-initiated senescence. Nature 2003; 426: 194-198. (Pubitemid 37442589)
68. Jones DP,. Extracellular redox state: refining the definition of oxidative stress in aging. Rejuvenat Res 2006; 9: 169-181. (Pubitemid 43856805)
69. Saretzki G, Von Zglinicki T,. Replicative aging, telomeres, and oxidative stress. Ann NY Acad Sci 2002; 959: 24-29. (Pubitemid 34457548)
70. Valdes AM, Andrew T, Gardner JP, et al,. Obesity, cigarette smoking, and telomere length in women. Lancet 2005; 366: 662-664. (Pubitemid 41176019)
71. Bodnar AG, Ouellette M, Frolkis M, et al,. Extension of life-span by introduction of telomerase into normal human cells. Science 1998; 279: 349-352. (Pubitemid 28063364)
72. Vaziri H, Benchimol S,. Reconstitution of telomerase activity in normal human cells leads to elongation of telomeres and extended replicative life span. Curr Biol 1998; 8: 279-282. (Pubitemid 28111458)
73. Ulaner GA, Giudice LC,. Developmental regulation of telomerase activity in human fetal tissues during gestation. Mol Hum Reprod 1997; 3: 769-773. (Pubitemid 127509610)
74. Karlseder J, Smogorzewska A, de Lange T,. Senescence induced by altered telomere state, not telomere loss. Science 2002; 295: 2446-2449. (Pubitemid 34270257)
75. Armanios MY, Chen JJ, Cogan JD, et al,. Telomerase mutations in families with idiopathic pulmonary fibrosis. N Engl J Med 2007; 356: 1317-1326. (Pubitemid 46513297)
76. Alder JK, Cogan JD, Brown AF, et al,. Ancestral mutation in telomerase causes defects in repeat addition processivity and manifests as familial pulmonary fibrosis. PLoS Genet 2011; 7: e1001352.
77. Tsakiri KD, Cronkhite JT, Kuan PJ, et al,. Adult-onset pulmonary fibrosis caused by mutations in telomerase. Proc Natl Acad Sci USA 2007; 104: 7552-7557. (Pubitemid 47185944)
78. Alder JK, Chen JJ, Lancaster L, et al,. Short telomeres are a risk factor for idiopathic pulmonary fibrosis. Proc Natl Acad Sci USA 2008; 105: 13051-13056.
79. Cronkhite JT, Xing C, Raghu G, et al,. Telomere shortening in familial and sporadic pulmonary fibrosis. Am J Respir Crit Care Med 2008; 178: 729-737.
80. Waisberg DR, Barbas-Filho JV, Parra ER, et al,. Abnormal expression of telomerase/apoptosis limits type II alveolar epithelial cell replication in the early remodeling of usual interstitial pneumonia/idiopathic pulmonary fibrosis. Hum Pathol 2009; 41: 385-391.
81. Driscoll B, Buckley S, Bui KC, et al,. Telomerase in alveolar epithelial development and repair. Am J Physiol Lung Cell Mol Physiol 2000; 279: L1191-1198. (Pubitemid 32009505)
82. Diaz de Leon A, Cronkhite JT, Katzenstein AL, et al,. Telomere lengths, pulmonary fibrosis and telomerase (TERT) mutations. PLoS One 2010; 5: e10680.
83. Alison MR, Sarraf CE,. Apoptosis: a gene-directed programme of cell death. J R Coll Physicians Lond 1992; 26: 25-35.
84. Lee HC, Wei YH,. Oxidative stress, mitochondrial DNA mutation, and apoptosis in aging. Exp Biol Med (Maywood) 2007; 232: 592-606.
85. Thannickal VJ, Horowitz JC,. Evolving concepts of apoptosis in idiopathic pulmonary fibrosis. Proc Am Thorac Soc 2006; 3: 350-356. (Pubitemid 43845412)
86. Pollack M, Phaneuf S, Dirks A, et al,. The role of apoptosis in the normal aging brain, skeletal muscle, and heart. Ann NY Acad Sci 2002; 959: 93-107. (Pubitemid 34457555)
87. Selman M, Rojas M, Mora AL, et al,. Aging and interstitial lung diseases: unraveling an old forgotten player in the pathogenesis of lung fibrosis. Semin Respir Crit Care Med 2010; 31: 607-617.
88. Chen W, Wu W,. The asymmetric, rectifier-like I-V curve of the Na/K pump transient currents in frog skeletal muscle fibers. Bioelectrochemistry 2002; 56: 199-202. (Pubitemid 34497004)
89. Breckenridge DG, Stojanovic M, Marcellus RC, et al,. Caspase cleavage product of BAP31 induces mitochondrial fission through endoplasmic reticulum calcium signals, enhancing cytochrome c release to the cytosol. J Cell Biol 2003; 160: 1115-1127. (Pubitemid 36443879)
90. Hagimoto N, Kuwano K, Nomoto Y, et al,. Apoptosis and expression of Fas/Fas ligand mRNA in bleomycin-induced pulmonary fibrosis in mice. Am J Respir Cell Mol Biol 1997; 16: 91-101. (Pubitemid 127497440)
91. Lee CG, Cho SJ, Kang MJ, et al,. Early growth response gene 1-mediated apoptosis is essential for transforming growth factor-β1-induced pulmonary fibrosis. J Exp Med 2004; 200: 377-389. (Pubitemid 39031256)
92. Gilhar A, Ullmann Y, Karry R, et al,. Aging of human epidermis: reversal of aging changes correlates with reversal of keratinocyte fas expression and apoptosis. J Gerontol A Biol Sci Med Sci 2004; 59: 411-415. (Pubitemid 38649930)
93. Barja G,. Free radicals and aging. Trends Neurosci 2004; 27: 595-600. (Pubitemid 39221395)
94. Kuwano K, Hagimoto N, Maeyama T, et al,. Mitochondria-mediated apoptosis of lung epithelial cells in idiopathic interstitial pneumonias. Lab Invest 2002; 82: 1695-1706. (Pubitemid 35462982)
95. Wallach-Dayan SB, Izbicki G, Cohen PY, et al,. Bleomycin initiates apoptosis of lung epithelial cells by ROS but not by Fas/FasL pathway. Am J Physiol Lung Cell Mol Physiol 2006; 290: L790-796. (Pubitemid 43725085)
96. Beere HM,. 'The stress of dying': the role of heat shock proteins in the regulation of apoptosis. J Cell Sci 2004; 117: 2641-2651. (Pubitemid 38997248)
97. Totan S, Echo A, Yuksel E,. Heat shock proteins modulate keloid formation. Eplasty 2011; 11: e21.
98. Tanaka K, Tanaka Y, Namba T, et al,. Heat shock protein 70 protects against bleomycin-induced pulmonary fibrosis in mice. Biochem Pharmacol 2010; 80: 920-931.
99. Chung L, Ng YC,. Age-related alterations in expression of apoptosis regulatory proteins and heat shock proteins in rat skeletal muscle. Biochim Biophys Acta 2006; 1762: 103-109. (Pubitemid 43021092)
100. Phaneuf S, Leeuwenburgh C,. Cytochrome c release from mitochondria in the aging heart: a possible mechanism for apoptosis with age. Am J Physiol Regul Integr Comp Physiol 2002; 282: R423-430. (Pubitemid 34664592)
101. Plataki M, Koutsopoulos AV, Darivianaki K, et al,. Expression of apoptotic and antiapoptotic markers in epithelial cells in idiopathic pulmonary fibrosis. Chest 2005; 127: 266-274. (Pubitemid 41724587)
102. Berger SL, Kouzarides T, Shiekhattar R, et al,. An operational definition of epigenetics. Genes Dev 2009; 23: 781-783.
103. Holliday R,. Mechanisms for the control of gene activity during development. Biol Rev Camb Phil Soc 1990; 65: 431-471.
104. Schwartz DA,. Epigenetics and environmental lung disease. Proc Am Thorac Soc 2010; 7: 123-125.
105. Berdasco M, Esteller M,. Aberrant epigenetic landscape in cancer: how cellular identity goes awry. Dev Cell 2010; 19: 698-711.
106. Oberdoerffer P, Michan S, McVay M, et al,. SIRT1 redistribution on chromatin promotes genomic stability but alters gene expression during aging. Cell 2008; 135: 907-918.
107. Jones PA, Takai D,. The role of DNA methylation in mammalian epigenetics. Science 2001; 293: 1068-1070. (Pubitemid 32758075)
108. Gravina S, Vijg J,. Epigenetic factors in aging and longevity. Pflugers Arch 2010; 459: 247-258.
109. Guilleret I, Yan P, Grange F, et al,. Hypermethylation of the human telomerase catalytic subunit (hTERT) gene correlates with telomerase activity. Int J Cancer 2002; 101: 335-341. (Pubitemid 34983188)
110. Pfeifer GP, Rauch TA,. DNA methylation patterns in lung carcinomas. Semin Cancer Biol 2009; 19: 181-187.
111. Rabinovich EI, Kapetanaki MG, Steinfeld I, et al,. Global methylation patterns in idiopathic pulmonary fibrosis. PLoS One 2012; 7: e33770.
112. Wang H, Cao R, Xia L, et al,. Purification and functional characterization of a histone H3-lysine 4-specific methyltransferase. Mol Cell 2001; 8: 1207-1217. (Pubitemid 34084993)
113. Fuks F, Burgers WA, Godin N, et al,. Dnmt3a binds deacetylases and is recruited by a sequence-specific repressor to silence transcription. EMBO J 2001; 20: 2536-2544. (Pubitemid 32452872)
114. Ooi SK, Qiu C, Bernstein E, et al,. DNMT3L connects unmethylated lysine 4 of histone H3 to de novo methylation of DNA. Nature 2007; 448: 714-717. (Pubitemid 47236857)
115. Cedar H, Bergman Y,. Linking DNA methylation and histone modification: patterns and paradigms. Nat Rev Genet 2009; 10: 295-304.
116. Rodenhiser D, Mann M,. Epigenetics and human disease: translating basic biology into clinical applications. CMAJ 2006; 174: 341-348. (Pubitemid 43235307)
117. Wilson VL, Smith RA, Ma S, et al,. Genomic 5-methyldeoxycytidine decreases with age. J Biol Chem 1987; 262: 9948-9951.
118. Ono T, Dean RG, Chattopadhyay SK, et al,. Dysdifferentiative nature of aging: age-dependent expression of MuLV and globin genes in thymus, liver and brain in the AKR mouse strain. Gerontology 1985; 31: 362-372. (Pubitemid 16247902)
119. Pollina EA, Brunet A,. Epigenetic regulation of aging stem cells. Oncogene 2011; 30: 3105-3126.
120. Berdasco M, Esteller M,. Hot topics in epigenetic mechanisms of aging: 2011. Aging Cell 2012; 11: 181-186.
121. Sanders YY, Pardo A, Selman M, et al,. Thy-1 promoter hypermethylation: a novel epigenetic pathogenic mechanism in pulmonary fibrosis. Am J Respir Cell Mol Biol 2008; 39: 610-618.
122. Sanders YY, Tollefsbol TO, Varisco BM, et al,. Epigenetic regulation of thy-1 by histone deacetylase inhibitor in rat lung fibroblasts. Am J Respir Cell Mol Biol 2010; 45: 16-23.
123. Coward WR, Watts K, Feghali-Bostwick CA, et al,. Defective histone acetylation is responsible for the diminished expression of cyclooxygenase 2 in idiopathic pulmonary fibrosis. Mol Cell Biol 2009; 29: 4325-4339.
124. Coward WR, Watts K, Feghali-Bostwick CA, et al,. Repression of IP-10 by interactions between histone deacetylation and hypermethylation in idiopathic pulmonary fibrosis. Mol Cell Biol 2010; 30: 2874-2886.
125. Hu B, Gharaee-Kermani M, Wu Z, et al,. Essential role of MeCP2 in the regulation of myofibroblast differentiation during pulmonary fibrosis. Am J Pathol 2011; 178: 1500-1508.
126. Cisneros J, Hagood J, Checa M, et al,. Hypermethylation-mediated silencing of p14ARF in fibroblasts from idiopathic pulmonary fibrosis. Am J Physiol Lung Cell Mol Physiol 2012; 303: L295-303.
127. Lee RC, Feinbaum RL, Ambros V., The C. elegans heterochronic gene lin-4 encodes small RNAs with antisense complementarity to lin-14. Cell 1993; 75: 843-854. (Pubitemid 24014542)
128. Kim VN, Han J, Siomi MC,. Biogenesis of small RNAs in animals. Nat Rev Mol Cell Biol 2009; 10: 126-139.
129. Vasudevan S,. Posttranscriptional upregulation by microRNAs. Wiley Interdiscip Rev RNA 2012; 3: 311-330.
130. Filipowicz W, Bhattacharyya SN, Sonenberg N,. Mechanisms of post-transcriptional regulation by microRNAs: are the answers in sight? Nat Rev Genet 2008; 9: 102-114.
131. Bartel DP,. MicroRNAs: target recognition and regulatory functions. Cell 2009; 136: 215-233.
132. Williams AE,. Functional aspects of animal microRNAs. Cell Mol Life Sci 2008; 65: 545-562. (Pubitemid 351297689)
133. Kozomara A, Griffiths-Jones S., miRBase: integrating microRNA annotation and deep-sequencing data. Nucleic Acids Res 2011; 39: D152-157.
134. Williams AE, Moschos SA, Perry MM, et al,. Maternally imprinted microRNAs are differentially expressed during mouse and human lung development. Dev Dyn 2007; 236: 572-580. (Pubitemid 46233301)
135. Izzotti A, Calin GA, Steele VE, et al,. Relationships of microRNA expression in mouse lung with age and exposure to cigarette smoke and light. FASEB J 2009; 23: 3243-3250.
136. Williams AE, Perry MM, Moschos SA, et al,. microRNA expression in the aging mouse lung. BMC Genom 2007; 8: 172.
137. Noren Hooten N, Abdelmohsen K, Gorospe M, et al,. microRNA expression patterns reveal differential expression of target genes with age. PLoS One 2010; 5: e10724.
138. Ji J, Shi J, Budhu A, et al,. MicroRNA expression, survival, and response to interferon in liver cancer. N Engl J Med 2009; 361: 1437-1447.
139. Chen C, Wu CQ, Zhang ZQ, et al,. Loss of expression of miR-335 is implicated in hepatic stellate cell migration and activation. Exp Cell Res 2011; 317: 1714-1725.
140. Milosevic J, Pandit KV, Jacobs R, et al,. The role of miR-154 microRNA family in IPF. Am J Respir Crit Care Med 2009; 179.
141. Aller MA, Blanco-Rivero J, Arias JI, et al,. The wound-healing response and upregulated embryonic mechanisms: brothers-in-arms forever. Exp Dermatol 2012; 21: 497-503.
142. Selman M, Pardo A, Kaminski N,. Idiopathic pulmonary fibrosis: aberrant recapitulation of developmental programs? PLoS Med 2008; 5: e62.
143. Vettori S, Gay S, Distler O,. Role of MicroRNAs in Fibrosis. Open Rheumatol J 2012; 6: 130-139.
144. Kato M, Putta S, Wang M, et al,. TGFβ activates Akt kinase through a microRNA-dependent amplifying circuit targeting PTEN. Nat Cell Biol 2009; 11: 881-889.
145. Liu G, Friggeri A, Yang Y, et al,. miR-21 mediates fibrogenic activation of pulmonary fibroblasts and lung fibrosis. J Exp Med 2010; 207: 1589-1597.
146. Wang B, Koh P, Winbanks C, et al,. miR-200a Prevents renal fibrogenesis through repression of TGFβ2 expression. Diabetes 2011; 60: 280-287.
147. Duisters RF, Tijsen AJ, Schroen B, et al,. miR-133 and miR-30 regulate connective tissue growth factor: implications for a role of microRNAs in myocardial matrix remodeling. Circ Res 2009; 104: 170-178, 176pp following 178.
148. van Almen GC, Verhesen W, van Leeuwen RE, et al,. MicroRNA-18 and microRNA-19 regulate CTGF and TSP-1 expression in age-related heart failure. Aging Cell 2011; 10: 769-779.
149. Maurer B, Stanczyk J, Jungel A, et al,. MicroRNA-29, a key regulator of collagen expression in systemic sclerosis. Arthritis Rheum 2010; 62: 1733-1743.
150. Muth M, Hussein K, Jacobi C, et al,. Hypoxia-induced down-regulation of microRNA-449a/b impairs control over targeted SERPINE1 (PAI-1) mRNA-a mechanism involved in SERPINE1 (PAI-1) overexpression. J Transl Med 2011; 9: 24.
151. Wang Q, Wang Y, Minto AW, et al,. MicroRNA-377 is up-regulated and can lead to increased fibronectin production in diabetic nephropathy. FASEB J 2008; 22: 4126-4135.
152. Pandit KV, Corcoran D, Yousef H, et al,. Inhibition and role of let-7d in idiopathic pulmonary fibrosis. Am J Respir Crit Care Med 2010; 182: 220-229.
153. Wang B, Herman-Edelstein M, Koh P, et al,. E-cadherin expression is regulated by miR-192/215 by a mechanism that is independent of the profibrotic effects of transforming growth factor-β. Diabetes 2010; 59: 1794-1802.
154. Mann J, Chu DC, Maxwell A, et al,. MeCP2 controls an epigenetic pathway that promotes myofibroblast transdifferentiation and fibrosis. Gastroenterology 2010; 138: 705-714, 714 e701-704.
155. Bhaskaran M, Xi D, Wang Y, et al,. Identification of microRNAs changed in the neonatal lungs in response to hyperoxia exposure. Physiol Genom 2012; 44: 970-980.
156. Pandit KV, Milosevic J, Kaminski N,. MicroRNAs in idiopathic pulmonary fibrosis. Transl Res 2011; 157: 191-199.