Roles of Proteoglycans and Glycosaminoglycans in Wound Healing and Fibrosis

International Journal of Cell Biology

Volumn 2015, Issue , 2015, Pages

  Ghatak, Shibnath ^a   Maytin, Edward V ^b   MacK, Judith A ^b   Hascall, Vincent C ^b   Atanelishvili, Ilia ^a   Moreno Rodriguez, Ricardo ^a   Markwald, Roger R ^a   Misra, Suniti ^a  

^a MEDICAL UNIVERSITY OF SOUTH CAROLINA   (United States)

^b LERNER RESEARCH INSTITUTE   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

CARBOHYDRATES AND CARBOHYDRATE DERIVATIVES; CYTOKINE; FRESOLIMUMAB; GLYCOSAMINOGLYCAN; GROWTH FACTOR; HERMES ANTIGEN; HYALURONIC ACID; PROTEOGLYCAN; SYNTHETASE;

BIOCHEMICAL ANALYSIS; CELL ACTIVATION; CELL INTERACTION; CELL MIGRATION; CELL PROLIFERATION; DEGRADATION; EXTRACELLULAR MATRIX; FIBROSIS; HOMEOSTASIS; HUMAN; INFLAMMATION; MESENCHYME CELL; NONHUMAN; PHYSIOLOGICAL PROCESS; PRIORITY JOURNAL; PROTEIN INTERACTION; REVIEW; TISSUE INJURY; TISSUE REPAIR; WOUND HEALING;

EID: 84942288147     PISSN: 16878876     EISSN: 16878884     Source Type: Journal    
DOI: 10.1155/2015/834893     Document Type: Review

References (260)

1. N. B. Menke, K. R. Ward, T. M. Witten, D. G. Bonchev, and R. F. Diegelmann, "Impaired wound healing," Clinics in Dermatology, vol. 25, no. 1, pp. 19-25, 2007.
2. M. J. Bissell, H. G. Hall, and G. Parry, "How does the extracellular matrix direct gene expression?" Journal of Theoretical Biology, vol. 99, no. 1, pp. 31-68, 1982.
3. R. O. Hynes, "Integrins: bidirectional, allosteric signaling machines," Cell, vol. 110, no. 6, pp. 673-687, 2002.
4. H. Ponta, L. Sherman, and P. A. Herrlich, "CD44: fromadhesion molecules to signalling regulators," Nature Reviews Molecular Cell Biology, vol. 4, no. 1, pp. 33-45, 2003.
5. R. V. Sionov and D. Naor, "Hyaluronan-independent lodgment of CD44+ lymphoma cells in lymphoid organs," International Journal of Cancer, vol. 71, no. 3, pp. 462-469, 1997.
6. S. Misra, V. C. Hascall, N. K. Karamanos, R. R. Markwald, and S. Ghatak, Delivery Systems Targeting Cancer at the Level of ECM, DeGruyter, Berlin, Germany, 2012.
7. S. Misra, P. Heldin, V. C. Hascall et al., "Hyaluronan-CD44 interactions as potential targets for cancer therapy," The FEBS Journal, vol. 278, no. 9, pp. 1429-1443, 2011.
8. M. Shimaoka and T. A. Springer, "Therapeutic antagonists and conformational regulation of integrin function," Nature Reviews: Drug Discovery, vol. 2, no. 9, pp. 703-716, 2003.
9. J. P. Iredale, "Tissue inhibitors of metalloproteinases in liver fibrosis," International Journal of Biochemistry and Cell Biology, vol. 29, no. 1, pp. 43-54, 1997.
10. A. M. El Nahas, E. C. Muchaneta-Kubara, M. Essawy, and O. Soylemezoglu, "Renal fibrosis: insights into pathogenesis and treatment," International Journal of Biochemistry and Cell Biology, vol. 29, no. 1, pp. 55-62, 1997.
11. R. P. Marshall, R. J. McAnulty, and G. J. Laurent, "The pathogenesis of pulmonary fibrosis: Is there a fibrosis gene?" International Journal of Biochemistry and Cell Biology, vol. 29, no. 1, pp. 107-120, 1997.
12. K. T. Weber, Y. Sun, and L. C. Katwa, "Myofibroblasts and local angiotensin II in rat cardiac tissue repair," International Journal of Biochemistry and Cell Biology, vol. 29, no. 1, pp. 31-42, 1997.
13. K. T. Weber, Y. Sun, L. C. Katwa, and J. P. M. Cleutjens, "Tissue repair and angiotensin II generated at sites of healing," Basic Research in Cardiology, vol. 92, no. 2, pp. 75-78, 1997.
14. S. Ghatak, G. S. Bogatkevich, I. Atnelishvilis et al., "Overexpression of c-Met and CD44v6 receptors contributes to autocrine TGF-β1 signaling in interstitial lung disease," The Journal of Biological Chemistry, vol. 289, no. 11, pp. 7856-7872, 2014.
15. M. B. Kahaleh, "The role of vascular endothelium in fibroblast activation and tissue fibrosis, particularly in scleroderma (systemic sclerosis) and pachydermoperiostosis (primary hypertrophic osteoarthropathy)," Clinical and Experimental Rheumatology, vol. 10, supplement 7, pp. 51-56, 1992.
16. R. A. F. Clark, J. M. Lanigan, P. DellaPelle, E. Manseau, H. F. Dvorak, and R. B. Colvin, "Fibronectin and fibrin provide a provisional matrix for epidermal cell migration during wound reepithelialization," Journal of Investigative Dermatology, vol. 79, no. 5, pp. 264-269, 1982.
17. K. M. Yamada and R. A. F. Clark, Provisional Matrix, Plenum Press, New York, NY, USA, 1996.
18. R. A. F. Clark, Wound Repair: Overview and General Considerations, Plenum Press, 1996.
19. G. Broughton II, J. E. Janis, and C. E. Attinger, "The basic science of wound healing," Plastic and Reconstructive Surgery, vol. 117, no. 7, pp. 12S-34S, 2006.
20. A. C. L. Campos, A. K. Groth, and A. B. Branco, "Assessment and nutritional aspects of wound healing," Current Opinion in Clinical Nutrition and Metabolic Care, vol. 11, no. 3, pp. 281-288, 2008.
21. A. Gosain and L. A. DiPietro, "Aging and wound healing," World Journal of Surgery, vol. 28, no. 3, pp. 321-326, 2004.
22. A. J. Meszaros, J. S. Reichner, and J. E. Albina, "Macrophageinduced neutrophil apoptosis," Journal of Immunology, vol. 165, no. 1, pp. 435-441, 2000.
23. D. M. Mosser and J. P. Edwards, "Exploring the full spectrum of macrophage activation," Nature Reviews. Immunology, vol. 8, no. 12, pp. 958-969, 2008.
24. J. E. Park and A. Barbul, "Understand ing the role of immune regulation in wound healing," American Journal of Surgery, vol. 187, no. 5, 2004.
25. M. E. Swift, A. L. Burns, K. L. Gray, and L. A. DiPietro, "Agerelated alterations in the inflammatory response to dermal injury," Journal of Investigative Dermatology, vol. 117, no. 5, pp. 1027-1035, 2001.
26. R. A. F. Clark, Overview and General Consideration of Wound Repair, Plenum, New York, NY, USA, 1988.
27. T. Iwayama and L. E. Olson, "Involvement of PDGF in fibrosis and scleroderma: recent insights from animal models and potential therapeutic opportunities," Current Rheumatology Reports, vol. 15, no. 2, 2013.
28. A. Leask, "Possible strategies for anti-fibrotic drug intervention in scleroderma," Journal of Cell Communication and Signaling, vol. 5, no. 2, pp. 125-129, 2011.
29. M. Trojanowska, "Role of PDGF in fibrotic diseases and systemic sclerosis," Rheumatology, vol. 47, pp. v2-4, 2008.
30. M. Eastwood, R. Porter, U. Khan, G. McGrouther, and R. Brown, "Quantitative analysis of collagen gel contractile forces generated by dermal fibroblasts and the relationship to cell morphology," Journal of Cellular Physiology, vol. 166, no. 1, pp. 33-42, 1996.
31. R. J. McAnulty, J. S. Campa, A. D. Cambrey, and G. J. Laurent, "The effect of transforming growth factor beta on rates of procollagen synthesis and degradation in vitro," Biochimica et Biophysica Acta-Molecular Cell Research, vol. 1091, no. 2, pp. 231-235, 1991.
32. A. B. Roberts, K. C. Flanders, U. I. Heine et al., "Transforming growth factor-beta: multifunctional regulator of differentiation and development," Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, vol. 327, no. 1239, pp. 145-154, 1990.
33. A. B. Roberts, U. I. Heine, K. C. Flanders, and M. B. Sporn, "Transforming growth factor-β. Major role in regulation of extracellular matrix," Annals of the New York Academy of Sciences, vol. 580, pp. 225-232, 1990.
34. M. Shah, D. M. Foreman, and M. W. J. Ferguson, "Neutralisation of TGF-beta 1 and TGF-beta 2 or exogenous addition of TGFbeta 3 to cutaneous rat wounds reduces scarring," Journal of Cell Science, vol. 108, no. 3, pp. 985-1002, 1995.
35. T. J. Franklin, "Therapeutic approaches to organ fibrosis," International Journal of Biochemistry and Cell Biology, vol. 29, no. 1, pp. 79-89, 1997.
36. J. Folkman and M. Klagsbrun, "Angiogenic factors," Science, vol. 235, no. 4787, pp. 442-447, 1987.
37. P. R. Atkison, E. R. Weidman, B. Bhaumick, and R. M. Bala, "Release of somatomedin-like activity by cultured WI-38 human fibroblasts," Endocrinology, vol. 106, no. 6, pp. 2006-2012, 1980.
38. W. N. Rom, P. Basset, G. A. Fells, T. Nukiwa, B. C. Trapnell, and R. G. Crystal, "Alveolarmacrophages release an insulin-like growth factor I-typemolecule," Journal of Clinical Investigation, vol. 82, no. 5, pp. 1685-1693, 1988.
39. R. H. Goldstein, C. F. Poliks, P. F. Pilch, B. D. Smith, and A. Fine, "Stimulation of collagen formation by insulin and insulinlike growth factor I in cultures of human lung fibroblasts," Endocrinology, vol. 124, no. 2, pp. 964-970, 1989.
40. N. K. Harrison, A. D. Cambrey, A. R. Myers et al., "Insulin-like growth factor-I is partially responsible for fibroblast proliferation induced by bronchoalveolar lavage fluid from patients with systemic sclerosis," Clinical Science, vol. 86, no. 2, pp. 141-148, 1994.
41. M. Laato, V. M. Kähäri, J. Niinikoski, and E. Vuorio, "Epidermal growth factor increases collagen production in granulation tissue by stimulation of fibroblast proliferation and not by activation of procollagen genes," Biochemical Journal, vol. 247, no. 2, pp. 385-388, 1987.
42. J.-P. Pienimäki, K. Rilla, C. Fülöp et al., "Epidermal growth factor activates hyaluronan synthase 2 in epidermal keratinocytes and increases pericellular and intracellular hyaluronan," The Journal of Biological Chemistry, vol. 276, no. 23, pp. 20428-20435, 2001.
43. J. G. Rheinwald and H. Green, "Epidermal growth factor and the multiplication of cultured human epidermal keratinocytes," Nature, vol. 265, no. 5593, pp. 421-424, 1977.
44. A. B. Schreiber, M. E. Winkler, and R. Derynck, "Transforming growth factor-α: a more potent angiogenic mediator than epidermal growth factor," Science, vol. 232, no. 4755, pp. 1250-1253, 1986.
45. G. L. Brown, L. B. Nanney, J. Griffen et al., "Enhancement of wound healing by topical treatment with epidermal growth factor," The New England Journal of Medicine, vol. 321, no. 2, pp. 76-79, 1989.
46. S. Cattaruzza and R. Perris, "Proteoglycan control of cell movement during wound healing and cancer spreading," Matrix Biology, vol. 24, no. 6, pp. 400-417, 2005.
47. N. Hattori, D. A. Carrino, M. E. Lauer et al., "Pericellular versican regulates the fibroblast-myofibroblast transition: a role for ADAMTS5 protease-mediated proteolysis," The Journal of Biological Chemistry, vol. 286, no. 39, pp. 34298-34310, 2011.
48. Z. Zhou, J. Wang, R. Cao et al., "Impaired angiogenesis, delayed wound healing and retarded tumor growth in Perlecan heparan sulfate-deficient mice," Cancer Research, vol. 64, no. 14, pp. 4699-4702, 2004.
49. K. Elenius, S. Vainio, M. Laato, M. Salmivirta, I. Thesleff, and M. Jalkanen, "Induced expression of syndecan in healing wounds," Journal of Cell Biology, vol. 114, no. 3, pp. 585-595, 1991.
50. M. A. Stepp, H. E. Gibson, P. H. Gala et al., "Defects in keratinocyte activation during wound healing in the syndecan-1-deficient mouse," Journal of Cell Science, vol. 115, no. 23, pp. 4517-4531, 2002.
51. F. Echtermeyer, M. Streit, S. Wilcox-Adelman et al., "Delayed wound repair and impaired angiogenesis in mice lacking syndecan-4," The Journal of Clinical Investigation, vol. 107, no. 2, pp. R9-R14, 2001.
52. Y. Yamaguchi, D. M. Mann, and E. Ruoslahti, "Negative regulation of transforming growth factor-β by the proteoglycan decorin," Nature, vol. 346, no. 6281, pp. 281-284, 1990.
53. Y. Isaka, D. K. Brees, K. Ikegaya et al., "Gene therapy by skeletal muscle expression of decorin prevents fibrotic disease in rat kidney," Nature Medicine, vol. 2, no. 4, pp. 418-423, 1996.
54. K. Fukushima, N. Badlani, A. Usas, F. Riano, F. H. Fu, and J. Huard, "The use of an antifibrosis agent to improve muscle recovery after laceration," The American Journal of Sports Medicine, vol. 29, no. 4, pp. 394-402, 2001.
55. M. Kolb, P. J. Margetts, P. J. Sime, and J. Gauldie, "Proteoglycans decorin and biglycan differentially modulate TGF-β-mediated fibrotic responses in the lung," The American Journal of Physiology-Lung Cellular and Molecular Physiology, vol. 280, no. 6, pp. L1327-L1334, 2001.
56. J. R. Couchman and C. A. Pataki, "An introduction to proteoglycans and their localization," Journal of Histochemistry and Cytochemistry, vol. 60, no. 12, pp. 885-897, 2012.
57. K. Prydz and K. T. Dalen, "Synthesis and sorting of proteoglycans," Journal of Cell Science, vol. 113, no. 2, pp. 193-205, 2000.
58. P. Martin, "Wound healing-aiming for perfect skin regeneration," Science, vol. 276, no. 5309, pp. 75-81, 1997.
59. J. Laterra, R. Ansbacher, and L. A. Culp, "Glycosaminoglycans that bind cold-insoluble globulin in cell-substratum adhesion sites ofmurine fibroblasts," Proceedings of the National Academy of Sciences of the United States of America, vol. 77, no. 11, pp. 6662-6666, 1980.
60. T. N. Wight, D. K. Heinegard, and V. C. Hascall, Proteoglycans: Structure and Function, Plenum Press, New York, NY, USA, 1991.
61. M. W. Lark and L. A. Culp, Cell-Matrix Interaction: Biochemistry of Close and Tight-Focal Adhesive Contacts of Fibroblasts to Extracellular Matrices, Mercel Decker, New York, NY, USA, 1987.
62. P. V. Peplow, "Glycosaminoglycan: a cand idate to stimulate the repair of chronic wounds," Thrombosis and Haemostasis, vol. 94, no. 1, pp. 4-16, 2005.
63. N. Ortéga, F. E. LFaqihi, and J. Plouët, "Control of vascular endothelial growth factor angiogenic activity by the extracellular matrix," Biology of the Cell, vol. 90, no. 5, pp. 381-390, 1998.
64. K. R. Taylor and R. L. Gallo, "Glycosaminoglycans and their proteoglycans: host-associatedmolecular patterns for initiation and modulation of inflammation," The FASEB Journal, vol. 20, no. 1, pp. 9-22, 2006.
65. C. A. de La, V. C. Motte, and J. Drazba, "Poly I:C induces mononuclear leukocyte-adhesive hyaluronan structures on colon smooth muscel cells: IalphaI and versican facilitiate adhesion," in Hyaluronan 2000, Woodhead Publishing, Wrexham, Wales, 2000.
66. V. C.Hascall, A. K.Majors, C. A. de laMotte et al., Intracellular hyaluronan: a new frontier for inflammation? Biochimica et Biophysica ActaGeneral Subjects, vol. 1673, no. 1-2, pp. 312, 2004.
67. W. Y. J. Chen and G. Abatangelo, "Functions of hyaluronan in wound repair," Wound Repair and Regeneration, vol. 7, no. 2, pp. 79-89, 1999.
68. N. S. Gandhi and R. L. Mancera, "The structure of glycosaminoglycans and their interactions with proteins," Chemical Biology and Drug Design, vol. 72, no. 6, pp. 455-482, 2008.
69. K. Meyer and J. W. Palmer, "The polysaccharide of the vitreous humor," The Journal of Biological Chemistry, vol. 107, no. 3, pp. 629-634, 1934.
70. P. Olczyk, K. Komosińska-Vassev, K. Winsz-Szczotka, K. Kuźnik-Trocha, and K. Olczyk, "Hyaluronan: structure, metabolism, functions, and role in wound healing," Postepy Higieny i Medycyny Dos̈wiadczalnej, vol. 62, pp. 651-659, 2008.
71. T. C. Laurent and J. R. E. Fraser, "Hyaluronan," The FASEB Journal, vol. 6, no. 7, pp. 2397-2404, 1992.
72. G. M. Campo, A. Avenoso, S. Campo, A. DAscola, G. Nastasi, and A. Calatroni, "Molecular size hyaluronan differentlymodulates toll-like receptor-4 in LPS-induced inflammation inmouse chondrocytes," Biochimie, vol. 92, no. 2, pp. 204-215, 2010.
73. M. Mohamadzadeh, H. DeGrendele, H. Arizpe, P. Estess, and M. Siegelman, "Proinflammatory stimuli regulate endothelial hyaluronan expression and CD44/HA-dependent primary adhesion," Journal of Clinical Investigation, vol. 101, no. 1, pp. 97-108, 1998.
74. P. L. Bollyky, S. P. Evanko, R. P. Wu et al., "Th1 cytokines promote T-cell binding to antigen-presenting cells via enhanced hyaluronan production and accumulation at the immune synapse," Cellular and Molecular Immunology, vol. 7, no. 3, pp. 211-220, 2010.
75. T. S. Wilkinson, S. Potter-Perigo, C. Tsoi, L. C. Altman, and T. N. Wight, "Pro-and anti-inflammatory factors cooperate to control hyaluronan synthesis in lung fibroblasts," American Journal of Respiratory Cell and Molecular Biology, vol. 31, no. 1, pp. 92-99, 2004.
76. M. A. Dentener, J. H. J. Vernooy, S. Hendriks, and E. F. M. Wouters, "Enhanced levels of hyaluronan in lungs of patients with COPD: relationship with lung function and local inflammation," Thorax, vol. 60, no. 2, pp. 114-119, 2005.
77. O. Nettelbladt and R. Hallgren, "Hyaluronan (hyaluronic acid) in bronchoalveolar lavage fluid during the development of bleomycin-induced alveolitis in the rat," American Review of Respiratory Disease, vol. 140, no. 4, pp. 1028-1032, 1989.
78. J. P. Freitas, P. Filipe, I. Emerit, P. Meunier, C. F. Manso, and F. Guerra Rodrigo, "Hyaluronic acid in progressive systemic sclerosis," Dermatology, vol. 192, no. 1, pp. 46-49, 1996.
79. H. Levesque, N. Baudot, B. Delpech, M. Vayssairat, A. Gancel, and H. Courtois, "Clinical correlations and prognosis based on hyaluronic acid serum levels in patients with progressive systemic sclerosis," British Journal of Dermatology, vol. 124, no. 5, pp. 423-428, 1991.
80. E. S. Bensadoun, A. K. Burke, J. C. Hogg, and C. R. Roberts, "Proteoglycan deposition in pulmonary fibrosis," American Journal of Respiratory and Critical Care Medicine, vol. 154, no. 6, part 1, pp. 1819-1828, 1996.
81. A. M. Cantin, P. Larivee, M. Martel, and R. Begin, "Hyaluronan (hyaluronic acid) in lung lavage of asbestos-exposed humans and sheep," Lung, vol. 170, no. 4, pp. 211-220, 1992.
82. E. Papakonstantinou, F. M. Kouri, G. Karakiulakis, I. Klagas, and O. Eickelberg, "Increased hyaluronic acid content in idiopathic pulmonary arterial hypertension," European Respiratory Journal, vol. 32, no. 6, pp. 1504-1512, 2008.
83. Y. Li, D. Jiang, J. Liang et al., "Severe lung fibrosis requires an invasive fibroblast phenotype regulated by hyaluronan and CD44," Journal of Experimental Medicine, vol. 208, no. 7, pp. 1459-1471, 2011.
84. E. Puré and C. A. Cuff, "A crucial role for CD44 in inflammation," Trends in Molecular Medicine, vol. 7, no. 5, pp. 213-221, 2001.
85. D. Jiang, J. Liang, J. Fan et al., "Regulation of lung injury and repair by Toll-like receptors and hyaluronan," Nature Medicine, vol. 11, no. 11, pp. 1173-1179, 2005.
86. V. C. Hascall, A.Wang, M. Tammi et al., The dynamic metabolism of hyaluronan regulates the cytosolic concentration of UDP-GlcNAc, Matrix Biology, vol. 35, pp. 1417, 2014.
87. J. Y. Lee and A. P. Spicer, "Hyaluronan: a multifunctional, megaDalton, stealthmolecule," Current Opinion in Cell Biology, vol. 12, no. 5, pp. 581-586, 2000.
88. R. H. Tammi, A. G. Passi, K. Rilla et al., "Transcriptional and post-translational regulation of hyaluronan synthesis," FEBS Journal, vol. 278, no. 9, pp. 1419-1428, 2011.
89. N. Itano and K. Kimata, "Mammalian hyaluronan synthases," IUBMB Life, vol. 54, no. 4, pp. 195-199, 2002.
90. N. Itano, T. Sawai, M. Yoshida et al., "Three isoforms of mammalian hyaluronan synthases have distinct enzymatic properties," The Journal of Biological Chemistry, vol. 274, no. 35, pp. 25085-25092, 1999.
91. P. H. Weigel, V. C. Hascall, and M. Tammi, "Hyaluronan synthases," The Journal of Biological Chemistry, vol. 272, no. 22, pp. 13997-14000, 1997.
92. J. Monslow, J. D. Williams, C. A. Guy et al., "Identification and analysis of the promoter region of the human hyaluronan synthase 2 gene," The Journal of Biological Chemistry, vol. 279, no. 20, pp. 20576-20581, 2004.
93. K. Saavalainen, S. Pasonen-Seppänen, T. W. Dunlop, R. Tammi, M. I. Tammi, and C. Carlberg, "The human hyaluronan synthase 2 gene is a primary retinoic acid and epidermal growth factor responding gene," The Journal of Biological Chemistry, vol. 280, no. 15, pp. 14636-14644, 2005.
94. S. Karvinen, S. Pasonen-Seppänen, J. M. T. Hyttinen et al., "Keratinocyte growth factor stimulates migration and hyaluronan synthesis in the epidermis by activation of keratinocyte hyaluronan synthases 2 and 3. ," The Journal of Biological Chemistry, vol. 278, no. 49, pp. 49495-49504, 2003.
95. S. Pasonen-Seppänen, S. Karvinen, K. Törrönen et al., "EGF upregulates, whereas TGF-β downregulates, the hyaluronan synthases Has2 and Has3 in organotypic keratinocyte cultures: correlations with epidermal proliferation and differentiation," Journal of Investigative Dermatology, vol. 120, no. 6, pp. 1038-1044, 2003.
96. J. Monslow, N. Sato, J. A. MacK, and E. V. Maytin, "Woundinginduced synthesis of hyaluronic acid in organotypic epidermal cultures requires the release of heparin-binding EGF and activation of the EGFR," Journal of Investigative Dermatology, vol. 129, no. 8, pp. 2046-2058, 2009.
97. S. M. Pasonen-Seppänen, E. V. Maytin, K. J. Törrönen et al., "All-trans retinoic acid-induced hyaluronan production and hyperplasia are partly mediated by EGFR signaling in epidermal keratinocytes," Journal of Investigative Dermatology, vol. 128, no. 4, pp. 797-807, 2008.
98. M. S. Pandey, E. N. Harris, J. A. Weigel, and P. H. Weigel, "The cytoplasmic domain of the hyaluronan receptor for endocytosis (hare) contains multiple endocytic motifs targeting coated pitmediated internalization," The Journal of Biological Chemistry, vol. 283, no. 31, pp. 21453-21461, 2008.
99. R. Tammi, K. Rilla, J.-P. Pienimäki et al., "Hyaluronan enters keratinocytes by a novel endocytic route for catabolism," The Journal of Biological Chemistry, vol. 276, no. 37, pp. 35111-35122, 2001.
100. B. Andre, C. Duterme, K. VanMoer, J. Mertens-Strijthagen, M. Jadot, and B. Flamion, "Hyal2 is a glycosylphosphatidylinositolanchored, lipid raft-associated hyaluronidase," Biochemical and Biophysical Research Communications, vol. 411, no. 1, pp. 175-179, 2011.
101. D. Vigetti, S. Deleonibus, P. Moretto et al., "Role of UDP-Nacetylglucosamine (GlcNAc) and O-GlcNacylation of hyaluronan synthase 2 in the control of chondroitin sulfate and hyaluronan synthesis," The Journal of Biological Chemistry, vol. 287, no. 42, pp. 35544-35555, 2012.
102. S. Ghatak, S. Misra, R. A. Norris et al., "Periostin induces intracellular cross-talk between kinases and hyaluronan in atrioventricular valvulogenesis," The Journal of Biological Chemistry, vol. 289, no. 12, pp. 8545-8561, 2014.
103. D. Vigetti, M. Viola, E. Karousou, G. de Luca, and A. Passi, "Metabolic control of hyaluronan synthases," Matrix Biology, vol. 35, pp. 8-13, 2014.
104. T. D. Camenisch, J. A. Schroeder, J. Bradley, S. E. Klewer, and J. A. McDonald, "Heart-valvemesenchyme formation is dependent on hyaluronan-augmented activation of ErbB2-ErbB3 receptors," Nature Medicine, vol. 8, no. 8, pp. 850-855, 2002.
105. T. D. Camenisch, A. P. Spicer, T. Brehm-Gibson et al., "Disruption of hyaluronan synthase-2 abrogates normal cardiac morphogenesis and hyaluronan-mediated transformation of epithelium to mesenchyme," Journal of Clinical Investigation, vol. 106, no. 3, pp. 349-360, 2000.
106. E. A. Craig, A. F. Austin, R. R. Vaillancourt, J. V. Barnett, and T. D. Camenisch, "TGFβ2-mediated production of hyaluronan is important for the induction of epicardial cell differentiation and invasion," Experimental Cell Research, vol. 316, no. 20, pp. 3397-3405, 2010.
107. J. McDonald, V. C. Hascall, K. Meyer, and J. W. Palmer, "Hyaluronan minireview series," The Journal of Biological Chemistry, vol. 277, no. 7, pp. 4575-4579, 2002.
108. J. A. Mack, R. J. Feldman, N. Itano et al., "Enhanced inflammation and accelerated wound closure following tetraphorbol ester application or full-thickness wounding in mice lacking hyaluronan synthases Has1 and Has3," Journal of Investigative Dermatology, vol. 132, no. 1, pp. 198-207, 2012.
109. P. Gribbon, B. C. Heng, and T. E. Hardingham, "Themolecular basis of the solution properties of hyaluronan investigated by confocal fluorescence recovery after photobleaching," Biophysical Journal, vol. 77, no. 4, pp. 2210-2216, 1999.
110. I. Ellis, J. Banyard, and S. L. Schor, "Differential response of fetal and adult fibroblasts to cytokines: cell migration and hyaluronan synthesis," Development, vol. 124, no. 8, pp. 1593-1600, 1997.
111. I. R. Ellis and S. L. Schor, "Differential effects of TGF-β1 on hyaluronan synthesis by fetal and adult skin fibroblasts: Implications for cell migration and wound healing," Experimental Cell Research, vol. 228, no. 2, pp. 326-333, 1996.
112. J. A. Brown, "The role of hyaluronic acid in wound healings proliferative phase," Journal of Wound Care, vol. 13, no. 2, pp. 48-51, 2004.
113. S. Ghatak, V. C. Hascall, N. K. Karamanos, R. R. Markwald, and S. Misra, Targeting the Tumor Microenvironment in Cancer Progression, DeGruyter, Berlin, Germany, 2012.
114. S. Ghatak, V. C. Hascall, R. R. Markwald, and S. Misra, "Stromal hyaluronan interaction with epithelial CD44 variants promotes prostate cancer invasiveness by augmenting expression and function of hepatocyte growth factor and and rogen receptor," The Journal of Biological Chemistry, vol. 285, no. 26, pp. 19821-19832, 2010.
115. K. Rilla, M. J. Lammi, R. Sironen et al., "Changed lamellipodial extension, adhesion plaques, and migration in epidermal keratinocytes containing constitutively expressed sense and antisense hyaluronan synthase 2 (Has2) genes," Journal of Cell Science, vol. 115, no. 18, pp. 3633-3643, 2002.
116. B. P. Toole, "Hyaluronan in morphogenesis," Seminars in Cell & Developmental Biology, vol. 12, no. 2, pp. 79-87, 2001.
117. S. Misra, S. Ghatak, N. Patil et al., "Novel dual cyclooxygenase and lipoxygenase inhibitors targeting hyaluronan-CD44v6 pathway and inducing cytotoxicity in colon cancer cells," Bioorganic and Medicinal Chemistry, vol. 21, no. 9, pp. 2551-2559, 2013.
118. S. Misra, V. Hascall, N. Karamanos, R. A. Markwald, and S. Ghatak, Targeting Tumor Microenvironment in Cancer Progression, De Gruyter, 2012.
119. S. Misra, V. C. Hascall, F. G. Berger, R. R. Markwald, and S. Ghatak, "Hyaluronan, CD44, and cyclooxygenase-2 in colon cancer," Connective Tissue Research, vol. 49, no. 3-4, pp. 219-224, 2008.
120. S. Misra, L. M. Obeid, Y. A. Hannun et al., "Hyaluronan constitutively regulates activation of COX-2-mediated cell survival activity in intestinal epithelial and colon carcinoma cells," The Journal of Biological Chemistry, vol. 283, no. 21, pp. 14335-14344, 2008.
121. C. A. Maxwell, J. J. Keats, M. Crainie et al., "RHAMM is a centrosomal protein that interacts with dynein and maintains spindle pole stability," Molecular Biology of the Cell, vol. 14, no. 6, pp. 2262-2276, 2003.
122. C. A. Maxwell, J. McCarthy, and E. Turley, "Cell-surface and mitotic-spindle RHAMM: moonlighting or dual oncogenic functions?" Journal of Cell Science, vol. 121, part 7, pp. 925-932, 2008.
123. E. A. Turley, "Hyaluronan and cell locomotion," Cancer and Metastasis Reviews, vol. 11, no. 1, pp. 21-30, 1992.
124. E. A. Turley, P. W. Noble, and L. Y. Bourguignon, "Signaling properties of hyaluronan receptors," The Journal of Biological Chemistry, vol. 277, no. 7, pp. 4589-4592, 2002.
125. S. Meran, D. W. Thomas, P. Stephens et al., "Hyaluronan facilitates transforming growth factor-beta1-mediated fibroblast proliferation," The Journal of Biological Chemistry, vol. 283, no. 10, pp. 6530-6545, 2008.
126. S. Meran, D. Thomas, P. Stephens et al., "Involvement of hyaluronan in regulation of fibroblast phenotype," The Journal of Biological Chemistry, vol. 282, no. 35, pp. 25687-25697, 2007.
127. H. Harada and M. Takahashi, "CD44-dependent intracellular and extracellular catabolism of hyaluronic acid by hyaluronidase-1 and -2," The Journal of Biological Chemistry, vol. 282, no. 8, pp. 5597-5607, 2007.
128. S. Meran, D. D. Luo, R. Simpson et al., "Hyaluronan facilitates transforming growth factor-beta1-dependent proliferation via CD44 and epidermal growth factor receptor interaction," The Journal of Biological Chemistry, vol. 286, no. 20, pp. 17618-17630, 2011.
129. A. C. Midgley, M. Rogers, M. B. Hallett et al., "Transforming growth factor-β1 (TGF-β1)-stimulated fibroblast to myofibroblast differentiation is mediated by hyaluronan (HA)-facilitated epidermal growth factor receptor (EGFR) and CD44 colocalization in lipid rafts," The Journal of Biological Chemistry, vol. 288, no. 21, pp. 14824-14838, 2013.
130. N. Aden, A. Nuttall, X. Shiwen et al., "Epithelial cells promote fibroblast activation via IL-1α in systemic sclerosis," Journal of Investigative Dermatology, vol. 130, no. 9, pp. 2191-2200, 2010.
131. A. Ghaffari, R. T. Kilani, and A. Ghahary, "Keratinocyte-conditioned media regulate collagen expression in dermal fibroblasts," Journal of Investigative Dermatology, vol. 129, no. 2, pp. 340-347, 2009.
132. P. Shephard, G. Martin, S. Smola-Hess, G. Brunner, T. Krieg, and H. Smola, "Myofibroblast differentiation is induced in keratinocyte-fibroblast co-cultures and is antagonistically regulated by endogenous transforming growth factor-β, and interleukin-1," American Journal of Pathology, vol. 164, no. 6, pp. 2055-2066, 2004.
133. G. Cheng, S. Swaidani, M. Sharma, M. E. Lauer, V. C. Hascall, and M. A. Aronica, "Hyaluronan deposition and correlation with inflammation in a murine ovalbumin model of asthma," Matrix Biology, vol. 30, no. 2, pp. 126-134, 2011.
134. A. B. Csóka, S. W. Scherer, and R. Stern, "Expression analysis of six paralogous human hyaluronidase genes clustered on chromosomes 3p21 and 7q31," Genomics, vol. 60, no. 3, pp. 356-361, 1999.
135. V. Atmuri, D. C. Martin, R. Hemming et al., "Hyaluronidase 3 (HYAL3) knockout mice do not display evidence of hyaluronan accumulation," Matrix Biology, vol. 27, no. 8, pp. 653-660, 2008.
136. H. Yoshida, A. Nagaoka, A. Kusaka-Kikushima et al., "KIAA1199, a deafness gene of unknown function, is a new hyaluronan binding protein involved in hyaluronan depolymerization," Proceedings of the National Academy of Sciences of the United States of America, vol. 110, no. 14, pp. 5612-5617, 2013.
137. T. B. Csóka, G. I. Frost, T. Wong, and R. Stern, "Purification and microsequencing of hyaluronidase isozymes from human urine," FEBS Letters, vol. 417, no. 3, pp. 307-310, 1997.
138. G. Lepperdinger, B. Strobl, and G. Kreil, "HYAL2, a human gene expressed in many cells, encodes a lysosomal hyaluronidase with a novel type of specificity," The Journal of Biological Chemistry, vol. 273, no. 35, pp. 22466-22470, 1998.
139. S. K. Rai, F.-M. Duh, V. Vigdorovich, A. Danilkovitch-Miagkova, M. I. Lerman, and A. D. Miller, "Cand idate tumor suppressor HYAL2 is a glycosylphosphatidylinositol (GPI)-anchored cell-surface receptor for jaagsiekte sheep retrovirus, the envelope protein of which mediates oncogenic transformation," Proceedings of the National Academy of Sciences of the United States of America, vol. 98, no. 8, pp. 4443-4448, 2001.
140. G. Kreil, "Hyaluronidases-a group of neglected enzymes," Protein Science, vol. 4, no. 9, pp. 1666-1669, 1995.
141. M. Slevin, J. Krupinski, J. Gaffney et al., "Hyaluronan-mediated angiogenesis in vascular disease: uncovering RHAMM and CD44 receptor signaling pathways," Matrix Biology, vol. 26, no. 1, pp. 58-68, 2007.
142. R. Stern, A. A. Asari, and K. N. Sugahara, "Hyaluronan fragments: an information-rich system," European Journal of Cell Biology, vol. 85, no. 8, pp. 699-715, 2006.
143. B. A. Neudecker, R. Stern, and M. K. Connolly, "Aberrant serum hyaluronan and hyaluronidase levels in scleroderma," British Journal of Dermatology, vol. 150, no. 3, pp. 469-476, 2004.
144. J. D. McNeil, O. W. Wiebkin, W. H. Betts, and L. G. Cleland , "Depolymerisation products of hyaluronic acid after exposure to oxygen-derived free radicals," Annals of the Rheumatic Diseases, vol. 44, no. 11, pp. 780-789, 1985.
145. D. C. West, I. N. Hampson, F. Arnold, and S. Kumar, "Angiogenesis induced by degradation products of hyaluronic acid," Science, vol. 228, no. 4705, pp. 1324-1336, 1985.
146. C. M. McKee, C. J. Lowenstein, M. R. Horton et al., "Hyaluronan fragments induce nitric-oxide synthase inmurine macrophages through a nuclear factor κB-dependent mechanism," The Journal of Biological Chemistry, vol. 272, no. 12, pp. 8013-8018, 1997.
147. C. M. McKee, M. B. Penno, M. Cowman et al., "Hyaluronan (HA) fragments induce chemokine gene expression in alveolar macrophages: the role of HA size and CD44," Journal of Clinical Investigation, vol. 98, no. 10, pp. 2403-2413, 1996.
148. B. Oertli, B. Beck-Schimmer, X. Fan, and R. P. Wüthrich, "Mechanisms of hyaluronan-induced up-regulation of ICAM-1 and VCAM-1 expression by murine kidney tubular epithelial cells: hyaluronan triggers cell adhesion molecule expression through a mechanism involving activation of nuclear factor-κB and activating protein-1," The Journal of Immunology, vol. 161, no. 7, pp. 3431-3437, 1998.
149. M. Slevin, J. Krupinski, S. Kumar, and J. Gaffney, "Angiogenic oligosaccharides of hyaluronan induce protein tyrosine kinase activity in endothelial cells and activate a cytoplasmic signal transduction pathway resulting in proliferation," Laboratory Investigation, vol. 78, no. 8, pp. 987-1003, 1998.
150. K. Ghazi, U. Deng-Pichon, J. M. Warnet, and P. Rat, "Hyaluronan fragments improve wound healing on in vitro cutaneous model through P2X7 purinoreceptor basal activation: role of molecular weight," PLoS ONE, vol. 7, no. 11, Article ID e48351, 2012.
151. D. C. West, D. M. Shaw, P. Lorenz, N. S. Adzick, and M. T. Longaker, "Fibrotic healing of adult and late gestation fetal wounds correlates with increased hyaluronidase activity and removal of hyaluronan," International Journal of Biochemistry and Cell Biology, vol. 29, no. 1, pp. 201-210, 1997.
152. D. C. West and M. Yaqoob, "Serum hyaluronan levels follow disease activity in vasculitis," Clinical Nephrology, vol. 48, no. 1, pp. 9-15, 1997.
153. D. J. Whitby and M. W. J. Ferguson, "The extracellularmatrix of lip wounds in fetal, neonatal and adult mice," Development, vol. 112, no. 2, pp. 651-668, 1991.
154. T. Sawai, N. Usui, K. Sando et al., "Hyaluronic acid of wound fluid in adult and fetal rabbits," Journal of Pediatric Surgery, vol. 32, no. 1, pp. 41-43, 1997.
155. D. Jiang, J. Liang, and P. W. Noble, "Hyaluronan in tissue injury and repair," Annual Review of Cell and Developmental Biology, vol. 23, pp. 435-461, 2007.
156. D. C. West and S. Kumar, "Hyaluronan and angiogenesis," Ciba Foundation Symposium, vol. 143, pp. 187-201, 1989.
157. P. Teder, R. W. Vandivier, D. Jiang et al., "Resolution of lung inflammation by CD44," Science, vol. 296, no. 5565, pp. 155-158, 2002.
158. C. B. Knudson and W. Knudson, "Hyaluronan-binding proteins in development, tissue homeostasis, and disease," The FASEB Journal, vol. 7, no. 13, pp. 1233-1241, 1993.
159. D. Naor, S. Nedvetzki, I. Golan, L. Melnik, and Y. Faitelson, "CD44 in cancer," Critical Reviews in Clinical Laboratory Sciences, vol. 39, no. 6, pp. 527-579, 2002.
160. D. Naor, S. B. Wallach-Dayan, M. A. Zahalka, and R. V. Sionov, "Involvement of CD44, a molecule with a thousand faces, in cancer dissemination," Seminars in Cancer Biology, vol. 18, no. 4, pp. 260-267, 2008.
161. D. Naor, R. V. Sionov, and D. Ish-Shalom, "CD44: structure, function, and associationwith the malignant process," Advances in Cancer Research, vol. 71, pp. 241-319, 1997.
162. E. A. Turley, L. Austen, K. Vandeligt, and C. Clary, "Hyaluronan and a cell-associated hyaluronan binding protein regulate the locomotion of ras-transformed cells," The Journal of Cell Biology, vol. 112, no. 5, pp. 1041-1047, 1991.
163. M. Zöller, "CD44: can a cancer-initiating cell profit from an abundantly expressed molecule?" Nature Reviews Cancer, vol. 11, no. 4, pp. 254-267, 2011.
164. D. Kohda, C. J. Morton, A. A. Parkar et al., "Solution structure of the link module: a hyaluronan-binding domain involved in extracellular matrix stability and cell migration," Cell, vol. 86, no. 5, pp. 767-775, 1996.
165. K. T. Dicker, L. A. Gurski, S. Pradhan-Bhatt, R. L. Witt, M. C. Farach-Carson, and X. Jia, "Hyaluronan: a simple polysaccharide with diverse biological functions," Acta Biomaterialia, vol. 10, no. 4, pp. 1558-1570, 2014.
166. A. J. Day and C. A. de la Motte, "Hyaluronan cross-linking: a protective mechanism in inflammation?" Trends in Immunology, vol. 26, no. 12, pp. 637-643, 2005.
167. S. Banerji, J. Ni, S.-X. Wang et al., "LYVE-1, a new homologue of theCD44 glycoprotein, is a lymph-specific receptor for hyaluronan," The Journal of Cell Biology, vol. 144, no. 4, pp. 789-801, 1999.
168. J. D. Kahmann, R. OBrien, J. M. Werner et al., "Localization and characterization of the hyaluronan-binding site on the link module from human TSG-6," Structure, vol. 8, no. 7, pp. 763-774, 2000.
169. G. Tzircotis, R. F. Thorne, and C. M. Isacke, "Chemotaxis towards hyaluronan is dependent on CD44 expression and modulated by cell type variation in CD44-hyaluronan binding," Journal of Cell Science, vol. 118, no. 21, pp. 5119-5128, 2005.
170. M. S. Pandey, B. A. Baggenstoss, J. Washburn, E. N. Harris, and P. H. Weigel, "Thehyaluronan receptor for endocytosis (HARE) activates NF-kappaB-mediated gene expression in response to 40-400-kDa, but not smaller or larger, hyaluronans," The Journal of Biological Chemistry, vol. 288, no. 20, pp. 14068-14079, 2013.
171. T. Ahrens, V. Assmann, C. Fieber et al., "CD44 is the principal mediator of hyaluronic-acid-induced melanoma cell proliferation," Journal of Investigative Dermatology, vol. 116, no. 1, pp. 93-101, 2001.
172. P. W. Noble, "Hyaluronan and its catabolic products in tissue injury and repair," Matrix Biology, vol. 21, no. 1, pp. 25-29, 2002.
173. B. P. Toole and V. C. Hascall, "Hyaluronan and tumor growth," American Journal of Pathology, vol. 161, no. 3, pp. 745-747, 2002.
174. M. W. Makgoba, M. E. Sanders, G. E. Ginther Luce et al., "ICAM-1 a ligand for LFA-1-dependent adhesion of B, T and myeloid cells," Nature, vol. 331, no. 6151, pp. 86-88, 1988.
175. G. R. Screaton, M. V. Bell, D. G. Jackson, F. B. Cornelis, U. Gerth, and J. I. Bell, "Genomic structure of DNA encoding the lymphocyte homing receptor CD44 reveals at least 12 alternatively spliced exons," Proceedings of the National Academy of Sciences of the United States of America, vol. 89, no. 24, pp. 12160-12164, 1992.
176. K. W. Lynch, "Consequences of regulated pre-mRNA splicing in the immune system," Nature Reviews Immunology, vol. 4, no. 12, pp. 931-940, 2004.
177. J. Cichy and E. Puré, "Oncostatin M and transforming growth factor-beta 1 induce post-translational modification and hyaluronan binding to CD44 in lung-derived epithelial tumor cells," The Journal of Biological Chemistry, vol. 275, no. 24, pp. 18061-18069, 2000.
178. A. Maiti, G. Maki, and P. Johnson, "TNF-alpha induction of CD44-mediated leukocyte adhesion by sulfation," Science, vol. 282, no. 5390, pp. 941-943, 1998.
179. L. E. Esford, A. Maiti, S. A. Bader, F. Tufaro, and P. Johnson, "Analysis ofCD44 interactionswith hyaluronan inmurine L cell fibroblasts deficient in glycosaminoglycan synthesis: a role for chondroitin sulfate," Journal of Cell Science, vol. 111, part 7, pp. 1021-1029, 1998.
180. S. Katoh, Z. Zheng, K. Oritani, T. Shimozato, and P. W. Kincade, "Glycosylation of CD44 negatively regulates its recognition of hyaluronan," Journal of Experimental Medicine, vol. 182, no. 2, pp. 419-429, 1995.
181. J. Lesley, N. English, A. Perschl, J. Gregoroff, and R. Hyman, "Variant cell lines selected for alterations in the function of the hyaluronan receptor CD44 show differences in glycosylation," Journal of Experimental Medicine, vol. 182, no. 2, pp. 431-437, 1995.
182. S. Misra, V. C. Hascall, C. de Giovanni, R. R. Markwald, and S. Ghatak, "Delivery of CD44 shRNA/nanoparticles within cancer cells: perturbation of hyaluronan/CD44v6 interactions and reduction in adenoma growth in Apc Min/+ MICE," The Journal of Biological Chemistry, vol. 284, no. 18, pp. 12432-12446, 2009.
183. E. Puré, R. L. Camp, D. Peritt, R. A. Panettieri Jr., A. L. Lazaar, and S. Nayak, "Defective phosphorylation and hyaluronate binding of CD44 with point mutations in the cytoplasmic domain," Journal of Experimental Medicine, vol. 181, no. 1, pp. 55-62, 1995.
184. T. P. Skelton, C. Zeng, A. Nocks, and I. Stamenkovic, "Glycosylation provides both stimulatory and inhibitory effects on cell surface and soluble CD44 binding to hyaluronan," The Journal of Cell Biology, vol. 140, no. 2, pp. 431-446, 1998.
185. L. Y. W. Bourguignon, P. A. Singleton, H. Zhu, and F. Diedrich, "Hyaluronan-mediated CD44 interaction with RhoGEF and Rho kinase promotes Grb2-associated binder-1 phosphorylation and phosphatidylinositol 3-kinase signaling leading to cytokine (macrophage-colony stimulating factor) production and breast tumor progression," The Journal of Biological Chemistry, vol. 278, no. 32, pp. 29420-29434, 2003.
186. L. Y. W. Bourguignon, H. Zhu, L. Shao, and Y. W. Chen, "CD44 interaction with Tiam1 promotes Rac1 signaling and hyaluronic acid-mediated breast tumor cell migration," The Journal of Biological Chemistry, vol. 275, no. 3, pp. 1829-1838, 2000.
187. L. Y. W. Bourguignon, H. Zhu, L. Shao, and Y.-W. Chen, "CD44 interaction with c-Src kinase promotes cortactin-mediated cytoskeleton function and hyaluronic acid-dependent ovarian tumor cell migration," The Journal of Biological Chemistry, vol. 276, no. 10, pp. 7327-7336, 2001.
188. S. Misra, S. Ghatak, and B. P. Toole, "Regulation of MDR1 expression and drug resistance by a positive feedback loop involving hyaluronan, phosphoinositide 3-kinase, and ErbB2," The Journal of Biological Chemistry, vol. 280, no. 21, pp. 20310-20315, 2005.
189. S. Misra, S. Ghatak, A. Zoltan-Jones, and B. P. Toole, "Regulation of multidrug resistance in cancer cells by hyaluronan," Journal of Biological Chemistry, vol. 278, no. 28, pp. 25285-25288, 2003.
190. S. Misra, B. P. Toole, and S. Ghatak, "Hyaluronan constitutively regulates activation of multiple receptor tyrosine kinases in epithelial and carcinoma cells," The Journal of Biological Chemistry, vol. 281, no. 46, pp. 34936-34941, 2006.
191. S. Ghatak, S. Misra, and B. P. Toole, "Hyaluronan oligosaccharides inhibit anchorage-independent growth of tumor cells by suppressing the phosphoinositide 3-kinase/Akt cell survival pathway," The Journal of Biological Chemistry, vol. 277, no. 41, pp. 38013-38020, 2002.
192. S. Ghatak, S. Misra, and B. P. Toole, "Hyaluronan constitutively regulates ErbB2 phosphorylation and signaling complex formation in carcinoma cells," The Journal of Biological Chemistry, vol. 280, no. 10, pp. 8875-8883, 2005.
193. K.-H. Heider, M. Hofmann, E. Hors et al., "Ahuman homologue of the rat metastasis-associated variant of CD44 is expressed in colorectal carcinomas and adenomatous polyps," Journal of Cell Biology, vol. 120, no. 1, pp. 227-233, 1993.
194. T. Ishida, "Immunohistochemical expression of the CD44 variant 6 in colorectal adenocarcinoma," Surgery Today, vol. 30, no. 1, pp. 28-32, 2000.
195. R. Neumayer, H. R. Rosen, A. Reiner et al., "CD44 expression in benign and malignant colorectal polyps," Diseases of the Colon and Rectum, vol. 42, no. 1, pp. 50-55, 1999.
196. V. J. M. Wielenga, K.-H. Heider, G. J. A. Offerhaus et al., "Expression of CD44 variant proteins in human colorectal cancer is related to tumor progression," Cancer Research, vol. 53, no. 20, pp. 4754-4756, 1993.
197. V. J. M. Wielenga, R. van der Neut, G. J. A. Offerhaus, and S. T. Pals, "CD44 Glycoproteins in colorectal cancer: expression, function, and prognostic value," Advances in Cancer Research, vol. 77, pp. 169-187, 2000.
198. V. J. M. Wielenga, R. vanderVoort, J. W. R. Mulder et al., "CD44 splice variants as prognostic markers in colorectal cancer," Scandinavian Journal of Gastroenterology, vol. 33, no. 1, pp. 82-87, 1998.
199. S. M. Chang, R. D. Xing, F. M. Zhang, and Y. Q. Duan, "Serum soluble CD44v6 levels in patients with oral and maxillofacial malignancy," Oral Diseases, vol. 15, no. 8, pp. 570-572, 2009.
200. B. Delpech, B. Chevallier, N. Reinhardt et al., "Serum hyaluronan (hyaluronic acid) in breast cancer patients," International Journal of Cancer, vol. 46, no. 3, pp. 388-390, 1990.
201. A. Josefsson, H. Adamo, P. Hammarsten et al., "Prostate cancer increases hyaluronan in surrounding nonmalignant stroma, and this response is associated with tumor growth and an unfavorable outcome," The American Journal of Pathology, vol. 179, no. 4, pp. 1961-1968, 2011.
202. T. Jung, W. Gross, and M. Zöller, "CD44v6 coordinates tumor matrix-triggered motility and apoptosis resistance," The Journal of Biological Chemistry, vol. 286, no. 18, pp. 15862-15874, 2011.
203. Q. Xie, R. Bradley, L. Kang et al., "Hepatocyte growth factor (HGF) autocrine activation predicts sensitivity to MET inhibition in glioblastoma," Proceedings of the National Academy of Sciences of the United States of America, vol. 109, no. 2, pp. 570-575, 2011.
204. R. D. Xing, S. M. Chang, Y. Q. Duan, F. M. Zhang, and F. S. Dong, "Examination of serum hyaluronic acid level in patients with oral and maxillofacial malignancy," Hua Xi Kou Qiang Yi Xue Za Zhi, vol. 22, no. 4, pp. 309-311, 2004.
205. Q. Yu and I. Stamenkovic, "Localization of matrix metalloproteinase 9 to the cell surface provides a mechanism for CD44-mediated tumor invasion," Genes and Development, vol. 13, no. 1, pp. 35-48, 1999.
206. M. C. Levesque and B. F. Haynes, "In vitro culture of human peripheral blood monocytes induces hyaluronan binding and up-regulates monocyte variant CD44 isoformexpression," Journal of Immunology, vol. 156, no. 4, pp. 1557-1565, 1996.
207. Y. Matsubara, S. Katoh, H. Taniguchi, M. Oka, J. Kadota, and S. Kohno, "Expression of CD44 variants in lung cancer and its relationship to hyaluronan binding," Journal of International Medical Research, vol. 28, no. 2, pp. 78-90, 2000.
208. J. P. Sleeman, S. Arming, J. F. Moll et al., "Hyaluronate-independent metastatic behavior of CD44 variant-expressing pancreatic carcinoma cells," Cancer Research, vol. 56, no. 13, pp. 3134-3141, 1996.
209. V. Orian-Rousseau, L. Chen, J. P. Sleeman, P. Herrlich, and H. Ponta, "CD44 is required for two consecutive steps in HGF/c-Met signaling," Genes and Development, vol. 16, no. 23, pp. 3074-3086, 2002.
210. U. Protin, T. Schweighoffer, W. Jochum, and F. Hilberg, "CD44-deficient mice develop normally with changes in subpopulations and recirculation of lymphocyte subsets," Journal of Immunology, vol. 163, no. 9, pp. 4917-4923, 1999.
211. A. Q. Rafi-Janajreh, D. Chen, R. Schmits et al., "Evidence for the involvement of CD44 in endothelial cell injury and induction of vascular leak syndrome by IL-2," Journal of Immunology, vol. 163, no. 3, pp. 1619-1627, 1999.
212. D. Chen, R. J. McKallip, A. Zeytun et al., "CD44-Deficient mice exhibit enhanced hepatitis after concanavalin A injection: evidence for involvement of CD44 in activation-induced cell death," Journal of Immunology, vol. 166, no. 10, pp. 5889-5897, 2001.
213. G. Abatangelo, M. Martelli, and P. Vecchia, "Healing of hyaluronic acid-enriched wounds: histological observations," Journal of Surgical Research, vol. 35, no. 5, pp. 410-416, 1983.
214. D. Foschi, F. Castoldi, E. Radaelli et al., "Hyaluronic acid prevents oxygen free-radical damage to granulation tissue: a study in rats," International Journal of Tissue Reactions, vol. 12, no. 6, pp. 333-339, 1990.
215. S. R. King, W. L. Hickerson, and K. G. Proctor, "Beneficial actions of exogenous hyaluronic acid on wound healing," Surgery, vol. 109, no. 1, pp. 76-84, 1991.
216. M. Nakamura, M. Hikida, and T. Nakano, "Concentration and molecular weight dependency of rabbit corneal epithelial wound healing on hyaluronan," Current Eye Research, vol. 11, no. 10, pp. 981-986, 1992.
217. C. Laurent, S. Hellstrom, and L.-E. Stenfors, "Hyaluronic acid reduces connective tissue formation in middle ears filled with absorbable gelatin sponge: an experimental study," The American Journal of Otolaryngology: Head and Neck Medicine and Surgery, vol. 7, no. 3, pp. 181-186, 1986.
218. E. A. Balasz and J. L. Denlinger, Clinical Uses of Hyaluronan, John Wiley & Sons, 1989.
219. M. T. Longaker, E. S. Chiu, N. S. Adzick, M. Stern, M. R. Harrison, and R. Stern, "Studies in fetal wound healing. V. A prolonged Presence of hyaluronic acid characterizes fetal wound fluid," Annals of Surgery, vol. 213, no. 4, pp. 292-296, 1991.
220. J. A. Mack, S. R. Abramson, Y. Ben et al., "Hoxb13 knockout adult skin exhibits high levels of hyaluronan and enhanced wound healing," The FASEB Journal, vol. 17, no. 10, pp. 1352-1354, 2003.
221. J. A. Mack, L. Li, N. Sato, V. C. Hascall, and E. V. Maytin, "Hoxb13 up-regulates transglutaminase activity and drives terminal differentiation in an epidermal organotypic model," The Journal of Biological Chemistry, vol. 280, no. 33, pp. 29904-29911, 2005.
222. J. A. MacK and E. V. Maytin, "Persistent inflammation and angiogenesis during wound healing in K14-directed hoxb13 transgenic mice," Journal of Investigative Dermatology, vol. 130, no. 3, pp. 856-865, 2010.
223. D. A. R. Burd, R. M. Greco, S. Regauer, M. T. Longaker, J. W. Siebert, and H. G. Garg, "Hyaluronanand woundhealing: a new perspective," British Journal of Plastic Surgery, vol. 44, no. 8, pp. 579-584, 1991.
224. P. Heldin, T. C. Laurent, and C.-H. Heldin, "Effect of growth factors on hyaluronan synthesis in cultured human fibroblasts," Biochemical Journal, vol. 258, no. 3, pp. 919-922, 1989.
225. B. A. Mast, R. F. Diegelmann, T. M. Krummel, and I. K. Cohen, "Hyaluronic acid modulates proliferation, collagen and protein synthesis of cultured fetal fibroblasts," Matrix, vol. 13, no. 6, pp. 441-446, 1993.
226. G. F. Pierce, T. A. Mustoe, J. Lingelbach et al., "Platelet-derived growth factor and transforming growth factor-beta enhance tissue repair activities by unique mechanisms," The Journal of Cell Biology, vol. 109, no. 1, pp. 429-440, 1989.
227. S. E. Lynch, J. C. Nixon, R. B. Colvin, and H. N. Antoniades, "Role of platelet-derived growth factor in wound healing: synergistic effects with other growth factors," Proceedings of the National Academy of Sciences of the United States of America, vol. 84, no. 21, pp. 7696-7700, 1987.
228. G. F. Pierce, J. E. Tarpley, J. Tseng et al., "Detection of plateletderived growth factor (PDGF)-AA in actively healing human wounds treated with recombinant PDGF-BB and absence of PDGF in chronic nonhealing wounds," The Journal of Clinical Investigation, vol. 96, no. 3, pp. 1336-1350, 1995.
229. A. L. Carre, A. W. James, L. MacLeod et al., "Interaction of wingless protein (Wnt), transforming growth factor-beta1, and hyaluronan production in fetal and postnatal fibroblasts," Plastic and Reconstructive Surgery, vol. 125, no. 1, pp. 74-88, 2010.
230. M. David-Raoudi, F. Tranchepain, B. Deschrevel et al., "Differential effects of hyaluronan and its fragments on fibroblasts: relation to wound healing," Wound Repair and Regeneration, vol. 16, no. 2, pp. 274-287, 2008.
231. R. C. Cabera, J. W. Siebert, Y. Eidelman, L. I. Gold, M. T. Longaker, and H. G. Garg, "The in vivo effect of hyaluronan associated protein-collagen complex on wound repair," Biochemistry and Molecular Biology International, vol. 37, no. 1, pp. 151-158, 1995.
232. H. Morrison, L. S. Sherman, J. Legg et al., "The NF2 tumor suppressor gene product, merlin, mediates contact inhibition of growth through interactions with CD44," Genes and Development, vol. 15, no. 8, pp. 968-980, 2001.
233. S. P. Thankamony and W. Knudson, "Acylation of CD44 and its association with lipid rafts are required for receptor and hyaluronan endocytosis," The Journal of Biological Chemistry, vol. 281, no. 45, pp. 34601-34609, 2006.
234. P. Heldin, Growth Factor Regulation of Hyaluronan Metabolism in Tumor Progression, 2011.
235. E. Bell, B. Ivarsson, and C. Merrill, "Production of a tissue-like structure by contraction of collagen lattices by human fibroblasts of different proliferative potential in vitro," Proceedings of the National Academy of Sciences of the United States of America, vol. 76, no. 3, pp. 1274-1278, 1979.
236. I. V. Yannas, E. Lee, D. P. Orgill, E. M. Skrabut, and G. F. Murphy, "Synthesis and characterization of a model extracellular matrix that induces partial regeneration of adult mammalian skin," Proceedings of the National Academy of Sciences of the United States of America, vol. 86, no. 3, pp. 933-937, 1989.
237. S. R. Slivka, L. K. Land een, F. Zeigler, M. P. Zimber, and R. L. Bartel, "Characterization, barrier function, and drug metabolism of an in vitro skin model," Journal of Investigative Dermatology, vol. 100, no. 1, pp. 40-46, 1993.
238. G. Galassi, P. Brun, M. Radice et al., "In vitro reconstructed dermis implanted in human wounds: degradation studies of the HA-based supporting scaffold," Biomaterials, vol. 21, no. 21, pp. 2183-2191, 2000.
239. D. A. Hollander, C. Soranzo, S. Falk, and J. Windolf, "Extensive traumatic soft tissue loss: reconstruction in severely injured patients using cultured hyaluronan-based three-dimensional dermal and epidermal autografts," Journal of Trauma-Injury, Infection and Critical Care, vol. 50, no. 6, pp. 1125-1136, 2001.
240. Y. Liu, A. Skardal, X. Z. Shu, and G. D. Prestwich, "Preventionof peritendinous adhesions using a hyaluronan-derived hydrogel film following partial-thickness flexor tendon injury," Journal of Orthopaedic Research, vol. 26, no. 4, pp. 562-569, 2008.
241. G. D. Prestwich, "Engineering a clinically-useful matrix for cell therapy," Organogenesis, vol. 4, no. 1, pp. 42-47, 2008.
242. G. D. Prestwich, "Evaluating drug efficacy and toxicology in three dimensions: using synthetic extracellular matrices in drug discovery," Accounts of Chemical Research, vol. 41, no. 1, pp. 139-148, 2008.
243. X. Z. Shu, Y. Liu, Y. Luo, M. C. Roberts, and G. D. Prestwich, "Disulfide cross-linked hyaluronan hydrogels," Biomacromolecules, vol. 3, no. 6, pp. 1304-1311, 2002.
244. T. D. Mehra, K. Ghosh, X. Z. Shu, G. D. Prestwich, and R. A. F. Clark, "Molecular stenting with a crosslinked hyaluronan derivative inhibits collagen gel contraction," Journal of Investigative Dermatology, vol. 126, no. 10, pp. 2202-2209, 2006.
245. J.-W. Kuo, D. A. Swann, and G. D. Prestwich, "Chemical modification of hyaluronic acid by carbodiimides," Bioconjugate Chemistry, vol. 2, no. 4, pp. 232-241, 1991.
246. E. J. Oh, S. W. Kang, B. S. Kim, G. Jiang, H. C. Il, and K. H. Sei, "Control of the molecular degradation of hyaluronic acid hydrogels for tissue augmentation," Journal of BiomedicalMaterials Research-Part A, vol. 86, no. 3, pp. 685-693, 2008.
247. K. L. Chao, L. Muthukumar, and O. Herzberg, "Structure of human hyaluronidase-1, a hyaluronan hydrolyzing enzyme involved in tumor growth and angiogenesis," Biochemistry, vol. 46, no. 23, pp. 6911-6920, 2007.
248. J. Entwistle, C. L. Hall, and E. A. Turley, "HA receptors: regulators of signalling to the cytoskeleton," Journal of Cellular Biochemistry, vol. 61, no. 4, pp. 569-577, 1996.
249. H. Lee, H. Mok, S. Lee, Y.-K. Oh, and T. G. Park, "Target-specific intracellular delivery of siRNA using degradable hyaluronic acid nanogels," Journal of Controlled Release, vol. 119, no. 2, pp. 245-252, 2007.
250. A. Gabrielli, E. V. Avvedimento, and T. Krieg, "Scleroderma," The New England Journal of Medicine, vol. 360, no. 19, pp. 1989-2003, 2009.
251. J. Varga and D. Abraham, "Systemic sclerosis: a prototypicmultisystem fibrotic disorder," The Journal of Clinical Investigation, vol. 117, no. 3, pp. 557-567, 2007.
252. M. S. Anscher, W. P. Peters, H. Reisenbichler, W. P. Petros, and R. L. Jirtle, "Transforming growth factor beta as a predictor of liver and lung fibrosis after autologous bone marrow transplantation for advanced breast cancer," The New England Journal of Medicine, vol. 328, no. 22, pp. 1592-1598, 1993.
253. K. Matsumoto and T. Nakamura, "Hepatocyte growth factor: renotropic role and potential therapeutics for renal diseases," Kidney International, vol. 59, no. 6, pp. 2023-2038, 2001.
254. A. Akhmetshina, P. Venalis, C. Dees et al., "Treatment with imatinib prevents fibrosis in different preclinical models of systemic sclerosis and induces regression of established fibrosis," Arthritis and Rheumatism, vol. 60, no. 1, pp. 219-224, 2009.
255. J. H. W. Distler, A. Jüngel, L. C. Huber et al., "Imatinib mesylate reduces production of extracellular matrix and prevents development of experimental dermal fibrosis," Arthritis and Rheumatism, vol. 56, no. 1, pp. 311-322, 2007.
256. H. A. Burgess, L. E. Daugherty, T. H. Thatcher et al., "PPARγ agonists inhibit TGF-β induced pulmonary myofibroblast differentiation and collagen production: implications for therapy of lung fibrosis," The American Journal of Physiology: Lung Cellular and Molecular Physiology, vol. 288, no. 6, pp. L1146-L1153, 2005.
257. L. Hecker, J. Cheng, and V. J. Thannickal, "Targeting NOX enzymes in pulmonary fibrosis," Cellular and Molecular Life Sciences, vol. 69, no. 14, pp. 2365-2371, 2012.
258. K. Burridge and M. Chrzanowska-Wodnicka, "Focal adhesions, contractility, and signaling," Annual Review of Cell and Developmental Biology, vol. 12, pp. 463-519, 1996.
259. M. Chrzanowska-Wodnicka and K. Burridge, "Rho-stimulated contractility drives the formation of stress fibers and focal adhesions," Journal of Cell Biology, vol. 133, no. 6, pp. 1403-1415, 1996.
260. Y. Zhou, X. Huang, L. Hecker et al., "Inhibition ofmechanosensitive signaling in myofibroblasts ameliorates experimental pulmonary fibrosis," Journal of Clinical Investigation, vol. 123, no. 3, pp. 1096-1108, 2013.