Hyaluronan and CD44: Strategic players for cell-matrix interactions during chondrogenesis and matrix assembly

Birth Defects Research Part C - Embryo Today: Reviews

Volumn 69, Issue 2, 2003, Pages 174-196

  Knudson, Cheryl B ^a  

^a RUSH MEDICAL COLLEGE   (United States)

Author keywords

Cartilage; CD44; Chondrocyte; Chondrogenesis; Hyaluronan; Matrix assembly

Indexed keywords

HERMES ANTIGEN; HYALURONIC ACID; HYALURONIC ACID BINDING PROTEIN; MORPHOGEN;

ANTIGEN EXPRESSION; CARTILAGE; CARTILAGE CELL; CELL INTERACTION; CELL STRUCTURE; CELL SURFACE; CHONDROGENESIS; CHONDROLYSIS; EMBRYO DEVELOPMENT; EMBRYO INDUCTION; EXTRACELLULAR MATRIX; HUMAN; MORPHOGENESIS; NONHUMAN; PRIORITY JOURNAL; REVIEW; SIGNAL TRANSDUCTION;

ANIMALS; ANTIGENS, CD44; CARTILAGE; CHONDROCYTES; CHONDROGENESIS; EXTRACELLULAR MATRIX; HUMANS; HYALURONIC ACID;

EID: 0141993878     PISSN: 1542975X     EISSN: None     Source Type: Journal    
DOI: 10.1002/bdrc.10013     Document Type: Review

References (236)

1. Adams JC, Watt FM. 1993. Regulation of development and differentiation by the extracellular matrix. Development 117:1183-1198.
2. Aguiar DJ, Knudson W, Knudson CB. 1999. Internalization of the hyaluronan receptor CD44 by chondrocytes. Exp Cell Res 252:292-302.
3. Ahrens PB, Solursh M, Reiter RS. 1977. Stage-related capacity for limb chondrogenesis in cell culture. Dev Biol 60: 69-82.
4. Arikawa-Hirasawa E, Watanabe E, Watanabe H, et al. 1999. Perlecan is essential for cartilage and cephalic development. Nat Genet 23:354-358.
5. Aruffo A, Stamenkovic I, Melnick M, et al. 1990. CD44 is the principal cell surface receptor for hyaluronate. Cell 61:1303-1313.
6. Aspberg A, Miura R, Bourdoulous S, et al. 1997. The c-type lectin domains of lecticans, a family of aggregating chondroitin sulfate proteoglycans, bind tenascin-R by protein-protein interactions independent of carbohydrate moiety. Proc Natl Acad Sci U S A 94:10116-10121.
7. Aspberg A, Adam S, Kostka G, et al. 1999. Fibulin-1 is a ligand for the c-type lectin domains of aggrecan and versican. J Biol Chem 274:20444-20449.
8. Aulthouse AL, Solursh M. 1987. The detection of a precartilage, blastema-specific marker. Dev Biol 120:377-384.
9. Aydelotte MB, Kuettner KE. 1988. Differences between subpopulations of cultured bovine articular chondrocytes. Connect Tissue Res 18:223-234.
10. Bajorath J, Greenfield B, Munro S, et al. 1998. Identification of CD44 residues important for hyaluronan binding and delineation of the binding site. J Biol Chem 273:338-343.
11. Banerji S, Ni J, Wang S-X, et al. 1999. LYVE-1, a new homologue of the CD44 glycoprotein, is a lymph-specific receptor for hyaluronan. J Cell Biol 144:789-801.
12. Bartolazzi A, Nocks A, Aruffo A, et al. 1996. Glycosylation of CD44 is implicated in CD44-mediated cell adhesion to hyaluronan. J Cell Biol 132:1199-1208.
13. Bayliss MT, Howat S, Davidson C, Dudhia J. 2000. The organization of aggrecan in human articular cartilage. J Biol Chem 275:6321-6327.
14. Bayliss MT, Howat S, Dudhia J, et al. 2001. Up-regulation and differential expression of the hyaluronan-binding protein TSG-6 in cartilage and synovium in rheumatoid arthritis and osteoarthritis. Osteoarthritis Cartilage 9: 42-48.
15. Bee JA, von der Mark K. 1990. An analysis of chick limb bud intercellular adhesion underlying the establishment of cartilage aggregates in suspension culture. J Cell Sci 96:527-536.
16. Benya PD, Padilla SR. 1986. Modulation of the rabbit chondrocyte phenotype by retinoic acid terminates type II collagen synthesis without inducing type I collagen: the modulated phenotype differs from that produced by subculture. Dev Biol 118:296-305.
17. Berg EL, Goldstein LA, Jutila MA, et al. 1989. Homing receptors and vascular addressins: cell adhesion molecules that direct lymphocyte traffic. Immunol Rev 108:5-18.
18. Bourguignon LYW, Zhu D, Zhu H. 1998. CD44 isoform-cytoskeleton interaction in oncogenic signaling and tumor progression. Front Biosci 3:637-649.
19. Buckwalter JA, Rosenberg LC. 1982. Electron micrographic studies of cartilage proteoglycans. J Biol Chem 257: 9830-9839.
20. Camenisch TD, Spicer AP, Brehm-Gibson T, et al. 2000. Disruption of hyaluronan synthase-2 abrogates normal cardiac morphogenesis and hyaluronan-mediated transformation of epithelium to mesenchyme. J Clin Invest 106:349-60.
21. Camp RL, Kraus TA, Pure E. 1991. Variations in the cytoskeletal interaction and posttranslational modification of the CD44 homing receptor in macrophages. J Cell Biol 115:1283-1292.
22. Chow G, Knudson CB, Homandberg G, Knudson W. 1995. Increased CD44 expression in bovine articular chondrocytes by catabolic cellular mediators. J Biol Chem 270:27734-27741.
23. Chow G, Nietfeld J, Knudson CB, Knudson W. 1998. Antisense inhibition of chondrocyte CD44 expression results in cartilage chondrolysis. Arthritis Rheum 41:1411-1419.
24. Clarris BJ, Fraser JRE. 1968. On the pericellular zone of some mammalian cells in vitro. Exp Cell Res 49:181-193.
25. Costell M, Gustafsson E, Aszodi A, et al. 1999. Perlecan maintains the integrity of cartilage and some basement membranes. J Cell Biol 147:1109-1122.
26. Csoka AB, Frost GI, Stern R. 2001. The six hyaluronidase-like genes in the human and mouse genomes. Matrix Biol 20:499-508.
27. Culty M, Nguyen HA, Underhill CB. 1992. The hyaluronan receptor (CD44) participates in the uptake and degradation of hyaluronan. J Cell Biol 116:1055-1062.
28. Daniels K, Solursh M. 1991. Modulation of chondrogenesis by the cytoskeleton and extracellular matrix. J Cell Sci 100:249-254.
29. Dealy CN, Seghatoleslami MR, Ferrari D, Kosher RA. 1997. FGF-stimulated outgrowth and proliferation of limb mesoderm is dependent on syndecan-3. Dev Biol 184:343-350.
30. De Arcangelis A, Mark M, Kreidberg J, et al. 1999. Synergistic activities of alpha3 and alpha6 integrins are required during apical ectodermal ridge formation and organogenesis in the mouse. Development 126:3957-68.
31. de Crombrugghe B, Lefebvre V, Nakashima K. 2001. Regulatory mechanisms in the pathways of cartilage and bone formation. Curr Opin Cell Biol 13: 721-727.
32. DeLise AM, Fischer L, Tuan RS. 2000. Cellular interactions and signaling in cartilage development. Osteoarthritis Cartilage 8:309-334.
33. DeLise AM, Tuan R. 2002. Alterations in the spatiotemporal expression pattern and function of n-cadherin inhibit cellular condensation and chondrogenesis of limb mesenchymal cells in vitro. J Cell Biochem 87:342-359.
34. Dieppe P, Cushnaghan J, McAlindon T. 1992. Epidemiology, clinical course, and outcome of knee osteoarthritis. In: Kuettner KE, Schleyerbach R, Peyron JG, Hascall VC, editors. Articular cartilage and osteoarthritis. New York: Raven Press. p 617-628.
35. Dingle JT. 1993. Mechanisms of cartilage destruction and repair: the outlook for therapeutic interventions. Clin Rheumatol 3:1-5.
36. Doege K, Rhodes C, Sasaki M, et al. 1990. Molecular biology of cartilage proteoglycan (aggrecan) and link protein. In: Sandell LJ, Boyd CD, editors. Extracellular matrix genes. San Diego: Academic Press, Inc. p 137-155.
37. Doege K, Hall L, McKinnon W, et al. 2002. A remote upstream element regulates tissue-specific expression of the rat aggrecan gene. J Biol Chem 277:13989-13997.
38. Drossopoulou G, Lewis KE, Sanz-Ezquerro JJ, et al. 2000. A model for anteroposterior patterning of the vertebrate limb based on sequential long- and short-range Shh signalling and BMP signalling. Development 127: 1337-1348.
39. Durr J, Goodman S, Potocnik A, et al. 1993. Localization of b1-integrins in human cartilage and their role in chondrocyte adhesion to collagen and fibronectin. Exp Cell Res 207:235-244.
40. Enomoto-Iwamoto M, Iwamoto M, Nakashima K, et al. 1997. Involvement of alpha5beta1 integrin in matrix interactions and proliferation of chondrocytes. J Bone Miner Res 12:1124-1132.
41. Flechtenmacher J, Huch K, Thonar EJMA, et al. 1996. Recombinant human osteogenic protein 1 is a potent stimulator of the synthesis of cartilage proteoglycans and collagens by human articular chondrocytes. Arthritis Rheum 39:478-488.
42. Fraser JRE, Alcorn D, Laurent TC, et al. 1985. Uptake of circulating hyaluronic acid by the rat liver. Cell Tissue Res 242:505-510.
43. Fraser JRE, Kimpton WG, Laurent TC, et al. 1988. Uptake and degradation of hyaluronan in lymphatic tissue. Biochem J 256:153-158.
44. Fraser JRE, Laurent TC. 1989. Turnover and metabolism of hyaluronan. In: Evered D, editor. The biology of hyaluronan. Chichester: John Wiley & Sons. p 41-59.
45. French MM, Smith SE, Akanbi K, et al. 1999. Expression of the heparan sulfate proteoglycan, perlecan, during mouse embryogenesis and perlecan chondrogenic activity in vivo. J Cell Biol 145:1103-1115.
46. Geduspan JS, Solursh M. 1993. Effects of the mesonephros and insulin-like growth factor i on chondrogenesis of limb explants. Dev Biol 156:500-508.
47. Giancotti FG, Ruoslahti E. 1999. Integrin signaling. Science 285:1028-1032.
48. Goldberg RL, Toole BP. 1984. Pericellular coat of chick embryo chondrocytes: structural role of hyaluronate. J Cell Biol 99:2114-2122.
49. Gould SE, Upholt WB, Kosher RA. 1992. Syndecan 3: a member of the syndecan family of membrane-intercalated proteoglycans that is expressed in high amounts at the onset of chicken limb cartilage differentiation. Proc Natl Acad Sci U S A 89:3271-3275.
50. Grover J, Roughley PJ. 1995. Expression of cell-surface proteoglycan mrna by human articular chondrocytes. Biochem J 309:963-968.
51. Gunthert U, Hofmann M, Rudy W, et al. 1991. A new variant of glycoprotein CD44 confers metastatic potential to rat carcinoma cells. Cell 65:13-24.
52. Gunthert U. 1993. CD44. A multitude of isoforms with diverse functions. Curr Top Microbiol Immunol 184:47.
53. Hall BK, Miyake T. 2000. All for one and one for all: condensations and the initiation of skeletal development. BioEssays 22:138-147.
54. Hamerman D, 1993. Aging and osteoarthritis: basic mechanisms. J Am Geriatr Soc 41:760-770.
55. Handler M, Yurchenco PD, Iozzo RV. 1997. Developmental expression of perlecan during murine embryogenesis. Dev Dyn 210:130-145.
56. Hardingham TE, Muir H. 1972. The specific interaction of hyaluronic acid with cartilage proteoglycans. Biochim Biophys Acta 279:401-405.
57. Hardingham TE, Muir H. 1973. Binding of oligosaccharides of hyaluronic acid to proteoglycans. Biochem J 135:905-908.
58. Hardingham TE. 1979. The role of link-protein in the structure of cartilage proteoglycan aggregates. Biochem J 177:237-247.
59. Hardwick C, Hoare K, Owens R, et al. 1992. Molecular cloning of a novel hyaluronan receptor that mediates tumor cell motility. J Cell Biol 117: 1343-1350.
60. Hascall VC, Sajdera SW. 1969. Protein polysaccharide complex from bovine nasal cartilage. The function of glycoprotein in the formation of aggregates. J Biol Chem 244:2384-2396.
61. Hascall VC, Heinegard D. 1974. Aggregation of cartilage proteoglycans, II. Oligosaccharide competitors of the proteoglycan-hyaluronic acid interaction. J Biol Chem 249:4242-4249.
62. Hascall VC. 1977. Interaction of cartilage proteoglycans with hyaluronic acid. J Supramol Struct 7:101-120.
63. Hauselmann HJ, Aydelotte MB, Schumacher BL, et al. 1992. Synthesis and turnover of proteoglycans by human and bovine adult articular chondrocytes cultured in alginate beads. Matrix 12:130-136.
64. Hauselmann HJ, Fernandes RJ, Mok SS, et al. 1994. Phenotypic stability of bovine articular chondrocytes after long-term culture in alginate beads. J Cell Sci 107:17-27.
65. Hay ED. 1991. Collagen and other matrix glycoproteins in embryogenesis. In: Hay ED, editor. Cell biology of the extracellular matrix. New York: Plenum Press. p 419-462.
66. Hay ED. 1989. Extracellular matrix, cell skeletons, and embryonic development. Am J Med Genet 34:14-29.
67. Haynes BF, Telen MJ, Hale LP, Denning SM. 1989. CD44-a molecule involved in leukocyte adherence and t-cell activation. Immunol Today 10: 423-428.
68. Haynes BF, Hua-Xin L, Patton KL. 1991. The transmembrane hyaluronate receptor (CD44): multiple functions, multiple forms. Cancer Cells 3:347-350.
69. Heinegard D, Paulsson M. 1984. Structure and metabolism of proteoglycans. In: Piez KA, Reddi AH, editors. Extracellular matrix biochemistry. New York: Elsevier. p 277-328.
70. Heinegard D, Lorenzo P, Sommarin Y. 1995. Articular cartilage matrix proteins. In: Kuettner KE, Goldberg VM, editors. Osteoarthritic disorders. Rosemont, IL: American Academy of Orthopaedic Surgeons. p 239-237.
71. Hoffmann A, Czichos S, Kaps C, et al. 2002. The t-box transcription factor brachyury mediates cartilage development in mesenchymal stem cell line C3H10T1/2. J Cell Sci 115:769-781.
72. Honda A, Noguchi N, Takehara H, et al. 1991. Cooperative enhancement of hyaluronic acid synthesis by combined use:of IGF-I and EGF, and inhibition by tyrosine kinase inhibitor genistein, in cultured mesothelial cells from rabbit pericardial cavity. J Cell Sci 98:91-98.
73. Horton WE, Yamada Y, Hassell JR. 1987. Retinoic acid rapidly reduces cartilage matrix synthesis by altering gene transcription in chondrocytes. Dev Biol 123:508-516.
74. Hua Q, Knudson CB, Knudson W. 1993. Internalization of hyaluronan by chondrocytes occurs via receptor-mediated endocytosis. J Cell Sci 106:365-375.
75. Hunziker EB. 1992. Articular cartilage structure in humans and experimental animals. In: Kuettner KE, Schleyerbach R, Peyron JG, Hascall VC, editors. Articular cartilage and osteoarthritis. New York: Raven Press. p 183-199.
76. Isacke CM. 1994. The role of the cytoplasmic domain in regulating CD44 function. J Cell Sci 107:2353-2359.
77. Itano N, Kimata K. 1996. Expression cloning and molecular characterization of has protein, eukaryotic hyaluronansynthase. J Biol Chem 271:9875-9878.
78. Itano N, Sawai T, Yoshida M, et al. 1999. Three isoforms of mammalian hyaluronan synthases have distinct enzymatic properties. J Biol Chem 274: 25085-25092.
79. Jackson DG, Buckley J, Bell JI. 1992. Multiple variants of the human lymphocyte homing receptor CD44 generated by insertions at a single site in the extracellular domain. J Biol Chem 267:4732-4739.
80. Jiang H, Knudson CB, Knudson W. 2001. Antisense inhibition of alternatively spliced CD44 variant in human articular chondrocytes promotes hyaluronaninternalization. Arthritis Rheum 44: 2599-2619.
81. Jiang H, Peterson RS, Wang W, et al. 2002. A requirement for the CD44 cytoplasmic domain for hyaluronan binding, pericellular matrix assembly and receptor mediated endocytosis in COS-7 cells. J Biol Chem 277:10531-10538.
82. Johnson HJ, Rosenberg L, Choi HU, et al. 1997. Characterization of epiphycan, a small proteoglycan with a leucine-rich repeat core protein. J Biol Chem 272:18709-18717.
83. Johnson J, Shinomura T, Eberspaecher H, et al. 1999. Expression and localization of PG-Lb/epiphycan during mouse development. Dev Dyn 216: 499-510.
84. Kalomiris EL, Bourguignon LYW. 1988. Mouse T lymphoma cells contain a transmembrane glycoprotein (gp85) that binds ankyrin. J Cell Biol 106:319-327.
85. Kalomiris EL, Bourguignon LYW. 1989. Lymphoma protein kinase c is associated with the transmembrane glycoprotein, gp85, and may function in gp85-ankyrin binding. J Biol Chem 264: 8113-8119.
86. Kimata K, Yasuteru O, Katsuko T, et al. 1986. A large chondroitin sulfate proteoglycan (PG-M) synthesized before chondrogenesis in the limb bud of chick embryo. J Biol Chem 261:13517-13525.
87. Knudson CB, Toole BP. 1985. Changes in the pericellular matrix during differentiation of limb bud mesoderm. Dev Biol 112:308-318.
88. Knudson CB, Toole BP. 1987. Hyaluronate-cell interactions during differentiation of chick embryo limb mesoderm. Dev Biol 124:82-90.
89. Knudson CB. 1993. Hyaluronan receptor-directed assembly of chondrocyte pericellular matrix. J Cell Biol 120:825-834.
90. Knudson CB, Knudson W. 1993. Hyaluronan-binding proteins in development, tissue homeostasis and disease. FASEB J 7:1233-1241.
91. Knudson CB, Munaim SI, Toole BP. 1995. Ectodermal stimulation of the production of hyaluronan-dependent pericellular matrix by embryonic limb mesodermal cells. Dev Dyn 204:186-191.
92. Knudson CB, Constable CT, Nofal GA. 1997. Changes in phosphorylation and actin-association of CD44 with receptor occupancy on articular chondrocytes. Trans Orthop Res Soc 22:34.
93. Knudson CB. 1998. Hyaluronan in embryonic cell adhesion and matrix assembly. In: Laurent TC, editor. The chemistry, biology and medical applications of hyaluronan and its derivatives. London: Portland Press. p 161-168.
94. Knudson CB, Nofal GA, Pamintuan L, Aguiar DJ. 1999. The chondrocyte pericellular matrix: a model for hyaluronan-mediated cell-matrix interactions. Biochem Soc Trans 27:142-147.
95. Knudson CB, Nofal GA, Chow G, Peterson RS. 2000a. CD44: the link between hyaluronan and the cytoskeleton. In: Kennedy JF, editor. Hyaluronan 2000. Cambridge, UK: Woodhead Publishing. p 331-340.
96. Knudson W, Knudson CB. 1991. Assembly of a chondrocyte-like pericellular matrix on non-chondrogenic cells. J Cell Sci 99:227-235.
97. Knudson W, Bartnik E, Knudson CB. 1993. Assembly of pericellular matrices by cos-7 cells transfected with CD44 homing receptor genes. Proc Natl Acad Sci U S A 90:4003-4007.
98. Knudson W, Aguiar DJ, Hua Q, Knudson CB. 1996. CD44-anchored hyaluronan-rich pericellular matrices: an ultrastructural and biochemical analysis. Exp Cell Res 228:216-228.
99. Knudson W, Casey B, Nishida Y, et al. 2000b. Hyaluronan oligosaccharides perturb cartilage matrix homeostasis and induce chondrocytic chondrolysis. Arthritis Rheum 43:1165-1174.
100. Knudson W, Loeser RF. 2002. CD44 and integrin matrix receptors participate in cartilage homeostasis. Cell Mol Life Sci 59:36-44.
101. Knudson W, Chow G, Knudson CB. 2002. CD44-mediated uptake and degradation of hyaluronan. Matrix Biol 21:15-23.
102. Knutson IR, Iida J, Fields GB, McCarthy JB. 1996. CD44/chondroitin sulfate proteoglycan and alpha 2 beta1 integrin mediate human melanoma cell migration on type iv collagen and invasion of basement membrane. Mol Biol Cell 7:383-396.
103. Kochhar DM, Penner ID, Hickey T. 1984. Retinoic acid enhances the displacement of newly synthesized hyaluronate from cell layer to culture medium during early phases of chondrogenesis. Cell Diff 14:213-221.
104. Kolettas E, Muir HI, Barrett JC, Hardingham TE. 2001. Chondrocyte phenotype and cell survival are regulated by culture conditions and by specific cytokines through the expression of sox-9 transcription factor. Rheumatology 40:1146-1156.
105. Kosher RA, Savage MP, Walker KH. 1981. A gradation of hyaluronate accumulation along the proximodistal axis of the embryonic chick limb bud. J Embryol Exp Morphol 63:85-98.
106. Kosher RA, Walker KH, Ledger PW. 1982. Temporal and spatial distribution of fibronectin during development of the embryonic chick limb bud. Cell Diff 11:217-228.
107. Kresse H, Schonherr E. 2001. Proteoglycans of the extracellular matrix and growth control. J Cell Physiol 189:266-274.
108. Kuettner KE. 1994. Osteoarthritis: cartilage integrity and homeostasis. In: Klippel JH, Dieppe PA, editors. Rheumatology. St. Louis, MO: Mosby Yearbook Europe Limited. p 6.1-6.16.
109. Kulyk WM, Kosher RA. 1987. Temporal and spatial analysis of hyaluronidase activity during development of the embryonic chick limb bud. Dev Biol 120:535-541.
110. Kurima K, Warman ML, Krishnan S, et al. 1998. A member of a family of sulfate-activating enzymes causes murine brachymorphism. Proc Natl Acad Sci U S A 95:8681-8685.
111. Lark MW, Bayne EK, Flanagan J, et al. 1997. Aggrecan degradation in human cartilage. J Clin Invest 100:93-105.
112. Laurent TC, Fraser JRE. 1986. The properties and turnover of hyaluronan. In: Evered D, Whelan J, editors. Functions of the proteoglycans. Ciba Foundation Symposium 124. Chichester, UK: Wiley Press. p 9-29.
113. Laurent TC, Fraser RE. 1992. Hyaluronan. FASEB J 6:2397-2404.
114. Laurent TC, Laurent UB, Fraser JRE. 1995. Functions of hyaluronan. Ann Rheum Dis 54:429-432.
115. Lazaar AL, Pure E. 1995. CD44: a model for regulated adhesion function. Immunologist 3:19-25.
116. Lee GM, Johnstone B, Jacobson K, Caterson B. 1993. The dynamic structure of the pericellular matrix on living cells. J Cell Biol 123:1899-1907.
117. Lee JY, Spicer AP. 2000. Hyaluronan: a multifunctional, megadalton, stealth molecule. Curr Opin Cell Biol 12:581-586.
118. Lesley J, Schulte R, Hyman R. 1990. Binding of hyaluronic acid to lymphoid cell lines is inhibited by monoclonal antibodies against pgp-1. Exp Cell Res 187:224-233.
119. Lesley J, Hyman R. 1998. CD44 structure and function. Front Biosci 3:616-630.
120. Lesley J, English N, Charles C, Hyman R. 2000a. The role of CD44 cytoplasmic domain and transmembrane domains in constitutive and inducible binding. Eur J Immunol 30:245-253.
121. Lesley J, Hascall VC, Tammi M, Hyman R. 2000b. Hyaluronan binding by cell surface CD44. J Biol Chem 275:26967-26975.
122. Lin CQ, Bissell MJ. 1993. Multi-faceted regulation of cell differentiation by extracellular matrix. FASEB J 7:737-743.
123. Loeser RF, Carlson CS, McGee MP. 1995. Expression of beta 1 integrins by cultured articular chondrocytes and in osteoarthritic cartilage. Exp Cell Res 217:248-257.
124. Maleski MP, Knudson CB. 1996a. Hyaluronan meditated aggregation of limb bud mesoderm and mesenchymal condensation during chondrogenesis. Exp Cell Res 225:55-66.
125. Maleski MP, Knudson CB. 1996b. Matrix accumulation and retention in embryonic cartilage and in vitro chondrogenesis. Connect Tissue Res 34:75-86.
126. Maroudas A. 1979. Physicochemical properties of articular cartilage. In: Freeman MAR, editor. Adult articular cartilage. London: Pitman. p 215-290.
127. Martinsen A, Skjak-Braek G, Smidsrod O. 1989. Alginate as immobilization material. I. Correlation between chemical and physical properties of alginate gel beads. Biotechnol Bioeng 33:79-89.
128. McCourt PAG, Ek B, Forsberg N, Gustafson S. 1994. Intercellular adhesion molecule-1 is a cell surface receptor for hyaluronan. J Biol Chem 269: 30081-30084.
129. McDonald JA. 1989. Receptors for extracellular matrix components. Am J Physiol 257:331-337.
130. McGuire PG, Castellot JJ Jr, Orkin RW. 1992. Size dependent hyaluronate degradation by cultured cells. Cell Physiol 133:267-276.
131. McQueeney K, Dealy CN. 2001. Roles of insulin-like growth factor-I (IGF-I) and IGF-1 binding protein-2 (IGFBP2) and -5 (IGFBP5) in developing chick limbs. Growth Hormone IGF Res 11:346-363.
132. Merino R, Rodriguez-Leon J, Macias D, et al. 1999. The bmp antagonist gremlin regulates outgrowth, chondrogenesis and programmed cell death in the developing limb. Development 126:5515-5522.
133. Milstone L, Hough-Monroe L, Kugelman L, et al. 1994. Epican, a heparan/chondroitin sulfate proteoglycan form of CD44, mediates cell-cell adhesion. J Cell Sci 107:3183-3190.
134. Miyake K, Underhill CB, Lesley J, Kincade PW. 1990. Hyaluronate can function as a cell adhesion molecule and CD44 participates in hyaluronate recognition. J Exp Med 172:69-75.
135. Moftah MZ, Downie SA, Bronstein NB, et al. 2002. Ectodermal FGFs induce perinodular inhibition of limb chondrogenesis in vitro and in vivo via FGF receptor 2. Dev Biol 249:270-282.
136. Mok SS, Masuda K, Hauselmann HJ, et al. 1994. Aggrecan synthesized by mature bovine chondrocytes suspended in alginate. J Biol Chem 269:33021-33027.
137. Natowicz MR, Short MP, Wang Y, et al. 1996. Clinical and biochemical manifestations of hyaluronidase deficiency. N Engl J Med 335:1029-1033.
138. Nemec RE, Toole BP, Knudson W. 1987. The cell surface hyaluronate binding sites of invasive human bladder carcinoma cells. Biochem Biophys Res Commun 149:249-257.
139. Nishida Y, Knudson CB, Nietfeld JJ, et al. 1999. Antisense inhibition of hyaluronan synthase-2 in human articular chondrocytes inhibits proteoglycan retention and matrix assembly. J Biol Chem 274:21893-21899.
140. Nishida Y, Knudson CB, Eger W, et al. 2000a. Osteogenic protein-1 stimulates cell-associated matrix assembly by normal human articular chondrocytes. Arthritis Rheum 43:206-214.
141. Nishida Y, Knudson CB, Kuettner KE, Knudson W. 2000b. Osteogenic protein-1 promotes the synthesis and retention of extracellular matrix within bovine articular cartilage and chondrocyte cultures. Osteoarthritis Cartilage 8:127-136.
142. Nofal GA, Knudson CB. 2002. Latrunculin and cytochalasin decrease chondrocyte matrix retention. J Histochem Cytochem 50:1313-1324.
143. Oberlender SA, Tuan RS. 1994. Expression and functional involvement of n-cadherin in embryonic limb chondrogenesis. Development 120:177-187.
144. Ohya T, Kaneko Y. 1970. Novel hyaluronidase from streptomyces. Biochem Biophys Acta 198:607-609.
145. Okamoto I, Kawano Y, Murakami D, et al. 2001. Proteolytic release of CD44 intracellular domain and its role in the CD44 signaling pathway. J Cell Biol 155:755-762.
146. Orkin RW, Knudson W, Toole BP. 1985. Loss of hyaluronate-dependent coat during myoblast fusion. Dev Biol 107:527-530.
147. Pacifici M, Koyama E, Iwamoto M, Gentili C. 2000. Development of articular cartilage: what do we know about it and how may it occur? Connect Tissue Res 41:175-184.
148. Paine-Saunders S, Viviano BL, Zupicich J, et al. 2000. Glypican-3 controls cellular responses to bmp4 in limb patterning and skeletal development. Dev Biol 225:179-187.
149. Paine-Saunders S, Viviano BL, Economides AN, Saunders S. 2002. Heparan sulfate proteoglycans retain noggin at the cell surface: a potential mechanism for shaping bone morphogenetic protein gradients. J Biol Chem 277:2089-2096.
150. Petit B, Masuda K, D'Souza A, et al. 1996. Characterization of cross-linked collagens sythesized by mature articular chondrocytes cultured in alginate beads: comparison of two distinct matrix compartments. Exp Cell Res 225:151-161.
151. Pirok EW III, Li H, Mensch JR, et al. 1997. Structural and functional analysis of the chick chondroitin sulfate proteoglycan (aggrecan) promoter and enhancer region. J Biol Chem 272:11566-11574.
152. Pizette S, Niswander L. 2000. Bmps are required at two steps of limb chondrogenesis: formation of prechondrogenic condensations and their differentiation into chondrocytes. Dev Biol 219:237-249.
153. Podgrabinska S, Braun P, Velasco P, et al. 2002. Molecular characterization of lymphatic endothelial cells. Proc Natl Acad Sci U S A 99:16069-16074.
154. Ponta H, Sherman L, Herrlich P. 2003. CD44: from adhesion molecules to signaling regulators. Nat Rev Mol Cell Biol 4:33-45.
155. Prehm P. 1984. Hyaluronate is synthesized at plasma membranes. Biochem J 220:597-600.
156. Pure E, Camp RL, Peritt D, et al. 1995. Defective phosporylation and hyaluronate binding of CD44 with point mutations in the cytoplasmic domain. J Exp Med 181:55-62.
157. Raja RH, McGary CT, Weigel PH. 1988. Affinity and distribution of surface and intracellular hyaluronic acid receptors in isolated rat liver endothelial cells. J Biol Chem 263:16661-16668.
158. Ramsden CA, Bankier A, Brown TJ, et al. 2000. A new disorder of hyaluronan metabolism associated with generalized folding and thickening of the skin. J Pediatr 136:62-68.
159. Rauch U, Karthikeyan L, Maurel P, et al. 1992. Cloning and primary structure of neurocan, a developmentally regulated, aggregating chondroitin sulfate proteoglycan of brain. J Biol Chem 267: 19536-19547.
160. Recklies A, White C, Melching L, Roughley P. 2001. Differential regulation and expression of hyaluronan synthases in human articular chondrocytes, synovial cells and osteosarcoma cells. Biochem J 354:17-24.
161. Reddi AH. 1998. Role of morphogenetic proteins in skeletal tissue engineering and regeneration. Nat Biotech 16:247-252.
162. Rittenhouse E, Dunn LC, Cookingham J, et al. 1978. Cartilage matrix deficiency (cmd): a new autosomal recessive lethal mutation in the mouse. J Embryol Exp Morphol 43:71-84.
163. Rousche KT, Knudson CB. 2002. Temporal expression of CD44 during embryonic chick limb development and modulation of its expression with retinoic acid. Matrix Biol 21:53-62.
164. Salter DM, Godolphin JL, Gourlay MS, et al. 1996. Analysis of human articular chondrocyte CD44 isoform expression and function in health and disease. Am J Pathol 179:396-402.
165. Salter DM, Hughes DE, Simpson R, Gardner DL. 1992. Integrin expression by human articular chondrocytes. Br J Rheum 31:231-234.
166. Sanz-Ezquerro JJ, Tickle C. 2001. "fingering" the vertebrate limb. Differentiation 69:91-99.
167. Schaller SA, Muneoka K. 2001. Inhibition of polarizing activity in the anterior limb bud is regulated by extracellular factors. Dev Biol 240:443-456.
168. Schenk RK, Eggli PS, Hunziker EB. 1986. Articular cartilage morphology. In: Kuettner KE, Schleyerbach R, Hascall VC, editors. Articular cartilage biochemistry. New York: Raven Press. p 3-22.
169. Schmid TM, Linsenmayer TF. 1985. Immunohistochemical localization of short chain cartilage collagen (type X) in avian tissues. J Cell Biol 100:598-605.
170. Schwartz NB, Pirok EW III, Mensch JR Jr, Domowicz MS. 1999. Domain organization, genomic structure, evolution, and regulation of expression of the aggrecan gene family. Prog Nucleic Acid Res Mol Biol 62:177-225.
171. Screaton GR, Bell MV, Jackson DG, et al. 1992. Genomic structure of DNA encoding the lymphocyte homing receptor CD44 reveals at least 12 alternatively spliced exons. Proc Natl Acad Sci U S A 89:12160-12164.
172. Seghatoleslami MR, Kosher RA. 1996. Inhibition of in vitro limb cartilage differentiation by syndecan-3 antibodies. Dev Dyn 207:114-119.
173. Sekiya I, Tsuji K, Koopman P, et al. 2000. Sox9 enhances aggrecan gene promoter/enhancer activity and is upregulated by retinoic acid in a cartilage-derived cell line, TC6. J Biol Chem 275:10738-10744.
174. Shamblott MJ, Bugg EM, Lawler AM, Gearhart JD. 2002. Craniofacial abnormalities resulting from targeted disruption of the murine Sim2 gene. Dev Dyn 224:373-380.
175. Sheikh H, Uff CR, Peck D, Isacke CM. 1995. Modulation of hyaluronan receptor, CD44, by its transmembrane and cytoplasmic domains. Biochem Soc Trans 23:831-835.
176. Sherman L, Wainwright D, Ponta H, Herrlich P. 1998. A splice variant of CD44 expressed in the apical ectodermal ridge presents fibroblast growth factors to limb mesenchymme and is required for limb outgrowth. Genes Dev 12:1058-1071.
177. Shimazu A, Nah HD, Kirsch T, et al. 1996. Syndecan-3 and the control of chondrocyte proliferation during endochondral ossification. Exp Cell Res 229:126-136.
178. Shinomura T, Nishida Y, Kimata K. 1992. A large chondroitin sulfate proteoglycan (pg-m) and cartilage differentiation. In: Kuettner KE, Schleyerbach R, Peyron JG, Hascall VC, editors. Articular cartilage and disease. New York: Raven Press. p 35-44.
179. Singley CT, Solursh M. 1981. The spatial distribution of hyaluronic acid and mesenchymal condensation in the embryonic chick wing. Dev Biol 84: 102-120.
180. Smedsrod B, Pertoft H, Eriksson S, et al. 1984. Studies in vitro on the uptake and degradation of sodium hyaluronate in rat liver endothelial cells. Biochem J 223:617-626.
181. Solursh M, Ahrens PB, Reiter RS. 1978. A tissue culture analysis of the steps in limb chondrogenesis. In Vitro 14: 51-61.
182. Solursh M, Reiter S. 1980. Evidence for histogenic interactions during in vitro limb chondrogenesis. Dev Biol 78:141-150.
183. Solursh M. 1984a. Cell and matrix interactions during limb chondrogenesis in vitro. In: Trelstad RL, editor. The role of extracellular matrix in development. New York: A.R. Liss. p 277-303.
184. Solursh M. 1984b. Ectoderm as a determinant of early tissue pattern in the limb bud. Cell Diff 15:17-24.
185. Spicer AP, Augustine ML, McDonald JA. 1996. Molecular cloning and characterization of a putative mouse hyaluronan synthase. J Biol Chem 271: 23400-23406.
186. Spicer AP, Seldin MF, Olsen AS, et al. 1997. Chromosomal localization of the human and mouse hyaluronan synthase genes. Genomics 41:493-497.
187. Sretavan DW, Feng L, Pure E, Reichardt LF. 1994. Embryonic neurons of the developing optic chiasm express L1 and CD44, cell surface molecules with opposing effects on retinal axon growth. Neuron 12:957-975.
188. Sugiyama Y, Shimada A, Sayo T, et al. 1998. J Invest Dermatol 110:16-121.
189. SundarRaj N, Fite D, Ledbetter S, et al. 1995. Perlecan is a component of cartilage matrix and promotes chondrocyte attachment. J Cell Sci 108:2663-2672.
190. Sy MS, Guo YJ, Stamenkovic I. 1991. Distinct effects of two CD44 isoforms on tumor growth in vivo. J Exp Med 174:859-866.
191. Taher TEI, Smit L, Griffioen AW, et al. 1996. Signaling through CD44 is mediated by tryrosine kinases: asssociation with p56lck in T lymphocytes. J Biol Chem 271:2863-2867.
192. Tavella S, Bellese G, Castagnola P, et al. 1997. Regulated expression of fibronectin, laminin and related integrin receptors during the early chondrocyte differentiation. J Cell Sci 110: 2261-2270.
193. Thonar EJ-MA, Bjornsson S, Kuettner KE. 1986. Age-related changes in cartilage proteoglycans. In: Kuettner KE, Schleyerbach R, Peyron JG, Hascall VC, editors. Articular cartilage biochemistry. New York: Raven Press. p 273-291.
194. Thorogood PV, Hinchliffe JR. 1975. An analysis of the condensation process during chondrogenesis in the embryonic chick hind limb. J Embryol Exp Morphol 33:581-606.
195. Tomasek JJ, Mazurkiewicz JE, Newman SA. 1982. Nonuniform distribution of fibronectin during avian limb development. Dev Biol 90:118-126.
196. Toole BP. 1972. Hyaluronate turnover during chondrogenesis in the developing chick limb and axial skeleton. Dev Biol 29:321-329.
197. Toole BP. 1981. Glycosaminoglycans in morphogenesis. In: Hay ED, editor. Cell biology of the extracellular matrix. New York: Plenum Press. p 259-294.
198. Toole BP. 1990. Hyaluronan and its binding proteins, the hyaladherins. Curr Opin Cell Biol 2:839-844.
199. Toole BP. 1991. Proteoglycans and hyaluronan in morphogenesis and differentiation. In: Hay ED, editor. Cell biology of extracellular matrix. New York: Plenum Press. p 305-339.
200. Toole BP, Banerjee SD, Turner R, et al. 1992. Hyaluronan-cell interactions in limb development. In: Hinchliffe J, Hurle J, Summerbell D, editors. Developmental patterning of the vertebrate limb. Plenum Press. p 1-9.
201. Toole BP, Goldberg RL, Chi-Rosso G, et al. 1984. Hyaluronate-cell interactions. In: Trelstad RL, editor. The role of the extracellular matrix in development. New York: A.R. Liss. p 43-46.
202. Toole BP, Munian SI, Wells S, Knudson CB. 1989. Hyaluronate-cell interactions and growth factor regulation of hyaluronate synthesis during limb development. In: Evered D, Whelan J, editors. The biology of hyaluronan. Ciba Foundation Symposium 143. Chichester, UK: John Wiley and Sons. p 138-149.
203. Triggs-Raine B, Salo TJ, Zhang H, et al. 1999. Mutations in HYAL1, a member of a tandemly distributed multigene family encoding disparate hyaluronidase activities, cause a newly described lysosomal disorder, mucopolysaccharidosis IX. Proc Natl Acad Sci U S A 96:6296-6300.
204. Tsukita S, Oishi K, Sato N, et al. 1994. ERM family members as molecular linkers between the cell surface glycoprotein CD44 and actin-based cytoskeletons. J Cell Biol 126:391-401.
205. Tucker AS, Al Khamis A, Ferguson CA, et al. 1999. Conserved regulation of mesenchymal gene expression by FGF-8 in face and limb development. Development 126:221-228.
206. Tuhkanen A, Tammi M, Tammi R. 1997. CD44 substituted with heparan sulfate and endo-beta-galactosidase-sensitive oligosaccharides: a major proteoglycan in adult human epidermis. J Invest Dermatol 109:213-218.
207. Turley EA, Belch AJ, Poppema S, Pilarski LM. 1993. Expression and function of a receptor for hyaluronan-mediated motility on normal and malignant B lymphocytes. Blood 81:446-453.
208. Turley EA, Noble PW, Bourguignon LY. 2002. Signaling properties of hyaluronan receptors. J Biol Chem 277:4589-4592.
209. Underhill CB, Dorfman A. 1978. The role of hyaluronic acid in intercellular adhesion of cultured mouse cells. Exp Cell Res 117:155-164.
210. Underhill CB, Toole BP. 1979. Binding of hyaluronate to the surface of cultured cells. J Cell Biol 82:475-484.
211. Underhill CB, Toole BP. 1980. Physical characteristics of hyaluronate binding to the surface of simian virus 40-transformed 3T3 cells. J Biol Chem 255: 4544-4549.
212. Underhill CB, Toole BP. 1981. Receptors for hyaluronate on the surface of parent and virus-transformed cell lines. Exp Cell Res 131:419-423.
213. Underhill CB. 1982. Interaction of hyaluronate with the surface of simian virus 40-transformed 3T3 cells: aggregation and binding studies. J Cell Sci 56:177-189.
214. Underhill CB, Toole BP. 1982. Transformation-dependent loss of the hyaluronate-containing coats of cultured cells. J Cell Physiol 110:123-128.
215. Underhill CB, Chi-Rosso G, Toole BP. 1983. Effect of detergent solubilization on the hyaluronanate-binding protein from membranes of simian virus 40-transformed 3T3 cells. J Biol Chem 258:8086-8091.
216. Underhill CB, Green SJ, Cologlio PM, Tarone G. 1987. The hyaluronate receptor is identical to a glycoprotein of 85,000mr (gp 85) as shown by a monoclonal antibody that interferes with binding activity. J Biol Chem 262: 13142-13146.
217. Underhill CB. 1992. CD44: the hyaluronan receptor. J. Cell Sci 103:293-298.
218. Underhill CB, Nguyen HA, Shizari M, Culty M. 1993. CD44 positive macrophages take up hyaluronan during lung development. Dev Biol 155:324-336.
219. Walther BT, Ohman R, Roseman S. 1973. A quantitative assay for intercellular adhesion. Proc Natl Acad Sci U S A 70:1569-1573.
220. Watanabe H, Nakata K, Kimata K, et al. 1997. Dwarfism and age-associated spinal degeneration of heterozygote cmd mice defective in aggrecan. Proc Natl Acad Sci U S A 94:6943-6947.
221. Watanabe H, Yamada Y, Kimata K. 1998. Roles of aggrecan, a large chondroitin sulfate proteoglycan, in cartilage structure and function. J Biochem 124:687-693.
222. Weber B, Rosel M, Arch R, et al. 1996. Transient expression of CD44 variant isoforms in the ontogeny of the rat: ectoderm-, endoderm- and mesoderm-derived cells express different exon combinations. Differentiation 60: 17-29.
223. Weigel PH, Hascall VC, Tammi M. 1997. Hyaluronansynthases. J Biol Chem 272: 13997-14000.
224. Weigel J, Raymond R, McGary C, et al. A blocking antibody to the hyaluronan (HA) receptor for endocytosis (HARE) inhibits HA clearance by perfused liver. J Biol Chem 278:9808-9812.
225. Weston AD, Rosen V, Chandraratna RA, Underhill TM. 2000. Regulation of skeletal progenitor differentiation by the bmp and retinoid signaling pathways. J Cell Biol 148:679-690.
226. Wheatley SC, Isacke CM, Crossley PH. 1993. Restricted expression of the hyaluronan receptor, CD44, during postimplantation mouse embryogenesis suggests key roles in tissue formation and patterning. Development 119:295-306.
227. Widelitz RB, Jiang T, Murray BA, Chuong C. 1993. Adhesion molecules in skeletogenesis. II. Neural cell adhesion molecules mediate precartilaginous mesenchymal condensations and enhance chondrogenesis. J Cell Physiol 156:399-411.
228. Woods VL, Schreck PJ, Gesink DS, et al. 1994. Integrin expression by human articular chondrocytes. Arthritis Rheum 37:537-544.
229. Yamada H, Watanabe K, Shimonaka M, Yamaguchi Y. 1994. Molecular cloning of brevican, a novel brain proteoglycan of the aggrecan/versican family. J Biol Chem 269:10119-10126.
230. Yamada KM. 1997. Integrin signaling. Matrix Biol 16:137-141.
231. Yannariello-Brown J, Weigel PH. 1992. Detergent solubilization of the endocytic Ca2+-independent hyaluronan receptor from rat liver endothelial cells and separation from a Ca2+-dependent hyaluronan-binding activity. Biochemistry 31:576-584.
232. Yu Q, Grammatikakis N, Toole BP. 1996. Expression of multiple CD44 isoforms in the apical ectodermal ridge of the embryonic mouse limb. Dev Dyn 207: 204-214.
233. Zebrower M, Kieras FJ, Brown WT. 1986. Urinary hyaluronic acid elevation in Hutchinson-Gilford progeria syndrome. Mech Ageing Dev 35:39-46.
234. Zhang Y, Cao L, Kiani CG, et al. 1998. The G3 domain of versican inhibits mesenchymal chondrogenesis via the epidermal growth factor-like motifs. J Biol Chem 273:33054-33063.
235. Zhou B, Weigel J, Saxena A, Weigel P. 2002. Molecular cloning and functional expression of the rat 175-kDa hyaluronan receptor for endocytosis. Mol Biol Cell 13:2853-2868.
236. Zimmermann DR, Ruoslahti E. 1989. Multiple domains of the large fibroblast proteoglycan, versican. EMBO J 8:2975-2981.