D. Howell, J. Pelletier, W. Mccarty, and . Koopman, Etiopathogenesis of osteoarthritis Arthritis and Allied Conditions, 1993.

H. Guenther, H. Guenther, E. Froesch, and H. Fleisch, Effect of insulin-like growth factor on collagen and glycosaminoglycan synthesis by rabbit articular chondrocytes in culture, Experientia, vol.41, issue.8
DOI : 10.1042/bj1960293

D. Mcquillan, C. Handley, M. Campbell, S. Bolis, V. Milway et al., Stimulation of proteoglycan biosynthesis by serum and insulin-like growth factor-I in cultured bovine articular cartilage, Biochemical Journal, vol.240, issue.2, pp.423-430, 1986.
DOI : 10.1042/bj2400423

G. Tesch, C. Handley, H. Cornell, and A. Herington, Effects of free and bound insulin-like growth factors on proteoglycan metabolism in articular cartilage explants, Journal of Orthopaedic Research, vol.68, issue.1, pp.14-22, 1992.
DOI : 10.1016/B978-0-12-655102-0.50008-4

D. Willis, . Jr, and J. Liberti, Post-receptor actions of somatomedin on chondrocyte collagen biosynthesis, Biochimica et Biophysica Acta (BBA) - Molecular Cell Research, vol.844, issue.1, pp.72-80, 1985.
DOI : 10.1016/0167-4889(85)90236-8

J. Tyler, Insulin-like growth factor 1 can decrease degradation and promote synthesis of proteoglycan in cartilage exposed to cytokines, Biochemical Journal, vol.260, issue.2, pp.543-548, 1989.
DOI : 10.1042/bj2600543

R. Rogachefsky, D. Dean, D. Howell, and R. Altman, Treatment of canine osteoarthritis with insulin-like growth factor-1 (IGF-1) and sodium pentosan polysulfate, Osteoarthritis and Cartilage, vol.1, issue.2, p.114, 1993.
DOI : 10.1016/S1063-4584(05)80025-1

S. Dore, J. Pelletier, J. Dibattista, G. Tardif, P. Brazeau et al., Human Osteoarthritic Chondrocytes Possess an Increased Number of Insulin-Like Growth Factor 1 Binding Sites but are Unresponsive to its Stimulation, Arthritis & Rheumatism, vol.127, issue.2, pp.253-263, 1994.
DOI : 10.1042/bj2600543

J. Schalkwijk, L. Joosten, W. Van-den-berg, and L. Van-de-putte, Chondrocyte nonresponsiveness to insulin-like growth factor 1 in experimental arthritis, Arthritis Rheum, vol.32, pp.894-900, 1989.

J. Martin, M. Ballesteros, and R. Baxter, Insulin-like growth factor-I (IGF-I) and transforming growth factor-beta 1 release IGF-binding protein-3 from human fibroblasts by different mechanisms., Endocrinology, vol.131, issue.4, pp.1703-1710, 1992.
DOI : 10.1210/endo.131.4.1382959

C. Rouyer-fessard, S. Gammeltoft, and M. Laburthe, Expression of two types of receptor for insulinlike growth factors in human colonic epithelium, Gastroenterology, vol.98, issue.3, pp.703-707, 1990.
DOI : 10.1016/0016-5085(90)90291-8

D. Morgan, J. Edman, D. Standring, V. Fried, M. Smith et al., Insulin-like growth factor II receptor as a multifunctional binding protein, Nature, vol.329, issue.6137, pp.301-307, 1987.
DOI : 10.1038/329301a0

D. Clemmons, IGF binding proteins: regulation of cellular actions, Growth Regul, vol.2, pp.80-87, 1992.

R. Altman, E. Asch, D. Bloch, G. Bole, D. Borenstein et al., Development of criteria for the classification and reporting of osteoarthritis: Classification of osteoarthritis of the knee, Arthritis & Rheumatism, vol.1, issue.8, pp.39-1049, 1986.
DOI : 10.1002/art.1780290816

B. Swodoba, R. Holmdahl, H. Stoss, . Van-der, and . Mark, Cellular heterogeneity in cultured human chondrocytes identified by antibodies specific for a2 (Xl) collagen chains, J Cell Biol, vol.109, pp.1363-1369, 1989.

K. Elima and E. Vuorio, Expression of mRNAs for collagens and other matrix components in dedifferentiating and redifferentiating human chondrocytes in culture, FEBS Letters, vol.242, issue.2, pp.195-198, 1989.
DOI : 10.1126/science.3201241

P. Smith, R. Krohn, G. Hermanson, A. Mallia, F. Gartner et al., Measurement of protein using bicinchoninic acid, Analytical Biochemistry, vol.150, issue.1, pp.76-85, 1985.
DOI : 10.1016/0003-2697(85)90442-7

U. Laemmli, Cleavage of Structural Proteins during the Assembly of the Head of Bacteriophage T4, Nature, vol.244, issue.5259, pp.680-685979, 1970.
DOI : 10.1101/SQB.1963.028.01.053

H. Towbin, T. Staehelin, and J. Gordon, Electrophoretic transfer of proteins from polyacrylamide gels to nitrocellulose sheets: procedure and some applications., Proceedings of the National Academy of Sciences, vol.76, issue.9, pp.4350-4354, 1979.
DOI : 10.1073/pnas.76.9.4350

W. Burnette, ???Western Blotting???: Electrophoretic transfer of proteins from sodium dodecyl sulfate-polyacrylamide gels to unmodified nitrocellulose and radiographic detection with antibody and radioiodinated protein A, Analytical Biochemistry, vol.112, issue.2, pp.195-203, 1981.
DOI : 10.1016/0003-2697(81)90281-5

J. Sambrook, E. Fritsch, and T. Maniates, Molecular Cloning: A Laboratory Manual. Cold Spring Harbor, 1989.

J. Fernandes, J. Martel-pelletier, I. Otterness, A. Lopez-anaya, F. Mineau et al., Effects of tenidap on canine experimental osteoarthritis i. morphologic and metalloprotease analysis, Arthritis & Rheumatism, vol.19, issue.9, pp.1290-1303, 1995.
DOI : 10.1042/bj2480265

A. Ullrich, A. Gray, A. Tam, T. Yang-feng, M. Tsubokawa et al., Insulin-like growth factor 1 receptor primaly structure: comparison with insulin receptor suggests structural determinants that define functional specificity, EMBO J, vol.5, pp.2503-2512, 1986.

G. Gilliland, S. Ferrin, K. Blanchard, and H. Bunn, Analysis of cytokine mRNA and DNA: detection and quantitation by competitive polymerase chain reaction., Proceedings of the National Academy of Sciences, vol.87, issue.7, pp.2725-2729, 1990.
DOI : 10.1073/pnas.87.7.2725

A. Wang, M. Doyle, and D. Mark, Quantitation of mRNA by the polymerase chain reaction., Proceedings of the National Academy of Sciences, vol.86, issue.24, pp.9717-9721, 1989.
DOI : 10.1073/pnas.86.24.9717

A. Oshima, C. Nolan, J. Kyle, J. Grubb, and W. Sly, The human cation-independent mannose 6-phosphate receptor: cloning and sequence of the full-length cDNA and expression of functional receptor in COS cells, J Biol Chem, vol.263, pp.2553-2562, 1988.

C. Binkert, J. Landwehr, J. Mary, J. Schwander, and G. Heinrich, Cloning, sequence analysis and expression of a cDNA encoding a novel insulin-like growth factor binding protein

W. Wood, G. Cachianes, W. Henzel, G. Winslow, S. Spencer et al., Cloning and Expression of the Growth Hormone-Dependent Insulin-Like Growth Factor-Binding Protein, Molecular Endocrinology, vol.2, issue.12, pp.1176-1185, 1988.
DOI : 10.1210/mend-2-12-1176

S. Shimasaki, F. Uchiyama, N. Shimonaka, and N. Ling, Molecular cloning of the cDNAs encoding a novel insulin-like growth factorbinding protein from rat and human, Mol, vol.4, pp.1451-1458, 1990.

S. Dore, T. Abribat, N. Rousseau, P. Brazeau, G. Tardif et al., Increased insulin-like growth factor 1 production by human osteoarthritic chondrocytes is not dependent on growth hormone action, Arthritis & Rheumatism, vol.74, issue.3, pp.413-419, 1995.
DOI : 10.1042/bj0620315

J. Middleton and J. Tyler, Upregulation of insulin-like growth factor I gene expression in the lesions of osteoarthritic human articular cartilage., Annals of the Rheumatic Diseases, vol.51, issue.4, pp.440-447, 1992.
DOI : 10.1136/ard.51.4.440

G. Condorelli, R. Bueno, and R. Smith, Two alternatively spliced forms of the human insulin-like growth factor 1 receptor have distinct biological activities and internalization kinetics, J Biol Chem, pp.2693510-8516, 1994.

B. Bhaumick and R. Bala, Differential effects of insulin-like growth factors I and II on growth, differentiation and glucoregulation in differentiating chondrocyte cells in culture, European Journal of Endocrinology, vol.125, issue.2, pp.201-211, 1991.
DOI : 10.1530/acta.0.1250201

J. Zapf, C. Schmid, H. Guler, M. Waldvogel, C. Hauri et al., Regulation of binding proteins for insulin-like growth factors (IGF) in humans. Increased expression of IGF binding protein 2 during IGF I treatment of healthy adults and in patients with extrapancreatic tumor hypoglycemia., Journal of Clinical Investigation, vol.86, issue.3, pp.952-961, 1990.
DOI : 10.1172/JCI114797

B. Bhaumick, Insulin-like growth factor (IGF) binding proteins and insulin-like growth factor secretion by cultured chondrocyte cells: identification, characterization and ontogeny during cell differentiation, Regulatory Peptides, vol.48, issue.1-2, pp.113-122, 1993.
DOI : 10.1016/0167-0115(93)90340-E

R. Olney, R. Smith, Y. Kee, and D. Wilson, Production and hormonal regulation of insulin-like growth factor binding proteins in bovine chondrocytes., Endocrinology, vol.133, issue.2, 1993.
DOI : 10.1210/endo.133.2.7688290

B. Froger-gaillard, P. Hossenlopp, A. M. Binoux, and M. , Production of Insulin-Like Growth Factors and Their Binding Proteins by Rabbit Articular Chondrocytes: Relationships with Cell Multiplication*, Endocrinology, vol.124, issue.5, pp.1242365-2372, 1989.
DOI : 10.1210/endo-124-5-2365

D. Latour, S. Mohan, T. Linkhart, D. Baylink, and D. Strong, Inhibitory Insulin-Like Growth Factor-Binding Protein: Cloning, Complete Sequence, and Physiological Regulation, Molecular Endocrinology, vol.4, issue.12, pp.1806-1814, 1990.
DOI : 10.1210/mend-4-12-1806

G. Ceda, P. Fielder, W. Henzel, A. Louie, S. Donovan et al., Differential Effects of Insulin-like Growth Factor (IGF)- I and IGF-II on the Expression of IGF Binding Proteins (IGFBPs) in a Rat Neuroblastoma Cell Line: Isolation and Characterization of Two Forms of IGFBP-4*, Endocrinology, vol.128, issue.6, pp.2815-2824, 1991.
DOI : 10.1210/endo-128-6-2815

P. Cheung, E. Smith, S. Shimasaki, N. Ling, and S. Chernausek, Characterization of an Insulin-Like Growth Factor Binding Protein (IGFBP-4) Produced by the B104 Rat Neuronal Cell Line: Chemical and Biological Properties and Differential Synthesis by Sublines*, Endocrinology, vol.129, issue.2, pp.1006-1015, 1991.
DOI : 10.1210/endo-129-2-1006

C. Camacho-hubner, W. Busby, . Jr, R. Mccusker, G. Wright et al., Identification of the forms of insulin-like growth factor-binding proteins produced by human fibroblasts and the mechanisms that regulate their secretion, J Biol Chem, vol.267, pp.11949-11956, 1992.

L. Bach, N. Thotakura, and M. Rechler, Human insulin-like growth factor binding protein-6 is O-glycosylated, Biochemical and Biophysical Research Communications, vol.186, issue.1, pp.59-62, 1993.
DOI : 10.1016/S0006-291X(05)80807-1

S. Amarnani, H. Merriman, T. Linkhart, D. Baylink, and S. Mohan, Autocrine regulators of MC3T3-E1 cell proliferation, Journal of Bone and Mineral Research, vol.255, issue.2, pp.157-165, 1993.
DOI : 10.1016/0304-4165(86)90169-8

E. Wang, J. Wang, E. Chin, J. Zhou, and C. Bondy, Cellular patterns of insulin-like growth factor system gene expression in murine chondrogenesis and osteogenesis., Endocrinology, vol.136, issue.6, pp.2741-2751, 1995.
DOI : 10.1210/endo.136.6.7750499

T. Nam, W. Busby, . Jr, and D. Clemmons, Human fibroblasts secrete a serine protease that cleaves insulin-like growth factor-binding protein-5., Endocrinology, vol.135, issue.4, pp.1385-1391, 1994.
DOI : 10.1210/endo.135.4.7523096

C. Tavera, T. Abribat, P. Reboul, S. Dore, P. Brazeau et al., IGF and IGF-binding protein system in the synovial fluid of osteoarthritic and rheumatoid arthritic patients, Osteoarthritis and Cartilage, vol.4, issue.4
DOI : 10.1016/S1063-4584(05)80104-9

URL : https://hal.archives-ouvertes.fr/hal-01705669

C. Conover, Glycosylation of Insulin-Like Growth Factor Binding Protein-3 (IGFBP-3) is not Required for Potentiation IGF-I Action: Evidence for Processing of Cell-Bound IGFBP-3*, Endocrinology, vol.129, issue.6, pp.3259-3268, 1991.
DOI : 10.1210/endo-129-6-3259

M. Ross, G. Francis, L. Szabo, J. Wallace, and F. Ballard, Insulin-like growth factor (1GF)-binding proteins inhibit the biological activities of IGF-1 and IGF-2 but not des-(1-3)-IGF-l, Biochem J

P. Schiltz, S. Mohan, and D. Baylink, Insulin-like growth factor binding protein-4 inhibits both basal and IGF-mediated chick pelvic cartilage growth in vitro, Journal of Bone and Mineral Research, vol.128, issue.3, pp.391-396, 1993.
DOI : 10.1016/0304-4165(88)90127-4

D. Mellow, J. Baxter, and R. , Growth hormone-dependent insulinlike growth factor (IGF) binding protein both inhibits and potentiates IGF-1-stimulated DNA synthesis in human skin fibroblasts, Biochem Biophys Res Commun, pp.156399-204, 1988.

C. Blat, J. Delbe, J. Villaudy, G. Chatelain, and A. Golde, Hare1 L Inhibitoly diffusible factor 45 bifunctional activity: as a cell growth inhibitor and as an insulin-like growth factor I-binding protein, J Biol Chem, pp.26412449-12454, 1989.

W. Blum, E. Jenne, F. Reppin, K. Kietzmann, M. Ranke et al., Insulin-Like Growth Factor I (IGF-I)-Binding Protein Complex Is a Better Mitogen than Free IGF-I*, Endocrinology, vol.125, issue.2
DOI : 10.1210/endo-125-2-766

T. Tsukazaki, T. Usa, T. Matsumoto, H. Enomoto, A. Ohtsuru et al., Effect of Transforming Growth Factor-?? on the Insulin-like Growth Factor-I Autocrine/Paracrine Axis in Cultured Rat Articular Chondrocytes, Experimental Cell Research, vol.215, issue.1, pp.9-16, 1994.
DOI : 10.1006/excr.1994.1307

L. Bale and C. Conover, Regulation of insulin-like growth factor binding protein-3 messenger ribonucleic acid expression by insulin-like growth factor I, Endocrinology, vol.131, pp.608-614, 1992.

Y. Yang, P. Pioli, G. Fiorelli, M. Brandi, and M. Rechler, Cyclic adenosine monophosphate stimulates insulin-like growth factor binding protein-4 and its messenger ribonucleic acid in a clonal endothelial cell line., Endocrinology, vol.133, issue.1, pp.343-351, 1993.
DOI : 10.1210/endo.133.1.7686482

T. Matsumoto, T. Tsukazaki, H. Enomoto, K. Iwasaki, and S. Yamashita, Effects of interleukin-1 beta on insulin-like growth factor-I autocrine/paracrine axis in cultured rat articular chondrocytes., Annals of the Rheumatic Diseases, vol.53, issue.2, pp.128-133, 1994.
DOI : 10.1136/ard.53.2.128

S. Scharla, D. Strong, S. Mohan, T. Chevalley, and T. Linkhart, Effect of tumor necrosis factor-?? on the expression of insulin-like growth factor I and insulin-like growth factor binding protein 4 in mouse osteoblasts, European Journal of Endocrinology, vol.131, issue.3, pp.293-301, 1994.
DOI : 10.1530/eje.0.1310293