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L. Barbour, S. Kick, J. Steiner, M. Loverde, L. Heddleston et al., A prospective study of heparin-induced osteoporosis in pregnancy using bone densitometry, American Journal of Obstetrics and Gynecology, vol.170, issue.3, pp.862-869, 1994.
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G. Bernas, Angiotherapeutics from natural products: from bench to clinics?, Clin Hemorheol Microcirc, vol.29, pp.199-203, 2003.

M. Bernfield, M. Gotte, P. Park, O. Reizes, M. Fitzgerald et al., Functions of Cell Surface Heparan Sulfate Proteoglycans, Annual Review of Biochemistry, vol.68, issue.1, pp.729-777, 1999.
DOI : 10.1146/annurev.biochem.68.1.729

S. Colliec-jouault, L. Chevolot, D. Helley, J. Ratiskol, A. Bros et al., Characterization, chemical modifications and in vitro anticoagulant properties of an exopolysaccharide produced by Alteromonas infernus, Biochimica et Biophysica Acta (BBA) - General Subjects, vol.1528, issue.2-3, pp.141-151, 2001.
DOI : 10.1016/S0304-4165(01)00185-4

M. Downey, L. Holliday, J. Aguirre, and T. Wronski, In vitro and in vivo evidence for stimulation of bone resorption by an EP4 receptor agonist and basic fibroblast growth factor: Implications for their efficacy as bone anabolic agents, Bone, vol.44, issue.2, pp.266-274, 2009.
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V. Eswarakumar, I. Lax, and J. Schlessinger, Cellular signaling by fibroblast growth factor receptors, Cytokine & Growth Factor Reviews, vol.16, issue.2, pp.139-149, 2005.
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G. Franz and A. S. , Structure-activity relationship of antithrombotic polysaccharide derivatives, International Journal of Biological Macromolecules, vol.17, issue.6, pp.311-314, 1995.
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J. Gallagher, Multiprotein signalling complexes: regional assembly on heparan sulphate, Biochemical Society Transactions, vol.34, issue.3, pp.438-441, 2006.
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J. Guezennec, P. Pignet, Y. Lijour, E. Gentric, J. Ratiskol et al., Sulphation and depolymerization of a bacterial exopolysaccharide from hydrothermal origin. Carbohydr Polymers, pp.19-24, 1998.

J. Guezennec, Deep-sea hydrothermal vents: A new source of innovative bacterial exopolysaccharides of biotechnological interest?, Journal of Industrial Microbiology and Biotechnology, vol.29, issue.4, pp.204-208, 2002.
DOI : 10.1038/sj.jim.7000298

L. Haupt, S. Murali, F. Mun, N. Teplyuk, L. Mei et al., The heparan sulfate proteoglycan (HSPG) glypican-3 mediates commitment of MC3T3-E1 cells toward osteogenesis, Journal of Cellular Physiology, vol.281, issue.3, pp.780-791, 2009.
DOI : 10.1002/jcp.21825

A. Irie, M. Takami, H. Kubo, N. Sekino-suzuki, K. Kasahara et al., Heparin enhances osteoclastic bone resorption by inhibiting osteoprotegerin activity, Bone, vol.41, issue.2, pp.165-174, 2007.
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R. Jackson, S. Murali, A. Van-wijnen, G. Stein, V. Nurcombe et al., Heparan sulfate regulates the anabolic activity of MC3T3-E1 preosteoblast cells by induction of Runx2, Journal of Cellular Physiology, vol.19, issue.1, pp.38-50, 2007.
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G. Karsenty, Minireview: Transcriptional Control of Osteoblast Differentiation, Endocrinology, vol.142, issue.7, pp.2731-2733, 2001.
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A. Santos, G. Van, and A. Itie, OPGL is a key regulator of osteoclastogenesis, lymphocyte development and lymph-node organogenesis, Nature, vol.397, pp.315-323, 1999.

V. Kram, E. Zcharia, O. Yacoby-zeevi, S. Metzger, T. Chajek-shaul et al., Heparanase is expressed in osteoblastic cells and stimulates bone formation and bone mass, Journal of Cellular Physiology, vol.16, issue.3, pp.784-792, 2006.
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A. Kumarasuriyar, I. Lee, V. Nurcombe, and S. Cool, De-sulfation of MG-63 cell glycosaminoglycans delays in vitro osteogenesis, up-regulates cholesterol synthesis and disrupts cell cycle and the actin cytoskeleton, Journal of Cellular Physiology, vol.284, issue.3, pp.572-583, 2009.
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K. Tat, S. Padrines, M. Theoleyre, S. Heymann, D. Fortun et al., IL-6, RANKL, TNF- alpha/IL-1: interrelations in bone resorption pathophysiology, Cytokine Growth Factor Rev, vol.15, pp.49-60, 2004.

K. Tat, S. Padrines, M. Théoleyre, S. Couillaud-battaglia, S. Heymann et al., OPG/membranous???RANKL complex is internalized via the clathrin pathway before a lysosomal and a proteasomal degradation, Bone, vol.39, issue.4, pp.706-715, 2006.
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F. Lamoureux, M. Baud-'huin, L. Duplomb, D. Heymann, and F. Redini, Proteoglycans: key partners in bone cell biology, BioEssays, vol.53, issue.8, pp.758-771, 2007.
DOI : 10.1002/bies.20612

F. Lamoureux, G. Picarda, L. Garrigue-antar, M. Baud-'huin, V. Trichet et al., Glycosaminoglycans as Potential Regulators of Osteoprotegerin Therapeutic Activity in Osteosarcoma, Cancer Research, vol.69, issue.2, pp.526-536, 2009.
DOI : 10.1158/0008-5472.CAN-08-2648

L. Ling, C. Dombrowski, K. Foong, L. Haupt, G. Stein et al., Synergism between Wnt3a and Heparin Enhances Osteogenesis via a Phosphoinositide 3-Kinase/Akt/RUNX2 Pathway, Journal of Biological Chemistry, vol.285, issue.34, 2010.
DOI : 10.1074/jbc.M110.122069

K. Manton, M. Sadasivam, S. Cool, and V. Nurcombe, Bone-specific heparan sulfates induce osteoblast growth arrest and downregulation of retinoblastoma protein, Journal of Cellular Physiology, vol.322, issue.1, pp.219-229, 2006.
DOI : 10.1002/jcp.20727

A. Rousselle and D. Heymann, Osteoclastic acidification pathways during bone resorption, Bone, vol.30, issue.4, pp.533-540, 2002.
DOI : 10.1016/S8756-3282(02)00672-5

R. Velasco, C. Colliec-jouault, S. Redini, F. Heymann, D. Padrines et al., Proteoglycans on bone tumor development, Drug Discovery Today, vol.15, issue.13-14, pp.553-560, 2010.
DOI : 10.1016/j.drudis.2010.05.009

W. Simonet, D. Lacey, C. Dunstan, M. Kelley, M. Chang et al., Osteoprotegerin: A Novel Secreted Protein Involved in the Regulation of Bone Density, Cell, vol.89, issue.2, pp.309-319, 1997.
DOI : 10.1016/S0092-8674(00)80209-3

I. Sutherland, Products of primary metabolism, Biotechnology Second, completely revised edition, vol.6, pp.613-657, 1996.

S. Theoleyre, K. Tat, S. Vusio, P. Blanchard, F. Gallagher et al., Characterization of osteoprotegerin binding to glycosaminoglycans by surface plasmon resonance: role in the interactions with receptor activator of nuclear factor kappaB ligand (RANKL) and RANK, Biochem. Biophys. Res. Commun, vol.34, pp.460-467, 2006.
URL : https://hal.archives-ouvertes.fr/hal-00313928

R. Rank, OPG: new therapeutic targets in bone tumours and associated osteolysis, Biochim Biophys Acta, vol.1704, pp.49-57

L. Xiao, T. Sobue, A. Esliger, M. Kronenberg, J. Coffin et al., Disruption of the Fgf2 gene activates the adipogenic and suppresses the osteogenic program in mesenchymal marrow stromal stem cells, Bone, vol.47, issue.2, pp.360-370, 2010.
DOI : 10.1016/j.bone.2010.05.021

H. Yasuda, N. Shima, N. Nakagawa, S. Mochizuki, K. Yano et al., Identity of Osteoclastogenesis Inhibitory Factor (OCIF) and Osteoprotegerin (OPG): A Mechanism by which OPG/OCIF Inhibits Osteoclastogenesis in Vitro, Endocrinology, vol.139, issue.3, pp.1329-1337, 2004.
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B. Zhao, T. Katagiri, H. Toyoda, T. Takada, T. Yanai et al., Heparin Potentiates the in Vivo Ectopic Bone Formation Induced by Bone Morphogenetic Protein-2, Journal of Biological Chemistry, vol.281, issue.32, pp.23246-23253, 2006.
DOI : 10.1074/jbc.M511039200

W. Ariyoshi, T. Takahashi, T. Kanno, H. Ichimiya, K. Shinmyouzu et al., Heparin inhibits osteoclastic differentiation and function, Journal of Cellular Biochemistry, vol.24, issue.6, pp.1707-1717, 2008.
DOI : 10.1002/jcb.21559

L. Barbour, S. Kick, J. Steiner, M. Loverde, L. Heddleston et al., A prospective study of heparin-induced osteoporosis in pregnancy using bone densitometry, American Journal of Obstetrics and Gynecology, vol.170, issue.3, pp.862-869, 1994.
DOI : 10.1016/S0002-9378(94)70299-3

G. Bernas, Angiotherapeutics from natural products: from bench to clinics?, Clin Hemorheol Microcirc, vol.29, pp.199-203, 2003.

M. Bernfield, M. Gotte, P. Park, O. Reizes, M. Fitzgerald et al., Functions of Cell Surface Heparan Sulfate Proteoglycans, Annual Review of Biochemistry, vol.68, issue.1, pp.729-777, 1999.
DOI : 10.1146/annurev.biochem.68.1.729

S. Colliec-jouault, L. Chevolot, D. Helley, J. Ratiskol, A. Bros et al., Characterization, chemical modifications and in vitro anticoagulant properties of an exopolysaccharide produced by Alteromonas infernus, Biochimica et Biophysica Acta (BBA) - General Subjects, vol.1528, issue.2-3, pp.141-151, 2001.
DOI : 10.1016/S0304-4165(01)00185-4

M. Downey, L. Holliday, J. Aguirre, and T. Wronski, In vitro and in vivo evidence for stimulation of bone resorption by an EP4 receptor agonist and basic fibroblast growth factor: Implications for their efficacy as bone anabolic agents, Bone, vol.44, issue.2, pp.266-274, 2009.
DOI : 10.1016/j.bone.2008.10.041

V. Eswarakumar, I. Lax, and J. Schlessinger, Cellular signaling by fibroblast growth factor receptors, Cytokine & Growth Factor Reviews, vol.16, issue.2, pp.139-149, 2005.
DOI : 10.1016/j.cytogfr.2005.01.001

G. Franz and A. S. , Structure-activity relationship of antithrombotic polysaccharide derivatives, International Journal of Biological Macromolecules, vol.17, issue.6, pp.311-314, 1995.
DOI : 10.1016/0141-8130(96)81837-X

J. Gallagher, Multiprotein signalling complexes: regional assembly on heparan sulphate, Biochemical Society Transactions, vol.34, issue.3, pp.438-441, 2006.
DOI : 10.1042/BST0340438

J. Guezennec, P. Pignet, Y. Lijour, E. Gentric, J. Ratiskol et al., Sulphation and depolymerization of a bacterial exopolysaccharide from hydrothermal origin. Carbohydr Polymers, pp.19-24, 1998.

J. Guezennec, Deep-sea hydrothermal vents: A new source of innovative bacterial exopolysaccharides of biotechnological interest?, Journal of Industrial Microbiology and Biotechnology, vol.29, issue.4, pp.204-208, 2002.
DOI : 10.1038/sj.jim.7000298

L. Haupt, S. Murali, F. Mun, N. Teplyuk, L. Mei et al., The heparan sulfate proteoglycan (HSPG) glypican-3 mediates commitment of MC3T3-E1 cells toward osteogenesis, Journal of Cellular Physiology, vol.281, issue.3, pp.780-791, 2009.
DOI : 10.1002/jcp.21825

A. Irie, M. Takami, H. Kubo, N. Sekino-suzuki, K. Kasahara et al., Heparin enhances osteoclastic bone resorption by inhibiting osteoprotegerin activity, Bone, vol.41, issue.2, pp.165-174, 2007.
DOI : 10.1016/j.bone.2007.04.190

R. Jackson, S. Murali, A. Van-wijnen, G. Stein, V. Nurcombe et al., Heparan sulfate regulates the anabolic activity of MC3T3-E1 preosteoblast cells by induction of Runx2, Journal of Cellular Physiology, vol.19, issue.1, pp.38-50, 2007.
DOI : 10.1002/jcp.20813

G. Karsenty, Minireview: Transcriptional Control of Osteoblast Differentiation, Endocrinology, vol.142, issue.7, pp.2731-2733, 2001.
DOI : 10.1210/endo.142.7.8306

A. Santos, G. Van, and A. Itie, OPGL is a key regulator of osteoclastogenesis, lymphocyte development and lymph-node organogenesis, Nature, vol.397, pp.315-323, 1999.

V. Kram, E. Zcharia, O. Yacoby-zeevi, S. Metzger, T. Chajek-shaul et al., Heparanase is expressed in osteoblastic cells and stimulates bone formation and bone mass, Journal of Cellular Physiology, vol.16, issue.3, pp.784-792, 2006.
DOI : 10.1002/jcp.20625

A. Kumarasuriyar, I. Lee, V. Nurcombe, and S. Cool, De-sulfation of MG-63 cell glycosaminoglycans delays in vitro osteogenesis, up-regulates cholesterol synthesis and disrupts cell cycle and the actin cytoskeleton, Journal of Cellular Physiology, vol.284, issue.3, pp.572-583, 2009.
DOI : 10.1002/jcp.21700

K. Tat, S. Padrines, M. Theoleyre, S. Heymann, D. Fortun et al., IL-6, RANKL, TNF- alpha/IL-1: interrelations in bone resorption pathophysiology, Cytokine Growth Factor Rev, vol.15, pp.49-60, 2004.

K. Tat, S. Padrines, M. Théoleyre, S. Couillaud-battaglia, S. Heymann et al., OPG/membranous???RANKL complex is internalized via the clathrin pathway before a lysosomal and a proteasomal degradation, Bone, vol.39, issue.4, pp.706-715, 2006.
DOI : 10.1016/j.bone.2006.03.016

F. Lamoureux, M. Baud-'huin, L. Duplomb, D. Heymann, and F. Redini, Proteoglycans: key partners in bone cell biology, BioEssays, vol.53, issue.8, pp.758-771, 2007.
DOI : 10.1002/bies.20612

F. Lamoureux, G. Picarda, L. Garrigue-antar, M. Baud-'huin, V. Trichet et al., Glycosaminoglycans as Potential Regulators of Osteoprotegerin Therapeutic Activity in Osteosarcoma, Cancer Research, vol.69, issue.2, pp.526-536, 2009.
DOI : 10.1158/0008-5472.CAN-08-2648

L. Ling, C. Dombrowski, K. Foong, L. Haupt, G. Stein et al., Synergism between Wnt3a and Heparin Enhances Osteogenesis via a Phosphoinositide 3-Kinase/Akt/RUNX2 Pathway, Journal of Biological Chemistry, vol.285, issue.34, 2010.
DOI : 10.1074/jbc.M110.122069

K. Manton, M. Sadasivam, S. Cool, and V. Nurcombe, Bone-specific heparan sulfates induce osteoblast growth arrest and downregulation of retinoblastoma protein, Journal of Cellular Physiology, vol.322, issue.1, pp.219-229, 2006.
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K. Manton, D. Leong, S. Cool, and V. Nurcombe, Disruption of Heparan and Chondroitin Sulfate Signaling Enhances Mesenchymal Stem Cell-Derived Osteogenic Differentiation via Bone Morphogenetic Protein Signaling Pathways, Stem Cells, vol.277, issue.11, pp.2845-2854, 2007.
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A. Rousselle and D. Heymann, Osteoclastic acidification pathways during bone resorption, Bone, vol.30, issue.4, pp.533-540, 2002.
DOI : 10.1016/S8756-3282(02)00672-5

R. Velasco, C. Colliec-jouault, S. Redini, F. Heymann, D. Padrines et al., Proteoglycans on bone tumor development, Drug Discovery Today, vol.15, issue.13-14, pp.553-560, 2010.
DOI : 10.1016/j.drudis.2010.05.009

W. Simonet, D. Lacey, C. Dunstan, M. Kelley, M. Chang et al., Osteoprotegerin: A Novel Secreted Protein Involved in the Regulation of Bone Density, Cell, vol.89, issue.2, pp.309-319, 1997.
DOI : 10.1016/S0092-8674(00)80209-3

I. Sutherland, Products of primary metabolism, Biotechnology Second, completely revised edition, vol.6, pp.613-657, 1996.

S. Theoleyre, K. Tat, S. Vusio, P. Blanchard, F. Gallagher et al., Characterization of osteoprotegerin binding to glycosaminoglycans by surface plasmon resonance: role in the interactions with receptor activator of nuclear factor kappaB ligand (RANKL) and RANK, Biochem. Biophys. Res. Commun, vol.34, pp.460-467, 2006.
URL : https://hal.archives-ouvertes.fr/hal-00313928

R. Rank, OPG: new therapeutic targets in bone tumours and associated osteolysis, Biochim Biophys Acta, vol.1704, pp.49-57

L. Xiao, T. Sobue, A. Esliger, M. Kronenberg, J. Coffin et al., Disruption of the Fgf2 gene activates the adipogenic and suppresses the osteogenic program in mesenchymal marrow stromal stem cells, Bone, vol.47, issue.2, pp.360-370, 2010.
DOI : 10.1016/j.bone.2010.05.021

H. Yasuda, N. Shima, N. Nakagawa, S. Mochizuki, K. Yano et al., Identity of Osteoclastogenesis Inhibitory Factor (OCIF) and Osteoprotegerin (OPG): A Mechanism by which OPG/OCIF Inhibits Osteoclastogenesis in Vitro, Endocrinology, vol.139, issue.3, pp.1329-1337, 2004.
DOI : 10.1210/en.139.3.1329

B. Zhao, T. Katagiri, H. Toyoda, T. Takada, T. Yanai et al., Heparin Potentiates the in Vivo Ectopic Bone Formation Induced by Bone Morphogenetic Protein-2, Journal of Biological Chemistry, vol.281, issue.32, pp.23246-23253, 2006.
DOI : 10.1074/jbc.M511039200