Glucocorticoid receptors and bone

Current Pharmaceutical Design

Volumn 16, Issue 32, 2010, Pages 3586-3592

  La Corte, Renato ^a,b   Trotta, Francesco ^a,b   Adami, Silvano ^c  

^a UNIVERSITY OF FERRARA   (Italy)

^b UNIVERSITY HOSPITAL OF FERRARA   (Italy)

^c UNIVERSITY OF VERONA   (Italy)

Author keywords

Glucocorticoid; Glucocorticoid receptors; Osteoblast; Osteoclast; Osteocyte

Indexed keywords

BISPHOSPHONIC ACID DERIVATIVE; CELL ADHESION MOLECULE; CHEMOKINE; COLONY STIMULATING FACTOR 1; CYTOKINE; DEXAMETHASONE; GAMMA INTERFERON; GLUCOCORTICOID; GLUCOCORTICOID RECEPTOR ALPHA; GLUCOCORTICOID RECEPTOR BETA; INTERLEUKIN 6; INTERSTITIAL COLLAGENASE; MESSENGER RNA; MITOGEN ACTIVATED PROTEIN KINASE; OSTEOCLAST DIFFERENTIATION FACTOR; OSTEOPROTEGERIN; SCLEROSTIN; SOMATOMEDIN C; STROMELYSIN; TRANSCRIPTION FACTOR; WNT PROTEIN;

AMINO TERMINAL SEQUENCE; APOPTOSIS; ARTICLE; BONE CELL; CARBOXY TERMINAL SEQUENCE; CELL FUNCTION; CELL INTERACTION; CELL MATURATION; CELL NUCLEUS; CELL PROLIFERATION; CELL SURVIVAL; CHROMOSOME 5Q; CYTOPLASM; DENDRITIC CELL; DNA BINDING; DRUG DNA BINDING; DRUG MECHANISM; DRUG RECEPTOR BINDING; EXON; GENETIC REGULATION; GENETIC TRANSCRIPTION; HORMONE STRUCTURE; HUMAN; NONHUMAN; OSTEOBLAST; OSTEOCLAST; OSTEOCLAST ACTIVITY; OSTEOCYTE; OSTEOLYSIS; PRIORITY JOURNAL; PROTEIN DOMAIN; PROTEIN EXPRESSION; PROTEIN LOCALIZATION; PROTEIN SECRETION; TRANSACTIVATION; TRANSREPRESSION;

BONE AND BONES; HUMANS; MOLECULAR STRUCTURE; RECEPTORS, GLUCOCORTICOID; TRANSCRIPTIONAL ACTIVATION;

EID: 78650873859     PISSN: 13816128     EISSN: None     Source Type: Journal    
DOI: 10.2174/138161210793797924     Document Type: Article

References (130)

1. Cooper MS, Bujalska I, Rabbitt E, et al. Modulation of 11beta-hydroxysteroid dehydrogenase isoenzymes by proinflammatory cy-tokines in osteoblasts: an autocrine switch from glucocorticoid in-activation to activation. J Bone Miner Res 2001; 16: 1037-44.
2. Lu NZ, Collins JB, Grissom SF, Cidlowsky JA. Selective regula-tion of bone cells apoptosis by translational isoforms of the gluco-corticoid receptor. Mol Cell Biol 2007; 27: 7143-60.
3. van der Laan S, Meijer OC. Pharmacology of glucocorticoids: Be-yond receptor. Eur J Pharmacol 2008; 585: 483-91.
4. Schäcke H, Schottelius A, Döcke WD, et al. Dissociation of trans-activation from transrepression by a selective glucocorticoid recep-tor leads to separation of therapeutic effects from side effects. ProcNatl Acad Sci USA 2004; 101: 227-32.
5. Lazar MA. Thyroid hormone receptors: multiple forms, multiplepossibilities. Endocrin Rev 1993; 14: 184-93.
6. Encio IJ, Detera-Wadleigh SD. The genomic structure of the hu-man glucocorticoid receptor. J Biol Chem 1991; 266: 8182-8.
7. Giguere V, Hollenberg SM, Rosenfeld MG, et al. Functional do-mains of the human glucocorticoid receptor. Cell 2001; 46: 645-52
8. Weinberg C, Hollenberg SM, Rosenfeld MG, et al. Domain struc-ture of human glucocorticoid receptor and its relationship to the v-erb-A oncogen product. Nature 1985; 318: 670-72.
9. Wikstrom AC. Glucocorticoid action and novel mechanisms ofsteroid resistance: role of glucocorticoid receptor-interacting pro-teins for glucocorticoid responsiveness. J Endocrinol 2003; 178:331-7.
10. Beaven S, Horner A, Bord S, Ireland D, Compstom J. Colocaliza-tion of glucocorticoid and mineralcorticoid receptor in humanbone. J Bone Miner Res 2001; 16: 1496-504.
11. Lu NZ, Cidlowski. Translational regulatory mechanisms generateN-terminal glucocorticoid receptor isoforms with unique transcrip-tional target genes. Mol Cell 2005; 18: 331-42.
12. Zhou J, Cidlowski JA. The human glucocorticoid receptor: Onegene, multiple proteins and diverse responses. Steroids 2005; 70:407-17.
13. Hollenberg SM, Weinberg C, Ong SM, et al. Primary structure andexpression of a functional human glucocorticoid receptor cDNA. Nature 1985; 318: 636-41.
14. Almawi WI, Melemedjian OK. Molecular mechanisms of gluco-corticoid antiproliferative effects: antagonism of transcription fac-tor activity by glucocorticoid receptor. J Leukoc Biol 2002; 71: 9-15.
15. Pratt WB, Morishima Y, Murphy M, Harrell M. Chaperoning ofglucocorticoid receptors. Handb Exp Pharmacol 2006; 172: 111-38.
16. Bovio A, Micossi I, Bianco E, Angeli A. Determinants of glucocor-ticoid action in the bone microenviroment. J Endocrinol Invest 2008; 31(suppl.7): 7-15.
17. Vottero A, Chrousos GP. Glucocortioid receptor beta: view. TrendsEndocrinol Metab. 1999; 10: 333-8.
18. Oakley RH, Webster JC, Sar M, Parker Jr. CR, Cidlowski JA. Ex-pression and subcellular distribution of the beta-isoform of the hu-man glucocorticoid receptor. Endocrinology 1997; 138: 5028-38.
19. Goecke A, Guerrero J. Glucocorticoid receptor beta in acute andchronic inflammatory conditions: clinical implications. Immunobi-ology 2006; 211: 85-96.
20. Hamid QA, Wenzel SE, Hauk PJ, et al. Increased glucocorticoidreceptor beta in airway cells of glucocorticoid-insensitive asthma. Am J Resp Crit Care Med 1999; 159(5 part 1): 1600-04.
21. Sousa AR, Lane SJ, Cidlowski JA, Staynov DZ, Lee TH. Gluco-corticoid resistance in asthma is associated with elevated in vivoexpression of the glucocorticoid receptor beta-isoform. J AllergyClin Immunol 1999; 105: 943-50.
22. Oakley RH, Sar M, Cidlowski JA. The human glucocorticoid re-ceptor beta isoform. Expression, biochemical properties, and puta-tive function. J Biol Chem 1996; 271: 9559-9.
23. Bamberger CM, Bamberger AM, de Castro M, Chrousos GP. Glu-cocorticoid receptor beta, a potential endogenous inhibitor of glu-cocorticoid action in humans. J Clin Invest 1995; 95: 2435-41.
24. Chikanza IC. Mechanisms of corticosteroid resistance in rheuma-toid arthritis: a putative role for the glucocorticoid receptor betaisoform. Ann NY Acad Sci 2002; 966: 39-48.
25. Honda M, Orii F, Ayabe T, et al. Expression of glucocorticoid re-ceptor beta in lymphocytes of patients with glucocorticoid-resistantulcerative colitis. Gastroenterology 2000; 118: 859-66.
26. Hauk PJ, Hamid QA, Chrousos GP, Leung DY. Induction of corti-costeroid insensitivity in human pbmcs by microbial superantigens. J Allergy Clin Immunol 2000; 105: 282-7
27. Leung DY, de Castro M. Szefler SJ, Chrousos GP. Mechanisms ofglucocorticoid-resistant asthma. Ann NY Acad Sci 1998; 840: 735-46.
28. Leung DY, Hamid Q, et al. Association of glucocorticoid insensi-tivity with increased expression of glucocorticoid receptor beta. J Exp Med 1997; 186: 1567-74.
29. Webster JC, Oakley RH, Jewell CM, Cidlowski JA. Proinflamma-tory cytokines regulate human glucocorticoid receptor gene expres-sion and lead to the accumulation of the dominant negative betaisoform: A mechanism for the generation of glucocorticoid resis-tance. Proc Natl Acad Sci USA 2001; 98: 6865-70.
30. Whorwood CB, Donovan SJ, Wood PJ, Phillips DI. Regulation ofglucocorticoid receptor alfa and beta isoforms and type 1 11beta-hydroxysteroid dehydrogenase expression in human skeletal mus-cle cells: a key role in the pathogenesis of insulin resistance. J ClinEndocrinol Metab 2001; 86: 2296-308.
31. Krett NL, Pillay S, Moalli PA, Greipp PR, Rosen ST. A variant ofglucocorticoid receptor messenger RNA is expressed in multiplemyeloma patients. Cancer Res 1995; 55: 2727-9.
32. Moalli PA, Pillay S, Krett NL, Rosen ST. Alternatively splicedglucocorticoid messenger RNAs in glucocorticoid-resistant humanmultiple myeloma cells. Cancer Res 1993; 53: 3877-9.
33. Beger C, Gerdes K, Lauten M, et al. Expression of heat-shock pro-tein 90 in glucocorticoid-sensitive and -resistant childhood acutelymphoblastic leukaemia. Br J Haematol 2003; 122: 245-5.
34. Huizenga NA, Koper JA, de Lange P, et al. Brinkmann, de Jong FH, Lamberts SWJ. A polymorphism in the glucocorticoid receptorgene may be associated with an increased sensitivity to glucocorti-coids in vivo. Clin Endocrinol Metab 1998: 83: 144-51.
35. Duma D, Jewell CM, Cidlowski JA. Multiple glucocorticoid recep-tor isoforms and mechanisms of post-translational modification. J Steroid Biochem Drug Saf 2007; 30: 755-63.
36. Cidlowski JA. Glucocorticoid and their actions in cells. Retina 2009; 29: S21-3
37. van Rossum EF, Koper JW, van den Beld AW, et al. Identificationof the BclI polymorphism in the glucocorticoid receptor gene: as-sociation with sensitivity to glucocorticoids in vivo and body massindex. Clin Endocrinol 2003; 59: 585-92.
38. Peng YM, Lei SF, Guo Y, et al. Sex-specific association of theglucorticoid receptor gene with extreme BMD. Bone Miner Res 2008; 23: 247-52.
39. Stahn C, Buttgereit F. Genomic and nongenomic effects of gluco-corticoids. Nat Clin Pract Rheumatol 2008; 4: 525-33.
40. Buttgereit F, Scheffold A. Rapid glucocorticoid effects on immunecells. Steroids 2002; 67: 529-34.
41. Cato AC, Nestl A, Mink S. Rapid actions of steroid receptors incellular signaling pathways. Sci STKE 2002 (138): re9.
42. Croxtall JD, Choudhury Q, Flower RJ. Glucocorticoids act withinminute to inhibit recruitment of signalling factors to activated EGFreceptors through a receptor-dependent, trascription-ndependentmechanism. Br J Pharmacol 2000; 130: 289-98.
43. Croxtall JD, van Hal PT, Choudhury Q, Gilroy DW, Flower RJ.Different glucocorticoids vary in their genomic and non-genomicmechanism of action in A549 cells. Br J Pharmacol 2002; 135:511-9.
44. Marchetti M. C, Di Marco B, Cifone G, Migliorati G, Riccardi C.Dexamethasone-induced apoptosis of thymocytes: role of glucocor-ticoid receptor associated Src kinase and caspase-8 activation.Blood 2003; 101, 585-93.
45. Stellato C. Post-transaltional and nongenomic effects of glucocorti-coids. Proc Am Thorac Soc 2004; 1: 255-63.
46. Löwenberg M, Stahn C, Hommes DW, Buttgereit F. Novel insightsinto mechanisms of glucocorticoid action and the development ofnew glucocorticoid receptor ligands. Steroids 2008; 73: 1025-9.
47. Robertson NM, Schulman G, Karnik S, Alnemri E, Litwack G.Demonstration of nuclear translocation of the mineralcorticoid re-ceptor (MR) using an anti-MR antibody and confocal laser scan-ning microscopy. Mol Endocrinol 1993; 7: 1226-39.
48. Lu NZ, Cidlowski JA. Glucocorticoid receptor isoforms generatetranscription specificity. Trends Cell Biol 2006; 16: 310-7.
49. Heitzer MD, Wolf IM, Sanchez ER, Witchel SF, DeFranco DB.Glucocorticoid receptor physiology. Rev Endocr Metab Disord 2007; 8: 321-30.
50. Ishaq M, DeGray G, Mou K, et al. Zap 70 signaling pathway medi-ates glucocorticoid receptor-dependent transcriptional activation:role in the regulation of annexin 1 expression in T cells. J Immunol 2007; 179: 3851-8.
51. Smoak KA, Cidlowski JA. Mechanisms of glucocorticoid receptorsignaling during inflammation. Mech Ageing Dev 2004; 125: 697-706.
52. Ohnaka K, Tanabe M, Kawate H, Nawata H, Takayanagi R. Glu-cocorticoid suppresses the canonical Wnt signal in cultured humanosteoblasts. Biophys Res Commun 2005; 1; 329(1): 177-181.
53. Smith E, Frenkel B. Glucocorticoids inhibit the transcriptional ac-tivity of LEF/TCF in differentiating osteoblasts in a glycogen syn-thase-3beta-dependent and -independent manner. J Biol Chem 2005; 280: 2388-94.
54. Kawano Y, Kypta R. Secreted antagonists of Wnt signalling path-way. J Cell Sci 2003; 116: 2627-34.
55. Yun SI, Yoon HG, Jeong SY, Chung YS.Glucocorticoid inducesapoptosis of osteoblast cells through the activation of glycogensynthase 3. J Bone Miner Metab 2009; 27: 140-8.
56. Heck S, Kullmann M, Gast A, et al. A distinct modulating domainin glucocorticoid receptor monomers in the repression of activity ofthe transcription factor AP-1. Endo J 1994; 13: 4087-95.
57. Sakai DD, Helms S, Carlstedt-Duke J, Gustafsson JA, RottmanFM, Yamamoto KR. Hormone-mediated repression: a negativeglucocorticoid response element from the bovine prolactin gene. Genes Dev 1988; 2: 1144-54.
58. De Bosscher K, Vanden Berghe W, Haegeman G, The interplaybetween the glucocorticoid-receptor and nuclear factor-kappaB oractivator protein-1: molecular mechanisms for gene repression. En-docr Rev 2003; 24: 488-522.
59. McKay LI, Cidlowki JA. Molecular control of immune/ inflamma-tory responses: interactions between nuclear factor-kB and steroidreceptor-signaling pathways. Endocr Rev 1999; 20: 435-59.
60. DeRijk RH, Schaaf M, de Kloet ER. Glucocorticoid receptor vari-ants: clinical implications. J Steroid Biochem Mol Biol 2002; 81:103-22.
61. Schaaf MJ, Cidlowski JA. Molecular mechanisms of glucocorticoidaction and resistance. J Steroid Biochem Mol Biol 2002 Dec; 83:37-48.
62. Subramaniam N, Cairns W, Okret S. Studies on the mechanism ofglucocorticoid-mediated repression from a negative glucocorticoidresponse element from the bovine prolactin gene. DNA Cell Biol 1997; 16: 153-63. Erratum in: DNA Cell Biol 1997; 16: 670.
63. Aeberli D, Leech M, Morand EF. Macrophage migration inhibitoryfactor and glucocorticoid sensitivity. Rheumatology 2006; 45: 937-943.
64. Morsink MC, Steenbergen PJ, Vos JB, et al. Acute activation ofhippocampal glucocorticoid receptors results in different waves ofgene expression throughtout time. J Neuroendocrinol 2006; 18:239-52.
65. Hofbauer LC, Gori F, Riggs BL, et al. Stimulation of osteoprote-gerin ligand and inhibition of osteoprotegerin production by gluco-corticoid in human osteoblastic lineage cells: Potential paracrinemechanisms of glucocorticoid-induced osteoporosis. Endocrinol-ogy 1999; 140: 4282-89.
66. Rubin J, Biskobing DM, Jadhav L, et al. Dexamethasone promotesexpression of membrane-bound macrophage colony-stimulatingfactor in murine osteoblast-like cells. Endocrinology 1998; 139:1006-12.
67. Sivagurunathan S, Muir MM, Brennan TC, Seale JP, Mason RS.Influence of glucocorticoid on human osteoclast generation and ac-tivity. J Bone Min Res 2005; 20: 390-98.
68. Vanderbilt JN, Miesfeld R, Maler BA, Yamamoto KR. Intracellularreceptor concentration limits glucocorticoid-dependent enhanceractivity. Endocrinology 1987; 1: 68-74.
69. Silva CM, Powell-Oliver FE, Jewell CM, Sar M, Allgood VE, Cid-lowski JA. Regulation of the human glucocorticoid receptor bylong-term and chronic treatment with glucocorticoid. Steroids 1994; 59: 436-442.
70. Seck JR, Walzer BR. Minireview: 11 -Hydroxysteroid dehydro-genase type1-a tissue specific amplifier of glucocorticoid action. Endocrinology 2001; 142: 1371-6.
71. Diederich S, Qinkler M, Burkhardt P, Grossmann C, Bahr V,Oelkers W. 11 Hydroxysteroid dehydrogenase isoforms: Tissuedistribution and implication for clinical medicine. Eur J Clin Invest 2000; 30: 21-27.
72. Cooper MS, Rabbitt EH, Goddard PE, Bartlett WA, Hewison M,Stewart PM. Autocrine activation of glucocorticoids in osteoblasts increase with age and glucocorticoid exposure. J Bone Min Res 2002; 17: 979-986.
73. Cooper MS, Walker EA, Bland R, Fraser WD, Hewison M, StewartPM. Expression and functional consequences of 11 hydroxyster-oid dehydrogenase activity in human bone. Bone 2000; 27: 375-81.
74. Jamieson PM, Chapman KE, Edwards CRW, Seckl JR. 11 Hy-droxysteroid dehydrogenase is an exclusive 11 reductase in pri-mary cultures of rat hepatocytes: Effect of physicochemical andhormonal manipulations. Endocrinology 1995; 136: 4754-61.
75. Canalis E, Delany AM. 11 -Hydroxysteroid dehydrogenase, anamplifer of glucocorticoid action in osteoblats. J Bon Min Res 2002; 17: 987-990.
76. Tomlinson JW, Walker EA, Bujalska IJ, et al. 11beta-hydroxysteroid dehydrogenase type 1: a tissue-specific regulator ofglucocorticoid response. Endocr Rev 2004; 25: 831-66.
77. Cooper MS, Rabbitt EH, Goddard PE, Bartlett WA, Hewison M,Stewart PM. Osteoblastic 11beta-hydroxysteroid dehydrogenasetype 1 activity increases with age and glucocorticoid exposure. JBone Miner Res 2002; 17: 979-86.
78. Pereira RC, Delany AM, Canalis E. CCAAT/enhancer binding pro-tein homologous protein (DDIT3) induces osteoblastic cell diffe-rentiation. Endocrinology 2004; 145: 1952-60.
79. Canalis E, Delany AM. 11beta-hydroxysteroid dehydrogenase, anamplifier of glucocorticoid action in osteoblasts. J Bone Miner Res 2002; 17: 987-90.
80. Canalis E. effect of cortisol on periosteal and non-periosteal colla-gen and DNA synthesis in cultured rat calvaria. Calcif Tissue Int 1984; 36: 158-66.
81. Canalis E, Effect of glucocorticoids on type I collagen synthesis,alkaline phosphatase activity, and deoxyribonucleic acid content incultured rat calvaria. Endocrinology 1983; 112; 931-9.
82. Dempster D. Bone histomorphometry in glucocorticoid-inducedosteoporosis. J Bone Min Res 1989; 4: 4137-141.
83. Canalis E. Mechanisms of glucocorticoid action in bone. Curr Os-teopor Rep 2005; 3: 98-102.
84. Sher LB, Harrison JR, Adams DJ, Kream BE. Impaired corticalbone acquisition and osteoblast differentiation in mice with os-teoblast-targeted disruption of glucocorticoid signalling. Calcif Tis-sue Int 2006; 79: 118-25.
85. Ito S, Suzuki N, Kato S, Takahashi T, Takagi M. Glucocorticoidsinduce the differentiation of a mesenchimal progenitor cell line,ROB-C26 into adipocytes and osteoblasts, but fail to induce termi-nal osteoblast differentiation. Bone 2007; 40: 84-92.
86. Wu Z, Bucher NLR, Farmer SR. Induction of perixosome prolifera-tor-activated receptor during the conversion of 3T3 fibroblastsinto adypocytes is mediated by C/EBP, C/EBP, and glucocorti-coids. Mol Cell Biol 1996; 6: 4128-36.
87. Wedel A, Ziegler-Heitbrock HW. The C/EBP family of transcrip-tion factors. Immunobiology 1995; 193: 171-85.
88. Delany AM, Gabbitas BY, Canalis E. Cortisol down regulates os-teoblast I procollagen mRNA by transcriptional and post-transcrip-tional mechanisms. J Cell Biochem 1995: 57: 488-494.
89. Bortner CD, Cidlowski JA. Cellular mechanisms for the repressionof apoptosis. Ann Rev Pharmacol Toxicol 2002; 42: 259-81.
90. Engelbrecht Y, de Wet H, Horsch K, Langeveldt CR, Hough FS,Hulley PA. Glucocorticoids induce rapid-up-regulation of mitogen-activated protein kinase phospatase-1 and dephosphorylation of ex-tracellular signal-regulated kinase and impair proliferation in hu-man and mouse osteoblast cell lines. Endocrinology 2003; 144:412-22.
91. Hulley PA, Gordon F, Houghs FS. Inhibition of mitogen-activatedprotein kinase activity and proliferation of and early osteoblast cellline (MBA 15.4) by dexamethasone: role of protein phosphatases. Endocrinology 1998; 139: 2423-31.
92. Ley R, Ewings KE, Hadfield K, Howes B, Balmanno K, Cook SJ.Extracellular signal-regulated kinases 1/2 are serum-stimulated"Bim(EL) kinases" that bind to the BH3-only protein (EL) causingits phosphorylation and turnover. J Biol Chem 2004; 279: 8837-47.
93. Tran SE, HOlmstrom TH, Ahonen M, Kahari VM, Eriksson JE.MAPK/ERK overrides the apoptotic signaling from Fas, TNF, andTRAIL receptors. J Biol Chem 2001; 276: 16484-90.
94. Conradie MM, de Wet H, Kotze DDR, Burrin JM, Hough FS, Hul-ley P. Vanadate prevents glucocorticoid-induced apoptosis of os-teoblasts in vitro and osteocytes in vivo. J Endocrinol 2007; 195:229-40.
95. Liu Y, Porta A, Xiaorong P, et al. Prevention of glucocorticoid-induced Apoptosis in osteocytes and osteoblasts by Calbindin-D28k. J Bone Min Res 2004; 19: 479-90.
96. Canalis E, Centrella M, Burch W, McCarthy TL. Insulin-likegrowth factor I mediates selective anabolic effects of parathyroidhormone in bone cultures. J Clin Invest 1989; 83: 60-65.
97. Okazaki R, Riggs BL, Conover CA. Glucocorticoid regulation ofinsulin-like growth factor-binding protein expression in normalhuman osteoblast-like cells. Endocrinology 1994; 1334: 126-32.
98. Gabbitas B, Pash JM, Delany AM, Canalis E. Cortisol inhibits thesynthesis of insulin-like growth factor-binding protein-5 in bonecell cultures by transcriptional mechanisms. J Biol Chem 1996;271: 9033-8.
99. Devlin RD, Du Z, Buccilli V, Jorgetti V, Canalis E. Transgenicmice overexpressing insulin-like growth factor binding protein-5display transiently decreased osteoblastic function and osteopenia. Endocrinology 2002; 142: 3955-62.
100. Lane NE, Sanchez S, Modin GW, Genant HK, Pierini E, ArnaudCD. Parathyroid hormone treatment can reverse corticosteroid-induced osteoporosis. Results of a randomised controlled clinicaltrial. J Clin Invest 1998; 102: 1627-33.
101. Canalis E, Bilezikian JP, Angeli A, Giustina A. Perspectives onglucocorticoid-induced osteoporosis. Bone 2004; 34: 593-8
102. Teitelbaum SL. Bone resorption by osteoclasts. Science 2000; 289:1504-8.
103. Hofbauer LC, Zeitz U, Schoppet M, et al. Prevetion of glucocorti-coid-induced bone loss in mice by inhibition of RANKL. ArthritisRheum 2009; 60: 1427-37.
104. Swanson C, Lorentzon M, Conaway HH, Lerner UH. Glucocorti-coid regulation of osteoclast differentiation and expression of re-ceptor activator of nuclear factor-kappaB (NF-kappaB) ligand, os-teoprotegerin, and receptor activator of NF-kappaB in mouse cal-varia bones. Endocrinology 2006; 147: 3613-22.
105. Delany AM, Jeffrey JJ, Rydziel S, Canalis E. Cortisol increasesinterstitial collagenase expression in osteoblasts by post-transcriptional mechanisms. J Biol Chem 1995; 270: 26607-12.
106. Seeman E. Bone modeling and remodeling. Crit Rev EukaryotGene Expr 2009; 19: 219-33.
107. Gu G, Hentunen TA, Nars M, Härkönen PL, Väänänen HK. Estro-gen protects primary osteoblasts against glucocorticoid-inducedapoptosis. Apoptosis 2005; 10: 583-95.
108. Weinstein RS, Chen JR, Powers CC, et al. Promotion of osteoclastsurvival and antagonism of bisphosphonate-induced osteoclastapoptosis by glucocorticoids. J Clin Invest 2002; 109: 1041-1048.
109. Jia D, O'Brien CA, Stewart SA, Manolagas SC, Weinstein RS, glu-cocorticoids act directly on osteoclasts to increase their life spanand reduce bone density. Endocrinology 2006; 147: 5592-5599.
110. Lane NE, Yao W, Balooch M, et al. Glucocorticoid-treated micehave localized changes in trabecular bone material properties andosteocyte lacunar size that are not observed in placebo-treated orestrogen-deficient mice. J Bone Miner Res 2006; 21: 466-76.
111. O'Brien CA, Jia D, Plotkin LI, et al. Glucocorticoids act directly onosteoblasts and osteocytes to induce their apoptosis and reducebone formation and strength. Endocrinology 2004; 145: 1835-41.
112. Plotkin LI, Lezcano V, Thostenson J, Weinstein RS, ManolagasSC, Bellido T. Connexin 43 is required for the anti-apoptotic effectof bisphosphonates on osteocytes and osteoblasts in vivo. J BoneMiner Res 2008; 23: 1712-1721
113. Baylink DJ, Wergedal JE. Bone formation by osteocytes. Am JPhysiol 1971; 221: 669-678.
114. Yao W, Cheng Z, Pham A, et al. Glucocorticoidinduced bone lossin mice can be reversed by the actions of parathyroid hormone andrisedronate on different pathways for bone formation and minerali-zation. Arthritis Rheum 2008; 58: 3485-3497.
115. Vaes BL, Dechering KJ, van Someren EP, et al. Microarray analy-sis reveals expression regulation of Wnt antagonists in differentiat-ing osteoblasts. Bone 2005; 36: 803-811.
116. van der Horst G, van der Werf SM, Farih-Sips H, van Bezooijen RL, Lowik CW, Karperien M. Down regulation of Wnt signalingby increased expression of Dickkopf-1 and -2 is a prerequisite forlate-stage osteoblast differentiation of KS-483 cells. J Bone MinerRes 2005; 20: 1867-77.
117. Poole KE, van bezooijen RL, Loveridge N, et al. Sclerostin is adelayed secreted product of osteocytes that inhibits bone formation. FASEB J 2005; 19: 1842-3.
118. Raisz LG, Trummel CL, Wener JA, Simmons H. Effect of gluco-corticoids on bone resorption in tissue culture. Endocrinology 1972; 90: 961-7.
119. Teitelbaum SL, Malone JD, Kahan AJ. Glucocorticoid enhance-ment of bone resorption by rat peritoneal macrophages in vitro. Endocrinology 1981; 108: 795-99.
120. Reid IR, Katz JM, Ibbertson HK, Gray DH. The effects of hydro-cortisone, parathyroid hormone and the bisphosphonate, APD, onbone resorption in neonatal mouse calvaria. Calcif Tissue Int 1986;38: 38-43.
121. Tobias J, Chambers TJ. Glucocorticoids impair bone resorptiveactivity and viability of osteoclasts disaggregated from neonatal ratlong bone. Endocrinology 1989; 125: 1290-95.
122. Lukert BP, Raisz LG. Glucocorticoid-induced osteoporosis: patho-genesis and management. Ann Intern Med 1990: 112: 352-64.
123. Udawaga N, Takahashi N, Akatsu T, et al. Origin of osteoclasts:Mature monocytes and macrophages are capable of differentiatinginto osteoclasts under a suitable microenviroment prepared by bonemarrow-derived stromal cells. Proc Natl Acad Sci USA 1990; 87:7260-64.
124. Elford PR, Felix R, Cecchini M, Trechsel U, Fleisch H. Murineosteoblast-like cells and the osteogenic cell MC3T3-E1 release amacrophage colony-stimulating activity in culture. Calcif Tissue Int 1987; 41: 151-6.
125. Dovio A, Perazzolo L, Saba L, et al. High-dose glucocorticoidsincrease serum levels of solubile IL-6 receptor alpha and its ratio tosolubile gp30: an additional mechanism for early increased boneresorption. Eur J Endocrinol 2006; 154: 745-51.
126. Takuma A, Kaneda T, Sato T, Ninomiya S, Kumegawa M, HakedaY. Dexamethasone enhances osteoclast formation synergisticallywith transforming growth factor-beta by stimulating the priming ofosteoclast progenitors for differentiation into osteoclasts. J BiolChem 2003; 278: 44667-74.
127. Kim HJ, Zhao H, Kitaura H, et al. Glucocorticoids and the osteo-clast. Ann N Y Acad Sci 2007; 1116: 355-9.
128. Dostert A, Heinzel T. Negative glucocorticoid receptor responseelements and their role in glucocorticoid action. Curr Pharm Des 2004; 10(23): 2807-16.
129. Gordeladze JO, Reseland JE, Drevon CA. Pharmacological inter-ference with transcriptional control of osteoblasts: a possible rolefor leptin and fatty acids in maintaining bone strength and bodylean mass. Curr Pharm Des 2001; 7(4): 275-90.
130. Sambrook PN. Glucocorticoid osteoporosis. Curr Pharm Des 2002;8(21): 1877-83.