Roles of Wnt signalling in bone growth, remodelling, skeletal disorders and fracture repair

Journal of Cellular Physiology

Volumn 215, Issue 3, 2008, Pages 578-587

  Macsai, Carmen E ^a,b   Foster, Bruce K ^a,b   Xian, Cory J ^a,b  

^a WOMEN S AND CHILDREN S HOSPITAL   (Australia)

^b UNIVERSITY OF ADELAIDE   (Australia)

Author keywords

[No Author keywords available]

Indexed keywords

WNT PROTEIN;

BONE ATROPHY; BONE DEVELOPMENT; BONE DISEASE; BONE GROWTH; BONE REGENERATION; BONE REMODELING; BONE TUMOR; CHONDROGENESIS; EMBRYO DEVELOPMENT; FRACTURE HEALING; GENE FUNCTION; HUMAN; NONHUMAN; OSSIFICATION; OSTEOBLAST; OSTEOCLAST; OSTEOSARCOMA; POSTNATAL DEVELOPMENT; PRIORITY JOURNAL; REVIEW; SIGNAL TRANSDUCTION; ANIMAL; FRACTURE; METABOLISM; PATHOLOGY; PHYSIOLOGY; WOUND HEALING;

ANIMALS; BONE DEVELOPMENT; BONE REMODELING; FRACTURES, BONE; HUMANS; SIGNAL TRANSDUCTION; WNT PROTEINS; WOUND HEALING;

EID: 42949117323     PISSN: 00219541     EISSN: None     Source Type: Journal    
DOI: 10.1002/jcp.21342     Document Type: Review

References (122)

1. Ai M, Heeger S, Bartels CF, Schelling DK. 2005a. Clinical and molecular findings in osteoporosis-pseudoglioma syndrome. Am J Hum Genet 77:741-753.
2. Ai M, Holmen SL, Van Hul W, Williams BO, Warman ML 2005b. Reduced affinity to and inhibition by DKK1 form a common mechanism by which high bone mass-associated missense mutations in LRP5 affect canonical Wnt signaling. Mol Cell Biol 25:4946-4955.
3. Andrade AC, Nilsson O, Barnes KM, Baron J. 2007. Wnt gene expression in the post-natal growth plate: Regulation with chondrocyte differentiation. Bone 40:1361-1369.
4. Arasapam G, Scherer M, Cool JC, Foster BK, Xian CJ. 2006. Roles of COX-2 and iNOS in the bony repair of the injured growth plate cartilage. J Cell Biochem 99:450-461.
5. Asagiri M, Takayanagi H. 2007. The molecular understanding of osteoclast differentiation. Bone 40:251-264.
6. Austin TW, Solar GP, Ziegler FC, Liem L, Matthews W. 1997. A role for the Wnt gene family in hematopoiesis: Expansion of multilineage progenitor cells. Blood 89:3624-3635.
7. Baig-Lewis S, Peterson-Nedry W, Wehrli M. 2007. Wingless/Wnt signal transduction requires distinct initiation and amplification steps that both depend on Arrow/LRP. Dev Biol 306:94-111.
8. Bain G, Muller T, Wang X, Papkoff J. 2003. Activated beta-catenin induces osteoblast differentiation of C3H10T1/2 cells and participates in BMP2 mediated signal transduction. Biochem Biophys Res Commun 301:84-91.
9. Baron R, Rawadi G. 2007. Wnt signaling and the regulation of bone mass. Curr Osteoporos Rep 5:73-80.
10. Bellido T. 2006. Downregulation of SOST/sclerostin by PTH: A novel mechanism of hormonal control of bone formation mediated by osteocytes. J Musculoskelet Neuronal Interact 6:358-359.
11. Bennett CN, Longo KA, Wright WS, Suva LJ, Lane TF, Hankenson KD, MacDougald OA. 2005. Regulation of osteoblastogenesis and bone mass by Wnt10b. Proc Natl Acad Sci USA 102:3324-3329.
12. Blair JM, Zheng Y, Dunstan CR. 2007. RANK ligand. Int J Biochem Cell Biol 39:1077-1081.
13. Bodine PV, Zhao W, Kharode YP, Bex FJ, Lambert AJ, Goad MB, Gaur T, Stein GS, Lian JB, Komm BS. 2004. The Wnt antagonist secreted frizzled-related protein-1 is a negative regulator of trabecular bone formation in adult mice. Mol Endocrinol 18:1222-1237.
14. Boland GM, Perkins G, Hall DJ, Tuan RS. 2004. Wnt 3a promotes proliferation and suppresses osteogenic differentiation of adult human mesenchymal stem cells. J Cell Biochem 93:1210-1230.
15. Boyden LM, Mao J, Belsky J, Mitzner L, Farhi A, Mitnick MA, Wu D, Insogna K, Lifton RP. 2002. High bone density due to a mutation in LDL-receptor-related protein 5. N Engl J Med 346:1513-1521.
16. Cadigan KM, Liu YI. 2006. Wnt signaling: Complexity at the surface. J Cell Sci 119:395-402.
17. Chen Y, Whetstone HC, Lin AC, Nadesan P, Wei Q, Poon R, Alman BA. 2007a. Beta-catenin signaling plays a disparate role in different phases of fracture repair: Implications for therapy to improve bone healing. PLoS Med 4:e249.
18. Chen Y, Whetstone HC, Youn A, Nadesan P, Chow EC, Lin AC, Alman BA. 2007b. Beta-catenin signaling pathway is crucial for bone morphogenetic protein 2 to induce new bone formation. J Biol Chem 282:526-533.
19. Chung R, Cool JC, Scherer MA, Foster BK, Xian CJ. 2006. Roles of neutrophil-mediated inflammatory response in the bony repair of injured growth plate cartilage in young rats. J Leukoc Biol 80:1272-1280.
20. Ciani L, Salinas PC. 2005. WNTs in the vertebrate nervous system: From patterning to neuronal connectivity. Nat Rev Neurosci 6:351-362.
21. Clines GA, Mohammad KS, Bao Y, Stephens OW, Suva LJ, Shaughnessy JD, Jr., Fox JW, Chirgwin JM, Guise TA. 2007. Dickkopf homolog 1 mediates endothelin-1-stimulated new bone formation. Mol Endocrinol 21:486-498.
22. Day TF, Guo X, Garrett-Beal L, Yang Y. 2005. Wnt/beta-catenin signaling in mesenchymal progenitors controls osteoblast and chondrocyte differentiation during vertebrate skeletogenesis. Dev Cell 8:739-750.
23. De Boer J, Siddappa R, Gaspar C, van Apeldoorn A, Fodde R, van Blitterswijk C. 2004a. Wnt signaling inhibits osteogenic differentiation of human mesenchymal stem cells. Bone 34:818-526.
24. De Boer J, Wang HJ, Van Blitterswijk C. 2004b. Effects of Wnt signaling on proliferation and differentiation of human mesenchymal stem cells. Tissue Eng 10:393-401.
25. Dealy CN, Roth A, Ferrari D, Brown AM, Kosher RA. 1993. Wnt-5a and Wnt-7a are expressed in the developing chick limb bud in a manner suggesting roles in pattern formation along the proximodistal and dorsoventral axes. Mech Dev 43:175-186.
26. Dejmek J, Safholm A, Kamp Nielsen C, Andersson T, Leandersson K. 2006. Wnt-5a/Ca2+-induced NFAT activity is counteracted by Wnt-5a/Yes-Cdc42-casein kinase 1alpha signaling in human mammary epithelial cells. Mol Cell Biol 26:6024-6036.
27. Diarra D, Stolina M, Polzer K, Zwerina J, Ominsky MS, Dwyer D, Korb A, Smolen J, Hoffmann M, Scheinecker C, van der Heide D, Landewe R, Lacey D, Richards WG, Schett G. 2007. Dickkopf-1 is a master regulator of joint remodeling. Nat Med 13:156-163.
28. Dihlmann S, von Knebel Doeberitz M. 2005. Wnt/beta-catenin-pathway as a molecular target for future anti-cancer therapeutics. Int J Cancer 113:515-524.
29. Dong Y, Drissi H, Chen M, Chen D, Zuscik MJ, Schwarz EM, O'Keefe RJ. 2005. Wnt-mediated regulation of chondrocyte maturation: Modulation by TGF-beta. J Cell Biochem 95:1057-1068.
30. Eisenberg LM, Eisenberg CA. 2006. Wnt signal transduction and the formation of the myocardium. Dev Biol 293:305-315.
31. Eisenberg LM, Eisenberg CA. 2007. Evaluating the role of Wnt signal transduction in promoting the development of the heart. Sci World J 7:161-176.
32. Endo Y, Rubin JS. 2007. Wnt signaling and neurite outgrowth: Insights and questions. Cancer Sci 98:1311-1317.
33. Fanto M, McNeill H. 2004. Planar polarity from flies to vertebrates. J Cell Sci 117:527-533.
34. French DM, Kaul RJ, D'Souza AL, Crowley CW, Bao M, Frantz GD, Filvaroff EH, Desnoyers L. 2004. WISP-1 is an osteoblastic regulator expressed during skeletal development and fracture repair. Am J Pathol 165:855-867.
35. Gardner JC, van Bezooijen RL, Mervis B, Hamdy NA, Lowik CW, Hamersma H, Beighton P, Papapoulos SE. 2005. Bone mineral density in sclerosteosis; affected individuals and gene carriers. J Clin Endocrinol Metab 90:6392-6395.
36. Glass DA II, Bialek P, Ahn JD, Starbuck M, Patel MS, Clevers H, Taketo MM, Long F, McMahon AP, Lang RA, Karsenty G. 2005. Canonical Wnt signaling in differentiated osteoblasts controls osteoclast differentiation. Dev Cell 8:751-764.
37. Gong Y, Slee RB, Fukai N, Rawadi G, Roman-Roman S, Reginato AM, Wang H, Cundy T, Glorieux FH, Lev D, Zacharin M, Oexle K, Marcelino J, Suwairi W, Heeger S, Sabatakos G, Apte S, Adkins WN, Allgrove J, Arslan-Kirchner M, Batch JA, Beighton P, Black GC, Boles RG, Boon LM, Borrone C, Brunner HG, Carle GF, Dallapiccola B, De Paepe A, Floege B, Halfhide ML, Hall B, Hennekam RC, Hirose T, Jans A, Juppner H, Kim CA, Keppler-Noreuil K, Kohlschuetter A, LaCombe D, Lambert M, Lemyre E, Letteboer T, Peltonen L, Ramesar RS, Romanengo M, Somer H, Steichen-Gersdorf E, Steinmann B, Sullivan B, Superti-Furga A, Swoboda W, van den Boogaard MJ, Van Hul W, Vikkula M, Votruba M, Zabel B, Garcia T, Baron R, Olsen BR, Warman ML. 2001. LDL receptor-related protein 5 (LRP5) affects bone accrual and eye development. Cell 107:513-523.
38. Gregory CA, Singh H, Perry AS, Prockop DJ. 2003. The Wnt signaling inhibitor dickkopf-1 is required for reentry into the cell cycle of human adult stem cells from bone marrow. J Biol Chem 278:28067-28078.
39. Habas R, Dawid IB. 2005. Dishevelled and Wnt signaling: Is the nucleus the final frontier? J Biol 4:2.
40. Hadjiargyrou M, Lombardo F, Zhao S, Ahrens W, Joo J, Ahn H, Jurman M, White DW, Rubin CT. 2002. Transcriptional profiling of bone regeneration. Insight into the molecular complexity of wound repair. J Biol Chem 277:30177-30182.
41. Hartikka H, Makitie O, Mannikko M, Doria AS, Daneman A, Cole WG, Ala-Kokko L, Sochett EB. 2005. Heterozygous mutations in the LDL receptor-related protein 5 (LRP5) gene are associated with primary osteoporosis in children. J Bone Miner Res 20:783-789.
42. Hausler KD, Horwood NJ, Chuman Y, Fisher JL, Ellis J, Martin TJ, Rubin JS, Gillespie MT. 2004. Secreted frizzled-related protein-1 inhibits RANKL-dependent osteoclast formation. J Bone Miner Res 19:1873-1881.
43. Hill TP, Spater D, Taketo MM, Birchmeier W, Hartmann C. 2005. Canonical Wnt/beta-catenin signaling prevents osteoblasts from differentiating into chondrocytes. Dev Cell 8:727-738.
44. Hoang BH, Kubo T, Healey JH, Sowers R, Mazza B, Yang R, Huvos AG, Meyers PA, Gorlick R. 2004. Expression of LDL receptor-related protein 5 (LRP5) as a novel marker for disease progression in high-grade osteosarcoma. Int J Cancer 109:106-111.
45. Holmen SL, Zylstra CR, Mukherjee A, Sigler RE, Faugere MC, Bouxsein ML, Deng L, Clemens TL, Williams BO. 2005. Essential role of beta-catenin in postnatal bone acquisition. J Biol Chem 280:21162-21168.
46. Huang W, Yang S, Shao J, Li YP. 2007. Signaling and transcriptional regulation in osteoblast commitment and differentiation. Front Biosci 12:3068-3092.
47. Hwang SG, Yu SS, Lee SW, Chun JS. 2005. Wnt-3a regulates chondrocyte differentiation via c-Jun/AP-1 pathway. FEBS Lett 579:4837-4842.
48. Iannotti JP. 1990. Growth plate physiology and pathology. Orthop Clin North Am 21:1-17.
49. Iglesias DM, Hueber PA, Chu L, Campbell R, Patenaude AM, Dziarmaga AJ, Quinlan J, Mohamed O, Dufort D, Goodyer PR. 2007. Canonical WNT signaling during kidney development. Am J Physiol Renal Physiol.
50. Imai K, Morikawa M, D'Armiento J, Matsumoto H, Komiya K, Okada Y. 2006. Differential expression of WNTs and FRPs in the synovium of rheumatoid arthritis and osteoarthritis. Biochem Biophys Res Commun 345:1615-1620.
51. Iozzo RV, Eichstetter I, Danielson KG. 1995. Aberrant expression of the growth factor Wnt-5A in human malignancy. Cancer Res 55:3495-3499.
52. Itasaki N, Jones CM, Mercurio S, Rowe A, Domingos PM, Smith JC, Krumlauf R. 2003. Wise, a context-dependent activator and inhibitor of Wnt signalling. Development 130:4295-4305.
53. Jackson A, Vayssiere B, Garcia T, Newell W, Baron R, Roman-Roman S, Rawadi G. 2005. Gene array analysis of Wnt-regulated genes in C3H10T1/2 cells. Bone 36:585-598.
54. Johnson ML. 2004. The high bone mass family - the role of Wnt/Lrp5 signaling in the regulation of bone mass. J Musculoskelet Neuronal Interact 4:135-138.
55. Karsenty G, Wagner EF. 2002. Reaching a genetic and molecular understanding of skeletal development. Dev Cell 2:389-406.
56. Kato M, Patel MS, Levasseur R, Lobov I, Chang BH, Glass DA II, Hartmann C, Li L, Hwang TH, Brayton CF, Lang RA, Karsenty G, Chan L. 2002. Cbfa1-independent decrease in osteoblast proliferation, osteopenia, and persistent embryonic eye vascularization in mice deficient in Lrp5, a Wnt coreceptor. J Cell Biol 157:303-314.
57. Kawakami Y, Wada N, Nishimatsu SI, Ishikawa T, Noji S, Nohno T. 1999. Involvement of Wnt-5a in chondrogenic pattern formation in the chick limb bud. Dev Growth Differ 41:29-40.
58. Kawano Y, Kypta R. 2003. Secreted antagonists of the Wnt signalling pathway. J Cell Sci 116:2627-2634.
59. Khosla S. 2001. Minireview: The OPG/RANKL/RANK system. Endocrinology 142:5050-5055.
60. Kikuchi A, Yamamoto H, Kishida S. 2007. Multiplicity of the interactions of Wnt proteins and their receptors. Cell Signal 19:659-671.
61. Koay MA, Brown MA. 2005. Genetic disorders of the LRP5-Wnt signalling pathway affecting the skeleton. Trends Mol Med 11:129-137.
62. Krishnan V, Bryant HU, Macdougald OA. 2006. Regulation of bone mass by Wnt signaling. J Clin Invest 116:1202-1209.
63. Kronenberg HM. 2006. PTHrP and skeletal development. Ann NY Acad Sci 1068:1-13.
64. Lamoureux F, Trichet V, Chipoy C, Blanchard F, Gouin F, Redini F. 2007. Recent advances in the management of osteosarcoma and forthcoming therapeutic strategies. Expert Rev Anticancer Ther 7:169-181.
65. Levasseur R, Lacombe D, de Vernejoul MC. 2005. LRP5 mutations in osteoporosis-pseudoglioma syndrome and high-bone-mass disorders. Joint Bone Spine 72:207-214.
66. Lian JB, Javed A, Zaidi SK, Lengner C, Montecino M, van Wijnen AJ, Stein JL, Stein GS. 2004. Regulatory controls for osteoblast growth and differentiation: Role of Runx/Cbfa/AML factors. Crit Rev Eukaryot Gene Expr 14:1-41.
67. Lin Z, Willers C, Xu J, Zheng MH. 2006. The chondrocyte: Biology and clinical application. Tissue Eng 12:1971-1984.
68. Little RD, Carulli JP, Del Mastro RG, Dupuis J, Osborne M, Folz C, Manning SP, Swain PM, Zhao SC, Eustace B, Lappe MM, Spitzer L, Zweier S, Braunschweiger K, Benchekroun Y, Hu X, Adair R, Chee L, FitzGerald MG, Tulig C, Caruso A, Tzellas N, Bawa A, Franklin B, McGuire S, Nogues X, Gong G, Allen KM, Anisowicz A, Morales AJ, Lomedico PT, Recker SM, Van Eerdewegh P, Recker RR, Johnson ML. 2002. A mutation in the LDL receptor-related protein 5 gene results in the autosomal dominant high-bone-mass trait. Am J Hum Genet 70:11-19.
69. Logan CY, Nusse R. 2004. The Wnt signaling pathway in development and disease. Annu Rev Cell Dev Biol 20:781-510.
70. Lowik CW, van Bezooijen RL. 2006. Wnt signaling is involved in the inhibitory action of sclerostin on BMP-stimulated bone formation. J Musculoskelet Neuronal Interact 6:357.
71. Lowik C, van Bezooijen RL. 2007. Sclerostin, a natural inhibitor of bone formation, fulfils criteria of an ideal drug target for osteoporosis. Bone 40:S114.
72. Luo Q, Kang Q, Si W, Jiang W, Park JK, Peng Y, Li X, Luu HH, Luo J, Montag AG, Haydon RC, He TC. 2004. Connective tissue growth factor (CTGF) is regulated by Wnt and bone morphogenetic proteins signaling in osteoblast differentiation of mesenchymal stem cells. J Biol Chem 279:55958-55968.
73. Macsai CE, Scherer MA, Foster BK, Zannettino A, Xian CJ. 2007. Expression of Wnt signalling molecules at the injured growth plate cartilage in young rats. Calcif Tissue Int 80:s54.
74. Mani A, Radhakrishnan J, Wang H, Mani MA, Nelson-Williams C, Carew KS, Mane S, Najmabadi H, Wu D, Lifton RP. 2007. LRP6 mutation in a family with early coronary disease and metabolic risk factors. Science 315:1278-1282.
75. Mao B, Niehrs C. 2003. Kremen2 modulates Dickkopf2 activity during Wnt/LRP6 signaling. Gene 302:179-183.
76. Mao B, Wu W, Davidson G, Marhold J, Li M, Mechler BM, Delius H, Hoppe D, Stannek P, Walter C, Glinka A, Niehrs C. 2002. Kremen proteins are Dickkopf receptors that regulate Wnt/beta-catenin signalling. Nature 417:664-667.
77. Miyamoto T, Suda T. 2003. Differentiation and function of osteoclasts. Keio J Med 52:1-7.
78. Nakanishi R, Shimizu M, Mori M, Akiyama H, Okudaira S, Otsuki B, Hashimoto M, Higuchi K, Hosokawa M, Tsuboyama T, Nakamura T. 2006. Secreted frizzled-related protein 4 is a negative regulator of peak BMD in SAMP6 mice. J Bone Miner Res 21:1713-1721.
79. Nam JS, Turcotte TJ, Smith PF, Choi S, Yoon JK. 2006. Mouse cristin/R-spondin family proteins are novel ligands for the Frizzled 8 and LRP6 receptors and activate beta-catenin-dependent gene expression. J Biol Chem 281:13247-13257.
80. Nusse R. 2005. Cell biology: Relays at the membrane. Nature 438:747-749.
81. Ominsky M, Stouch B, Doellgast G, Gong J, Gao Y, Haldankar R, Winters A, Chen Q, Graham K, Zhou L, Hale M, Henry A, Lightwood D, Moore A, Popplewell A, Robinson M, Vlasseros F, Jolette J, Smith SY, Kostenuik PJ, Simonet W, Lacey DL, Paszty C. 2006a. Administration of sclerostin monoclonal antibodies to female Cynomologous monkeys results in increased bone formation, bone mineral density, and bone strength. J Bone Miner Res 21:S44.
82. Ominsky M, Warmongton KS, Asuncion FJ, Tan HL, Grisanti MS, Geng Z, Stephens P, Henry A, Lawson A, Lightwood D, Perkins V, Kirby P, Moore A, Popplewell A, Robinson M, Li X, Kostenuik PJ, Simonet WS, Lacey DL, Paszty C. 2006b. Sclerostin monoclonal antibody treatment increases bone strength in aged osteopenic ovariectomized rats. J Bone Miner Res 21:S44.
83. Oshima T, Abe M, Asano J, Hara T, Kitazoe K, Sekimoto E, Tanaka Y, Shibata H, Hashimoto T, Ozaki S, Kido S, Inoue D, Matsumoto T. 2005. Myeloma cells suppress bone formation by secreting a soluble Wnt inhibitor, sFRP-2. Blood 106:3160-3165.
84. Ott SM. 2005. Sclerostin and Wnt signalling - The pathway to bone strength. J Clin Endocrinol Metab 90:6741-6743.
85. Patel A, Pollitt R, Dalton A, Saarinen A, Bishop NJ. 2007. Children with recurrent fractures and low bone mass or idiopathic juvenile osteoporosis may have abnormalities in the canonical Wnt-signalling pathway through LRP-5. Bone 40:S71-S72.
86. Pogue R, Lyons K. 2006. BMP signaling in the cartilage growth plate. Curr Top Dev Biol 76:1-48.
87. Pongracz JE, Stockley RA. 2006. Wnt signalling in lung development and diseases. Respir Res 7:15.
88. Prunier C, Hocevar BA, Howe PH. 2004. Wnt signaling: Physiology and pathology. Growth Factors 22:141-150.
89. Qiang Y, Chen Y, Brown N, Rudikoff S, Barlogie B, Shaughnessy JD, Jr. 2007. Dkk1 blocks Wnt-Induced osteoprotegrin production in osteoblast progenitors in multiple myeloma. Bone 40:S144.
90. Ross SE, Hemati N, Longo KA, Bennett CN, Lucas PC, Erickson RL, MacDougald OA. 2000. Inhibition of adipogenesis by Wnt signaling. Science 289:950-953
91. Saarinen A, Mayranpaa MK, Lehesjoki AE, Makitie O. 2007. Heterozygous LRP5 mutations in children with fractures. Bone 40:S139-S140.
92. Schoppet M, Preissner KT, Hofbauer LC. 2002. RANK ligand and osteoprotegerin: Paracrine regulators of bone metabolism and vascular function. Arterioscler Thromb Vasc Biol 22:549-553.
93. Semenov MV, He X. 2006. LRP5 mutations linked to high bone mass diseases cause reduced LRP5 binding and inhibition by SOST. J Biol Chem 281:38276-38284.
94. Semenov MV, Tamai K, Brott BK, Kuhl M, Sokol S, He X. 2001. Head inducer Dickkopf-1 is a ligand for Wnt coreceptor L RP6. Curr Biol 11:951-961.
95. Semenov M, Tamai K, He X. 2005. SOST is a ligand for LRP5/LRP6 and a Wnt signaling inhibitor. J Biol Chem 280:26770-26775.
96. Sen M, Lauterbach K, El-Gabalawy H, Firestein GS, Corr M, Carson DA. 2000. Expression and function of wingless and frizzled homologsin rheumatoid arthritis. Proc Natl Acad Sci USA 97:2791-2796.
97. Sen M, Chamorro M, Reifert J, Corr M, Carson DA. 2001. Blockade of Wnt-5A/frizzled 5 signaling inhibits rheumatoid synoviocyte activation. Arthritis Rheum 44:772-781.
98. Shinohara M, Takayanagi H. 2007. Novel osteoclast signaling mechanisms. Curr Osteoporos Rep 5:67-72.
99. Shoback D. 2007. Update in osteoporosis and metabolic bone disorders. J Clin Endocrinol Metab 92:747-753.
100. Silvestrini G, Ballanti P, Leopizzi M, Sebastiani M, Berni S, Di Vito M, Bonucci E. 2007. Effects of intermittent parathyroid hormone (PTH) administration on SOST mRNA and protein in rat bone. J Mol Histol 38: 261-269.
101. Spencer GJ, Utting JC, Etheridge SL, Arnett TR, Genever PG. 2006. Wnt signalling in osteoblasts regulates expression of the receptor activator of NFkappaB ligand and inhibits osteoclastogenesis in vitro. J Cell Sci 119:1283-1296.
102. Takemaru K, Yamaguchi S, Lee YS, Zhang Y, Carthew RW, Moon RT. 2003. Chibby, a nuclear beta-catenin-associated antagonist of the Wnt/Wingless pathway. Nature 422:905-909.
103. Tamamura Y, Otani T, Kanatani N, Koyama E, Kitagaki J, Komori T, Yamada Y, Costantini F, Wakisaka S, Pacifici M, Iwamoto M, Enomoto-Iwamoto M. 2005. Developmental regulation of Wnt/beta-catenin signals is required for growth plate assembly, cartilage integrity, and endochondral ossification. J Biol Chem 280:19185-19195.
104. Tian E, Zhan F, Walker R, Rasmussen E, Ma Y, Barlogie B, Shaughnessy JD, Jr. 2003. The role of the Wnt-signaling antagonist DKK1 in the development of osteolytic lesions in multiple myeloma. N Engl J Med 349:2483-2494.
105. Tufan AC, Tuan RS. 2001. Wnt regulation of limb mesenchymal chondrogenesis is accompanied by altered N-cadherin-related functions. FASEB J 15:1436-1438.
106. Vaes BL, Dechering KJ, van Someren EP, Hendriks JM, van de Ven CJ, Feijen A, Mummery CL, Reinders MJ, Olijve W, van Zoelen EJ, Steegenga WT. 2005. Microarray analysis reveals expression regulation of Wnt antagonists in differentiating osteoblasts. Bone 36:803-811.
107. van Bezooijen RL, Papapoulos SE, Lowik CW. 2005a. Bone morphogenetic proteins and their antagonists: The sclerostin paradigm. J Endocrinol Invest 28:15-17.
108. van Bezooijen RL, ten Dijke P, Papapoulos SE, Lowik CW. 2005b. SOST/sclerostin, an osteocyte-derived negative regulator of bone formation. Cytokine Growth Factor Rev. 16:319-327.
109. van Bezooijen RL, Svensson JP, Eefting D, Visser A, van der Horst G, Karperien M, Quax PH, Vrieling H, Papapoulos SE, ten Dijke P, Lowik CW. 2007. Wnt but not BMP signaling is involved in the inhibitory action of sclerostin on BMP-stimulated bone formation. J Bone Miner Res 22:19-28.
110. van der Horst G, van der Werf SM, Farih-Sips H, van Bezooijen RL, Lowik CW, Karperien M. 2005. Downregulation of Wnt signaling by increased expression of Dickkopf-1 and -2 is a prerequisite for late-stage osteoblast differentiation of KS483 cells. J Bone Miner Res 20:1867-1877.
111. Wada T, Nakashima T, Hiroshi N, Penninger JM. 2006. RANKL-RANK signaling in osteoclastogenesis and bone disease. Trends Mol Med 12:17-25.
112. Westendorf JJ, Kahler RA, Schroeder TM. 2004. Wnt signaling in osteoblasts and bone diseases. Gene 341:19-39.
113. Winkler DG, Sutherland MK, Geoghegan JC, Yu C, Hayes T, Skonier JE, Shpektor D, Jonas M, Kovacevich BR, Staehling-Hampton K, Appleby M, Brunkow ME, Latham JA. 2003. Osteocyte control of bone formation via sclerostin, a novel BMP antagonist. EMBO J 22:6267-6276.
114. Winkler DG, Sutherland MS, Ojala E, Turcott E, Geoghegan JC, Shpektor D, Skonier JE, Yu C, Latham JA. 2005. Sclerostin inhibition of Wnt-3a-induced C3H10T1/2 cell differentiation is indirect and mediated by bone morphogenetic proteins. J Biol Chem 280:2498-2502.
115. Xian CJ. 2007. Roles of epidermal growth factor family in the regulation of postnatal somatic growth. Endocr Rev 28:284-296.
116. Xian CJ, Zhou FH, McCarty RC, Foster BK. 2004. Intramembranous ossification mechanism for bone bridge formation at the growth plate cartilage injury site. J Orthop Res 22:417-426.
117. Xu Q, Wang Y, Dabdoub A, Smallwood PM, Williams J, Woods C, Kelley MW, Jiang L, Tasman W, Zhang K, Nathans J. 2004. Vascular development in the retina and inner ear: Control by Norrin and Frizzled-4, a high-affinity ligand-receptor pair. Cell 116:883-895.
118. Yang Y, Topol L, Lee H, Wu J. 2003. Wnt5a and Wnt5b exhibit distinct activities in coordinating chondrocyte proliferation and differentiation. Development 130:1003-1015.
119. Yoshino K, Rubin JS, Higinbotham KG, Uren A, Anest V, Plisov SY, Perantoni AO. 2001. Secreted Frizzled-related proteins can regulate metanephric development. Mech Dev 102:45-55.
120. Zhong N, Gersch RP, Hadjiargyrou M. 2006. Wnt signaling activation during bone regeneration and the role of Dishevelled in chondrocyte proliferation and differentiation. Bone 39:5-16.
121. Zhou FH, Foster BK, Zhou XF, Cowin AJ, Xian CJ. 2006. TNF-alpha mediates p38 MAP kinase activation and negatively regulates bone formation at the injured growth plate in rats. J Bone Miner Res 21:1075-1088.
122. Zhu M, Chen M, Zuscik M, Boyce BF, O'Keefe RJ, Chen D. 2007. B-catenin signaling is required for postnatal cartilage development. Bone 40:S147-S148.