Molecular Bases of the Regulation of Bone Remodeling by the Canonical Wnt Signaling Pathway

Current Topics in Developmental Biology

Volumn 73, Issue , 2006, Pages 43-84

  Glass II, Donald A ^a   Karsenty, Gerard ^a  

^a BAYLOR COLLEGE OF MEDICINE   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

AXIN2 PROTEIN, HUMAN; BETA CATENIN; BONE MORPHOGENETIC PROTEIN; CYTOSKELETON PROTEIN; FRIZZLED PROTEIN; GLYCOGEN SYNTHASE KINASE 3; GLYCOGEN SYNTHASE KINASE 3 BETA; KREMEN PROTEIN, MOUSE; LIPOPROTEIN RECEPTOR RELATED PROTEIN 5; LOW DENSITY LIPOPROTEIN RECEPTOR RELATED PROTEIN; LRP6 PROTEIN, HUMAN; MEMBRANE PROTEIN; PROTEIN TYROSINE KINASE; ROR2 PROTEIN, MOUSE; SOST PROTEIN, MOUSE; T CELL FACTOR PROTEIN; WNT PROTEIN;

ANIMAL; BONE REMODELING; DRUG ANTAGONISM; DRUG POTENTIATION; GENETIC MARKER; GENETICS; HUMAN; METABOLISM; OSTEOPOROSIS; PATHOPHYSIOLOGY; PHYSIOLOGY; REVIEW; SIGNAL TRANSDUCTION;

ANIMALS; BETA CATENIN; BONE MORPHOGENETIC PROTEINS; BONE REMODELING; CYTOSKELETAL PROTEINS; FRIZZLED RECEPTORS; GENETIC MARKERS; GLYCOGEN SYNTHASE KINASE 3; HUMANS; LDL-RECEPTOR RELATED PROTEINS; MEMBRANE PROTEINS; OSTEOPOROSIS; RECEPTOR PROTEIN-TYROSINE KINASES; SIGNAL TRANSDUCTION; TCF TRANSCRIPTION FACTORS; WNT PROTEINS;

EID: 33745034435     PISSN: 00702153     EISSN: None     Source Type: Book Series    
DOI: 10.1016/S0070-2153(05)73002-7     Document Type: Review

References (231)

1. Aberle H., Bauer A., Stappert J., Kispert A., and Kemler R. Beta-catenin is a target for the ubiquitin-proteasome pathway. EMBO J. 16 (1997) 3797-3804
2. Afzal A.R., Rajab A., Fenske C.D., Oldridge M., Elanko N., Ternes-Pereira E., Tuysuz B., Murday V.A., Patton M.A., Wilkie A.O., and Jeffery S. Recessive Robinow syndrome, allelic to dominant brachydactyly type B, is caused by mutation of ROR2. Nat. Genet. 25 (2000) 419-422
3. Ai M., Holmen S.L., Van Hul W., Williams B.O., and Warman M.L. 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 (2005) 4946-4955
4. Ali A., Hoeflich K.P., and Woodgett J.R. Glycogen synthase kinase-3: Properties, functions, and regulation. Chem. Rev. 101 (2001) 2527-2540
5. Avissar S., Schreiber G., Danon A., and Belmaker R.H. Lithium inhibits adrenergic and cholinergic increases in GTP binding in rat cortex. Nature 331 (1988) 440-442
6. Babij P., Zhao W., Small C., Kharode Y., Yaworsky P.J., Bouxsein M.L., Reddy P.S., Bodine P.V., Robinson J.A., Bhat B., Marzolf J., Moran R.A., et al. High bone mass in mice expressing a mutant LRP5 gene. J. Bone Miner. Res. 18 (2003) 960-974
7. Baeg G.H., Lin X., Khare N., Baumgartner S., and Perrimon N. Heparan sulfate proteoglycans are critical for the organization of the extracellular distribution of Wingless. Development 128 (2001) 87-94
8. Bafico A., Gazit A., Pramila T., Finch P.W., Yaniv A., and Aaronson S.A. Interaction of frizzled related protein (FRP) with Wnt ligands and the frizzled receptor suggests alternative mechanisms for FRP inhibition of Wnt signaling. J. Biol. Chem. 274 (1999) 16,180-16,187
9. Bafico A., Liu G., Yaniv A., Gazit A., and Aaronson S.A. Novel mechanism of Wnt signalling inhibition mediated by Dickkopf-1 interaction with LRP6/Arrow. Nat. Cell Biol. 7 (2001) 683-686
10. Bain G., Muller T., Wang X., and Papkoff J. Activated beta-catenin induces osteoblast differentiation of C3H10T1/2 cells and participates in BMP2 mediated signal transduction. Biochem. Biophys. Res. Commun. 301 (2003) 84-91
11. Baker J.C., Beddington R.S., and Harland R.M. Wnt signaling in Xenopus embryos inhibits bmp4 expression and activates neural development. Genes Dev. 13 (1999) 3149-3159
12. Balemans W., Ebeling M., Patel N., Van Hul E., Olson P., Dioszegi M., Lacza C., Wuyts W., Van Den Ende J., Willems P., Paes-Alves A.F., Hill S., et al. Increased bone density in sclerosteosis is due to the deficiency of a novel secreted protein (SOST). Hum. Mol. Genet. 10 (2001) 537-543
13. Behrens J., Jerchow B.A., Wurtele M., Grimm J., Asbrand C., Wirtz R., Kuhl M., Wedlich D., and Birchmeier W. Functional interaction of an axin homolog, conductin, with beta-catenin, APC, and GSK3beta. Science 280 (1998) 596-599
14. Behrens J., von Kries J.P., Kuhl M., Bruhn L., Wedlich D., Grosschedl R., and Birchmeier W. Functional interaction of beta-catenin with the transcription factor LEF-1. Nature 382 (1996) 638-642
15. Beighton P., Durr L., and Hamersma H. The clinical features of sclerosteosis: A review of the manifestations in twenty-five affected individuals. Ann. Intern. Med. 84 (1976) 393-397
16. Bennett C.N., Longo K.A., Wright W.S., Suva L.J., Lane T.F., Hankenson K.D., and MacDougald O.A. Regulation of osteoblastogenesis and bone mass by Wnt10b. Proc. Natl. Acad. Sci. USA 102 (2005) 3324-3329
17. Berndt T., Craig T.A., Bowe A.E., Vassiliadis J., Reczek D., Finnegan R., Jan De Beur S.M., Schiavi S.C., and Kumar R. Secreted frizzled-related protein 4 is a potent tumor-derived phosphaturic agent. J. Clin. Invest. 112 (2003) 785-794
18. Bhanot P., Brink M., Samos C.H., Hsieh J.C., Wang Y., Macke J.P., Andrew D., Nathans J., and Nusse R. A new member of the frizzled family from Drosophila functions as a Wingless receptor. Nature 382 (1996) 225-230
19. Billiard J., Way D.S., Seestaller-Wehr L.M., Moran R.A., Mangine A., and Bodine P.V. The orphan receptor tyrosine kinase Ror2 modulates canonical Wnt signaling in osteoblastic cells. Mol. Endocrinol. 19 (2005) 90-101
20. Billin A.N., Thirlwell H., and Ayer D.E. Beta-catenin-histone deacetylase interactions regulate the transition of LEF1 from a transcriptional repressor to an activator. Mol. Cell. Biol. 20 (2000) 6882-6890
21. Bodine P.V., Billiard J., Moran R.A., Ponce-de-Leon H., McLarney S., Mangine A., Scrimo M.J., Bhat R.A., Stauffer B., Green J., Stein G.S., Lian J.B., et al. The Wnt antagonist secreted frizzled-related protein-1 controls osteoblast and osteocyte apoptosis. J. Cell Biochem 96 (2005) 1212-1230
22. Bodine P.V., Zhao W., Kharode Y.P., Bex F.J., Lambert A.J., Goad M.B., Gaur T., Stein G.S., Lian J.B., and Komm B.S. The Wnt antagonist secreted frizzled-related protein-1 is a negative regulator of trabecular bone formation in adult mice. Mol. Endocrinol. 18 (2004) 1222-1237
23. Boyden L.M., Mao J., Belsky J., Mitzner L., Farhi A., Mitnick M.A., Wu D., Insogna K., and Lifton R.P. High bone density due to a mutation in LDL-receptor-related protein 5. N. Engl. J. Med. 346 (2002) 1513-1521
24. Bradbury J.M., Niemeyer C.C., Dale T.C., and Edwards P.A. Alterations of the growth characteristics of the fibroblast cell line C3H 10T1/2 by members of the Wnt gene family. Oncogene 9 (1994) 2597-2603
25. Brannon M., Brown J.D., Bates R., Kimelman D., and Moon R.T. XCtBP is a XTcf-3 co-repressor with roles throughout Xenopus development. Development 126 (1999) 3159-3170
26. Brantjes H., Roose J., van De Wetering M., and Clevers H. All Tcf HMG box transcription factors interact with Groucho-related co-repressors. Nucleic Acids Res. 29 (2001) 1410-1419
27. Brault V., Moore R., Kutsch S., Ishibashi M., Rowitch D.H., McMahon A.P., Sommer L., Boussadia O., and Kemler R. Inactivation of the beta-catenin gene by Wnt1-Cre-mediated deletion results in dramatic brain malformation and failure of craniofacial development. Development 128 (2001) 1253-1264
28. Bruhn L., Munnerlyn A., and Grosschedl R. ALY, a context-dependent coactivator of LEF-1 and AML-1, is required for TCRalpha enhancer function. Genes Dev. 11 (1997) 640-653
29. Cadigan K.M., Fish M.P., Rulifson E.J., and Nusse R. Wingless repression of Drosophila frizzled 2 expression shapes the Wingless morphogen gradient in the wing. Cell 93 (1998) 767-777
30. Cadigan K.M., Jou A.D., and Nusse R. Wingless blocks bristle formation and morphogenetic furrow progression in the eye through repression of Daughterless. Development 129 (2002) 3393-3402
31. Canalis E. Insulin like growth factors and the local regulation of bone formation. Bone 14 (1993) 273-276
32. Carlsson P., Waterman M.L., and Jones K.A. The hLEF/TCF-1 alpha HMG protein contains a context-dependent transcriptional activation domain that induces the TCR alpha enhancer in T cells. Genes Dev. 7 (1993) 2418-2430
33. Carron C., Pascal A., Djiane A., Boucaut J.C., Shi D.L., and Umbhauer M. Frizzled receptor dimerization is sufficient to activate the Wnt/beta-catenin pathway. J. Cell Sci. 116 (2003) 2541-2550
34. Cattelino A., Liebner S., Gallini R., Zanetti A., Balconi G., Corsi A., Bianco P., Wolburg H., Moore R., Oreda B., Kemler R., and Dejana E. The conditional inactivation of the beta-catenin gene in endothelial cells causes a defective vascular pattern and increased vascular fragility. J. Cell Biol. 162 (2003) 1111-1122
35. Chalecka-Franaszek E., and Chuang D.M. Lithium activates the serine/threonine kinase Akt-1 and suppresses glutamate-induced inhibition of Akt-1 activity in neurons. Proc. Natl. Acad. Sci. USA 96 (1999) 8745-8750
36. Chan S.D., Karpf D.B., Fowlkes M.E., Hooks M., Bradley M.S., Vuong V., Bambino T., Liu M.Y., Arnaud C.D., Strewler G.J., and Nissenson R.A. Two homologs of the Drosophila polarity gene frizzled (fz) are widely expressed in mammalian tissues. J. Biol. Chem. 267 (1992) 25202-25207
37. Cheng S.L., Lecanda F., Davidson M.K., Warlow P.M., Zhang S.F., Zhang L., Suzuki S., St John T., and Civitelli R. Human osteoblasts express a repertoire of cadherins, which are critical for BMP-2-induced osteogenic differentiation. J. Bone Miner Res. 13 (1998) 633-644
38. Chiu W.S., McManus J.F., Notini A.J., Cassady A.I., Zajac J.D., and Davey R.A. Transgenic mice that express Cre recombinase in osteoclasts. Genesis 39 (2004) 178-185
39. Cho E.A., and Dressler G.R. TCF-4 binds beta-catenin and is expressed in distinct regions of the embryonic brain and limbs. Mech. Dev. 77 (1998) 9-18
40. Chung Y.S., Baylink D.J., Srivastava A.K., Amaar Y., Tapia B., Kasukawa Y., and Mohan S. Effects of secreted frizzled-related protein 3 on osteoblasts in vitro. J. Bone Miner. Res. 19 (2004) 1395-1402
41. Day T.F., Guo X., Garrett-Beal L., and Yang Y. Wnt/beta-catenin signaling in mesenchymal progenitors controls osteoblast and chondrocyte differentiation during vertebrate skeletogenesis. Dev. Cell 8 (2005) 739-750
42. de Jong D.S., van Zoelen E.J., Bauerschmidt S., Olijve W., and Steegenga W.T. Microarray analysis of bone morphogenetic protein, transforming growth factor beta, and activin early response genes during osteoblastic cell differentiation. J. Bone Miner. Res. 17 (2002) 2119-2129
43. DeChiara T.M., Kimble R.B., Poueymirou W.T., Rojas J., Masiakowski P., Valenzuela D.M., and Yancopoulos G.D. Ror2, encoding a receptor-like tyrosine kinase, is required for cartilage and growth plate development. Nat. Genet. 24 (2000) 271-274
44. Derfoul A., Carlberg A.L., Tuan R.S., and Hall D.J. Differential regulation of osteogenic marker gene expression by Wnt-3a in embryonic mesenchymal multipotential progenitor cells. Differentiation 72 (2004) 209-223
45. Dong Y., Lathrop W., Weaver D., Qiu Q., Cini J., Bertolini D., and Chen D. Molecular cloning and characterization of LR3, a novel LDL receptor family protein with mitogenic activity. Biochem. Biophys. Res. Commun. 251 (1998) 784-790
46. Ducy P., Zhang R., Geoffroy V., Ridall A.L., and Karsenty G. Osf2/Cbfa1: A transcriptional activator of osteoblast differentiation. Cell 89 (1997) 747-754
47. El-Tanani M., Platt-Higgins A., Rudland P.S., and Campbell F.C. Ets gene PEA3 cooperates with beta-catenin-Lef-1 and c-Jun in regulation of osteopontin transcription. J. Biol. Chem. 279 (2004) 20794-20806
48. Farr III G.H., Ferkey D.M., Yost C., Pierce S.B., Weaver C., and Kimelman D. Interaction among GSK-3, GBP, axin, and APC in Xenopus axis specification. J. Cell Biol. 148 (2000) 691-702
49. Ferron M., and Vacher J. Targeted expression of Cre recombinase in macrophages and osteoclasts in transgenic mice. Genesis 41 (2005) 138-145
50. Frame S., Cohen P., and Biondi R.M. A common phosphate binding site explains the unique substrate specificity of GSK3 and its inactivation by phosphorylation. Mol. Cell 7 (2001) 1321-1327
51. Fujino T., Asaba H., Kang M.J., Ikeda Y., Sone H., Takada S., Kim D.H., Ioka R.X., Ono M., Tomoyori H., Okubo M., Murase T., et al. Low-density lipoprotein receptor-related protein 5 (LRP5) is essential for normal cholesterol metabolism and glucose-induced insulin secretion. Proc. Natl. Acad. Sci. USA 100 (2003) 229-234
52. Gaur T., Lengner C.J., Hovhannisyan H., Bhat R.A., Bodine P.V., Komm B.S., Javed A., van Wijnen A.J., Stein J.L., Stein G.S., and Lian J.B. Canonical Wnt signaling promotes osteogenesis by directly stimulating RUNX2 gene expression. J. Biol. Chem. 280 (2005) 33132-33140
53. Gazzerro E., Pereira R.C., Jorgetti V., Olson S., Economides A.N., and Canalis E. Skeletal overexpression of gremlin impairs bone formation and causes osteopenia. Endocrinology 146 (2005) 655-665
54. Giese K., Amsterdam A., and Grosschedl R. DNA-binding properties of the HMG domain of the lymphoid-specific transcriptional regulator LEF-1. Genes Dev. 5 (1991) 2567-2578
55. Giese K., and Grosschedl R. LEF-1 contains an activation domain that stimulates transcription only in a specific context of factor-binding sites. EMBO J. 12 (1993) 4667-4676
56. Giese K., Kingsley C., Kirshner J.R., and Grosschedl R. Assembly and function of a TCR alpha enhancer complex is dependent on LEF-1-induced DNA bending and multiple protein-protein interactions. Genes Dev. 9 (1995) 995-1008
57. Giese K., Pagel J., and Grosschedl R. Functional analysis of DNA bending and unwinding by the high mobility group domain of LEF-1. Proc. Natl. Acad. Sci. USA 94 (1997) 12,845-12,850
58. Glass II D.A., Bialek P., Ahn J.D., Starbuck M., Patel M.S., Clevers H., Taketo M.M., Long F., McMahon A.P., Lang R.A., and Karsenty G. Canonical Wnt signaling in differentiated osteoblasts controls osteoclast differentiation. Dev. Cell 8 (2005) 751-764
59. Glinka A., Wu W., Delius H., Monaghan A.P., Blumenstock C., and Niehrs C. Dickkopf-1 is a member of a new family of secreted proteins and functions in head induction. Nature 391 (1998) 357-362
60. Glise B., Miller C.A., Crozatier M., Halbisen M.A., Wise S., Olson D.J., Vincent A., and Blair S.S. Shifted, the Drosophila ortholog of Wnt inhibitory factor-1, controls the distribution and movement of Hedgehog. Dev. Cell 8 (2005) 255-266
61. Golan T., Yaniv A., Bafico A., Liu G., and Gazit A. The human Frizzled 6 (HFz6) acts as a negative regulator of the canonical Wnt. beta-catenin signaling cascade. J. Biol. Chem. 279 (2004) 14,879-14,888
62. Gong Y., Slee R.B., Fukai N., Rawadi G., Roman-Roman S., Reginato A.M., Wang H., Cundy T., Glorieux F.H., Lev D., Zacharin M., Oexle K., et al. LDL receptor-related protein 5 (LRP5) affects bone accrual and eye development. Cell 107 (2001) 513-523
63. Gong Y., Vikkula M., Boon L., Liu J., Beighton P., Ramesar R., Peltonen L., Somer H., Hirose T., Dallapiccola B., De Paepe A., Swoboda W., et al. Osteoporosis-pseudoglioma syndrome, a disorder affecting skeletal strength and vision, is assigned to chromosome region 11q12-13. Am. J. Hum. Genet. 59 (1996) 146-151
64. Gonzalez-Sancho J.M., Brennan K.R., Castelo-Soccio L.A., and Brown A.M. Wnt proteins induce dishevelled phosphorylation via an LRP5/6-independent mechanism, irrespective of their ability to stabilize beta-catenin. Mol. Cell. Biol. 24 (2004) 4757-4768
65. Gregory C.A., Singh H., Perry A.S., and Prockop D.J. 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 (2003) 28,067-28,078
66. Habas R., Dawid I.B., and He X. Coactivation of Rac and Rho by Wnt/Frizzled signaling is required for vertebrate gastrulation. Genes Dev. 17 (2003) 295-309
67. Hadjiargyrou M., Lombardo F., Zhao S., Ahrens W., Joo J., Ahn H., Jurman M., White D.W., and Rubin C.T. Transcriptional profiling of bone regeneration: Insight into the molecular complexity of wound repair. J. Biol. Chem. 277 (2002) 30,177-30,182
68. Haegel H., Larue L., Ohsugi M., Fedorov L., Herrenknecht K., and Kemler R. Lack of beta-catenin affects mouse development at gastrulation. Development 121 (1995) 3529-3537
69. Hamersma H., Gardner J., and Beighton P. The natural history of sclerosteosis. Clin. Genet. 63 (2003) 192-197
70. Hartmann C., and Tabin C.J. Dual roles of Wnt signaling during chondrogenesis in the chicken limb. Development 127 (2000) 3141-3159
71. Hausler K.D., Horwood N.J., Chuman Y., Fisher J.L., Ellis J., Martin T.J., Rubin J.S., and Gillespie M.T. Secreted frizzled-related protein-1 inhibits RANKL-dependent osteoclast formation. J. Bone Miner. Res. 19 (2004) 1873-1881
72. Hay E., Faucheu C., Suc-Royer I., Touitou R., Stiot V., Vayssiere B., Baron R., Roman-Roman S., and Rawadi G. Interaction between LRP5 and Frat1 mediates the activation of the Wnt canonical pathway. J. Biol. Chem. 280 (2005) 13,616-13,623
73. He T.C., Sparks A.B., Rago C., Hermeking H., Zawel L., da Costa L.T., Morin P.J., Vogelstein B., and Kinzler K.W. Identification of c-MYC as a target of the APC pathway. Science 281 (1998) 1509-1512
74. He X., Saint-Jeannet J.P., Woodgett J.R., Varmus H.E., and Dawid I.B. Glycogen synthase kinase-3 and dorsoventral patterning in Xenopus embryos. Nature 374 (1995) 617-622
75. Heaney C., Shalev H., Elbedour K., Carmi R., Staack J.B., Sheffield V.C., and Beier D.R. Human autosomal recessive osteopetrosis maps to 11q13, a position predicted by comparative mapping of the murine osteosclerosis (oc) mutation. Hum. Mol. Genet. 7 (1998) 1407-1410
76. Hecht A., Vleminckx K., Stemmler M.P., van Roy F., and Kemler R. The p300/CBP acetyltransferases function as transcriptional coactivators of beta-catenin in vertebrates. EMBO J. 19 (2000) 1839-1850
77. Hens J.R., Wilson K.M., Dann P., Chen X., Horowitz M.C., and Wysolmerski J.J. TOPGAL mice show that the canonical Wnt signaling pathway is active during bone development and growth and is activated by mechanical loading in vitro. J. Bone Miner. Res. 20 (2005) 1103-1113
78. Hey P.J., Twells R.C.J., Phillips M.S., Nakagawa Y., Brown S.D., Kawaguchi Y., Cox R., Xie G., Dugan V., Hammond H., Metzker M.L., Todd J.A., et al. Cloning of a novel member of the low-density lipoprotein receptor family. Gene 216 (1998) 103-111
79. Hill T.P., Spater D., Taketo M.M., Birchmeier W., and Hartmann C. Canonical Wnt/beta-catenin signaling prevents osteoblasts from differentiating into chondrocytes. Dev. Cell 8 (2005) 727-738
80. Hoang B., Moos Jr. M., Vukicevic S., and Luyten F.P. Primary structure and tissue distribution of FRZB, a novel protein related to Drosophila frizzled, suggest a role in skeletal morphogenesis. J. Biol. Chem. 271 (1996) 26,131-26,137
81. Hoang B.H., Kubo T., Healey J.H., Sowers R., Mazza B., Yang R., Huvos A.G., Meyers P.A., and Gorlick R. Expression of LDL receptor-related protein 5 (LRP5) as a novel marker for disease progression in high-grade osteosarcoma. Int. J. Cancer 109 (2004) 106-111
82. Hoeflich K.P., Luo J., Rubie E.A., Tsao M.S., Jin O., and Woodgett J.R. Requirement for glycogen synthase kinase-3beta in cell survival and NF-kappaB activation. Nature 406 (2000) 86-90
83. Holmen S.L., Giambernardi T.A., Zylstra C.R., Buckner-Berghuis B.D., Resau J.H., Hess J.F., Glatt V., Bouxsein M.L., Ai M., Warman M.L., and Williams B.O. Decreased BMD and limb deformities in mice carrying mutations in both Lrp5 and Lrp6. J. Bone Miner. Res. 19 (2004) 2033-2040
84. Holmen S.L., Zylstra C.R., Mukherjee A., Sigler R.E., Faugere M.C., Bouxsein M.L., Deng L., Clemens T.L., and Williams B.O. Essential role of beta-catenin in postnatal bone acquisition. J. Biol. Chem. 280 (2005) 21,162-21,168
85. Hovanes K., Li T.W., Munguia J.E., Truong T., Milovanovic T., Lawrence Marsh J., Holcombe R.F., and Waterman M.L. Beta-catenin-sensitive isoforms of lymphoid enhancer factor-1 are selectively expressed in colon cancer. Nat. Genet. 28 (2001) 53-57
86. Hovanes K., Li T.W., and Waterman M.L. The human LEF-1 gene contains a promoter preferentially active in lymphocytes and encodes multiple isoforms derived from alternative splicing. Nucleic Acids Res. 28 (2000) 1994-2003
87. Hsieh J.C., Kodjabachian L., Rebbert M.L., Rattner A., Smallwood P.M., Samos C.H., Nusse R., Dawid I.B., and Nathans J. A new secreted protein that binds to Wnt proteins and inhibits their activities. Nature 398 (1999) 431-436
88. Hsu S.C., Galceran J., and Grosschedl R. Modulation of transcriptional regulation by LEF-1 in response to Wnt-1 signaling and association with beta-catenin. Mol. Cell. Biol. 18 (1998) 4807-4818
89. Hu H., Hilton M.J., Tu X., Yu K., Ornitz D.M., and Long F. Sequential roles of hedgehog and Wnt signaling in osteoblast development. Development 132 (2005) 49-60
90. Huber O., Korn R., McLaughlin J., Ohsugi M., Herrmann B.G., and Kemler R. Nuclear localization of beta-catenin by interaction with transcription factor LEF-1. Mech. Dev. 59 (1996) 3-10
91. Huelsken J., Vogel R., Erdmann B., Cotsarelis G., and Birchmeier W. Beta-catenin controls hair follicle morphogenesis and stem cell differentiation in the skin. Cell 105 (2001) 533-545
92. Hussein S.M., Duff E.K., and Sirard C. Smad4 and beta-catenin co-activators functionally interact with lymphoid-enhancing factor to regulate graded expression of Msx2. J. Biol. Chem. 278 (2003) 48,805-48,814
93. Ikeda S., Kishida S., Yamamoto H., Murai H., Koyama S., and Kikuchi A. Axin, a negative regulator of the Wnt signaling pathway, forms a complex with GSK-3beta catenin and promotes GSK-3beta-dependent phosphorylation of beta-catenin. EMBO J. 17 (1998) 1371-1384
94. Itasaki N., Jones C.M., Mercurio S., Rowe A., Domingos P.M., Smith J.C., and Krumlauf R. Wise, a context-dependent activator and inhibitor of Wnt signalling. Development 130 (2003) 4295-4305
95. Jackson A., Vayssiere B., Garcia T., Newell W., Baron R., Roman-Roman S., and Rawadi G. Gene array analysis of Wnt-regulated genes in C3H10T1/2 cells. Bone 36 (2005) 585-598
96. James I.E., Kumar S., Barnes M.R., Gress C.J., Hand A.T., Dodds R.A., Connor J.R., Bradley B.R., Campbell D.A., Grabill S.E., Williams K., Blake S.M., et al. FrzB-2: A human secreted frizzled-related protein with a potential role in chondrocyte apoptosis. Osteoarthr. Cartil. 8 (2000) 452-463
97. Jamora C., DasGupta R., Kocieniewski P., and Fuchs E. Links between signal transduction, transcription and adhesion in epithelial bud development. Nature 422 (2003) 317-322
98. Jeon H., Meng W., Takagi J., Eck M.J., Springer T.A., and Blacklow S.C. Implications for familial hypercholesterolemia from the structure of the LDL receptor YWTD-EGF domain pair. Nat. Struct. Biol. 8 (2001) 499-504
99. Jiang J., and Struhl G. Regulation of the Hedgehog and Wingless signalling pathways by the F-box/WD40-repeat protein Slimb. Nature 391 (1998) 493-496
100. Johnson M.L., Gong G., Kimberling W., Recker S.M., Kimmel D.B., and Recker R.B. Linkage of a gene causing high bone mass to human chromosome 11 (11q12-13). Am. J. Hum. Genet. 60 (1997) 1326-1332
101. Kahler R.A., and Westendorf J.J. Lymphoid enhancer factor-1 and beta-catenin inhibit Runx2-dependent transcriptional activation of the osteocalcin promoter. J. Biol. Chem. 278 (2003) 11,937-11,944
102. Kalajzic I., Staal A., Yang W.P., Wu Y., Johnson S.E., Feyen J.H., Krueger W., Maye P., Yu F., Zhao Y., Kuo L., Gupta R.R., et al. Expression profile of osteoblast lineage at defined stages of differentiation. J. Biol. Chem. 280 (2005) 24,618-24,626
103. Kato M., Patel M.S., Levasseur R., Lobov I., Chang B.H., Glass II D.A., Hartmann C., Li L., Hwang T.H., Brayton C.F., Lang R.A., Karsenty G., et al. Cbfa1-independent decrease in osteoblast proliferation, osteopenia, and persistent embryonic eye vascularization in mice deficient in Lrp5, a Wnt coreceptor. J. Cell Biol. 157 (2002) 303-314
104. Kawano Y., and Kypta R. Secreted antagonists of the Wnt signalling pathway. J. Cell Sci. 116 (2003) 2627-2634
105. Klein P.S., and Melton D.A. A molecular mechanism for the effect of lithium on development. Proc. Natl. Acad. Sci. USA 93 (1996) 8455-8459
106. Kokubu C., Heinzmann U., Kokubu T., Sakai N., Kubota T., Kawai M., Wahl M.B., Galceran J., Grosschedl R., Ozono K., and Imai K. Skeletal defects in ringelschwanz mutant mice reveal that Lrp6 is required for proper somitogenesis and osteogenesis. Development 131 (2004) 5469-5480
107. Koller D.L., Rodriguez L.A., Christian J.C., Slemenda C.W., Econs M.J., Hui S.L., Morin P., Conneally P.M., Joslyn G., Curran M.E., Peacock M., Johnston C.C., et al. Linkage of a QTL contributing to normal variation in bone mineral denisty to chromosome 11q12-13. J. Bone Miner. Res. 13 (1998) 1903-1908
108. Komori T., Yagi H., Nomura S., Yamaguchi A., Sasaki K., Deguchi K., Shimizu Y., Bronson R.T., Gao Y.H., Inada M., Sato M., Okamoto R., et al. Targeted disruption of Cbfa1 results in a complete lack of bone formation owing to maturational arrest of osteoblasts. Cell 89 (1997) 755-764
109. Korinek V., Barker N., Moerer P., van Donselaar E., Huls G., Peters P.J., and Clevers H. Depletion of epithelial stem-cell compartments in the small intestine of mice lacking Tcf-4. Nat. Genet. 19 (1998) 379-383
110. Kramps T., Peter O., Brunner E., Nellen D., Froesch B., Chatterjee S., Murone M., Zullig S., and Basler K. Wnt/wingless signaling requires BCL9/legless-mediated recruitment of pygopus to the nuclear beta-catenin-TCF complex. Cell 109 (2002) 47-60
111. Kratochwil K., Dull M., Farinas I., Galceran J., and Grosschedl R. Lef1 expression is activated by BMP-4 and regulates inductive tissue interactions in tooth and hair development. Genes Dev. 10 (1996) 1382-1394
112. Krupnik V.E., Sharp J.D., Jiang C., Robison K., Chickering T.W., Amaravadi L., Brown D.E., Guyot D., Mays G., Leiby K., Chang B., Duong T., et al. Functional and structural diversity of the human Dickkopf gene family. Gene 238 (1999) 301-313
113. Kuhl M., Sheldahl L.C., Malbon C.C., and Moon R.T. Ca(2+)/calmodulin-dependent protein kinase II is stimulated by Wnt and Frizzled homologs and promotes ventral cell fates in Xenopus. J. Biol. Chem. 275 (2000) 12,701-12,711
114. Kulkarni N.H., Halladay D.L., Miles R.R., Gilbert L.M., Frolik C.A., Galvin R.J., Martin T.J., Gillespie M.T., and Onyia J.E. Effects of parathyroid hormone on Wnt signaling pathway in bone. J. Cell Biochem. 95 (2005) 1178-1190
115. Kwee M.L., Balemans W., Cleiren E., Gille J.J., Van Der Blij F., Sepers J.M., and Van Hul W. An autosomal dominant high bone mass phenotype in association with craniosynostosis in an extended family is caused by an LRP5 missense mutation. J. Bone Miner. Res. 20 (2005) 1254-1260
116. Labbe E., Letamendia A., and Attisano L. Association of Smads with lymphoid enhancer binding factor 1/T cell-specific factor mediates cooperative signaling by the transforming growth factor-beta and wnt pathways. Proc. Natl. Acad. Sci. USA 97 (2000) 8358-8363
117. Lacey D.L., Timms E., Tan H.L., Kelley M.J., Dunstan C.R., Burgess T., Elliott R., Colombero A., Elliott G., Scully S., Hsu H., Sullivan J., et al. Osteoprotegerin ligand is a cytokine that regulates osteoclast differentiation and activation. Cell 93 (1998) 165-176
118. Levanon D., Goldstein R.E., Bernstein Y., Tang H., Goldenberg D., Stifani S., Paroush Z., and Groner Y. Transcriptional repression by AML1 and LEF-1 is mediated by the TLE/Groucho corepressors. Proc. Natl. Acad. Sci. USA 95 (1998) 11,590-11,595
119. Li L., Yuan H., Weaver C.D., Mao J., Farr III G.H., Sussman D.J., Jonkers J., Kimelman D., and Wu D. Axin and Frat1 interact with dvl and GSK, bridging Dvl to GSK in Wnt-mediated regulation of LEF-1. EMBO J. 18 (1999) 4233-4240
120. Li X., Liu P., Liu W., Maye P., Zhang J., Zhang Y., Hurley M., Guo C., Boskey A., Sun L., Harris S.E., Rowe D.W., et al. Dkk2 has a role in terminal osteoblast differentiation and mineralized matrix formation. Nat. Genet. 37 (2005) 945-952
121. Li X., Zhang Y., Kang H., Liu W., Liu P., Zhang J., Harris S.E., and Wu D. Sclerostin binds to LRP5/6 and antagonizes canonical Wnt signaling. J. Biol. Chem. 280 (2005) 19,883-19,887
122. Li Y., Lu W., He X., Schwartz A.L., and Bu G. LRP6 expression promotes cancer cell proliferation and tumorigenesis by altering beta-catenin subcellular distribution. Oncogene 23 (2004) 9129-9135
123. Lin X., and Perrimon N. Dally cooperates with Drosophila Frizzled 2 to transduce Wingless signalling. Nature 400 (1999) 281-284
124. Little R.D., Carulli J.P., Del Mastro R.G., Dupuis J., Osborne M., Folz C., Manning S.P., Swain P.M., Zhao S.C., Eustace B., Lappe M.M., Spitzer L., et al. A mutation in the LDL receptor-related protein 5 gene results in the autosomal dominant high-bone-mass trait. Am. J. Hum. Genet. 70 (2002) 11-19
125. Liu C., Li Y., Semenov M., Han C., Baeg G.H., Tan Y., Zhang Z., Lin X., and He X. Control of beta-catenin phosphorylation/degradation by a dual-kinase mechanism. Cell 108 (2002) 837-847
126. Love J.J., Li X., Case D.A., Giese K., Grosschedl R., and Wright P.E. Structural basis for DNA bending by the architectural transcription factor LEF-1. Nature 376 (1995) 791-795
127. Lucas J.J., Hernandez F., Gomez-Ramos P., Moran M.A., Hen R., and Avila J. Decreased nuclear beta-catenin, tau hyperphosphorylation and neurodegeneration in GSK-3beta conditional transgenic mice. EMBO J. 20 (2001) 27-39
128. Luo Q., Kang Q., Si W., Jiang W., Park J.K., Peng Y., Li X., Luu H.H., Luo J., Montag A.G., Haydon R.C., and He T.C. 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 (2004) 55,958-55,968
129. Mansukhani A., Ambrosetti D., Holmes G., Cornivelli L., and Basilico C. Sox2 induction by FGF and FGFR2 activating mutations inhibits Wnt signaling and osteoblast differentiation. J. Cell Biol. 168 (2005) 1065-1076
130. Mao B., and Niehrs C. Kremen2 modulates Dickkopf2 activity during Wnt/LRP6 signaling. Gene 302 (2003) 179-183
131. Mao B., Wu W., Davidson G., Marhold J., Li M., Mechler B.M., Delius H., Hoppe D., Stannek P., Walter C., Glinka A., and Niehrs C. Kremen proteins are Dickkopf receptors that regulate Wnt/beta-catenin signalling. Nature 417 (2002) 664-667
132. Mao B., Wu W., Li Y., Hoppe D., Stannek P., Glinka A., and Niehrs C. LDL-receptor-related protein 6 is a receptor for Dickkopf proteins. Nature 411 (2001) 321-325
133. Mao J., Wang J., Liu B., Pan W., Farr III G.H., Flynn C., Yuan H., Takada S., Kimelman D., Li L., and Wu D. Low-density lipoprotein receptor-related protein-5 binds to Axin and regulates the canonical Wnt signaling pathway. Mol. Cell 7 (2001) 801-809
134. Mariadason J.M., Bordonaro M., Aslam F., Shi L., Kuraguchi M., Velcich A., and Augenlicht L.H. Down-regulation of beta-catenin TCF signaling is linked to colonic epithelial cell differentiation. Cancer Res. 61 (2001) 3465-3471
135. May L.G., and Gay C.V. Multiple G-protein involvement in parathyroid hormone regulation of acid production by osteoclasts. J. Cell Biochem. 64 (1997) 161-170
136. Mbalaviele G., Sheikh S., Stains J.P., Salazar V.S., Cheng S.L., Chen D., and Civitelli R. Beta-catenin and BMP-2 synergize to promote osteoblast differentiation and new bone formation. J. Cell Biochem. 94 (2005) 403-418
137. Merrill B.J., Pasolli H.A., Polak L., Rendl M., Garcia-Garcia M.J., Anderson K.V., and Fuchs E. Tcf3: A transcriptional regulator of axis induction in the early embryo. Development 131 (2004) 263-274
138. Mi K., and Johnson G.V. Role of the intracellular domains of LRP5 and LRP6 in activating the Wnt canonical pathway. J. Cell Biochem. 95 (2005) 328-338
139. Miller J.R. The Wnts. Genome Biol. 3 (2001) 1-15
140. Miyagishi M., Fujii R., Hatta M., Yoshida E., Araya N., Nagafuchi A., Ishihara S., Nakajima T., and Fukamizu A. Regulation of Lef-mediated transcription and p53-dependent pathway by associating beta-catenin with CBP/p300. J. Biol. Chem. 275 (2000) 35170-35175
141. Molenaar M., van de Wetering M., Oosterwegel M., Peterson-Maduro J., Godsave S., Korinek V., Roose J., Destree O., and Clevers H. XTcf-3 transcription factor mediates beta-catenin-induced axis formation in Xenopus embryos. Cell 86 (1996) 391-399
142. Monaghan H., Bubb V.J., Sirimujalin R., Millward-Sadler S.J., and Salter D.M. Adenomatous polyposis coli (APC), beta-catenin, and cadherin are expressed in human bone and cartilage. Histopathology 39 (2001) 611-619
143. Moon R.T., Bowerman B., Boutros M., and Perrimon N. The promise and perils of Wnt signaling through beta-catenin. Science 296 (2002) 1644-1646
144. Mukhopadhyay M., Shtrom S., Rodriguez-Esteban C., Chen L., Tsukui T., Gomer L., Dorward D.W., Glinka A., Grinberg A., Huang S.P., Niehrs C., Belmonte J.C., and Westphal H. Dickkopf1 is required for embryonic head induction and limb morphogenesis in the mouse. Dev. Cell 1 (2001) 423-434
145. Mundy G.R., Garrett R., Harris S., Chan J.K., Chen D., Rossini G., Boyce B., Zhao M., and Gutierrez G. Stimulation of bones formation in vitro and in rodents by statins. Science 286 (1999) 1946-1949
146. Nakamura T., Aoki S., Kitajima K., Takahashi T., and Matsumoto K. Molecular cloning and characterization of Kremen, a novel kringle-containing transmembrane protein. Biochim. Biophys. Acta 1518 (2001) 63-72
147. Nakashima K., Zhou X., Kunkel G., Zhang Z., Deng J.M., Behringer R.R., and de Crombrugghe B. The novel zinc finger-containing transcription factor osterix is required for osteoblast differentiation and bone formation. Cell 108 (2002) 17-29
148. Nishita M., Hashimoto M.K., Ogata S., Laurent M.N., Ueno N., Shibuya H., and Cho K.W. Interaction between Wnt and TGF-beta signalling pathways during formation of Spemann's organizer. Nature 403 (2000) 781-785
149. Noordermeer J., Klingensmith J., Perrimon N., and Nusse R. Dishevelled and armadillo act in the wingless signalling pathway in Drosophila. Nature 367 (1994) 80-83
150. Ohnaka K., Tanabe M., Kawate H., Nawata H., and Takayanagi R. Glucocorticoid suppresses the canonical Wnt signal in cultured human osteoblasts. Biochem. Biophys. Res. Commun. 329 (2005) 177-181
151. Ohnaka K., Taniguchi H., Kawate H., Nawata H., and Takayanagi R. Glucocorticoid enhances the expression of dickkopf-1 in human osteoblasts: Novel mechanism of glucocorticoid-induced osteoporosis. Biochem. Biophys. Res. Commun. 318 (2004) 259-264
152. Oishi I., Suzuki H., Onishi N., Takada R., Kani S., Ohkawara B., Koshida I., Suzuki K., Yamada G., Schwabe G.C., Mundlos S., and Shibuya H. The receptor tyrosine kinase Ror2 is involved in non-canonical Wnt5a/JNK signalling pathway. Genes Cells 8 (2003) 645-654
153. Oldridge M., Fortuna A.M., Maringa M., Propping P., Mansour S., Pollitt C., DeChiara T.M., Kimble R.B., Valenzuela D.M., Yancopoulos G.D., and Wilkie A.O. Dominant mutations in ROR2, encoding an orphan receptor tyrosine kinase, cause brachydactyly type B. Nat. Genet. 24 (2000) 275-278
154. Oosterwegel M., van de Wetering M., and Clevers H. HMG box proteins in early T-cell differentiation. Thymus 22 (1993) 67-81
155. Oosterwegel M., van de Wetering M., Timmerman J., Kruisbeek A., Destree O., Meijlink F., and Clevers H. Differential expression of the HMG box factors TCF-1 and LEF-1 during murine embryogenesis. Development 118 (1993) 439-448
156. Orford K., Crockett C., Jensen J.P., Weissman A.M., and Byers S.W. Serine phosphorylation-regulated ubiquitination and degradation of beta-catenin. J. Biol. Chem. 272 (1997) 24735-24738
157. Oshima T., Abe M., Asano J., Hara T., Kitazoe K., Sekimoto E., Tanaka Y., Shibata H., Hashimoto T., Ozaki S., Kido S., Inoue D., et al. Myeloma cells suppress bone formation by secreting a soluble Wnt inhibitor, sFRP-2. Blood 106 (2005) 3160-3165
158. Otto F., Thornell A.P., Crompton T., Denzel A., Gilmour K.C., Rosewell I.R., Stamp G.W., Beddington R.S., Mundlos S., Olsen B.R., Selby P.B., and Owen M.J. Cbfa1, a candidate gene for cleidocranial dysplasia syndrome, is essential for osteoblast differentiation and bone development. Cell 89 (1997) 765-771
159. Parr B.A., and McMahon A.P. Dorsalizing signal Wnt-7a required for normal polarity of D-V and A-P axes of mosue limb. Nature 374 (1995) 350-353
160. Payre F., Vincent A., and Carreno S. Ovo/svb integrates Wingless and DER pathways to control epidermis differentiation. Nature 400 (1999) 271-275
161. Peifer M., and Polakis P. Wnt signaling in oncogenesis and embryogenesis-a look outside the nucleus. Science 287 (2000) 1606-1609
162. Peifer M., Sweeton D., Casey M., and Wieschaus E. Wingless signal and zeste-white 3 kinase trigger opposing changes in the intracellular distribution of armadillo. Development 120 (1994) 369-380
163. Piccolo S., Agius E., Leyns L., Bhattacharyya S., Grunz H., Bouwmeester T., and De Robertis E.M. The head inducer Cerberus is a multifunctional antagonist of Nodal, BMP and Wnt signals. Nature 397 (1999) 707-710
164. Piepenburg O., Vorbruggen G., and Jackle H. Drosophila segment borders result from unilateral repression of hedgehog activity by wingless signaling. Mol. Cell 6 (2000) 203-209
165. Pierce S.B., and Kimelman D. Overexpression of Xgsk-3 disrupts anterior ectodermal patterning in Xenopus. Dev. Biol. 175 (1996) 256-264
166. Pinson K.I., Brennan J., Monkley S., Avery B.J., and Skarnes W.C. An LDL-receptor-related protein mediates Wnt signalling in mice. Nature 407 (2000) 535-538
167. Prieve M.G., and Waterman M.L. Nuclear localization and formation of beta-catenin-lymphoid enhancer factor 1 complexes are not sufficient for activation of gene expression. Mol. Cell. Biol. 19 (1999) 4503-4515
168. Qi H., Aguiar D.J., Williams S.M., La Pean A., Pan W., and Verfaillie C.M. Identification of genes responsible for osteoblast differentiation from human mesodermal progenitor cells. Proc. Natl. Acad. Sci. USA 100 (2003) 3305-3310
169. Rawadi G., Vayssiere B., Dunn F., Baron R., and Roman-Roman S. BMP-2 controls alkaline phosphatase expression and osteoblast mineralization by a Wnt autocrine loop. J. Bone Miner. Res. 18 (2003) 1842-1853
170. Reddi A.H. Bone morphogenetic proteins: An unconventional approach to isolation of first mammalian morphogens. Cytokines Growth Factor Rev. 8 (1997) 11-20
171. Riggs B.L., Khosla S., and Melton III L.J. A unitary model for involutional osteoporosis: Estrogen deficiency causes both type I and type II osteoporosis in postmenopausal women and contributes to bone loss in aging men. J. Bone Miner. Res. 13 (1998) 763-773
172. Roman-Roman S., Shi D.L., Stiot V., Hay E., Vayssiere B., Garcia T., Baron R., and Rawadi G. Murine Frizzled-1 behaves as an antagonist of the canonical Wnt/beta-catenin signaling. J. Biol. Chem. 279 (2004) 5725-5733
173. Roose J., Huls G., van Beest M., Moerer P., van der Horn K., Goldschmeding R., Logtenberg T., and Clevers H. Synergy between tumor suppressor APC and the beta-catenin-Tcf4 target Tcf1. Science 285 (1999) 1923-1926
174. Ross S.E., Hemati N., Longo K.A., Bennett C.N., Lucas P.C., Erickson R.L., and MacDougald O.A. Inhibition of adipogenesis by Wnt signaling. Science 289 (2000) 950-953
175. Schmid C., Guler H.P., Rowe D., and Froesch E.R. Insulin-like growth factor I regulates type I procollagen messenger ribonucleic acid steady state levels in bone of rats. Endocrinology 125 (1989) 1575-1580
176. Schwabe G.C., Tinschert S., Buschow C., Meinecke P., Wolff G., Gillessen-Kaesbach G., Oldridge M., Wilkie A.O., Komec R., and Mundlos S. Distinct mutations in the receptor tyrosine kinase gene ROR2 cause brachydactyly type B. Am. J. Hum. Genet. 67 (2000) 822-831
177. Semenov M., Tamai K., and He X. SOST is a ligand for LRP5/LRP6 and a Wnt signaling inhibitor. J. Biol. Chem. 280 (2005) 26,770-26,775
178. Semenov M.V., Tamai K., Brott B.K., Kuhl M., Sokol S., and He X. Head inducer Dickkopf-1 is a ligand for Wnt coreceptor LRP6. Curr. Biol. 12 (2001) 951-961
179. Shibamoto S., Winer J., Williams M., and Polakis P. A blockade in Wnt signaling is activated following the differentiation of F9 teratocarcinoma cells. Exp. Cell Res. 292 (2004) 11-20
180. Shulman J.M., Perrimon N., and Axelrod J.D. Frizzled signaling and the developmental control of cell polarity. Trends Genet. 14 (1998) 452-458
181. Siegfried E., Chou T.B., and Perrimon N. Wingless signaling acts through zeste-white 3, the Drosophila homolog of glycogen synthase kinase-3, to regulate engrailed and establish cell fate. Cell 71 (1992) 1167-1179
182. Siegfried E., Wilder E.L., and Perrimon N. Components of wingless signalling in Drosophila. Nature 367 (1994) 76-80
183. Simonet S., Lacey D.L., Dunstan C.R., Kelley M., Chang M.S., Luthy R., Nguyen H., Wooden S., Bennett T., and Boone T. Osteoprotegerin: A novel secreted protein involved in the regulation of bone density. Cell 89 (1997) 309-319
184. Slusarski D.C., Yang-Snyder J., Busa W.B., and Moon R.T. Modulation of embryonic intracellular Ca2+ signaling by Wnt-5A. Dev. Biol. 182 (1997) 114-120
185. Smith E., Coetzee G.A., and Frenkel B. Glucocorticoids inhibit cell cycle progression in differentiating osteoblasts via glycogen synthase kinase-3beta. J. Biol. Chem. 277 (2002) 18,191-18,197
186. Smith E., and Frenkel B. Glucocorticoids inhibit the transcriptional activity of LEF/TCF in differentiating osteoblasts in a glycogen synthase kinase-3beta-dependent and -independent manner. J. Biol. Chem. 280 (2005) 2,388-2,394
187. Stambolic V., Ruel L., and Woodgett J.R. Lithium inhibits glycogen synthase kinase-3 activity and mimics wingless signalling in intact cells. Curr. Biol. 6 (1996) 1664-1668
188. Stambolic V., and Woodgett J.R. Mitogen inactivation of glycogen synthase kinase-3 beta in intact cells via serine 9 phosphorylation. Biochem. J. 303 Pt. 3 (1994) 701-704
189. Sun Y., Kolligs F.T., Hottiger M.O., Mosavin R., Fearon E.R., and Nabel G.J. Regulation of beta-catenin transformation by the p300 transcriptional coactivator. Proc. Natl. Acad. Sci. USA 97 (2000) 12,613-12,618
190. Takada S., Stark K.L., Shea M.J., Vassileva G., McMahon J.A., and McMahon A.P. Wnt-3a regulates somite and tailbud formation in the mouse embryo. Genes Development 8 (1994) 174-189
191. Tamai K., Semenov M., Kato Y., Spokony R., Liu C., Katsuyama Y., Hess F., Saint-Jeannet J.P., and He X. LDL-receptor-related proteins in Wnt signal transduction. Nature 407 (2000) 530-535
192. Tamai K., Zeng X., Liu C., Zhang X., Harada Y., Chang Z., and He X. A mechanism for Wnt coreceptor activation. Mol. Cell 13 (2004) 149-156
193. Tetsu O., and McCormick F. Beta-catenin regulates expression of cyclin D1 in colon carcinoma cells. Nature 398 (1999) 422-426
194. Thomas K.R., and Capecchi M.R. Targeted disruption of the murine int-1 proto-oncogene resulting in severe abnormalities in midbrain and cerebellar development. Nature 346 (1990) 847-850
195. Tian E., Zhan F., Walker R., Rasmussen E., Ma Y., Barlogie B., and Shaughnessy Jr. J.D. The role of the Wnt-signaling antagonist DKK1 in the development of osteolytic lesions in multiple myeloma. N. Engl. J. Med. 349 (2003) 2483-2494
196. Toomes C., Bottomley H.M., Jackson R.M., Towns K.V., Scott S., Mackey D.A., Craig J.E., Jiang L., Yang Z., Trembath R., Woodruff G., Gregory-Evans C.Y., et al. Mutations in LRP5 or FZD4 underlie the common familial exudative vitreoretinopathy locus on chromosome 11q. Am. J. Hum. Genet. 74 (2004) 721-730
197. Travis A., Amsterdam A., Belanger C., and Grosschedl R. LEF-1, a gene encoding a lymphoid-specific protein with an HMG domain, regulates T-cell receptor alpha enhancer function [corrected]. Genes Dev. 5 (1991) 880-894
198. Tsuda M., Kamimura K., Nakato H., Archer M., Staatz W., Fox B., Humphrey M., Olson S., Futch T., Kaluza V., Siegfried E., Stam L., and Selleck S.B. The cell-surface proteoglycan Dally regulates Wingless signalling in Drosophila. Nature 400 (1999) 276-280
199. Tuan R.S. Cellular signaling in developmental chondrogenesis: N-cadherin, Wnts, and BMP-2. J. Bone Joint Surg. Am. 85-A Suppl. 2 (2003) 137-141
200. Tutter A.V., Fryer C.J., and Jones K.A. Chromatin-specific regulation of LEF-1-beta-catenin transcription activation and inhibition in vitro. Genes Dev. 15 (2001) 3342-3354
201. Tylzanowski P., Bossuyt W., Thomas J.T., and Luyten F.P. Characterization of Frzb-Cre transgenic mouse. Genesis 40 (2004) 200-204
202. Umbhauer M., Djiane A., Goisset C., Penzo-Mendez A., Riou J.F., Boucaut J.C., and Shi D.L. The C-terminal cytoplasmic Lys-thr-X-X-X-Trp motif in frizzled receptors mediates Wnt/beta-catenin signalling. EMBO J. 19 (2000) 4944-4954
203. Vaes B.L., Dechering K.J., Feijen A., Hendriks J.M., Lefevre C., Mummery C.L., Olijve W., van Zoelen E.J., and Steegenga W.T. Comprehensive microarray analysis of bone morphogenetic protein 2-induced osteoblast differentiation resulting in the identification of novel markers for bone development. J. Bone Miner. Res. 17 (2002) 2106-2118
204. Vaes B.L., Dechering K.J., van Someren E.P., Hendriks J.M., van de Ven C.J., Feijen A., Mummery C.L., Reinders M.J., Olijve W., van Zoelen E.J., and Steegenga W.T. Microarray analysis reveals expression regulation of Wnt antagonists in differentiating osteoblasts. Bone 36 (2005) 803-811
205. van Bokhoven H., Celli J., Kayserili H., van Beusekom E., Balci S., Brussel W., Skovby F., Kerr B., Percin E.F., Akarsu N., and Brunner H.G. Mutation of the gene encoding the ROR2 tyrosine kinase causes autosomal recessive robinow syndrome. Nat. Genet. 25 (2000) 423-426
206. Van de Wetering M., Castrop J., Korinek V., and Clevers H. Extensive alternative splicing and dual promoter usage generate Tcf-1 protein isoforms with differential transcription control properties. Mol. Cell. Biol. 16 (1996) 745-752
207. van de Wetering M., Cavallo R., Dooijes D., van Beest M., van Es J., Loureiro J., Ypma A., Hursh D., Jones T., Bejsovec A., Peifer M., Mortin M., and Clevers H. Armadillo coactivates transcription driven by the product of the Drosophila segment polarity gene dTCF. Cell 88 (1997) 789-799
208. van de Wetering M., and Clevers H. Sequence-specific interaction of the HMG box proteins TCF-1 and SRY occurs within the minor groove of a Watson-Crick double helix. EMBO J. 11 (1992) 3039-3044
209. van Genderen C., Okamura R.M., Farinas I., Quo R.G., Parslow T.G., Bruhn L., and Grosschedl R. Development of several organs that require inductive epithelial- mesenchymal interactions is impaired in LEF-1-deficient mice. Genes Dev. 8 (1994) 2691-2703
210. Van Wesenbeeck L., Cleiren E., Gram J., Beals R.K., Benichou O., Scopelliti D., Key L., Renton T., Bartels C., Gong Y., Warman M.L., De Vernejoul M.C., et al. Six novel missense mutations in the LDL receptor-related protein 5 (LRP5) gene in different conditions with an increased bone density. Am. J. Hum. Genet. 72 (2003) 763-771
211. Veeman M.T., Axelrod J.D., and Moon R.T. A second canon: Functions and mechanisms of beta-catenin-independent Wnt signaling. Dev. Cell 5 (2003) 367-377
212. Verbeek S., Izon D., Hofhuis F., Robanus-Maandag E., te Riele H., van de Wetering M., Oosterwegel M., Wilson A., MacDonald H.R., and Clevers H. An HMG-box-containing T-cell factor required for thymocyte differentiation. Nature 374 (1995) 70-74
213. Wang Q.M., Fiol C.J., DePaoli-Roach A.A., and Roach P.J. Glycogen synthase kinase-3 beta is a dual specificity kinase differentially regulated by tyrosine and serine/threonine phosphorylation. J. Biol. Chem. 269 (1994) 14,566-14,574
214. Wehrli M., Dougan S.T., Caldwell K., O'Keefe L., Schwartz S., and Vaizel-Ohayon D. Arrow encodes an LDL-receptor-related protein essential for Wingless signalling. Nature 407 (2000) 527-530
215. Westendorf J.J., Kahler R.A., and Schroeder T.M. Wnt signaling in osteoblasts and bone diseases. Gene 341 (2004) 19-39
216. Willert J., Epping M., Pollack J.R., Brown P.O., and Nusse R. A transcriptional response to Wnt protein in human embryonic carcinoma cells. BMC Dev. Biol. 2 (2002) 8
217. Willert K., Brown J.D., Danenberg E., Duncan A.W., Weissman I.L., Reya T., Yates III J.R., and Nusse R. Wnt proteins are lipid-modified and can act as stem cell growth factors. Nature 423 (2003) 448-452
218. Winkler D.G., Sutherland M.S., Ojala E., Turcott E., Geoghegan J.C., Shpektor D., Skonier J.E., Yu C., and Latham J.A. Sclerostin inhibition of Wnt-3a-induced C3H10T1/2 cell differentiation is indirect and mediated by bone morphogenetic proteins. J. Biol. Chem. 280 (2005) 2498-2502
219. Wodarz A., and Nusse R. Mechanisms of Wnt signaling in development. Annu. Rev. Cell Dev. Biol. 14 (1998) 59-88
220. Wong B.R., Josien R., and Choi Y. TRANCE is a TNF family member that regulates dendritic cell and osteoclast function. J. Leukoc. Biol. 65 (1999) 715-724
221. Wu W., Glinka A., Delius H., and Niehrs C. Mutual antagonism between dickkopf1 and dickkopf2 regulates Wnt/beta-catenin signalling. Curr. Biol. 10 (2000) 1611-1614
222. Yang X., Matsuda K., Bialek P., Jacquot S., Masuoka H.C., Schinke T., Li L., Brancorsini S., Sassone-Corsi P., Townes T.M., Hanauer A., and Karsenty G. ATF4 is a substrate of RSK2 and an essential regulator of osteoblast biology: Implication for Coffin-Lowry Syndrome. Cell 117 (2004) 387-398
223. Yang X., van Beest M., Clevers H., Jones T., Hursh D.A., and Mortin M.A. Decapentaplegic is a direct target of dTcf repression in the Drosophila visceral mesoderm. Development 127 (2000) 3695-3702
224. Yang Y. Wnts and wing: Wnt signaling in vertebrate limb development and musculoskeletal morphogenesis. Birth Defects Res. C Embryo Today 69 (2003) 305-317
225. Yang Y., and Niswander L. Interaction between the signaling molecules WNT7a and SHH during vertebrate limb development: Dorsal signals regulate anteroposterior patterning. Cell 80 (1995) 939-947
226. Yasuda H., Shima N., Nakagawa N., Yamaguchi K., Kinosaki M., Mochizuki S., Tomoyasu A., Yano K., Goto M., Murakami A., Tsuda E., Morinaga T., et al. Osteoclast differentiation factor is a ligand for osteoprotegerin/osteoclastogenesis-inhibitory factor and is identical to TRANCE/RANKL. Proc. Natl. Acad. Sci. USA 95 (1998) 3597-3602
227. Yoshida H., Hayashi S., Kunisada T., Ogawa M., Nishikawa S., Okamura H., Sudo T., Schulz L.D., and Nishikawa S. The murine mutation osteopetrosis is in the coding region of the macrophage colony stimulating factor gene. Nature 16 (1990) 391-396
228. Yoshida Y., Tanaka S., Umemori H., Minowa O., Usui M., Ikematsu N., Hosoda E., Imamura T., Kuno J., Yamashita T., Miyazono K., Noda M., et al. Negative regulation of BMP/Smad signaling by Tob in osteoblasts. Cell 103 (2000) 1085-1097
229. Yu H.M., Jerchow B., Sheu T.J., Liu B., Costantini F., Puzas J.E., Birchmeier W., and Hsu W. The role of Axin2 in calvarial morphogenesis and craniosynostosis. Development 132 (2005) 1995-2005
230. Zhang Y., Wang Y., Li X., Zhang J., Mao J., Li Z., Zheng J., Li L., Harris S., and Wu D. The LRP5 high-bone-mass G171V mutation disrupts LRP5 interaction with Mesd. Mol. Cell. Biol. 24 (2004) 4677-4684
231. Zhou P., Byrne C., Jacobs J., and Fuchs E. Lymphoid enhancer factor 1 directs hair follicle patterning and epithelial cell fate. Genes Dev. 9 (1995) 700-713