• 1
    Canalis E. Growth factor control of bone mass. J Cell Biochem. 2009; 108: 76977.
  • 2
    Firth SM, Baxter RC. Cellular actions of the insulin-like growth factor binding proteins. Endocr Rev. 2002; 23: 82454.
  • 3
    Wheatcroft SB, Kearney MT. IGF-dependent and IGF-I independent actions of IGF-binding protein-1 and -2: implications for metabolic homeostasis. Trends Endocrinol Metab. 2009; 20: 15362.
  • 4
    Jones JL, Clemmons DR. Insulin-like growth factors and their binding proteins: biological actions. Endocr Rev. 1995; 16: 334.
  • 5
    Hoeflich A, Nedbal S, Blum WF, Erhard M, Lahm H, Brem G, Kolb HJ, Wanke R, Wolf E. Growth inhibition in giant growth hormone transgenic mice by overexpression of insulin-like growth factor-binding protein-2. Endocrinology. 2001; 142: 186998.
  • 6
    Khosla S, Hassoun AA, Baker BK, Liu F, Zein NN, Whyte MP, Reasner CA, Nippoldt TB, Tiegs RD, Hintz RL, Conover CA. Insulin-like growth factor system abnormalities in hepatitis C-associated osteosclerosis. Potential insights into increasing bone mass in adults. J Clin Invest. 1998; 101: 216573.
  • 7
    Conover CA, Johnstone EW, Turner RT, Evans GL, Ballard FJ, Doran PM, Khosla S. Subcutaneous administration of insulin-like growth factor (IGF)-II/IGF binding protein-2 complex stimulates bone formation and prevents loss of bone mineral density in a rat model of disuse osteoporosis. Growth Horm IGF Res. 2002; 12: 17883.
  • 8
    Kawai M, Breggia AC, DeMambro VE, Shen X, Canalis E, Bouxsein ML, Beamer WG, Clemmons DR, Rosen CJ. The heparin-binding domain of IGFBP-2 has insulin-like growth factor binding-independent biologic activity in the growing skeleton. J Biol Chem. 2011; 286: 1467080.
  • 9
    DeMambro VE, Clemmons DR, Horton LG, Bouxsein ML, Wood TL, Beamer WG, Canalis E, Rosen CJ. Gender-specific changes in bone turnover and skeletal architecture in IGFBP-2 null mice. Endocrinology. 2008; 149: 205161.
  • 10
    Cantley LC. The phosphoinositide 3-kinase pathway. Science. 2002; 296: 16557.
  • 11
    Mandel CC, Ghosh-Choudhury G, Ghosh-Choudhury N. Phosphatidylinositol 3 kinase/Akt signaling relay cooperates with smad in bone morphogenetic protein-2-induced colony stimulating factor-1 (CSF-1) expression and osteoclast differentiation. Endocrinology. 2009; 150: 498998.
  • 12
    Munugalavadla V, Vemula S, Sims EC, Krishnan S, Chen S, Yan J, Li H, Niziolek PJ, Takemoto C, Robling AG, Yang FC, Kapur R. The p85alpha subunit of class IA phosphatidylinositol 3-kinase regulates the expression of multiple genes involved in osteoclast maturation and migration. Mol Cell Biol. 2008; 28: 718298.
  • 13
    Kwak HB, Lee MS, Kim HS, Cho HJ, Kim JW, Lee ZH, Oh J. Proteasome inhibitors induce osteoclast survival by activating the Akt pathway. Biochem Biophys Res Commun. 2008; 377: 16.
  • 14
    Lee SE, Woo KM, Kim SY, Kim HM, Kwack K, Lee ZH, Kim HH. The phosphatidylinositol 3-kinase, p38, and extracellular signal-regulated kinase pathways are involved in osteoclast differentiation. Bone. 2002; 30: 717.
  • 15
    Lee SE, Chung WJ, Kwak HB, Chung CH, Kwack KB, Lee ZH, Kim HH. Tumor necrosis factor-alpha supports the survival of osteoclasts through the activation of Akt and ERK. J Biol Chem. 2001; 276: 493439.
  • 16
    Russo VC, Schutt BS, Andaloro E, Ymer SI, Hoeflich A, Ranke MB, Bach LA, Werther GA. Insulin-like growth factor binding protein-2 to extracellular matrix plays a critical role in neuroblastoma cell proliferation, migration and invasion. Endocrinology. 2005; 146: 444555.
  • 17
    Wood TL, Rogler LE, Czick ME, Schuller AG, Pintar JE. Selective alterations in organ size in mice with a targeted disruption of the insulin-like growth factor binding protein-2 gene. Mol Endocrinol. 2000; 14: 147282.
  • 18
    Maile LA, Badley-Clarke J, Clemmons DR. The association between integrin-associated protein and SHPS-1 regulates insulin-like growth factor-I receptor signaling in vascular smooth muscle cells. Mol Biol Cell. 2003; 14: 351928.
  • 19
    Perks CM, Vernon EG, Rosendahl AH, Tonge D, Holly JM. IGF-II and IGFBP-2 differentially regulate PTEN in human breast cancer cells. Oncogene. 2007; 26: 596677.
  • 20
    Mehrian-Shai R, Chen CD, Shi T, Horvath S, Nelson SF, Reichardt JK, Sawyers CL. Insulin growth factor-binding protein 2 is a candidate biomarker for PTEN status and PI3K/Akt pathway activation in glioblastoma and prostate cancer. Proc Natl Acad Sci USA. 2007; 104: 55638.
  • 21
    Vazquez F, Ramaswamy S, Nakamura N, Sellers WR. Phosphorylation of the PTEN tail regulates protein stability and function. Mol Cell Biol. 2000; 20: 50108.
  • 22
    Al-Khouri AM, Ma Y, Togo SH, Williams S, Mustelin T. Cooperative phosphorylation of the tumor suppressor phosphatase and tensin homologue (PTEN) by casein kinases and glycogen synthase kinase 3beta. J Biol Chem. 2005; 280: 35192202.
  • 23
    Wang Y, Nishida S, Elalieh HZ, Long RK, Halloran BP, Bikle DD. Role of IGF-1 signaling in regulating osteoclastogenesis. J Bone Miner Res. 2006; 21: 13508.
  • 24
    Parker A, Rees C, Clarke J, Busby WH Jr, Clemmons DR. Binding of insulin-like growth factor (IGF)-binding protein-5 to smooth-muscle cell extracellular matrix is a major determinant of the cellular response to IGF-I. Mol Biol Cell. 1998; 9: 238392.
  • 25
    Yang YW, Yanagishita M, Rechler MM. Heparin inhibition of insulin-like growth factor-binding protein-3 binding to human fibroblasts and rat glioma cells: role of heparan sulfate proteoglycans. Endocrinology. 1996; 137: 436371.
  • 26
    Arai T, Busby W Jr, Clemmons DR. Binding of insulin-like growth factor (IGF) I or II to IGF-binding protein-2 enables it to bind to heparin and extracellular matrix. Endocrinology. 1996; 137: 45715.
  • 27
    Kuang Z, Yao S, Keizer DW, Wang CC, Bach LA, Forbes BE, Wallace JC, Norton RS. Structure, dynamics and heparin binding of the C-terminal domain of insulin-like growth factor-binding protein-2 (IGFBP-2). J Mol Biol. 2006; 364: 690704.
  • 28
    Torres J, Pulido R. The tumor suppressor PTEN is phosphorylated by the protein kinase CK2 at its C terminus. Implications for PTEN stability to proteasome-mediated degradation. J Biol Chem. 2001; 276: 9936.
  • 29
    Shehata M, Schnabl S, Demirtas D, Hilgarth M, Hubmann R, Ponath E, Badmya S, Lehner C, Hoelbl A, Duechler M, Gaiger A, Zielinski C, Schwarzmeier JD, Jaeger U. Reconstitution of PTEN activity by CK2 inhibitors and interference with the PI3-K/Akt cascade counteract the antiapoptotic effect of human stromal cells in chronic lymphocytic leukemia. Blood. 2010; 116: 251321.
  • 30
    Padmanabhan S, Mukhopadhyay A, Narasimhan SD, Tesz G, Czech MP, Tissenbaum HA. A PP2A regulatory subunit regulates C. Elegans insulin/IGF-1 signaling by modulating AKT-1 phosphorylation. Cell. 2009; 136: 93951.
  • 31
    Shen X, Xi G, Radhakrishnan Y, Clemmons DR. PDK1 recruitment to the SHPS-1 signaling complex enhances insulin-like growth factor-I-stimulated AKT activation and vascular smooth muscle cell survival. J Biol Chem. 2010; 285: 2941624.