• 1
    Orwoll ES, Bauer DC, Vogt TM, Fox KM 1996 Axial bone mass in older women. Study of Osteoporotic Fractures Research Group. Ann Intern Med 124: 187196.
  • 2
    Orwoll ES, Klein RF 1995 Osteoporosis in men. Endocr Rev 16: 87116.
  • 3
    Vanderschueren D, Vandenput L, Boonen S, Lindberg MK, Bouillon R, Ohlsson C 2004 Androgens and bone. Endocr Rev 25: 389425.
  • 4
    Colvard DS, Eriksen EF, Keeting PE, Wilson EM, Lubahn DB, French FS, Riggs BL, Spelsberg TC 1989 Identification of androgen receptors in normal human osteoblast-like cells. Proc Natl Acad Sci USA 86: 854857.
  • 5
    Kasperk C, Helmboldt A, Borcsok I, Heuthe S, Cloos O, Niethard F, Ziegler R 1997 Skeletal site-dependent expression of the androgen receptor in human osteoblastic cell populations. Calcif Tissue Int 61: 464473.
  • 6
    Abu EO, Horner A, Kusec V, Triffitt JT, Compston JE 1997 The localization of androgen receptors in human bone. J Clin Endocrinol Metab 82: 34933497.
  • 7
    van der Eerden BC, van Til NP, Brinkmann AO, Lowik CW, Wit JM, Karperien M 2002 Gender differences in expression of androgen receptor in tibial growth plate and metaphyseal bone of the rat. Bone 30: 891896.
  • 8
    Turner RT, Wakley GK, Hannon KS 1990 Differential effects of androgens on cortical bone histomorphometry in gonadectomized male and female rats. J Orthop Res 8: 612617.
  • 9
    Vanderschueren D, Jans I, van Herck E, Moermans K, Verhaeghe J, Bouillon R 1994 Time-related increase of biochemical markers of bone turnover in androgen-deficient male rats. Bone Miner 26: 123131.
  • 10
    Vanderschueren D, Van Herck E, Suiker AM, Visser WJ, Schot LP, Bouillon R 1992 Bone and mineral metabolism in aged male rats: Short and long term effects of androgen deficiency. Endocrinology 130: 29062916.
  • 11
    Kasperk CH, Wergedal JE, Farley JR, Linkhart TA, Turner RT, Baylink DJ 1989 Androgens directly stimulate proliferation of bone cells in vitro. Endocrinology 124: 15761578.
  • 12
    Kasperk CH, Wakley GK, Hierl T, Ziegler R 1997 Gonadal and adrenal androgens are potent regulators of human bone cell metabolism in vitro. J Bone Miner Res 12: 464471.
  • 13
    Huber DM, Bendixen AC, Pathrose P, Srivastava S, Dienger KM, Shevde NK, Pike JW 2001 Androgens suppress osteoclast formation induced by RANKL and macrophage-colony stimulating factor. Endocrinology 142: 38003808.
  • 14
    Chen Q, Kaji H, Sugimoto T, Chihara K 2001 Testosterone inhibits osteoclast formation stimulated by parathyroid hormone through androgen receptor. FEBS Lett 491: 9193.
  • 15
    Pederson L, Kremer M, Judd J, Pascoe D, Spelsberg TC, Riggs BL, Oursler MJ 1999 Androgens regulate bone resorption activity of isolated osteoclasts in vitro. Proc Natl Acad Sci USA 96: 505510.
  • 16
    Kawano H, Sato T, Yamada T, Matsumoto T, Sekine K, Watanabe T, Nakamura T, Fukuda T, Yoshimura K, Yoshizawa T, Aihara K, Yamamoto Y, Nakamichi Y, Metzger D, Chambon P, Nakamura K, Kawaguchi H, Kato S 2003 Suppressive function of androgen receptor in bone resorption. Proc Natl Acad Sci USA 100: 94169421.
  • 17
    Mason RA, Morris HA 1997 Effects of dihydrotestosterone on bone biochemical markers in sham and oophorectomized rats. J Bone Miner Res 12: 14311437.
  • 18
    Rennie PS, Bruchovsky N, Leco KJ, Sheppard PC, McQueen SA, Cheng H, Snoek R, Hamel A, Bock ME, MacDonald BS, Nickel BE, Chang C, Liao S, CaHini PA, Matusik RJ 1993 Characterization of two cis-acting DNA elements involved in the androgen regulation of the probasin gene. Mol Endocrinol 7: 2336.
  • 19
    Brinkmann AO, Blok LJ, de Ruiter PE, Doesburg P, Steketee K, Berrevoets CA, Trapman J 1999 Mechanisms of androgen receptor activation and function. J Steroid Biochem Mol Biol 69: 307313.
  • 20
    Kousteni S, Bellido T, Plotkin LI, O'Brien CA, Bodenner DL, Han L, Han K, DiGregorio GB, Katzenellenbogen JA, Katzenellenbogen BS, Roberson PK, Weinstein RS, Jilka RL, Manolagas SC 2001 Nongenotropic, sex-nonspecific signaling through the estrogen or androgen receptors: Dissociation from transcriptional activity. Cell 104: 719730.
  • 21
    Heinlein CA, Chang C 2002 The roles of androgen receptors and androgen-binding proteins in nongenomic androgen actions. Mol Endocrinol 16: 21812187.
  • 22
    MacLean HE, Warne GL, Zajac JD 1995 Defects of androgen receptor function: From sex reversal to motor neurone disease. Mol Cell Endocrinol 112: 133141.
  • 23
    Notini AJ, Davey RA, McManus JF, Bate KL, Zajac JD 2005 Genomic actions of the androgen receptor are required for normal male sexual differentiation in a mouse model. J Mol Endocrinol 35: 547555.
  • 24
    Kousteni S, Chen JR, Bellido T, Han L, Ali AA, O'Brien CA, Plotkin L, Fu Q, Mancino AT, Wen Y, Vertino AM, Powers CC, Stewart SA, Ebert R, Parfitt AM, Weinstein RS, Jilka RL, Manolagas SC 2002 Reversal of bone loss in mice by nongenotropic signaling of sex steroids. Science 298: 843846.
  • 25
    Syed F, Khosla S 2005 Mechanisms of sex steroid effects on bone. Biochem Biophys Res Commun 328: 688696.
  • 26
    Liu F, Woitge HW, Braut A, Kronenberg MS, Lichtler AC, Mina M, Kream BE 2004 Expression and activity of osteoblast-targeted Cre recombinase transgenes in murine skeletal tissues. Int J Dev Biol 48: 645653.
  • 27
    Davey RA, Hahn CN, May BK, Morris HA 2000 Osteoblast gene expression in rat long bones: Effects of ovariectomy and dihydrotestosterone on mRNA levels. Calcif Tissue Int 67: 7579.
  • 28
    Baldock PA, Moore RJ, Durbridge TC, Morris HA 1999 Comparison of three methods for estimation of bone resorption following ovariectomy in the distal femur and the proximal tibia of the rat. Bone 24: 597602.
  • 29
    Page K 1997 Bone and preparation of bone sections. Bancroft AS (ed.) Theory and Practice of Histological Techniques. Churchill Livingstone, London, UK, 223248.
  • 30
    Baldock PA, Sainsbury A, Couzens M, Enriquez RF, Thomas GP, Gardiner EM, Herzog H 2002 Hypothalamic Y2 receptors regulate bone formation. J Clin Invest 109: 915921.
  • 31
    Meinel L, Fajardo R, Hofmann S, Langer R, Chen J, Snyder B, Vunjak-Novakovic G, Kaplan D 2005 Silk implants for the healing of critical size bone defects. Bone 37: 688698.
  • 32
    Hildebrand T 1997 A new method for the model-independent assessment of thickness in three-dimensional images. J Microsc 185: 6775.
  • 33
    Hildebrand T, Laib A, Muller R, Dequeker J, Ruegsegger P 1999 Direct three-dimensional morphometric analysis of human cancellous bone: Microstructural data from spine, femur, iliac crest, and calcaneus. J Bone Miner Res 14: 11671174.
  • 34
    Spaliviero JA, Jimenez M, Allan CM, Handelsman DJ 2004 Luteinizing hormone receptor-mediated effects on initiation of spermatogenesis in gonadotropin-deficient (hpg) mice are replicated by testosterone. Biol Reprod 70: 3238.
  • 35
    van Casteren JI, Schoonen WG, Kloosterboer HJ 2000 Development of time-resolved immunofluorometric assays for rat follicle-stimulating hormone and luteinizing hormone and application on sera of cycling rats. Biol Reprod 62: 886894.
  • 36
    Jimenez M, Spaliviero JA, Grootenhuis AJ, Verhagen J, Allan CM, Handelsman DJ 2005 Validation of an ultrasensitive and specific immunofluorometric assay for mouse follicle-stimulating hormone. Biol Reprod 72: 7885.
  • 37
    Yarbrough WG, Quarmby VE, Simental JA, Joseph DR, Sar M, Lubahn DB, Olsen KL, French FS, Wilson EM 1990 A single base mutation in the androgen receptor gene causes androgen insensitivity in the testicular feminized rat. J Biol Chem 265: 88938900.
  • 38
    Vanderschueren D, Van Herck E, Suiker AM, Visser WJ, Schot LP, Chung K, Lucas RS, Einhorn TA, Bouillon R 1993 Bone and mineral metabolism in the androgen-resistant (testicular feminized) male rat. J Bone Miner Res 8: 801809.
  • 39
    Kuiper GG, Carlsson B, Grandien K, Enmark E, Haggblad J, Nilsson S, Gustafsson JA 1997 Comparison of the ligand binding specificity and transcript tissue distribution of estrogen receptors alpha and beta. Endocrinology 138: 863870.
  • 40
    Turner RT, Evans GL, Dobnig H 2000 The high-affinity estrogen receptor antagonist ICI 182,780 has no effect on bone growth in young male rats. Calcif Tissue Int 66: 461464.
  • 41
    Ferguson VL, Ayers RA, Bateman TA, Simske SJ 2003 Bone development and age-related bone loss in male C57BL/6J mice. Bone 33: 387398.
  • 42
    Morko J, Kiviranta R, Hurme S, Rantakokko J, Vuorio E 2005 Differential turnover of cortical and trabecular bone in transgenic mice overexpressing cathepsin K. Bone 36: 854865.
  • 43
    Yeh S, Tsai MY, Xu Q, Mu XM, Lardy H, Huang KE, Lin H, Yeh SD, Altuwaijri S, Zhou X, Xing L, Boyce BF, Hung MC, Zhang S, Gan L, Chang C 2002 Generation and characterization of androgen receptor knockout (ARKO) mice: An in vivo model for the study of androgen functions in selective tissues. Proc Natl Acad Sci USA 99: 1349813503.
  • 44
    Lindberg MK, Alatalo SL, Halleen JM, Mohan S, Gustafsson JA, Ohlsson C 2001 Estrogen receptor specificity in the regulation of the skeleton in female mice. J Endocrinol 171: 229236.
  • 45
    Moverare S, Venken K, Eriksson AL, Andersson N, Skrtic S, Wergedal J, Mohan S, Salmon P, Bouillon R, Gustafsson JA, Vanderschueren D, Ohlsson C 2003 Differential effects on bone of estrogen receptor alpha and androgen receptor activation in orchidectomized adult male mice. Proc Natl Acad Sci USA 100: 1357313578.
  • 46
    Sims NA, Clement-Lacroix P, Minet D, Fraslon-Vanhulle C, Gaillard-Kelly M, Resche-Rigon M, Baron R 2003 A functional androgen receptor is not sufficient to allow estradiol to protect bone after gonadectomy in estradiol receptor-deficient mice. J Clin Invest 111: 13191327.
  • 47
    Nagy A, Moens C, Ivanyi E, Pawling J, Gertsenstein M, Hadjantonakis AK, Pirity M, Rossant J 1998 Dissecting the role of N-myc in development using a single targeting vector to generate a series of alleles. Curr Biol 8: 661664.
  • 48
    Kulessa H, Hogan BL 2002 Generation of a loxP flanked bmp4loxP-lacZ allele marked by conditional lacZ expression. Genesis 32: 6668.
  • 49
    Holdcraft RW, Braun RE 2004 Androgen receptor function is required in Sertoli cells for the terminal differentiation of haploid spermatids. Development 131: 459467.
  • 50
    Davey RA, MacLean HE, McManus JF, Findlay DM, Zajac JD 2004 Genetically modified animal models as tools for studying bone and mineral metabolism. J Bone Miner Res 19: 882892.
  • 51
    Milatiner D, Halle D, Huerta M, Margalioth EJ, Cohen Y, Ben-Chetrit A, Gal M, Mimoni T, Eldar-Geva T 2004 Associations between androgen receptor CAG repeat length and sperm morphology. Hum Reprod 19: 14261430.
  • 52
    Masuyama A, Ouchi Y, Sato F, Hosoi T, Nakamura T, Orimo H 1992 Characteristics of steroid hormone receptors in cultured MC3T3-E1 osteoblastic cells and effect of steroid hormones on cell proliferation. Calcif Tissue Int 51: 376381.
  • 53
    Wiren KM, Zhang X, Semirale A, Price C, Woo A, Jepsen KJ 2006 Targeted Androgen Action in Mature Versus Preosteoblasts Eliminates Periosteal Stimulation and Reduces Bone Quality during Growth. J Bone Miner Res 21: S1: S5.
  • 54
    Mizuno Y, Hosoi T, Inoue S, Ikegami A, Kaneki M, Akedo Y, Nakamura T, Ouchi Y, Chang C, Orimo H 1994 Immunocytochemical identification of androgen receptor in mouse osteoclast-like multinucleated cells. Calcif Tissue Int 54: 325326.
  • 55
    Noble B, Routledge J, Stevens H, Hughes I, Jacobson W 1999 Androgen receptors in bone-forming tissue. Horm Res 51: 3136.