Emerging Roles of Steroid-binding Globulins

Key words

Sex hormone binding globulin - non-genomic steroid effects - corticosteroid-binding globulin - androgen-binding protein - mitogen-activated protein kinase - SHBG internalization - reproductive behavior - estradiol receptors - progestin receptors

References

• 1 Selye H. The anesthetic effect of steroid hormones.  Proc Soc Exp Biol Med. 1941;  46 116-121
• 2 Morrow A L, Pace J R, Purdy R H, Paul S M. Characterization of Steroid Interactions with gamma-aminobutyric acid receptor-gated chloride channels: Evidence for multiple steroid recognition sites.  Molecular Pharmacology. 1990;  37 263-270
• 3 Bitran D, Hilvers R J, Frye C A, Erskine M S. Chronic anabolic-androgenic steroid treatment affects brain GABA(A) receptor-gated chloride ion transport.  Life Sci. 1996;  58 573-583
• 4 Towle A C, Sze P Y. Steroid binding to synaptic plasma membrane: differential binding of glucocorticoids and gonadal steroids.  J Steroid Biochem. 1983;  18 135-143
• 5 Szego C M. Mechanisms of hormone action: parallels in receptor-mediated signal propagation for steroid and peptide effectors.  Life Sci. 1984;  35 2383-2396
• 6 Pietras R J, Szego C M. Estrogen receptors in uterine plasma membrane.  J Steroid Biochem. 1979;  11 1471-1483
• 7 Pietras R J, Szego C M. Specific binding sites for oestrogen at the outer surfaces of isolated endometrial cells.  Nature. 1977;  265 69-72
• 8 Pietras R J, Szego C M. Surface modifications evoked by estradiol and diethylstilbestrol in isolated endometrial cells: evidence from lectin probes and extracellular release of lysosomal protease.  Endocrinology. 1975;  97 1445-1454
• 9 Pietras R J, Szego C M. Endometrial cell calcium and oestrogen action.  Nature. 1975;  253 357-359
• 10 Pietras R J, Szego C M. Steroid hormone-responsive, isolated endometrial cells.  Endocrinology. 1975;  96 946-954
• 11 McEwen B S. Steroid Hormone Actions on the Brain: When is the Genome Involved?.  Hormones and Behavior. 1994;  28 396-405
• 12 Schumacher M, Baulieu E E. Neurosteroids: synthesis and functions in the central and peripheral nervous systems.  Ciba Found Symp. 1995;  191 90-106
• 13 Schumacher M, Coirini H, Robert F, Guennoun R, El-Etr M. Genomic and membrane actions of progesterone: implications for reproductive physiology and behavior.  Behav Brain Res. 1999;  105 37-52
• 14 Caldwell J D. Central oxytocin and female sexual behavior.  Ann N Y Acad Sci. 1992;  652 166-179
• 15 Caldwell J D, Walker C H, O’Rourke S T, Faggin B M, Morris M, Mason G A. Analogies between oxytocin systems of the uterus and brain.  Horm Metab Res. 1996;  28 65-74
• 16 Willnow T E, Nykjaer A, Herz J. Lipoprotein receptors: new roles for ancient proteins.  Nat Cell Biol. 1999;  1 E157-E162
• 17 Sendemir E, Herbert Z, Caldwell J D, Jirikowski G F. Localization of sex hormone binding globulin in the hypothalamus of late pregnant and early lactating rats.  Hormone and Metabolic Research. 2006;  38 218-223
• 18 Herbert Z, Jirikowski G F, Petrusz P, Englof I, Caldwell J D. Distribution of androgen binding globulin in the rat hypothalamo-hypophyseal system, co-localization with oxytocin.  Brain Research. 2003;  992 151-158
• 19 Schock H W, Herbert Z, Sigusch H, Figulla H R, Jirikowski G F, Lotze U. Expression of androgen-binding protein (ABP) in human cardiac myocytes.  Hormones and Metabolic Research. 2006;  38 224-228
• 20 Gao G, Herbert Z, Kong J, Wu D, Gabrielson N, Jirikowski G F, Caldwell J D. Estradiol Control of Estradiol Binding Proteins and Levels in the Medial Preoptic Area, Medial Hypothalamus and Pituitary.  Neuroendocrinology. 2003;  78 61-71
• 21 Selva D M, Hammond G L. Human Sex Hormone Binding Globulin is expressed in Testicular Germ Cells and not in Sertoli Cells.  Hormone and Metabolic Research. 2006;  38 229-234
• 22 Thomas P, Zhu Y, Pace M. Progestin membrane receptors involved in the meiotic maturation of teleost oocytes: a review with some new findings.  Steroids. 2002;  67 511-517
• 23 Zhu Y, Rice C D, Pang Y, Pace M, Thomas P. Cloning, expression and characterization of a novel membrane progestin receptor and evidence it is an intermediary in meiotic maturation of fish oocytes.  Proc Nat Acad Sci. 2003;  100 2231-2236
• 24 Zhu Y, Bond J, Thomas P. Identification, classification and partial characterization of gene in human and other vertebrates homologous to a novel fish membrane progestin receptor.  Proc Nat Acad Sci. 2003;  100 2237-2242
• 25 Fortunati N, Catalano M. Sex hormone-binding globulin (SHBG) and estradiol cross-talk in breast cancer cells.  Hormone and Metabolic Research. 2006;  38 235-239
• 26 Lewis J G, Mopert B, Shand B I, Doogue M P, Soule S G, Frampton C M, Elder P A. Plasma variation of corticosteroid-binding globulin and sex hormone-binding globulin.  Hormone and Metabolic Research. 2006;  38 240-244
• 27 Mopert B, Herbert Z, Caldwell J D, Jirikowski G F. Expression of Corticosterone Binding Globulin CBG in the rat hypothalamus.  Hormone and Metabolic Research. 2006;  38 245-251
• 28 Herbert Z, Pollak E I, Caldwell J D, Jirikowski G F. Subcellular localization of SHBG in rat hypothalamus and pituitary.  Society for Neurosci Abs. 2003;  25 610.3
• 29 Sivukhina E V, Jirikowski G F, Bernstein H G, Lewis J G, Herbert Z. Expression of corticosteroid-binding protein in the human hypothalamus, co-localization with oxytocin and vasopressin.  Hormone and Metabolic Research. 2006;  in press
• 30 Breuner C W, Lynn S E, Julian G E, Cornelius J M, Heidinger B J, Love O P, Sprague R S, Wada H, Whitman B A. Plasma binding globulins and the acute stress response.  Hormone and Metabolic Research. 2006;  38 259-267
• 31 Redondo C, Burke B J, Findlay J B. The Retinol Binding Protein System: A potential paradigm for steroid-binding globulins?.  Hormone and Metabolic Research. 2006;  38 268-277
• 32 Nykjaer A, Dragun D, Walther D, Vorum H, Jacobsen C, Herz J, Melsen F, Christensen E I, Willnow T E. An endocytic pathway essential for renal uptake and activation of the steroid 25-(OH) vitamin D3.  Cell. 1999;  96 507-515
• 33 Christensen E I, Moskaug J O, Vorum H, Jacobsen C, Gundersen T E, Nykjaer A, Blomhoff R, Willnow T E, Moestrup S K. Evidence for an essential role of megalin in transepithelial transport of retinol.  J Am Soc Nephrol. 1999;  10 685-695
• 34 Hammes A, Andreassen T K, Spoelgen R, Raila J, Hubner N, Schulz H, Metzger J, Schweigert F J, Luppa P B, Nykjaer A, Willnow T E. Role of Endocytosis in Cellular Uptake of Sex Steroids.  Cell. 2005;  122 751-762
• 35 Becchis M, Sullivan P M, Ordronneau P, Petrusz P, Joseph D R. Distribution of immunoreactive androgen-binding protein/sex hormone-binding globulin in tissues of the fetal rat.  Steroids. 1996;  61 392-400
• 36 Caldwell J D. A sexual arousability model involving steroid effects at the plasma membrane.  Neuroscience & Biobehavioral Reviews. 2002;  26 13-30
• 37 Caldwell J D, Walker C H, Pedersen C A, Barakat A S, Mason G A. Estrogen increases affinity of oxytocin receptors in the medial preoptic area-anterior hypothalamus.  Peptides. 1994;  15 1079-1084
• 38 Caldwell J D, Walker C H, Rivkina A, Pedersen C A, Mason G A. Radioligand assays for oestradiol and progesterone conjugated to protein reveal evidence for a common membrane binding site in the medial preoptic area-anterior hypothalamus and differential modulation by cholera toxin and GTPgammaS.  J Neuroendocrinol. 1999;  11 409-417
• 39 Caldwell J D, Walker C H, Faggin B M, Carr R B, Pedersen C A, Mason G A. Characterization of progesterone-3-[¹²⁵I-BSA] binding sites in the medial preoptic area and anterior hypothalamus.  Brain Res. 1995;  693 225-232
• 40 Mylonas C C, Woods 3rd  L C , Thomas P, Zohar Y. Endocrine profiles of female striped bass (Morone saxatilis) in captivity, during postvitellogenesis and induction of final oocyte maturation via controlled-release GnRHa-delivery systems.  Gen Comp Endocrinol. 1998;  110 276-289
• 41 Pinter J, Thomas P. The ovarian progestogen receptor in the spotted seatrout, Cynoscion nebulosus, demonstrates steroid specificity different from progesterone receptors in other vertebrates.  J Steroid Biochem Mol Biol. 1997;  60 113-119
• 42 Thomas P. Chemical interference with genomic and nongenomic actions of steroids in fishes: role of receptor binding.  Mar Environ Res. 2000;  50 127-134
• 43 Filardo E J. Epidermal growth factor receptor (EGFR) transactivation by estrogen via the G-protein-coupled receptor, GPR30: a novel signaling pathway with potential significance for breast cancer.  J Steroid Biochem Mol Biol. 2004;  80 231-238
• 44 Filardo E J, Quinn J A, Frackelton A R, Bland K I. Estrogen action via the G protein-coupled receptor, GPR30: stimulation of adenylyl cyclase and cAMP-mediated attenuation of the epidermal growth factor receptor-to-MAPK signaling axis.  Mol Endocrinol. 2002;  16 70-84
• 45 Filardo E J, Quinn J A, Bland K I, Frackelton A R. Estrogen-induced activation of Erk-1 and Erk-2 requires the G protein-coupled receptor homolog, GPR30, and occurs via trans-activation of the epidermal growth factor receptor through release of HB-EGF.  Mol Endocrinol. 2000;  14 1649-1660
• 46 Thomas P, Pang Y, Filardo E J, Dong J. Identity of an Estrogen Membrane Receptor Coupled to a G-protein in Human Breast Cancer Cells.  Endocrinology. 2004;  2 624-632
• 47 Alexander N J, Kim H K, Blye R P, Blackmore P F. Steroid specificity of the human sperm membrane progesterone receptor.  Steroids. 1996;  61 116-125
• 48 Alexander N J, Kim H K, Blye R B, Blackmore P F. Steroid specificity of the human sperm membrane progesterone receptor.  Steroids. 1996;  61 116-125
• 49 Falkenstein E, Heck M, Gerdes D, Grube D, Christ M, Weigel M, Buddhikot M, Meizel S, Wehling M. Specific progesterone binding to a membrane protein and related nongenomic effects on Ca2+-fluxes in sperm.  Endocrinology. 1999;  140 5999-6002
• 50 Cork R J, Taylor M, Smith L D, Robinson K R. Microinjected GTP-gamma-s inhibits progesterone-induced maturation of Xenopus oocytes.  Developmental Biology. 1990;  141 447-450
• 51 Sadler S E, Maller J L. Inhibition of Xenopus Oocyte Adenylate Cyclase by Progesterone: A novel mechanism of action. In: Cooper DMF, Seamon KB (eds) Advances in Cyclic Nucleotide and Protein Phosphorylation Research. New York; Raven Press 1985: 179-194
• 52 Kostellow A B, Ma G-Y, Morrill G A. Steroid action at the plasma membrane: progesterone stimulation of phosphatidylcholine-specific phospholipase C following release of the prophase block in amphibian oocytes.  Molec Cell Endocrinology. 1993;  92 33-44
• 53 Yoshikuni M, Nagahama Y. Involvement of an Inhibitory G-Protein in the Signal Transduction Pathway of Maturation-Inducing Hormone (17?,20?-dihydroxy-4-pregnen-3-one) Action in Rainbow Trout (Oncorhynchus mykiss) Oocytes.  Developmental Biology. 1994;  166 615-622
• 54 Kalinowski R, Berlot C H, Jones T LZ, Ross L F, Jaffe L A, Mehlmann L M. Maintenance of meiotic prophase arrest in vertebrate oocytes by a Gs protein-mediated pathway.  Developmental Biology. 2004;  267 1-13
• 55 Gallo C J, Hand A R, Jones T L, Jaffe L A. Stimulation of Xenopus oocyte maturation by inhibition of the G-protein alpha S subunit, a component of the plasma membrane and yolk platelet membranes.  Journal of Cellular Biology. 1995;  130 275-284
• 56 Lyons T J, Villa N Y, Regalla L M, Kupchak B R, Vagstad A, Eide D J. Metalloregulation of yeast membrane steroid receptor homologs.  Proc Natl Acad Sci U S A. 2004;  101 5506-5511
• 57 Gerard A. Endocytosis of androgen-binding protein (ABP) by spermatogenic cells.  J Steroid Biochem Mol Biol. 1995;  53 533-542
• 58 Zhu Y, Bond J, Thomas P. Identification, classification, and partial characterization of genes in humans and other vertebrates homologous to a fish membrane progestin receptor.  Proc Natl Acad Sci U S A. 2003;  100 2237-2242
• 59 Thomas P, Pang Y, Zhu Y, Detweiler C, Doughty K. Multiple rapid progestin actions and progestin membrane receptor subtypes in fish.  Steroids. 2004;  69 567-573
• 60 Petitti N, Etgen A M. Alpha 1-adrenoceptor augmentation of beta-stimulated cAMP formation is enhanced by estrogen and reduced by progesterone in rat hypothalamic slices.  J Neurosci. 1990;  10 2842-2849
• 61 Petitti N, Etgen A M. Progesterone depression of norepinephrine-stimulated cAMP accumulation in hypothalamic slices.  Brain Res Mol Brain Res. 1989;  5 109-119
• 62 Karkanias G B, Etgen A M. A thermodynamic analysis of estrogen regulation of alpha 2- adrenoceptor binding.  Brain Res. 1995;  674 26-32
• 63 Karkanias G B, Petitti N, Etgen A M. Progesterone attenuation of alpha 1-adrenergic receptor stimulation of phosphoinositol hydrolysis in hypothalamus of estrogen-primed female rats.  Endocrinology. 1995;  136 1993-1999
• 64 Etgen A M, Karkanias G B. Estrogen regulation of noradrenergic signaling in the hypothalamus.  Psychoneuroendocrinology. 1994;  19 603-610
• 65 Karkanias G B, Etgen A M. Estradiol reduction of the agonist high affinity form of the alpha 2-adrenoceptor in the hypothalamus of female rats: identification as the alpha 2D subtype.  Mol Pharmacol. 1994;  45 509-516
• 66 Mermelstein P G, Becker J B, Surmeier D J. Estradiol reduces calcium currents in rat neostriatal neurons via a membrane receptor.  J Neurosci. 1996;  16 595-604
• 67 Xiao L, Becker J B. Effects of estrogen agonists on amphetamine-stimulated striatal dopamine release.  Synapse. 1998;  29 379-391
• 68 Toran-Allerand C D, Singh M, Setalo Jr G . Novel Mechanisms of Estrogen Action in the Brain: New Players in an Old Story.  Front in Neuroendocrinology. 1999;  20 97-121
• 69 Singh M, Setalo Jr G , Guan X, Frail D E, Toran-Allerand C D. Estrogen-induced activation of the mitogen-activated protein kinase cascade in the cerebral cortex of estrogen receptor-alpha knock-out mice.  J Neurosci. 2000;  20 1694-1700
• 70 Singh M, Setalo Jr G , Guan X, Warren M, Toran-Allerand C D. Estrogen-induced activation of mitogen-activated protein kinase in cerebral cortical explants: convergence of estrogen and neurotrophin signaling pathways.  J Neurosci. 1999;  19 1179-1188
• 71 Chambliss K L, Shaul P W. Estrogen modulation of endothelial nitric oxide synthase.  Endocrine Reviews. 2002;  23 665-686
• 72 Razandi M, Pedram A, Greene G, Levin E R. Cell Membrane and nuclear estrogen receptors originate from a single transcript: studies of ER-alpha and ER-beta expressed in chinese hamster ovary cells.  Molecular Endocrinology. 1999;  13 307-319
• 73 Razandi M, Pedram A, Levin E R. Plasma Membrane Estrogen Receptors Signal to Antiapoptosis in Breast Cancer.  Molecular Endocrinology. 2000;  14 1434-1447
• 74 Watters J J, Campbell J S, Cunningham M J, Krebs E G, Dorsa D. Rapid membrane effects of steroids in neuroblastoma cells: effects of estrogen on mitogen activated protein kinase signalling cascade and c-fos immediate early gene transcription.  Endocrinology. 1997;  138 4030-4033
• 75 Wade C B, Robinson S, Shapiro R A, Dorsa D M. Estrogen receptor (ER)alpha and ERbeta exhibit unique pharmacologic properties when coupled to activation of the mitogen-activated protein kinase pathway.  Endocrinology. 2001;  142 2336-2342
• 76 Linford N, Wade C, Dorsa D. The rapid effects of estrogen are implicated in estrogen- mediated neuroprotection.  J Neurocytol. 2000;  29 367-374
• 77 Fitzpatrick J L, Mize A L, Wade C B, Harris J A, Shapiro R A, Dorsa D M. Estrogen-mediated neuroprotection against beta-amyloid toxicity requires expression of estrogen receptor alpha or beta and activation of the MAPK pathway.  J Neurochem. 2002;  82 674-682
• 78 Mize A L, Shapiro R A, Dorsa D M. Estrogen receptor-mediated neuroprotection from oxidative stress requires activation of the mitogen-activated protein kinase pathway.  Endocrinology. 2003;  144 306-312
• 79 Wade C B, Dorsa D M. Estrogen activation of cyclic adenosine 5’-monophosphate response element-mediated transcription requires the extracellularly regulated kinase/mitogen-activated protein kinase pathway.  Endocrinology. 2003;  144 832-838
• 80 Ramirez V D, Zheng J. Membrane sex-steroid receptors in the brain.  Front Neuroendocrinol. 1996;  17 402-439
• 81 Kipp J L, Ramirez V D. Estradiol and testosterone have opposite effects of microtubule polymerization.  Neuroendocrinology. 2003;  77 258-272
• 82 Gu Q, Moss R L. Novel mechanism for non-genomic action of 17beta-estradiol on kainate-induced currents in isolated rat CA1 hippocampal neurones.  J of Physiology. 1998;  506 745-754
• 83 Moss R L, Gu Q, Dudley C A. Estrogen: Nontranscriptional signaling pathway.  Recent Progress in Hormone Research. 1997;  52 33-69
• 84 Wong M, Moss R L. Patch-clamp analysis of direct steroidal modulation of glutamate receptor-channels.  J of Neuroendocrinology. 1994;  6 347-355
• 85 Gu Q, Korach K S, Moss R L. Rapid action of 17beta-estradiol on kainate-induced currents in hippocampal neurons lacking intracellular estrogen receptors.  Endocrinology. 1999;  140 660-666
• 86 Lagrange A H, Ronnekliev O K, Kelly M J. Modulation of G protein-coupled receptors by an estrogen receptor that activates protein kinase A.  Molecular Pharmacology. 1997;  51 605-612
• 87 Lagrange A H, Ronnekleiv O K, Kelly M J. Estradiol-17 beta and mu-opioid peptides rapidly hyperpolarize GnRH neurons: a cellular mechanism of negative feedback?.  Endocrinology. 1995;  136 2341-2344
• 88 Lagrange A H, Ronnekleiv O K, Kelly M J. The potency of mu-opioid hyperpolarization of hypothalamic arcuate neurons is rapidly attenuated by 17 beta-estradiol.  J Neurosci. 1994;  14 6196-6204
• 89 Kelly M J, Lagrange A H, Wagner E J, Ronnekleiv O K. Rapid effects of estrogen to modulate G protein-coupled receptors via activation of protein kinase A and protein kinase C pathways.  Steroids. 1999;  64 64-75
• 90 Sundaram M, van Aalten D M, Findlay J B, Sivaprasadarao A. The transfer of transthyretin and receptor-binding properties from the plasma retinol-binding protein to the epididymal retinoic acid-binding protein.  Biochem J. 2002;  362 265-271
• 91 Sivaprasadarao A, Sundaram M, Findlay J B. Interactions of retinol-binding protein with transthyretin and its receptor.  Methods Mol Biol. 1998;  89 155-163
• 92 Sundaram M, Sivaprasadarao A, De Sousa M M, Findlay J B. The transfer of retinol from serum retinol-binding protein to cellular retinol-binding protein is mediated by a membrane receptor.  J Biol Chem. 1998;  273 3336-3342
• 93 Reventos J, Sullivan P M, Joseph D R, Gordon J W. Tissue-specific expression of the rat androgen-binding protein/sex hormone-binding globulin gene in transgenic mice.  Mol Cell Endocrinol. 1993;  96 69-73
• 94 Joseph D R, Sullivan P M, Wang Y M, Millhorn D E, Bayliss D M. Complex structure and regulation of the ABP/SHBG gene.  J Steroid Biochem Mol Biol. 1991;  40 771-775
• 95 Joseph D R, Power G A, Petrusz P. Expression and distribution of androgen-binding protein/sex hormone-binding globulin in the female rodent reproductive system.  Biology of Reproduction. 1997;  56 14-20
• 96 Joseph D R, Adamson M C, Kozak C A. Genetic mapping of the gene for androgen-binding protein/sex hormone-binding globulin to mouse chromosome 11.  Cytogenet Cell Genet. 1991;  56 122-124
• 97 Pardridge W M, Eisenberg J, Fierer G, Musto N A. Developmental changes in brain and serum binding of testosterone and in brain capillary uptake of testosterone-binding serum proteins in the rabbit.  Brain Res. 1988;  466 245-253
• 98 Porto C S, Gunsalus G L, Bardin C W, Phillips D M, Musto N A. Receptor-mediated endocytosis of an extracellular steroid-binding protein (TeBG) in MCF-7 human breast cancer cells.  Endocrinology. 1991;  129 436-445
• 99 Sakiyama R, Pardridge W M, Musto N A. Influx of testosterone-binding globulin (TeBG) and TeBG-bound sex steroid hormone into rat testis and prostate.  J Clin Endocrin Metabolism. 1988;  67 98-103
• 100 Gueant J L, Fremont S, Felden F, Nicolas J P, Gerard A, Leheup B, Gerard H, Grignon G. Evidence that androgen-binding protein endocytosis in vitro is receptor mediated in principal cells of the rat epididymis.  J Mol Endocrinol. 1991;  7 113-122
• 101 Meyer S, Brumm C, Stegner H E, Sinnecker G H. Intracellular sex hormone-binding globulin (SHBG) in normal and neoplastic breast tissue - an additional marker for hormone dependency?.  Exp Clin Endocrinol. 1994;  102 334-340
• 102 Porto C S, Lazari M F, Abreu L C, Bardin C W, Gunsalus G L. Receptors for androgen-binding proteins: internalization and intracellular signalling.  J Steroid Biochem Mol Biol. 1995;  53 561-565
• 103 Sinnecker G, Hiort O, Kwan P W, De Lellis R A. Immunohistochemical localization of sex hormone-binding globulin in normal and neoplastic breast tissue.  Horm Metab Res. 1990;  22 47-50
• 104 Allera A, Wildt L. Glucocorticoid-recognizing and -effector sites in rat liver plasma membrane. Kinetics of corticosterone uptake by isolated membrane vesicles-II. Comparative influx and efflux.  J Steroid Biochem Mol Biol. 1992;  42 757-771
• 105 Miska W, Fehl P, Henkel R. Biochemical and immunological characterization of the acrosome reaction-inducing substance (ARIS) of hFF.  Biochem Biophys Res Commun. 1994;  199 125-129
• 106 Krupenko S A, Krupenko N I, Danzo B J. Interaction of sex hormone-binding globulin with plasma membranes from the rat epididymis and other tissues.  J Steroid Biochem Mol Biol. 1994;  51 115-124
• 107 Wong A S, Lui W Y, Hui I T, Lee W M. Rabbit sex hormone-binding globulin: expression in the liver and testis during postnatal development and structural characterization by truncated proteins.  Int J Androl. 2001;  24 165-174
• 108 Krieg M, Bartsch S, Herzer S, Becker H, Voigt K D. Quantification of androgen binding, androgen tissue levels, and sex hormone-binding globulin in prostate, muscle and plasma of patients with benign prostatic hypertrophy.  Acta Endocrinologica. 1977;  86 200-215
• 109 Bordin S, Petra P H. Immunocytochemical localization of the sex steroid-binding protein of plasma in tissues of the adult monkey Macaca nemestrina.  Proc Natl Acad Sci USA. 1980;  77 5678-5682
• 110 Hryb D J, Nakhla A M, Kahn S M, St. George J, Levy N C, Romas N A, Rosner W. Sex hormone-binding globulin in the human prostate is locally synthesized and may act as an autocrine/paracrine factor.  J Biol Chem. 2002;  277 26 618-26 622
• 111 Wang Y M, Bayliss D A, Millhorn D E, Petrusz P, Joseph D R. The androgen-binding protein gene is expressed in male and female rat brain.  Endocrinology. 1990;  127 3124-3130
• 112 Jirikowski G F, Herbert Z, Caldwell J D. Characterization and localization of sex hormone binding globulin in rat hypothalamus and pituitary.  Society for Neuroscience Abstracts. 2002;  24 73.14
• 113 Herbert Z, Pollak E I, Molnar L, Caldwell J D, Jirikowski G F. Co-transport of sex hormone binding globulin/SHBG with oxytocin in transport vesicle of the hypothalamo-hypophyseal system.  Hormone and Metabolic Research. 2006;  38 290-292
• 114 Herbert Z, Weigel S, Marshall A, Caldwell J D, Petrusz P, Peuckert C, Jirikowski G F. Coexpression of androgen-binding protein (ABP) and oxytocin in the male reproductive tract.  Anatomia, Histologia and Embrylogia. 2005;  34 293
• 115 Caldwell J D, Moe B D, Hoang J, Nguyen T. Sex hormone binding globulin stimulates female sexual receptivity.  Brain Res. 2000;  874 24-29
• 116 Caldwell J D, Hoefle S, Englof I. Sex Hormone Binding Globulin Facilitates female sexual receptivity except when coupled to Dihydrotestosterone.  Brain Research. 2002;  948 102-107
• 117 Caldwell J D. Evidence of Sex Hormone Binding Globulin Binding sites in the Medial Preoptic Area and Hypothalamus.  Hormone and Metabolic Research. 2001;  33 7-9
• 118 Krupenko N I, Avvakumov G V, Strel’chyonok O A. Binding of human sex hormone-binding globulin-androgen complexes to the placental syncytiotrophoblast membrane.  Biochem Biophys Res Commun. 1990;  171 1279-1283
• 119 Avvakumov G V, Zhuk N I, Strel’chyonok O A. Subcellular distribution and selectivity of the protein-binding component of the recognition system for sex-hormone-binding protein-estradiol complex in human decidual endometrium.  Biochim Biophys Acta. 1986;  881 489-498
• 120 Strel'chyonok O A, Avvakumov G V, Survilo L J. A recognition system for sex-hormone binding protein-estradiol complex in human decidual endometrium plasma membranes.  Biochim Biophys Acta. 1984;  802 459-466
• 121 Catalano M G, Comba A, Fazzari A, Benedusipagliano E, Sberverglieri M, Revelli A, Massobrio M, Frairia R, Fortunati N. Sex steroid binding protein receptor (SBP R) is related to a reduced proliferation rate in human breast cancer.  Breast Cancer Research and Treatment. 1997;  42 227-234
• 122 Becchis M, Frairia R, Ferrera P, Fazzari A, Ondei S, Alfarano A, Coluccia C, Biglia N, Sismondi P, Fortunati N. The additionally glycosylated variant of human sex hormone-binding globulin (SHBG) is linked to estrogen-dependence of breast cancer.  Breast Cancer Res Treat. 1999;  54 101-107
• 123 Fortunati N, Becchis M, Catalano M G, Comba A, Ferrera P, Raineri M, Berta L, Frairia R. Sex hormone-binding globulin, its membrane receptor, and breast cancer: a new approach to the modulation of estradiol action in neoplastic cells.  J Steroid Biochem Mol Biol. 1999;  69 473-479
• 124 Frairia R, Fortunati N, Revelli A, Guidetti D, Cavaglia S, Massobrio M. Binding of sex steroid binding protein to plasma membranes of human testis.  J Steroid Biochem Mol Biol. 1994;  51 319-322
• 125 Fortunati N, Raineri M, Cignetti A, Hammond G L, Frairia R. Control of the membrane sex hormone-binding globulin-receptor (SHBG-R) in MCF-7 cells: effect of locally produced SHBG.  Steroids. 1998;  63 282-284
• 126 Fortunati N, Comba A, Becchis M, Catalano M G, Fazzari A, Fissore F, Frairia R. MCF-7 cell progesterone receptor (PGR) is additionally modulated by sex steroid binding protein (SBP) and its membrane receptor (SBP-R) through cAMP and PKA.  Ann N Y Acad Sci. 1996;  784 453-457
• 127 Fortunati N, Fissore F, Fazzari A, Becchis M, Comba A, Catalano M G, Berta L, Frairia R. Sex steroid binding protein exerts a negative control on estradiol action in MCF-7 cells (human breast cancer) through cyclic adenosine 3’,5’-monophosphate and protein kinase A.  Endocrinology. 1996;  137 686-692
• 128 Fortunati N, Fissore F, Comba A, Becchis M, Catalano M G, Fazzari A, Berta L, Frairia R. Sex steroid-binding protein and its membrane receptor in estrogen- dependent breast cancer: biological and pathophysiological impact.  Horm Res. 1996;  45 202-206
• 129 Fissore F, Fortunati N, Comba A, Fazzari A, Gaidano G, Berta L, Frairia R. The receptor-mediated action of sex steroid binding protein (SBP, SHBG): accumulation of cAMP in MCF-7 cells under SBP and estradiol treatment.  Steroids. 1994;  59 661-667
• 130 Nakhla A M, Khan M S, Rosner W. Biologically active steroids activate receptor-bound human sex hormone-binding globulin to cause LNCaP cells to accumulate adenosine 3’,5’-monophosphate.  J Clin Endocrinol Metab. 1990;  71 398-404
• 131 Nakhla A M, Leonard J, Hryb D J, Rosner W. Sex hormone-binding globulin receptor signal transduction proceeds via a G protein.  Steroids. 1999;  64 213-216
• 132 Hryb D J, Khan M S, Romas N A, Rosner W. The control of the interaction of sex hormone-binding globulin with its receptor by steroid hormones.  J Biol Chem. 1990;  265 6048-6054
• 133 Caldwell J D, Morris M A, Walker C H, Carr R B, Faggin B M, Mason G A. Estradiol conjugated to BSA releases oxytocin from synaptosome- containing homogenates from the medial preoptic area-hypothalamus.  Horm Metab Res. 1996;  28 119-121
• 134 Panayiotis E S, Deecher D C, Suhadolnik L, Mallis L M, Frail D E. Differential effects of estradiol and estradiol-BSA conjugates.  Endocrinology. 1999;  140 5455-5458
• 135 Caldwell J D, Song Y, Englof I, Hoefle S, Key M, Morris M. 5alpha-reduced androgens Block estradiol-BSA Stimulated Release of Oxytocin.  Brain Research. 2003;  976 259-261
• 136 Siiteri P K, Murai J T, Hammond G L, Nisker J A, Raymoure W J, Kuhn R W. The serum transport of steroid hormones.  Recent Prog Horm Res. 1982;  38 457-510
• 137 Nykjaer A, Fyfe J C, Kozyraki R, Leheste J R, Jacobsen C, Nielsen M S, Verroust P J, Aminoff M, de la Chapelle A, Moestrup S K, Ray R, Gliemann J, Willnow T E, Christensen E I. Cubilin dysfunction causes abnormal metabolism of the steroid hormone 25(OH) vitamin D(3).  Proc Natl Acad Sci U S A. 2001;  98 13 895-13 900
• 138 Nykjaer A, Lee R, Teng K K, Jansen P, Madsen P O, Nielsen M S, Jacobsen C, Kliemannel M, Schwarz E, Willnow T E, Hempstead B L, Petersen C M. Sortilin is essential for proNGF-induced neuronal cell death.  Nature. 2004;  427 843-848
• 139 Chau P L, van Aalten D M, Bywater R P, Findlay J B. Functional concerted motions in the bovine serum retinol-binding protein.  J Comput Aided Mol Des. 1999;  13 11-20
• 140 Sivaprasadarao A, Findlay J B. Structure-function studies on human retinol-binding protein using site-directed mutagenesis.  Biochem J. 1994;  300 437-442
• 141 Aspinall S R, Stamp S, Davison A, Shenton B K, Lennard T W. The proliferative effects of 5-androstene-3 beta,17 beta-diol and 5 alpha-dihydrotestosterone on cell cycle analysis and cell proliferation in MCF7, T47D and MDAMB231 breast cancer cell lines.  J Steroid Biochem Mol Biol. 2004;  88 37-51
• 142 Friedman A E. The Estradiol-Dihydrotestosterone model of prostate cancer.  Theor Biol Med Model. 2005;  2 10
• 143 Greeve M A, Allan M K, Harvey J M, Bentel J M. Inhibition of MCF-7 breast cancer cell proliferation by 5alpha-dihydrotestosterone; a role for p21(Cip1/Waf1).  J Mol Endocrinol. 2004;  32 793-810
• 144 Lund T D, Munson D J, Haldy M E, Handa R J. Androgen inhibits, while oestrogen enhances, restraint-induced activation of neuropeptide neurones in the paraventricular nucleus of the hypothalamus.  J Neuroendocrinol. 2004;  16 272-278
• 145 Lund T D, Munson D J, Haldy M E, Handa R J. Dihydrotestosterone may inhibit hypothalamo-pituitary-adrenal activity by acting through estrogen receptor in the male mouse.  Neurosci Lett. 2004;  365 43-47
• 146 Riley L G, Hirano T, Grau E G. Estradiol-17beta and dihydrotestosterone differentially regulate vitellogenin and insulin-like growth factor-I production in primary hepatocytes of the tilapia Oreochromis mossambicus.  Comp Biochem Physiol C Toxicol Pharmacol. 2004;  138 177-186
• 147 Rosner W, Hryb D J, Khan M S, Nakhla A M, Romas N A. Sex hormone-binding globulin: anatomy and physiology of a new regulatory system.  J Steroid Biochem Mol Biol. 1991;  40 813-820
• 148 Rosner W, Hryb D J, Khan M S, Nakhla A M, Romas N A. Sex hormone-binding globulin. Binding to cell membranes and generation of a second messenger.  J Androl. 1992;  13 101-106
• 149 Rosner W, Hryb D J, Khan M S, Nakhla A M, Romas N A. Androgen and estrogen signaling at the cell membrane via G-proteins and cyclic adenosine monophosphate.  Steroids. 1999;  64 100-106
• 150 Mystkowski P, Schwartz M W. Gonadal steroids and energy homeostasis in the leptin era.  Nutrition. 2000;  16 937-946
• 151 Wade G N. Sex steroids and energy balance: sites and mechanisms of action.  Ann N Y Acad Sci. 1986;  474 389-399
• 152 Caldwell J D, Jirikowski G F, Greer E R, Pedersen C A. Medial preoptic area oxytocin and female sexual receptivity.  Behav Neurosci. 1989;  103 655-662
• 153 Caldwell J D, Jirikowski G F, Greer E R, Stumpf W E, Pedersen C A. Ovarian steroids and sexual interaction alter oxytocinergic content and distribution in the basal forebrain.  Brain Res. 1988;  446 236-244
• 154 Caldwell J D, Walker C H, Pedersen C A, Mason G A. Sexual activity decreases oxytocin receptor densities in the thymus.  Life Sci. 1993;  52 1781-1786
• 155 Caldwell J D, Walker C H, Pedersen C A, Mason G A. Effects of steroids and mating on central oxytocin receptors.  Ann N Y Acad Sci. 1992;  652 433-436