Extra-adrenal glucocorticoids and mineralocorticoids: Evidence for local synthesis, regulation, and function

American Journal of Physiology - Endocrinology and Metabolism

Volumn 301, Issue 1, 2011, Pages

  Taves, Matthew D ^a   Gomez Sanchez, Celso E ^b,c   Soma, Kiran K ^a  

^a UNIVERSITY OF BRITISH COLUMBIA   (Canada)

^b UNIVERSITY OF MISSISSIPPI MEDICAL CENTER   (United States)

^c UNIVERSITY OF MISSISSIPPI SCHOOL OF MEDICINE   (United States)

Author keywords

Aldosterone; Brain; Bursa of fabricius; Corticosterone; Cortisol; Heart; Immunosteroids; Intestine; Neurosteroids; Skin; Stress; Thymus

Indexed keywords

ALDOSTERONE; ALDOSTERONE SYNTHASE; CHOLESTEROL; CHOLESTEROL MONOOXYGENASE (SIDE CHAIN CLEAVING); COLLAGEN; CORTICOSTERONE; CORTICOSTERONE DERIVATIVE; CORTISONE; CORTODOXONE; CYP11B1 PROTEIN; CYTOCHROME P450 17; DEHYDROCORTICOSTERONE; DEOXYCORTICOSTERONE; GLUCOCORTICOID; HYDROCORTISONE; MINERALOCORTICOID; PREGNENOLONE; PREGNENOLONE DERIVATIVE; PROGESTERONE; PROGESTERONE DERIVATIVE; PROTEIN; STAR PROTEIN; T LYMPHOCYTE RECEPTOR; UNCLASSIFIED DRUG;

ADRENAL GLAND; APOPTOSIS; AUTOIMMUNITY; B LYMPHOCYTE; BEHAVIOR; BLOOD PRESSURE REGULATION; BLOOD VOLUME; BRAIN; CELL MATURATION; CELLULAR IMMUNITY; CONCENTRATION (PARAMETERS); DOWN REGULATION; DRUG SEEKING BEHAVIOR; HEART; HOMEOSTASIS; HORMONE ACTION; HORMONE RELEASE; HORMONE SYNTHESIS; HUMAN; HYPOTHALAMUS HYPOPHYSIS ADRENAL SYSTEM; IMMUNOCOMPETENT CELL; INFLAMMATION; INTESTINE; LYMPHOID ORGAN; NONHUMAN; PRIORITY JOURNAL; PROTEIN EXPRESSION; RENIN ANGIOTENSIN ALDOSTERONE SYSTEM; REVIEW; SKIN; SODIUM BALANCE; STRESS; TH1 CELL; THYMOCYTE; THYMUS; TISSUE REPAIR;

ADRENAL GLANDS; ANIMALS; FEEDBACK, PHYSIOLOGICAL; GLUCOCORTICOIDS; HUMANS; LYMPHOID TISSUE; METABOLIC NETWORKS AND PATHWAYS; MINERALOCORTICOIDS; MODELS, BIOLOGICAL; TISSUE DISTRIBUTION;

EID: 79959581081     PISSN: 01931849     EISSN: 15221555     Source Type: Journal    
DOI: 10.1152/ajpendo.00100.2011     Document Type: Review

References (122)

1. Ahmad N, Romero DG, Gomez-Sanchez EP, Gomez-Sanchez CE. Do human vascular endothelial cells produce aldosterone? Endocrinology 145: 3626-3629, 2004.
2. Aird F, Clevenger CV, Prystowsky MB, Redei EVA. Corticotropinreleasing factor mRNA in rat thymus and spleen. Proc Natl Acad Sci USA 90: 7104-7108, 1993.
3. Atanasov AG, Leiser D, Roesselet C, Noti M, Corazza N, Schoonjans K, Brunner T. Cell cycle-dependent regulation of extra-adrenal glucocorticoid synthesis in murine intestinal epithelial cells. FASEB J 22: 4117-4125, 2008.
4. Batanero E, de Leeuw FE, Jansen GH, van Wichen DF, Huber J, Schuurman HJ. The neural and neuro-endocrine component of the human thymus. II. Hormone immunoreactivity. Brain Behav Immun 6: 249-264, 1992.
5. Beyenburg S, Watzka M, Clusmann H, Blumcke I, Bidlingmaier F, Elger CE, Stoffel-Wagner B. Messenger RNA of steroid 21-hydroxylase (CYP21) is expressed in the human hippocampus. Neurosci Lett 308: 111-114, 2001.
6. Brewer JA, Kanagawa O, Sleckman BP, Muglia LJ. Thymocyte apoptosis induced by T cell activation is mediated by glucocorticoids in vivo. J Immunol 169: 1837-1843, 2002.
7. Byrd J. The effect of the humoral immune system-derived bursal anti-steroidogenic peptide (BASP) on corticosteroid biosynthesis in avian, porcine and canine adrenal cortical cells. Comp Biochem Physiol 108: 221-227, 1994.
8. Casal AJ, Silvestre JS, Delcayre C, Capponi AM. Expression and modulation of steroidogenic acute regulatory protein messenger ribonucleic acid in rat cardiocytes and after myocardial infarction. Endocrinology 144: 1861-1868, 2003.
9. Chai W, Hofland J, Jansen PM, Garrelds IM, de Vries R, van den Bogaerdt AJ, Feelders RA, de Jong FH, Danser AHJ. Steroidogenesis vs. steroid uptake in the heart: do corticosteroids mediate effects via cardiac mineralocorticoid receptors? J Hypertens 28: 1044-1053, 2010.
10. Chen Y, Qiao S, Tuckermann J, Okret S, Jondal M. Thymus-derived glucocorticoids mediate androgen effects on thymocyte homeostasis. FASEB J 24: 5043-5051, 2010.
11. Cima I, Corazza N, Dick B, Fuhrer A, Herren S, Jakob S, Ayuni E, Mueller C, Brunner T. Intestinal epithelial cells synthesize glucocorticoids and regulate T cell activation. J Exp Med 200: 1635-1646, 2004.
12. Compagnone NA, Mellon SH. Neurosteroids: biosynthesis and function of these novel neuromodulators. Front Neuroendocrinol 56: 1-56, 2000.
13. Coste A, Dubuquoy L, Barnouin R, Annicotte JS, Magnier B, Notti M, Corazza N, Antal MC, Metzger D, Desreumaux P, Brunner T, Auwerx J, Schoonjans K. LRH-1-mediated glucocorticoid synthesis in enterocytes protects against inflammatory bowel disease. Proc Natl Acad Sci USA 104: 13098-13103, 2007.
14. Croft AP, O'Callaghan MJ, Shaw SG, Connolly G, Jacquot C, Little HJ. Effects of minor laboratory procedures, adrenalectomy, social defeat or acute alcohol on regional brain concentrations of corticosterone. Brain Res 1238: 12-22, 2008.
15. Davies E, Mackenzie SM. Extra-adrenal production of corticosteroids. Clin Exp Pharmacol Physiol 30: 437-445, 2003.
16. Dhabhar FS. Enhancing versus suppressive effects of stress on immune function: implications for immunoprotection and immunopathology. Neuroimmunomodulation 16: 300-317, 2009.
17. Do Rego JL, Seong JY, Burel D, Leprince J, Luu-The V, Tsutsui K, Tonon MC, Pelletier G, Vaudry H. Neurosteroid biosynthesis: enzymatic pathways and neuroendocrine regulation by neurotransmitters and neuropeptides. Front Neuroendocrinol 30: 259-301, 2009.
18. Dostal DE, Rothblum KN, Chernin MI, Cooper GR, Baker KM. Intracardiac detection of angiotensinogen and renin: a localized reninangiotensin system in neonatal rat heart. Am J Physiol Cell Physiol 263: C838-C850, 1992.
19. Dumont M, Luu-The V, Dupont E, Pelletier G, Labrie F. Characterization, expression, and immunohistochemical localization of 3betahydroxysteroid dehydrogenas/Δ5-Δ4 isomerase in human skin. J Invest Dermatol 99: 415-421, 1992.
20. Falkenhahn M, Franke F, Bohle RM, Zhu YC, Stauss HM, Bachmann S, Danilov S, Unger T. Cellular distribution of angiotensinconverting enzyme after myocardial infarction. Hypertension 25: 219-226, 1995.
21. Fink T, Schafer MKH, di Sebastiano P, Frieß H, Buchler M, Weihe E. Differential opioid gene expression in neurons and neuroendocrine cells of rat and human gastrointestinal tract. Regul Pept 53: S149-S150, 1994.
22. Franchini A, Ottaviani E. Immunoreactive POMC-derived peptides and cytokines in the chicken thymus and bursa of Fabricius microenvironments: age-related changes. J Neuroendocrinol 11: 685-692, 1999.
23. Funder JW. Cardiac synthesis of aldosterone: going, going, gone? Endocrinology 145: 4793-4795, 2004.
24. Garvy BA, King LE, Telford WG, Morford LA, Fraker PJ. Chronic elevation of plasma corticosterone causes reductions in the number of cycling cells of the B lineage in murine bone marrow and induces apoptosis. Immunology 80: 587-592, 1993.
25. Geerling JC, Loewy AD. Aldosterone in the brain. Am J Physiol Renal Physiol 297: F559-F576, 2009.
26. Gomez-Sanchez CE, Zhou MY, Cozza EN, Morita H, Foecking MF, Gomez-Sanchez EP. Aldosterone biosynthesis in the rat brain. Endocrinology 138: 3369-3373, 1997.
27. Gomez-Sanchez CE, Zhou MY, Cozza EN, Morita H, Eddleman FC, Gomez-Sanchez EP. Corticosteroid synthesis in the central nervous system. Endocr Res 22: 463-470, 1996.
28. Gomez-Sanchez EP. Brain mineralocorticoid receptors: orchestrators of hypertension and end-organ disease. Curr Opin Nephrol Hypertens 13: 191-196, 2004.
29. Gomez-Sanchez EP, Ahmad N, Romero DG, Gomez-Sanchez CE. Origin of aldosterone in the rat heart. Endocrinology 145: 4796-4802, 2004.
30. Gomez-Sanchez EP, Ahmad N, Romero DG, Gomez-Sanchez CE. Is aldosterone synthesized within the rat brain? Am J Physiol Endocrinol Metab 288: E342-E346, 2005.
31. Gomez-Sanchez EP, Gomez-Sanchez CM, Plonczynski M, Gomez-Sanchez CE. Aldosterone synthesis in the brain contributes to Dahl salt-sensitive rat hypertension. Exp Physiol 95: 120-130, 2010.
32. Gomez-Sanchez EP, Samuel J, Vergara G, Ahmad N. Effect of 3β-hydroxysteroid dehydrogenase inhibition by trilostane on blood pressure in the Dahl salt-sensitive rat. Am J Physiol Regul Integr Comp Physiol 288: R389-R393, 2005.
33. Gunnar MR, Cheatham CL. Brain and behavior interface: stress and the developing brain. Infant Ment Health J 24: 195-211, 2003.
34. Hatakeyama H, Miyamori I, Fujita T, Takeda Y, Takeda R, Yamamoto H. Vascular aldosterone: biosynthesis and a link to angiotensin II-induced hypertrophy of vascular smooth muscle cells. J Biol Chem 269: 24316-24320, 1994.
35. Hatakeyama H, Miyamori I, Takeda Y, Yamamoto H, Mabuchi H. The expression of steroidogenic enzyme genes in human vascular cells. Biochem Mol Biol Int 40: 639-645, 1996.
36. + in Wistar rats. Am J Physiol Regul Integr Comp Physiol 295: R166-R172, 2008.
37. Huang BS, White RA, Jeng AY, Leenen FHH. Role of central nervous system aldosterone synthase and mineralocorticoid receptors in saltinduced hypertension in Dahl salt-sensitive rats. Am J Physiol Regul Integr Comp Physiol 296: R994-R1000, 2009.
38. Ito N, Ito T, Kromminga A, Bettermann A, Takigawa M, Kees F, Straub RH, Paus R. Human hair follicles display a functional equivalent of the hypothalamic-pituitary-adrenal axis and synthesize cortisol. FASEB J 19: 1332-1334, 2005.
39. Iwata M, Hanaoka S, Sato K. Rescue of thymocytes and T cell hybridomas from glucocorticoid-induced apoptosis by stimulation via the T cell receptor/CD3 complex: a possible in vitro model for positive selection of the T cell repertoire. Eur J Immunol 21: 643-648, 1991.
40. Janeway CA, Travers P, Walport M, Shlomchik MJ. Immunobiology: the Immune System in Health and Disease. New York: Garland, 2005.
41. Kayes-Wandover KM, White PC. Steroidogenic enzyme gene expression in the human heart. J Clin Endocrinol Metab 85: 2519-2525, 2000.
42. Keeney DS, Ikeda Y, Waterman MR, Parker KL. Cholesterol sidechain cleavage cytochrome P450 gene expression in the primitive gut of the mouse embryo does not require steroidogenic factor 1. Mol Endocrinol 9: 1091-1098, 1995.
43. Kimoto T, Ishii H, Higo S, Hojo Y, Kawato S. Semicomprehensive analysis of the postnatal age-related changes in the mRNA expression of sex steroidogenic enzymes and sex steroid receptors in the male rat hippocampus. Endocrinology 151: 5795-5806, 2010.
44. King LB, Vacchio MS, Dixon K, Hunziker R, Margulies DH, Ashwell JD. A targeted glucocorticoid receptor antisense transgene increases thymocyte apoptosis and alters thymocyte development. Immunity 3: 647-656, 1995.
45. Kishimoto W, Hiroi T, Shiraishi M, Osada M, Imaoka S, Kominami S, Igarashi T, Funae Y. Cytochrome P450 2D catalyze steroid 21- hydroxylation in the brain. Endocrinology 145: 699-705, 2004.
46. Lechner O, Dietrich H, Wiegers GJ, Vacchio MS, Wick G. Glucocorticoid production in the chicken bursa and thymus. Int Immunol 13: 769-776, 2001.
47. Lechner O, Wiegers GJ, Oliveira-Dos-Santos AJ, Dietrich H, Recheis H, Waterman M, Boyd R, Wick G. Glucocorticoid production in the murine thymus. Eur J Immunol 30: 337-346, 2000.
48. Little HJ, Croft AP, O'Callaghan MJ, Brooks SP, Wang G, Shaw SG. Selective increases in regional brain glucocorticoid: a novel effect of chronic alcohol. Neuroscience 156: 1017-1027, 2008.
49. Lowenberg M, Tuynman J, Bilderbeek J, Gaber T, Buttgereit F, van Deventer S, Peppelenbosch M, Hommes D. Rapid immunosuppressive effects of glucocorticoids mediated through Lck and Fyn. Blood 106: 1703-1710, 2005.
50. Lowenberg M, Verhaar AP, Bilderbeek J, van Marle J, Buttgereit F, Peppelenbosch MP, van Deventer SJ, Hommes DW. Glucocorticoids cause rapid dissociation of a T-cell-receptor-associated protein complex containing LCK and FYN. EMBO Rep 7: 1023-1029, 2006.
51. MacKenzie SM, Clark CJ, Fraser R, Gomez-Sanchez CE, Connell JM, Davies E. Expression of 11beta-hydroxylase and aldosterone synthase genes in the rat brain. J Mol Endocrinol 24: 321-328, 2000.
52. MacKenzie SM, Fraser R, Connell JMC, Davies E. Local reninangiotensin systems and their interactions with extra-adrenal corticosteroid production. J Renin Angiotensin Aldosterone Syst 3: 214-221, 2002.
53. Mellon SH, Miller WL. Extraadrenal steroid 21-hydroxylation is not mediated by P450c21. J Clin Invest 84: 1497-1502, 1989.
54. Mellon SH, Deschepper CF. Neurosteroid biosynthesis: genes for adrenal steroidogenic enzymes are expressed in the brain. Brain Res 629: 283-292, 1993.
55. Miller AH, Spencer RL, Stein M, McEwen BS. Adrenal steroid receptor binding in spleen and thymus after stress or dexamethasone. Am J Physiol Endocrinol Metab 259: E405-E412, 1990.
56. Mizuno Y, Yoshimura M, Yasue H, Sakamoto T, Ogawa H, Kugiyama K, Harada E, Nakayama M, Nakamura S, Ito T, Shimasaki Y, Saito Y, Nakao K. Aldosterone production is activated in failing ventricle in humans. Circulation 103: 72-77, 2001.
57. Mueller M, Atanasov A, Cima I, Corazza N, Schoonjans K, Brunner T. Differential regulation of glucocorticoid synthesis in murine intestinal epithelial versus adrenocortical cell lines. Endocrinology 148: 1445-1453, 2007.
58. Mueller M, Cima I, Noti M, Fuhrer A, Jakob S, Dubuquoy L, Schoonjans K, Brunner T. The nuclear receptor LRH-1 critically regulates extra-adrenal glucocorticoid synthesis in the intestine. J Exp Med 203: 2057-2062, 2006.
59. Newman AEM, Pradhan DS, Soma KK. Dehydroepiandrosterone and corticosterone are regulated by season and acute stress in a wild songbird: jugular versus brachial plasma. Endocrinology 149: 2537-2545, 2008.
60. Norton JM, Wira CR. Dose-related effects of the sex hormones and cortisol on the growth of the bursa of Fabricius in chick embryos. J Steroid Biochem 8: 985-987, 1977.
61. Noti M, Corazza N, Mueller C, Berger B, Brunner T. TNF suppresses acute intestinal inflammation by inducing local glucocorticoid synthesis. J Exp Med 207: 1057-1066, 2010.
62. Noti M, Corazza N, Tuffin G, Schoonjans K, Brunner T. Lipopolysaccharide induces intestinal glucocorticoid synthesis in a TNFalphadependent manner. FASEB J 24: 1340-1346, 2010.
63. Noti M, Sidler D, Brunner T. Extra-adrenal glucocorticoid synthesis in the intestinal epithelium: more than a drop in the ocean? Semin Immunopathol 31: 237-248, 2009.
64. Ohtani T, Ohta M, Yamamoto K, Mano T, Sakata Y, Nishio M, Takeda Y, Yoshida J, Miwa T, Okamoto M, Masuyama T, Nonaka Y, Hori M. Elevated cardiac tissue level of aldosterone and mineralocorticoid receptor in diastolic heart failure: beneficial effects of mineralocorticoid receptor blocker. Am J Physiol Regul Integr Comp Physiol 292: R946-R954, 2007.
65. Ottaviani E, Franchini A, Franceschi C. Evolution of neuroendocrine thymus: studies on POMC-derived peptides, cytokines and apoptosis in lower and higher vertebrates. J Neuroimmunol 72: 67-74, 1997.
66. Ottaviani E, Franchini A, Franceschi C. Presence of immunoreactive corticotropin-releasing hormone and cortisol molecules in invertebrate haemocytes and lower and higher vertebrate thymus. Histochem J 30: 61-67, 1998.
67. Passier RC, Smits JF, Verluyten MJ, Daemen MJ. Expression and localization of renin and angiotensinogen in rat heart after myocardial infarction. Am J Physiol Heart Circ Physiol 271: H1040-H1048, 1996.
68. Pazirandeh A, Jondal M, Okret S. Glucocorticoids delay age-associated thymic involution through directly affecting the thymocytes. Endocrinology 145: 2392-2401, 2004.
69. Pazirandeh A, Xue Y, Rafter I, Sjovall J, Jondal M, Okret S. Paracrine glucocorticoid activity produced by mouse thymic epithelial cells. FASEB J 13: 893-901, 1999.
70. Pazirandeh A, Xue Y, Prestegaard T, Jondal M, Okret S. Effects of altered glucocorticoid sensitivity in the T cell lineage on thymocyte and T cell homeostasis. FASEB J 16: 727-729, 2002.
71. Pitt B, Zannad F, Remme WJ, Cody R, Castaigne A, Perez A, Palensky J, Wittes J. The effect of spironolactone on morbidity and mortality in patients with severe heart failure. Randomized Aldactone Evaluation Study Investigators. N Engl J Med 341: 709-717, 1999.
72. Pradhan DS, Newman AEM, Wacker DW, Wingfield JC, Schlinger BA, Soma KK. Aggressive interactions rapidly increase androgen synthesis in the brain during the non-breeding season. Horm Behav 57: 381-389, 2010.
73. Provost PR, Tremblay Y. Genes involved in the adrenal pathway of glucocorticoid synthesis are transiently expressed in the developing lung. Endocrinology 146: 2239-2245, 2005.
74. Purton JF, Boyd RL, Cole TJ, Godfrey DI. Intrathymic T-cell development and selection proceeds normally in the absence of glucocorticoid receptor signaling. Immunity 13: 179-186, 2000.
75. Qiao S, Chen L, Okret S, Jondal M. Age-related synthesis of glucocorticoids in thymocytes. Exp Cell Res 314: 3027-3035, 2008.
76. Qiao S, Okret S, Jondal M. Thymocyte-synthesized glucocorticoids play a role in thymocyte homeostasis and are down-regulated by adrenocorticotropic hormone. Endocrinology 150: 4163-4169, 2009.
77. Raza K, Hardy R, Cooper MS. The 11beta-hydroxysteroid dehydrogenase enzymes-arbiters of the effects of glucocorticoids in synovium and bone. Rheumatology 49: 2016-2023, 2010.
78. Remage-Healey L, London SE, Schlinger BA. Birdsong and the neural production of steroids. J Chem Neuroanat 39: 72-81, 2010.
79. Revskoy S, Halasz I, Redei E. Corticotropin-releasing hormone and proopiomelanocortin gene expression is altered selectively in the male rat fetal thymus by maternal alcohol consumption. Endocrinology 138: 389-396, 1997.
80. Robel P, Baulieu EE. Neurosteroids: biosynthesis and function. Trends Endocrinol Metab 5: 1-8, 1994.
81. Rogoff D, Gomez-Sanchez CE, Foecking MF, Wortsman J, Slominski A. Steroidogenesis in the human skin: 21-hydroxylation in cultured keratinocytes. J Steroid Biochem Mol Biol 78: 77-81, 2001.
82. Roloff B, Fechner K, Slominski A, Furkert J, Botchkarev VA, Bulfone-Paus S, Zipper J, Krause E, Paus R. Hair cycle-dependent expression of corticotropin-releasing factor (CRF) and CRF receptors in murine skin. FASEB J 12: 287-297, 1998.
83. Sakai RR, Nicolaidis S, Epstein AN. Salt appetite is suppressed by interference with angiotensin II and aldosterone. Am J Physiol Regul Integr Comp Physiol 251: R762-R768, 1986.
84. Sapolsky RM, Romero LM, Munck AU. How do glucocorticoids influence stress responses? Integrating permissive, suppressive, stimulatory, and preparative actions. Endocr Rev 21: 55-89, 2000.
85. Schmidt KL, Chin EH, Shah AH, Soma KK. Cortisol and corticosterone in immune organs and brain of European starlings: developmental changes, effects of restraint stress, comparison with zebra finches. Am J Physiol Regul Integr Comp Physiol 297: R42-R51, 2009.
86. Schmidt KL, Malisch JL, Breuner CW, Soma KK. Corticosterone and cortisol binding sites in plasma, immune organs and brain of developing zebra finches: intracellular and membrane-associated receptors. Brain Behav Immun 24: 908-918, 2010.
87. Schmidt KL, Pradhan DS, Shah AH, Charlier TD, Chin EH, Soma KK. Neurosteroids, immunosteroids, and the Balkanization of endocrinology. Gen Comp Endocrinol 157: 266-274, 2008.
88. Schmidt KL, Soma KK. Cortisol and corticosterone in the songbird immune and nervous systems: local vs. systemic levels during development. Am J Physiol Regul Integr Comp Physiol 295: R103-R110, 2008.
89. Schmidt M, Enthoven L, van der Mark M, Levine S, de Kloet ER, Oitzl MS. The postnatal development of the hypothalamic-pituitaryadrenal axis in the mouse. Int J Dev Neurosci 21: 125-132, 2003.
90. Silvestre JS, Robert V, Heymes C, Aupetit-Faisant B, Mouas C, Moalic JM, Swynghedauw B, Delcayre C. Myocardial production of aldosterone and corticosterone in the rat. J Biol Chem 273: 4883-4891, 1998.
91. Silvestre JS, Heymes C, Oubenaissa A, Robert V, Aupetit-Faisant B, Carayon A, Swynghedauw B, Delcayre C. Activation of cardiac aldosterone production in rat myocardial infarction: effect of angiotensin II receptor blockade and role in cardiac fibrosis. Circulation 99: 2694-2701, 1999.
92. Simard J, Couet J, Durocher F, Labrie Y, Sanchez R, Breton N, Turgeon C, Labrie F. Structure and tissue-specific expression of a novel member of the rat 3β-hydroxysteroid dehydrogenase/Δ5-Δ4 isomerase (3β-HSD) family. J Biol Chem 268: 19659-19668, 1993.
93. Slominski A, Wortsman J. Neuroendocrinology of the skin. Endocr Rev 21: 457-487, 2000.
94. Slominski A, Ermak G, Mihm M. ACTH receptor, CYP11A1, CYP17 and CYP21A2 genes are expressed in the skin. J Clin Endocrinol Metab 81: 2746-2749, 1996.
95. Slominski A, Gomez-Sanchez CE, Foecking MF, Wortsman J. Active steroidogenesis in the normal rat skin. Biochim Biophys Acta 1474: 1-4, 2000.
96. Slominski A, Zbytek B, Semak I, Sweatman T, Wortsman J. CRH stimulates POMC activity and corticosterone production in dermal fibroblasts. J Neuroimmunol 162: 97-102, 2005.
97. Slominski A, Zbytek B, Szczesniewski A, Semak I, Kaminski J, Sweatman T, Wortsman J. CRH stimulation of corticosteroids production in melanocytes is mediated by ACTH. Am J Physiol Endocrinol Metab 288: E701-E706, 2005.
98. Slominski A, Zbytek B, Szczesniewski A, Wortsman J. Cultured human dermal fibroblasts do produce cortisol. J Invest Dermatol 126: 1177-1178, 2006.
99. Slominski A, Zjawiony J, Wortsman J, Semak I, Stewart J, Pisarchik A, Sweatman T, Marcos J, Dunbar C, Tuckey RC. A novel pathway for sequential transformation of 7-dehydrocholesterol and the expression of the P450scc system in mammalian skin. Eur J Biochem 271: 4178-4188, 2004.
100. Soma KK. Testosterone and aggression: Berthold, birds and beyond. J Neuroendocrinol 18: 543-551, 2006.
101. Stromstedt M, Waterman MR. Messenger RNAs encoding steroidogenic enzymes are expressed in rodent brain. Mol Brain Res 34: 75-88, 1995.
102. Takeda R, Hatakeyama H, Takeda Y, Iki K, Miyamori I, Sheng WP, Yamamoto H, Blair IA. Aldosterone biosynthesis and action in vascular cells. Steroids 60: 120-124, 1995.
103. Takeda Y, Miyamori I, Yoneda T, Iki K, Hatakeyama H, Blair IA, Hsieh FY, Takeda R. Synthesis of corticosterone in the vascular wall. Endocrinology 135: 2283-2286, 1994.
104. Taves MD, Schmidt KL, Ruhr IM, Kapusta K, Prior NH, Soma KK. Steroid concentrations in plasma, whole blood and brain: effects of saline perfusion to remove blood contamination from brain. PLoS ONE 5: e15727, 2010.
105. Teofoli P, Frezzolini A, Puddu P, De Pita O, Mauviel A, Lotti T. The role of proopiomelanocortin-derived peptides in skin fibroblast and mast cell functions. Ann NY Acad Sci 885: 268-276, 1999.
106. Thiboutot D, Jabara S, McAllister JM, Sivarajah A, Gilliland K, Cong Z, Clawson G. Human skin is a steroidogenic tissue: steroidogenic enzymes and cofactors are expressed in epidermis, normal sebocytes, and an immortalized sebocyte cell line (SEB-1). J Invest Dermatol 120: 905-914, 2003.
107. Tsutamoto T, Wada A, Maeda K, Mabuchi N, Hayashi M, Tsutsui T, Ohnishi M, Sawaki M, Fujii M, Matsumoto T, Horie H, Sugimoto Y, Kinoshita M. Spironolactone inhibits the transcardiac extraction of aldosterone in patients with congestive heart failure. J Am Coll Cardiol 36: 838-844, 2000.
108. Tsutsui K, Ukena K, Usui M, Sakamoto H, Takase M. Novel brain function: biosynthesis and actions of neurosteroids in neurons. Neurosci Res 36: 261-273, 2000.
109. Vacchio MS, Ashwell JD. Thymus-derived glucocorticoids regulate antigen-specific positive selection. J Exp Med 185: 2033-2038, 1997.
110. Vacchio MS, Papadopoulos V, Ashwell JD. Steroid production in the thymus: implications for thymocyte selection. J Exp Med 179: 1835-1846, 1994.
111. van Kats JP, Danser AH, van Meegen JR, Sassen LM, Verdouw PD, Schalekamp MA. Angiotensin production by the heart: a quantitative study in pigs with the use of radiolabeled angiotensin infusions. Circulation 98: 73-81, 1998.
112. Vukelic S, Stojadinovic O, Pastar I, Rabach M, Krzyzanowska A, Lebrun E, Davis SC, Resnik S, Brem H, Tomic-Canic M. Cortisol synthesis in epidermis is induced by IL-1 and tissue injury. J Biol Chem 286: 10265-10275, 2011.
113. Wada H, Salvante KG, Wagner E, Williams TD, Breuner CW. Ontogeny and individual variation in the adrenocortical response of zebra finch (Taeniopygia guttata) nestlings. Physiol Biochem Zool 82: 325-331, 2009.
114. Wingfield JC, Hahn TP. Testosterone and territorial behaviour in sedentary and migratory sparrows. Anim Behav 47: 77-89, 1994.
115. Xue C, Siragy HM. Local renal aldosterone system and its regulation by salt, diabetes, and angiotensin II type 1 receptor. Hypertension 46: 584-590, 2005.
116. Yamamoto N, Yasue H, Mizuno Y, Yoshimura M, Fujii H, Nakayama M, Harada E, Nakamura S, Ito T, Ogawa H. Aldosterone is produced from ventricles in patients with essential hypertension. Hypertension 39: 958-962, 2002.
117. Ye P, Kenyon CJ, MacKenzie SM, Jong AS, Miller C, Gray GA, Wallace A, Ryding AS, Mullins JJ, McBride MW, Graham D, Fraser R, Connell JMC, Davies E. The aldosterone synthase (CYP11B2) and 11beta-hydroxylase (CYP11B1) genes are not expressed in the rat heart. Endocrinology 146: 5287-5293, 2005.
118. Ye P, Kenyon CJ, MacKenzie SM, Seckl JR, Fraser R, Connell JMC, Davies E. Regulation of aldosterone synthase gene expression in the rat adrenal gland and central nervous system by sodium and angiotensin II. Endocrinology 144: 3321-3328, 2003.
119. Ye P, Kenyon CJ, Mackenzie SM, Nichol K, Seckl JR, Fraser R, Connell JMC, Davies E. Effects of ACTH, dexamethasone, and adrenalectomy on 11beta-hydroxylase (CYP11B1) and aldosterone synthase (CYP11B2) gene expression in the rat central nervous system. J Endocrinol 196: 305-311, 2008.
120. Young MJ, Clyne CD, Cole TJ, Funder JW. Cardiac steroidogenesis in the normal and failing heart. J Clin Endocrinol Metab 86: 5121-5126, 2001.
121. Yu L, Romero DG, Gomez-Sanchez CE, Gomez-Sanchez EP. Steroidogenic enzyme gene expression in the human brain. Mol Cell Endocrinol 190: 9-17, 2002.
122. Zmijewski MA, Sharma RK, Slominski A. Expression of molecular equivalent of hypothalamic-pituitary-adrenal axis in adult retinal pigment epithelium. J Endocrinol 193: 157-169, 2007.