11β-Hydroxysteroid Dehydrogenase Type 1: A Tissue-Specific Regulator of Glucocorticoid Response

Endocrine Reviews - Tập 25 Số 5 - Trang 831-866 - 2004
Jeremy Tomlinson1, Elizabeth A. Walker1, Iwona Bujalska1, Norman R. Draper1, Gareth G. Lavery1, Mark S. Cooper1, Martin Hewison1, Paul M. Stewart1
1Endocrinology, Division of Medical Sciences, University of Birmingham, Queen Elizabeth Hospital, Edgbaston, Birmingham B15 2TH, United Kingdom

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White, 1997, 11 β-Hydroxysteroid dehydrogenase and the syndrome of apparent mineralocorticoid excess., Endocr Rev, 18, 135

Munck, 1992, The ups and downs of glucocorticoid physiology, Permissive and suppressive effects revisited. Mol Cell Endocrinol, 90, C1, 10.1016/0303-7207(92)90091-J

Marver, 1984, Evidence of corticosteroid action along the nephron, Am J Physiol, 246, F111

Cope, 1958, The production rate of cortisol in man., Lancet, 1, 1020

Esteban, 1991, Daily cortisol production rate in man determined by stable isotope dilution/mass spectrometry., J Clin Endocrinol Metab, 72, 39, 10.1210/jcem-72-1-39

Peterson, 1960, The metabolism of corticosterone in man., J Clin Invest, 39, 741, 10.1172/JCI104091

Jones, 1959, Aldosterone secretion and metabolism in normal men and women and in pregnancy., Acta Endocrinol (Copenh), 30, 321, 10.1530/acta.0.0300321

Monder, 1984, 11β-Hydroxysteroid dehydrogenase: fact or fancy?, Steroids, 44, 383, 10.1016/S0039-128X(84)80001-X

Monder, 1993, 11β-Hydroxysteroid dehydrogenase., Vitam Horm, 47, 187, 10.1016/S0083-6729(08)60447-1

McGuire, 1959, The multiplicity and specificity of δ4-3-ketosteroid hydrogenases., Arch Biochem Biophys, 82, 476, 10.1016/0003-9861(59)90146-8

Cope, 1972, 80

Shackleton, 1993, Mass spectrometry in the diagnosis of steroid-related disorders and hypertension research., J Steroid Biochem Mol Biol, 45, 127, 10.1016/0960-0760(93)90132-G

Monder, 1980, Cortoic acids: explorations at the frontier of corticosteroid metabolism., Recent Prog Horm Res, 36, 345

Fukushima, 1960, Metabolic transformation of hydrocortisone-4-C14 in normal men., J Biol Chem, 235, 2246, 10.1016/S0021-9258(18)64606-X

Tortorella, 1999, In vivo evidence that human adrenal glands possess 11 β-hydroxysteroid dehydrogenase activity., Life Sci, 65, 2823, 10.1016/S0024-3205(99)00551-2

Meulenberg, 1990, The effect of pretreatment of saliva on steroid hormone concentrations., J Clin Chem Clin Biochem, 28, 923

Meulenberg, 1990, The effect of oral contraceptive use and pregnancy on the daily rhythm of cortisol and cortisone., Clin Chim Acta, 190, 211, 10.1016/0009-8981(90)90175-R

Kendall, 1971

Osinski, 1960, Steroid 11β-ol dehydrogenase in human placenta., Nature, 187, 777, 10.1038/187777a0

Jenkins, 1966, The metabolism of cortisol by human extrahepatic tissues., J Endocrinol, 34, 51, 10.1677/joe.0.0340051

Bush, 1969, 11β-Hydroxysteroid dehydrogenase: contrast between studies in vivo and studies in vitro., Adv Biosci, 3, 23

Hellman, 1971, Renal capture and oxidation of cortisol in man., J Clin Endocrinol, 33, 52, 10.1210/jcem-33-1-52

Srivastava, 1973, Plasma cortisone concentration as measured by radioimmunoassay., J Clin Endocrinol Metab, 36, 937, 10.1210/jcem-36-5-937

Whitworth, 1989, The kidney is the major site of cortisone production in man., Clin Endocrinol (Oxf), 31, 355, 10.1111/j.1365-2265.1989.tb01259.x

Walker, 1992, Glucocorticoids and blood pressure: a role for the cortisol/cortisone shuttle in the control of vascular tone in man., Clin Sci, 83, 171, 10.1042/cs0830171

Palermo, 1996, Urinary free cortisone and the assessment of 11β-hydroxysteroid dehydrogenase activity in man., Clin Endocrinol (Oxf), 45, 605, 10.1046/j.1365-2265.1996.00853.x

Weaver, 1994, The effect of growth hormone replacement on cortisol metabolism and glucocorticoid sensitivity in hypopituitary adults., Clin Endocrinol (Oxf), 41, 639, 10.1111/j.1365-2265.1994.tb01830.x

Weaver, 1998, Sexual dimorphism in 11 beta hydroxysteroid dehydrogenase activity and its relation to fat distribution and insulin sensitivity; a study in hypopituitary subjects., Clin Endocrinol (Oxf), 49, 13, 10.1046/j.1365-2265.1998.00494.x

Toogood, 2000, Modulation of cortisol metabolism by low-dose growth hormone replacement in elderly hypopituitary patients., J Clin Endocrinol Metab, 85, 1727

Trainer, 2001, Modulation of cortisol metabolism by the growth hormone receptor antagonist pegvisomant in patients with acromegaly., J Clin Endocrinol Metab, 86, 2989

Persson, 1993, Dual relationships of xylitol and alcohol dehydrogenases in families of two protein types., FEBS Lett, 324, 9, 10.1016/0014-5793(93)81522-2

Walter, 1992, Molecular characterization of two Clostridium acetobutylicum ATCC 824 butanol dehydrogenase isozyme genes., J Bacteriol, 174, 7149, 10.1128/jb.174.22.7149-7158.1992

Bohren, 1989, The aldo-keto reductase superfamily. cDNAs and deduced amino acid sequences of human aldehyde and aldose reductases., J Biol Chem, 264, 9547, 10.1016/S0021-9258(18)60566-6

Nordling, 2002, Medium-chain dehydrogenases/reductases (MDR). Family characterizations including genome comparisons and active site modeling., Eur J Biochem, 269, 4267, 10.1046/j.1432-1033.2002.03114.x

Persson, 1991, Characteristics of short-chain alcohol dehydrogenases and related enzymes., Eur J Biochem, 200, 537, 10.1111/j.1432-1033.1991.tb16215.x

Baker, 1996, Unusual evolution of 11β- and 17β-hydroxysteroid and retinol dehydrogenases., Bioessays, 18, 63, 10.1002/bies.950180112

Jornvall, 1995, Short-chain dehydrogenases/reductases (SDR)., Biochemistry, 34, 6003, 10.1021/bi00018a001

Persson, 2003, Coenzyme-based functional assignments of short-chain dehydrogenases/reductases (SDRs), Chem Biol Interact, 143–144, 271, 10.1016/S0009-2797(02)00223-5

Krozowski, 1994, The short-chain alcohol dehydrogenase superfamily: variations on a common theme., J Steroid Biochem Mol Biol, 51, 125, 10.1016/0960-0760(94)90084-1

Oppermann, 2001, Forms and functions of human SDR enzymes, Chem Biol Interact, 130–132, 699, 10.1016/S0009-2797(00)00301-X

Oppermann, 2003, Short-chain dehydrogenases/reductases (SDR): the 2002 update, Chem Biol Interact, 143–144, 247, 10.1016/S0009-2797(02)00164-3

Stewart, 1999, 11β-Hydroxysteroid dehydrogenase., Vitam Horm, 57, 249, 10.1016/S0083-6729(08)60646-9

Filling, 2002, Critical residues for structure and catalysis in short-chain dehydrogenases/reductases., J Biol Chem, 277, 25677, 10.1074/jbc.M202160200

Tanabe, 1998, Roles of the Ser146, Tyr159, and Lys163 residues in the catalytic action of 7α-hydroxysteroid dehydrogenase from Escherichia coli., J Biochem (Tokyo), 124, 634, 10.1093/oxfordjournals.jbchem.a022159

Tanaka, 1996, Crystal structures of the binary and ternary complexes of 7α-hydroxysteroid dehydrogenase from Escherichia coli., Biochemistry, 35, 7715, 10.1021/bi951904d

Grundy, 1997, Hidden Markov model analysis of motifs in steroid dehydrogenases and their homologs., Biochem Biophys Res Commun, 231, 760, 10.1006/bbrc.1997.6193

Duax, 2000, Steroid dehydrogenase structures, mechanism of action, and disease., Vitam Horm, 58, 121, 10.1016/S0083-6729(00)58023-6

Duax, 1996, The fascinating complexities of steroid-binding enzymes., Curr Opin Struct Biol, 6, 813, 10.1016/S0959-440X(96)80012-1

Tsigelny, 1995, Structures stabilizing the dimer interface on human 11 β-hydroxysteroid dehydrogenase types 1 and 2 and human 15-hydroxyprostaglandin dehydrogenase and their homologs., Biochem Biophys Res Commun, 217, 859, 10.1006/bbrc.1995.2851

Agarwal, 1989, Cloning and expression of rat cDNA encoding corticosteroid 11β-dehydrogenase., J Biol Chem, 264, 18939, 10.1016/S0021-9258(19)47248-7

Monder, 1989, Evidence for kinetically distinct forms of corticosteroid 11β-dehydrogenase in rat liver microsomes., J Steroid Biochem, 32, 77, 10.1016/0022-4731(89)90017-4

Lakshmi, 1988, Purification and characterization of the corticosteroid 11β-dehydrogenase component of the rat liver 11β-hydroxysteroid dehydrogenase complex., Endocrinology, 123, 2390, 10.1210/endo-123-5-2390

Nobel, 2002, Purification of full-length recombinant human and rat type 1 11β-hydroxysteroid dehydrogenases with retained oxidoreductase activities., Protein Expr Purif, 26, 349, 10.1016/S1046-5928(02)00547-8

Ozols, 1995, Lumenal orientation and post-translational modifications of the liver microsomal 11β-hydroxysteroid dehydrogenase., J Biol Chem [Erratum (1995) 270:10360], 270, 2305, 10.1074/jbc.270.5.2305

Tannin, 1991, The human gene for 11β-hydroxysteroid dehydrogenase. Structure, tissue distribution, and chromosomal localization., J Biol Chem, 266, 16653, 10.1016/S0021-9258(18)55351-5

Rajan, 1995, Cloning, sequencing and tissue-distribution of mouse 11β-hydroxysteroid dehydrogenase-1 cDNA., J Steroid Biochem Mol Biol, 52, 141, 10.1016/0960-0760(94)00159-J

Moore, 1993, Structure and function of the hepatic form of 11β-hydroxysteroid dehydrogenase in the squirrel monkey, an animal model of glucocorticoid resistance., Endocrinology, 133, 368, 10.1210/endo.133.1.8319583

Yang, 1992, Cloning of an ovine 11β-hydroxysteroid dehydrogenase complementary deoxyribonucleic acid: tissue and temporal distribution of its messenger ribonucleic acid during fetal and neonatal development., Endocrinology, 131, 2120, 10.1210/endo.131.5.1425412

Pu, 2000, Guinea pig 11β-hydroxysteroid dehydrogenase type 1: primary structure and catalytic properties., Steroids, 65, 148, 10.1016/S0039-128X(99)00098-7

Shafqat, 2003, Comparative enzymology of 11β-hydroxysteroid dehydrogenase type 1 from glucocorticoid resistant (Guinea pig) versus sensitive (human) species., J Biol Chem, 278, 2030, 10.1074/jbc.M210135200

Maser, 2002, 11 β-Hydroxysteroid dehydrogenase type 1 from human liver: dimerization and enzyme cooperativity support its postulated role as glucocorticoid reductase., Biochemistry, 41, 2459, 10.1021/bi015803t

Maser, 2003, Purification, characterization and NNK carbonyl reductase activities of 11β-hydroxysteroid dehydrogenase type 1 from human liver: enzyme cooperativity and significance in the detoxification of a tobacco-derived carcinogen, Chem Biol Interact, 143–144, 435, 10.1016/S0009-2797(02)00180-1

Condon, 1997, Ontogeny and sexual dimorphic expression of mouse type 2 11β-hydroxysteroid dehydrogenase., Mol Cell Endocrinol, 127, 121, 10.1016/S0303-7207(97)04000-8

Maser, 1994, 11β-Hydroxysteroid dehydrogenase mediates reductive metabolism of xenobiotic carbonyl compounds., Biochem Pharmacol, 47, 1805, 10.1016/0006-2952(94)90309-3

Duperrex, 1993, Rat liver 11β-hydroxysteroid dehydrogenase complementary deoxyribonucleic acid encodes oxoreductase activity in a mineralocorticoid-responsive toad bladder cell line., Endocrinology, 132, 612, 10.1210/endo.132.2.8425481

Low, 1994, ’Liver-type’ 11β-hydroxysteroid dehydrogenase cDNA encodes reductase but not dehydrogenase activity in intact mammalian COS-7 cells., J Mol Endocrinol, 13, 167, 10.1677/jme.0.0130167

Jamieson, 1995, 11β-Hydroxysteroid dehydrogenase is an exclusive 11β-reductase in primary cultures of rat hepatocytes: effect of physicochemical and hormonal manipulations., Endocrinology, 136, 4754, 10.1210/endo.136.11.7588203

Hammami, 1991, Regulation of 11β-hydroxysteroid dehydrogenase activity in human skin fibroblasts: enzymatic modulation of glucocorticoid action., J Clin Endocrinol Metab, 73, 326, 10.1210/jcem-73-2-326

Bujalska, 1997, Does central obesity reflect “Cushing’s disease of the omentum”?, Lancet, 349, 1210, 10.1016/S0140-6736(96)11222-8

Bujalska, 1999, Differentiation of adipose stromal cells: the roles of glucocorticoids and 11β-hydroxysteroid dehydrogenase., Endocrinology, 140, 3188, 10.1210/endo.140.7.6868

Hundertmark, 1995, Correlation of surfactant phosphatidylcholine synthesis and 11β-hydroxysteroid dehydrogenase in fetal lung., Endocrinology, 136, 2573, 10.1210/endo.136.6.7750479

Rajan, 1996, 11β-Hydroxysteroid dehydrogenase in cultured hippocampal cells reactivates inert 11-dehydrocorticosterone, potentiating neurotoxicity., J Neurosci, 16, 65, 10.1523/JNEUROSCI.16-01-00065.1996

Stewart, 1994, Human kidney 11 β-hydroxysteroid dehydrogenase is a high affinity nicotinamide adenine dinucleotide-dependent enzyme and differs from the cloned type I isoform., J Clin Endocrinol Metab, 79, 480

Gao, 1997, Hormonal regulation of oxidative and reductive activities of 11β-hydroxysteroid dehydrogenase in rat Leydig cells., Endocrinology, 138, 156, 10.1210/endo.138.1.4837

Phillips, 1989, Corticosteroid 11β-dehydrogenase in rat testis., Endocrinology, 125, 209, 10.1210/endo-125-1-209

Wang, 2002, Expression of 11β-hydroxylase in rat Leydig cells., Endocrinology, 143, 621, 10.1210/endo.143.2.8638

Leckie, 1998, 11β-Hydroxysteroid dehydrogenase is a predominant reductase in intact rat Leydig cells., J Endocrinol, 159, 233, 10.1677/joe.0.1590233

Bujalska, 2002, A switch in dehydrogenase to reductase activity of 11β-hydroxysteroid dehydrogenase type 1 upon differentiation of human omental adipose stromal cells., J Clin Endocrinol Metab, 87, 1205

Jellinck, 1999, 11β-Hydroxysteroid dehydrogenase functions reversibly as an oxidoreductase in the rat hippocampus in vivo., J Steroid Biochem Mol Biol, 71, 139, 10.1016/S0960-0760(99)00133-8

Seckl, 2001, Minireview: 11β-hydroxysteroid dehydrogenase type 1—a tissue-specific amplifier of glucocorticoid action., Endocrinology, 142, 1371, 10.1210/endo.142.4.8114

Morris, 2003, The functional roles of 11β-HSD1: vascular tissue, testis and brain., Mol Cell Endocrinol, 203, 1, 10.1016/S0303-7207(03)00094-7

Walker, 2001, Functional expression, characterization, and purification of the catalytic domain of human 11-β-hydroxysteroid dehydrogenase type 1., J Biol Chem, 276, 21343, 10.1074/jbc.M011142200

Agarwal, 1990, Expression of 11β-hydroxysteroid dehydrogenase using recombinant vaccinia virus., Mol Endocrinol, 4, 1827, 10.1210/mend-4-12-1827

Draper, 2003, Mutations in the genes encoding 11β-hydroxysteroid dehydrogenase type 1 and hexose-6-phosphate dehydrogenase interact to cause cortisone reductase deficiency., Nat Genet, 34, 434, 10.1038/ng1214

Odermatt, 1999, The N-terminal anchor sequences of 11β-hydroxysteroid dehydrogenases determine their orientation in the endoplasmic reticulum membrane., J Biol Chem, 274, 28762, 10.1074/jbc.274.40.28762

Obeid, 1992, Tyr-179 and Lys-183 are essential for enzymatic activity of 11β-hydroxysteroid dehydrogenase., Biochem Biophys Res Commun, 188, 222, 10.1016/0006-291X(92)92373-6

Mercer, 1993, Characterization of 11β-HSD1B gene expression and enzymatic activity., Mol Cell Endocrinol, 92, 247, 10.1016/0303-7207(93)90015-C

Blum, 2001, Human 11β-hydroxysteroid dehydrogenase 1/carbonyl reductase: additional domains for membrane attachment, Chem Biol Interact, 130–132, 749, 10.1016/S0009-2797(00)00305-7

Agarwal, 1995, Mutations in putative glycosylation sites of rat 11β-hydroxysteroid dehydrogenase affect enzymatic activity., Biochim Biophys Acta, 1248, 70, 10.1016/0167-4838(95)00005-F

Blum, 2000, Human 11β-hydroxysteroid dehydrogenase type 1 is enzymatically active in its nonglycosylated form., Biochem Biophys Res Commun, 276, 428, 10.1006/bbrc.2000.3491

Blum, 2000, Human 11β-hydroxysteroid dehydrogenase 1/carbonyl reductase: recombinant expression in the yeast Pichia pastoris and Escherichia coli., Toxicology, 144, 113, 10.1016/S0300-483X(99)00197-3

Latif, 1997, Selective inhibition of sheep kidney 11β-hydroxysteroid dehydrogenase isoform 2 activity by 5α-reduced (but not 5β) derivatives of adrenocorticosteroids., Steroids, 62, 230, 10.1016/S0039-128X(96)00208-5

Diederich, 2000, In the search for specific inhibitors of human 11β-hydroxysteroid-dehydrogenases (11β-HSDs): chenodeoxycholic acid selectively inhibits 11β-HSD-I., Eur J Endocrinol, 142, 200, 10.1530/eje.0.1420200

Sreerama, 1994, Identification of the class-3 aldehyde dehydrogenases present in human MCF-7/0 breast adenocarcinoma cells and normal human breast tissue., Biochem Pharmacol, 48, 617, 10.1016/0006-2952(94)90294-1

Maser, 1997, Role of type-1 11β-hydroxysteroid dehydrogenase in detoxification processes., Eur J Biochem, 249, 365, 10.1111/j.1432-1033.1997.00365.x

Winzer, 2002, Sex-specific biotransformation and detoxification after xenobiotic exposure of primary cultured hepatocytes of European flounder (Platichthys flesus L.)., Aquat Toxicol, 59, 17, 10.1016/S0166-445X(01)00213-2

Szotakova, 2000, Reduction of the potential anticancer drug oracin in the rat liver in-vitro., J Pharm Pharmacol, 52, 495, 10.1211/0022357001774273

Hult, 2001, Novel enzymological profiles of human 11β-hydroxysteroid dehydrogenase type 1, Chem Biol Interact, 130–132, 805, 10.1016/S0009-2797(00)00236-2

Oppermann, 2000, Molecular and structural aspects of xenobiotic carbonyl metabolizing enzymes. Role of reductases and dehydrogenases in xenobiotic phase I reactions., Toxicology, 144, 71, 10.1016/S0300-483X(99)00192-4

Bannenberg, 2003, 11β-Hydroxysteroid dehydrogenase type 1: tissue-specific expression and reductive metabolism of some anti-insect agent azole analogues of metyrapone, Chem Biol Interact, 143–144, 449, 10.1016/S0009-2797(02)00183-7

Maser, 1998, 11Beta-hydroxysteroid dehydrogenase responsible for carbonyl reduction of the tobacco-specific nitrosamine 4-(methylnitrosamino)-1-(3-pyridyl)-1-butanone in mouse lung microsomes., Cancer Res, 58, 2996

Finckh, 2001, Expression and NNK reducing activities of carbonyl reductase and 11β-hydroxysteroid dehydrogenase type 1 in human lung, Chem Biol Interact, 130–132, 761, 10.1016/S0009-2797(00)00306-9

Maser, 1995, Xenobiotic carbonyl reduction and physiological steroid oxidoreduction. The pluripotency of several hydroxysteroid dehydrogenases., Biochem Pharmacol, 49, 421, 10.1016/0006-2952(94)00330-O

Wsol, 2003, Stereochemical aspects of carbonyl reduction of the original anticancer drug oracin by mouse liver microsomes and purified 11β-hydroxysteroid dehydrogenase type 1, Chem Biol Interact, 143–144, 459, 10.1016/S0009-2797(02)00221-1

Monder, 1989, Licorice inhibits corticosteroid 11β-dehydrogenase of rat liver and kidney: in vivo and in vitro studies., Endocrinology, 124, 1046, 10.1210/endo-125-2-1046

Whorwood, 1993, Licorice inhibits 11β-hydroxysteroid dehydrogenase messenger ribonucleic acid levels and potentiates glucocorticoid hormone action., Endocrinology, 132, 2287, 10.1210/endo.132.6.8504732

Albiston, 1994, Cloning and tissue distribution of the human 11β-hydroxysteroid dehydrogenase type 2 enzyme, Mol Cell Endocrinol, 105, R11, 10.1016/0303-7207(94)90176-7

Bush, 1968, Metabolism of 11-oxygenated steroids. Metabolism in vitro by preparations of liver., Biochem J, 107, 239, 10.1042/bj1070239

Yang, 1994, Evidence for distinct isoforms of 11β-hydroxysteroid dehydrogenase in the ovine liver and kidney., J Steroid Biochem Mol Biol, 49, 245, 10.1016/0960-0760(94)90016-7

Stewart, 1987, Mineralocorticoid activity of liquorice: 11β-hydroxysteroid dehydrogenase deficiency comes of age., Lancet, 2, 821, 10.1016/S0140-6736(87)91014-2

Stewart, 1990, Mineralocorticoid activity of carbenoxolone: contrasting effects of carbenoxolone and liquorice on 11β-hydroxysteroid dehydrogenase activity in man., Clin Sci, 78, 49, 10.1042/cs0780049

Murphy, 1981, Specificity of human 11β-hydroxysteroid dehydrogenase., J Steroid Biochem, 14, 807, 10.1016/0022-4731(81)90019-4

Frey, 1987, Kinetics and dynamics of prednisolone., Endocr Rev, 8, 453, 10.1210/edrv-8-4-453

Ferrari, 1996, Substrate and inhibitor specificity of the cloned human 11β-hydroxysteroid dehydrogenase type 2 isoform, Am J Physiol, 270, E900

Diederich, 2002, 11β-hydroxysteroid dehydrogenase types 1 and 2: an important pharmacokinetic determinant for the activity of synthetic mineralo- and glucocorticoids., J Clin Endocrinol Metab, 87, 5695, 10.1210/jc.2002-020970

Buhler, 1994, Endogenous inhibitors of 11β-OHSD: existence and possible significance., Steroids, 59, 131, 10.1016/0039-128X(94)90090-6

Morita, 1996, 11β-Hydroxysteroid dehydrogenase type 2 complementary deoxyribonucleic acid stably transfected into Chinese hamster ovary cells: specific inhibition by 11 α-hydroxyprogesterone., Endocrinology, 137, 2308, 10.1210/endo.137.6.8641180

Souness, 1996, 11α- And 11 β-hydroxyprogesterone, potent inhibitors of 11β-hydroxysteroid dehydrogenase, possess hypertensinogenic activity in the rat., Hypertension, 27, 421, 10.1161/01.HYP.27.3.421

Alberts, 2002, Selective inhibition of 11β-hydroxysteroid dehydrogenase type 1 decreases blood glucose concentrations in hyperglycaemic mice., Diabetologia, 45, 1528, 10.1007/s00125-002-0959-6

Barf, 2002, Arylsulfonamidothiazoles as a new class of potential antidiabetic drugs. Discovery of potent and selective inhibitors of the 11β-hydroxysteroid dehydrogenase type 1., J Med Chem, 45, 3813, 10.1021/jm025530f

Alberts, 2003, Selective inhibition of 11β-hydroxysteroid dehydrogenase type 1 improves hepatic insulin sensitivity in hyperglycemic mice strains., Endocrinology, 144, 4755, 10.1210/en.2003-0344

Monder, 1990, Corticosteroid 11β-dehydrogenase of rat tissues: immunological studies., Endocrinology, 126, 2435, 10.1210/endo-126-5-2435

Davies, 1997, Cloning and expression of the 11β-hydroxysteroid dehydrogenase type 1 gene in the baboon., Mol Cell Endocrinol, 127, 201, 10.1016/S0303-7207(97)04008-2

Kong, 2002, Absence of type 1 11β-hydroxysteroid dehydrogenase enzyme in koala liver., Comp Biochem Physiol C Toxicol Pharmacol, 131, 39, 10.1016/S1532-0456(01)00278-2

Blum, 2003, Functional characterization of the human 11β-hydroxysteroid dehydrogenase 1B (11β-HSD 1B) variant., Biochemistry, 42, 4108, 10.1021/bi027425d

Yang, 1995, Identification and tissue distribution of a novel variant of 11β-hydroxysteroid dehydrogenase 1 transcript., J Steroid Biochem Mol Biol, 55, 247, 10.1016/0960-0760(95)00176-Z

Krozowski, 1992, Tissue-specific expression of an 11β-hydroxysteroid dehydrogenase with a truncated N-terminal domain. A potential mechanism for differential intracellular localization within mineralocorticoid target cells., J Biol Chem, 267, 2569, 10.1016/S0021-9258(18)45918-2

Maser, 1996, The identification of 11β-hydroxysteroid dehydrogenase as carbonyl reductase of the tobacco-specific nitrosamine 4-(methylnitrosamino)-1-(3-pyridyl)-1-butanone., Eur J Biochem, 238, 484, 10.1111/j.1432-1033.1996.0484z.x

Oppermann, 1995, Cloning and primary structure of murine 11β-hydroxysteroid dehydrogenase/microsomal carbonyl reductase., Eur J Biochem, 227, 202, 10.1111/j.1432-1033.1995.tb20377.x

Caramelli, 2001, Lack of mutations of type 1 11β-hydroxysteroid dehydrogenase gene in patients with abdominal obesity., Endocr Res, 27, 47, 10.1081/ERC-100107169

Draper, 2002, Association studies between microsatellite markers within the gene encoding human 11β-hydroxysteroid dehydrogenase type 1 and body mass index, waist to hip ratio, and glucocorticoid metabolism., J Clin Endocrinol Metab, 87, 4984, 10.1210/jc.2001-011375

Moisan, 1992, Differential promoter usage by the rat 11β-hydroxysteroid dehydrogenase gene., Mol Endocrinol, 6, 1082

Williams, 2000, C/EBP regulates hepatic transcription of 11beta -hydroxysteroid dehydrogenase type 1. A novel mechanism for cross-talk between the C/EBP and glucocorticoid signaling pathways., J Biol Chem, 275, 30232, 10.1074/jbc.M001286200

McKnight, 1989, Is CCAAT/enhancer-binding protein a central regulator of energy metabolism?, Genes Dev, 3, 2021, 10.1101/gad.3.12b.2021

Ramos, 1996, Glucocorticoid-stimulated CCAAT/enhancer-binding protein α expression is required for steroid-induced G1 cell cycle arrest of minimal-deviation rat hepatoma cells., Mol Cell Biol, 16, 5288, 10.1128/MCB.16.10.5288

Wang, 1995, Impaired energy homeostasis in C/EBPα knockout mice., Science, 269, 1108, 10.1126/science.7652557

Screpanti, 1995, Lymphoproliferative disorder and imbalanced T-helper response in C/EBPβ-deficient mice., EMBO J, 14, 1932, 10.1002/j.1460-2075.1995.tb07185.x

Kotelevtsev, 1997, 11β-Hydroxysteroid dehydrogenase type 1 knockout mice show attenuated glucocorticoid-inducible responses and resist hyperglycemia on obesity or stress., Proc Natl Acad Sci USA, 94, 14924, 10.1073/pnas.94.26.14924

Masuzaki, 2001, A transgenic model of visceral obesity and the metabolic syndrome., Science, 294, 2166, 10.1126/science.1066285

Rask, 2001, Tissue-specific dysregulation of cortisol metabolism in human obesity., J Clin Endocrinol Metab, 86, 1418, 10.1210/jcem.86.3.7453

Smith, 1982, Influence of corticosteroids on glycogen content and steroid 11-reductase activity in lung and liver of the fetal and newborn rat., Biol Neonate, 42, 201, 10.1159/000241600

Hundertmark, 1994, Gestational age dependence of 11β-hydroxysteroid dehydrogenase and its relationship to the enzymes of phosphatidylcholine synthesis in lung and liver of fetal rat., Biochim Biophys Acta, 1210, 348, 10.1016/0005-2760(94)90239-9

Diaz, 1998, Distinct ontogeny of glucocorticoid and mineralocorticoid receptor and 11β-hydroxysteroid dehydrogenase types I and II mRNAs in the fetal rat brain suggest a complex control of glucocorticoid actions., J Neurosci, 18, 2570, 10.1523/JNEUROSCI.18-07-02570.1998

Moisan, 1990, 11β-Hydroxysteroid dehydrogenase bioactivity and messenger RNA expression in rat forebrain: localization in hypothalamus, hippocampus, and cortex., Endocrinology, 127, 1450, 10.1210/endo-127-3-1450

Maser, 1994, Ontogenic pattern of carbonyl reductase activity of 11β-hydroxysteroid dehydrogenase in mouse liver and kidney., Xenobiotica, 24, 109, 10.3109/00498259409043225

Langlois, 1995, Differential expression of 11β-hydroxysteroid dehydrogenase 1 and 2 in the developing ovine fetal liver and kidney., J Endocrinol, 147, 405, 10.1677/joe.0.1470405

Yang, 1995, Developmental and glucocorticoid regulation of pituitary 11β-hydroxysteroid dehydrogenase 1 gene expression in the ovine fetus and lamb., J Mol Endocrinol, 14, 109, 10.1677/jme.0.0140109

Kim, 1995, Characterization of 11β-hydroxysteroid dehydrogenase activity in fetal and adult ovine tissues., Reprod Fertil Dev, 7, 377, 10.1071/RD9950377

Yang, 1997, Cellular localization and developmental regulation of 11β-hydroxysteroid dehydrogenase type 1 (11β-HSD1) gene expression in the ovine placenta., Placenta, 18, 503, 10.1016/0143-4004(77)90003-0

Murphy, 1978, Cortisol production and inactivation by the human lung during gestation and infancy., J Clin Endocrinol Metab, 47, 243, 10.1210/jcem-47-2-243

Abramovitz, 1982, Cortisol-cortisone interconversion in human fetal lung: contrasting results using explant and monolayer cultures suggest that 11β-hydroxysteroid dehydrogenase (EC 1.1.1.146) comprises two enzymes., J Clin Endocrinol Metab, 54, 563, 10.1210/jcem-54-3-563

Stewart, 1995, Type 2 11 β-hydroxysteroid dehydrogenase messenger ribonucleic acid and activity in human placenta and fetal membranes: its relationship to birth weight and putative role in fetal adrenal steroidogenesis., J Clin Endocrinol Metab, 80, 885

Jinno, 2001, Failure of cortisone acetate therapy in 21-hydroxylase deficiency in early infancy., Pediatr Int, 43, 478, 10.1046/j.1442-200X.2001.01432.x

Dimitriou, 2003, Adrenocortical activity in healthy children is associated with fat mass., Am J Clin Nutr, 77, 731, 10.1093/ajcn/77.3.731

Toogood, 2000, Sexual dimorphism of cortisol metabolism is maintained in elderly subjects and is not oestrogen dependent., Clin Endocrinol (Oxf), 52, 61, 10.1046/j.1365-2265.2000.00874.x

Low, 1994, Sexual dimorphism of hepatic 11β-hydroxysteroid dehydrogenase in the rat: the role of growth hormone patterns., J Endocrinol, 143, 541, 10.1677/joe.0.1430541

Albiston, 1995, Sex- and tissue-specific regulation of 11β-hydroxysteroid dehydrogenase mRNA., Mol Cell Endocrinol, 109, 183, 10.1016/0303-7207(95)03501-W

Finken, 1999, Cortisol metabolism in healthy young adults: sexual dimorphism in activities of A-ring reductases, but not 11β-hydroxysteroid dehydrogenases., J Clin Endocrinol Metab, 84, 3316

Stewart, 1991, Localization of renal 11β-dehydrogenase by in situ hybridization: autocrine not paracrine protector of the mineralocorticoid receptor., Endocrinology, 128, 2129, 10.1210/endo-128-4-2129

Yau, 1991, 11β-Hydroxysteroid dehydrogenase mRNA expression in rat kidney, Am J Physiol, 260, F764

Castello, 1989, Immunohistochemical localization of 11-hydroxysteroid dehydrogenase in rat kidney with monoclonal antibody., Ren Physiol Biochem, 12, 320

Edwards, 1988, Localisation of 11β-hydroxysteroid dehydrogenase-tissue specific protector of the mineralocorticoid receptor., Lancet, 2, 986, 10.1016/S0140-6736(88)90742-8

Rundle, 1989, The intrarenal localization of mineralocorticoid receptors and 11β-dehydrogenase: immunocytochemical studies., Endocrinology, 125, 1700, 10.1210/endo-125-3-1700

Escher, 1997, Tumor necrosis factor α and interleukin 1β enhance the cortisone/cortisol shuttle., J Exp Med, 186, 189, 10.1084/jem.186.2.189

Aziz, 1996, Aberrant 11β-hydroxysteroid dehydrogenase-1 activity in the cpk mouse: implications for regulation by the Ke 6 gene., Endocrinology, 137, 5581, 10.1210/endo.137.12.8940387

Burton, 1983, Inactivation of corticosteroids in intestinal mucosa by 11β-hydroxysteroid: NADP oxidoreductase (EC 1.1.1.146)., Am J Gastroenterol, 78, 627

Whorwood, 1994, Epithelial cell localization of type 2 11β-hydroxysteroid dehydrogenase in rat and human colon., Endocrinology, 135, 2533, 10.1210/endo.135.6.7988441

Sheppard, 1996, Specific nuclear localization of 11-dehydrocorticosterone in rat colon: evidence for a novel corticosteroid receptor., Endocrinology, 137, 3274, 10.1210/endo.137.8.8754750

Teelucksingh, 1990, Potentiation of hydrocortisone activity in skin by glycyrrhetinic acid, Lancet, 335, l060, 10.1016/0140-6736(90)92633-S

Ricketts, 1998, Immunohistochemical localization of type 1 11β-hydroxysteroid dehydrogenase in human tissues., J Clin Endocrinol Metab, 83, 1325

Ricketts, 1998, Regulation of 11β-hydroxysteroid dehydrogenase type 1 in primary cultures of rat and human hepatocytes., J Endocrinol, 156, 159, 10.1677/joe.0.1560159

Jamieson, 2000, 11β-Hydroxysteroid dehydrogenase type 1 is a predominant 11β-reductase in the intact perfused rat liver., J Endocrinol, 165, 685, 10.1677/joe.0.1650685

Lax, 1978, The hormonal regulation of hepatic microsomal 11β-hydroxysteroid dehydrogenase activity in the rat., Acta Endocrinol (Copenh), 89, 352, 10.1530/acta.0.0890352

Low, 1993, Regulation of 11β-hydroxysteroid dehydrogenase by sex steroids in vivo: further evidence for the existence of a second dehydrogenase in rat kidney., J Endocrinol, 139, 27, 10.1677/joe.0.1390027

Voice, 1996, 11β-Hydroxysteroid dehydrogenase type 1 expression in 2S FAZA hepatoma cells is hormonally regulated: a model system for the study of hepatic glucocorticoid metabolism., Biochem J, 317, 621, 10.1042/bj3170621

Whorwood, 1993, Tissue specific effects of thyroid hormone on 11β-hydroxysteroid dehydrogenase gene expression., J Steroid Biochem Mol Biol, 46, 539, 10.1016/0960-0760(93)90180-5

Zumoff, 1983, Influence of thyroid function on the in vivo cortisol in equilibrium cortisone equilibrium in man., J Steroid Biochem, 18, 437, 10.1016/0022-4731(83)90062-6

Taniyama, 1993, Urinary cortisol metabolites in the assessment of peripheral thyroid hormone action: application for diagnosis of resistance to thyroid hormone., Thyroid, 3, 229, 10.1089/thy.1993.3.229

Sampath-Kumar, 1997, Metyrapone is a competitive inhibitor of 11β-hydroxysteroid dehydrogenase type 1 reductase., J Steroid Biochem Mol Biol, 62, 195, 10.1016/S0960-0760(97)00027-7

Stewart, 1993, 11β-Hydroxysteroid dehydrogenase deficiency and glucocorticoid status in patients with alcoholic and non-alcoholic chronic liver disease., J Clin Endocrinol Metab, 76, 748

Escher, 1998, Down-regulation of hepatic and renal 11β-hydroxysteroid dehydrogenase in rats with liver cirrhosis., Gastroenterology, 114, 175, 10.1016/S0016-5085(98)70645-6

Quirk, 1990, Epithelial and adipose cells isolated from mammary glands of pregnant and lactating rats differ in 11 β-hydroxysteroid dehydrogenase activity., J Steroid Biochem Mol Biol, 37, 529, 10.1016/0960-0760(90)90397-4

Tomlinson, 2001, Regulation of expression of 11β-hydroxysteroid dehydrogenase type 1 in adipose tissue: tissue-specific induction by cytokines., Endocrinology, 142, 1982, 10.1210/endo.142.5.8168

Handoko, 2000, Insulin attenuates the stimulatory effects of tumor necrosis factor α on 11β-hydroxysteroid dehydrogenase 1 in human adipose stromal cells., J Steroid Biochem Mol Biol, 72, 163, 10.1016/S0960-0760(00)00029-7

Friedberg, 2003, Modulation of 11β-hydroxysteroid dehydrogenase type 1 in mature human subcutaneous adipocytes by hypothalamic messengers., J Clin Endocrinol Metab, 88, 385, 10.1210/jc.2002-020510

Hauner, 1987, Glucocorticoids and insulin promote the differentiation of human adipocyte precursor cells into fat cells., J Clin Endocrinol Metab, 64, 832, 10.1210/jcem-64-4-832

Rogatsky, 1997, Glucocorticoid receptor-mediated cell cycle arrest is achieved through distinct cell-specific transcriptional regulatory mechanisms., Mol Cell Biol, 17, 3181, 10.1128/MCB.17.6.3181

Sanchez, 1993, Glucocorticoids reversibly arrest rat hepatoma cell growth by inducing an early G1 block in cell cycle progression., Cell Growth Differ, 4, 215

Rabbitt, 2002, Prereceptor regulation of glucocorticoid action by 11β-hydroxysteroid dehydrogenase: a novel determinant of cell proliferation., FASEB J, 16, 36, 10.1096/fj.01-0582com

Gregoire, 1991, Glucocorticoids induce a drastic inhibition of proliferation and stimulate differentiation of adult rat fat cell precursors., Exp Cell Res, 196, 270, 10.1016/0014-4827(91)90261-R

Tomlinson, 2002, Expression of 11β-hydroxysteroid dehydrogenase type 1 in adipose tissue is not increased in human obesity., J Clin Endocrinol Metab, 87, 5630, 10.1210/jc.2002-020687

Yang, 1997, 11 β-Hydroxysteroid dehydrogenase 1 activity and gene expression in human adipose stromal cells: effect on aromatase activity., J Steroid Biochem Mol Biol, 60, 247, 10.1016/S0960-0760(96)00187-2

Napolitano, 1998, 11β-hydroxysteroid dehydrogenase 1 in adipocytes: expression is differentiation-dependent and hormonally regulated., J Steroid Biochem Mol Biol, 64, 251, 10.1016/S0960-0760(97)00200-8

Jessen, 2002, Expression profiling during adipocyte differentiation of 3T3–L1 fibroblasts., Gene, 299, 95, 10.1016/S0378-1119(02)01017-X

Berger, 2001, Peroxisome proliferator-activated receptor-γ ligands inhibit adipocyte 11β-hydroxysteroid dehydrogenase type 1 expression and activity., J Biol Chem, 276, 12629, 10.1074/jbc.M003592200

Stulnig, 2002, Liver X receptors downregulate 11β-hydroxysteroid dehydrogenase type 1 expression and activity., Diabetes, 51, 2426, 10.2337/diabetes.51.8.2426

Sasaki, 1984, Multihormonal regulation of phosphoenolpyruvate carboxykinase gene transcription. The dominant role of insulin., J Biol Chem, 259, 15242, 10.1016/S0021-9258(17)42541-5

Jamieson, 1999, Tissue- and temporal-specific regulation of 11β-hydroxysteroid dehydrogenase type 1 by glucocorticoids in vivo., J Steroid Biochem Mol Biol, 68, 245, 10.1016/S0960-0760(99)00037-0

Walker, 1995, Carbenoxolone increases hepatic insulin sensitivity in man: a novel role for 11-oxosteroid reductase in enhancing glucocorticoid receptor activation., J Clin Endocrinol Metab, 80, 3155

Andrews, 2003, Effects of the 11β-hydroxysteroid dehydrogenase inhibitor carbenoxolone on insulin sensitivity in men with type 2 diabetes., J Clin Endocrinol Metab, 88, 285, 10.1210/jc.2002-021194

Andrews, 2002, Abnormal cortisol metabolism and tissue sensitivity to cortisol in patients with glucose intolerance., J Clin Endocrinol Metab, 87, 5587, 10.1210/jc.2002-020048

Rask, 2002, Tissue-specific changes in peripheral cortisol metabolism in obese women: increased adipose 11β-hydroxysteroid dehydrogenase type 1 activity., J Clin Endocrinol Metab, 87, 3330

Stewart, 1999, Cortisol metabolism in human obesity: impaired cortisone→cortisol conversion in subjects with central adiposity., J Clin Endocrinol Metab, 84, 1022

Liu, 2003, Leptin activation of corticosterone production in hepatocytes may contribute to the reversal of obesity and hyperglycemia in leptin-deficient ob/ob mice., Diabetes, 52, 1409, 10.2337/diabetes.52.6.1409

Kerstens, 2000, Lack of relationship between 11β-hydroxysteroid dehydrogenase setpoint and insulin sensitivity in the basal state and after 24 h of insulin infusion in healthy subjects and type 2 diabetic patients., Clin Endocrinol (Oxf), 52, 403, 10.1046/j.1365-2265.2000.00975.x

Paterson, 2004, Metabolic syndrome without obesity: hepatic overexpression of 11β-hydroxysteroid dehydrogenase type 1 in transgenic mice., Proc Natl Acad Sci USA, 101, 7088, 10.1073/pnas.0305524101

Livingstone, 2000, Understanding the role of glucocorticoids in obesity: tissue-specific alterations of corticosterone metabolism in obese Zucker rats., Endocrinology, 141, 560, 10.1210/endo.141.2.7297

Livingstone, 2000, Mechanisms of dysregulation of 11β-hydroxysteroid dehydrogenase type 1 in obese Zucker rats., J Endocrinol, 167, 533, 10.1677/joe.0.1670533

Mattsson, 2003, Obese Zucker rats have reduced mineralocorticoid receptor and 11β-hydroxysteroid dehydrogenase type 1 expression in hippocampus—implications for dysregulation of the hypothalamic-pituitary-adrenal axis in obesity., Endocrinology, 144, 2997, 10.1210/en.2002-221015

Morton, 2004, Down-regulation of adipose 11β-hydroxysteroid dehydrogenase type 1 by high-fat feeding in mice: a potential adaptive mechanism counteracting metabolic disease., Endocrinology, 145, 2707, 10.1210/en.2003-1674

Morton, 2001, Improved lipid and lipoprotein profile, hepatic insulin sensitivity, and glucose tolerance in 11β-hydroxysteroid dehydrogenase type 1 null mice., J Biol Chem, 276, 41293, 10.1074/jbc.M103676200

Masuzaki, 2003, Transgenic amplification of glucocorticoid action in adipose tissue causes high blood pressure in mice., J Clin Invest, 112, 83, 10.1172/JCI17845

Fraser, 1999, Cortisol effects on body mass, blood pressure, and cholesterol in the general population., Hypertension, 33, 1364, 10.1161/01.HYP.33.6.1364

Marin, 1992, Cortisol secretion in relation to body fat distribution in obese premenopausal women., Metabolism, 41, 882, 10.1016/0026-0495(92)90171-6

Paulmyer-Lacroix, 2002, Expression of the mRNA coding for 11β-hydroxysteroid dehydrogenase type 1 in adipose tissue from obese patients: an in situ hybridization study., J Clin Endocrinol Metab, 87, 2701

Csabi, 2000, Urinary cortisol to cortisone metabolites in hypertensive obese children., J Endocrinol Invest, 23, 435, 10.1007/BF03343752

Reynolds, 2001, Altered control of cortisol secretion in adult men with low birth weight and cardiovascular risk factors., J Clin Endocrinol Metab, 86, 245

Andrew, 2002, Glucocorticoid metabolism and the metabolic syndrome: associations in an elderly cohort., Exp Clin Endocrinol Diabetes, 110, 284, 10.1055/s-2002-34591

Andrew, 1998, Obesity and gender influence cortisol secretion and metabolism in man., J Clin Endocrinol Metab, 83, 1806, 10.1210/jcem.83.5.4951

Tiosano, 2003, 11β-Hydroxysteroid dehydrogenase activity in hypothalamic obesity., J Clin Endocrinol Metab, 88, 379, 10.1210/jc.2002-020511

Weidenfeld, 1982, In vitro metabolism of cortisol by human abdominal adipose tissue., J Steroid Biochem, 17, 357, 10.1016/0022-4731(82)90211-4

Katz, 1999, An in vivo study of the cortisol-cortisone shuttle in subcutaneous abdominal adipose tissue., Clin Endocrinol (Oxf), 50, 63, 10.1046/j.1365-2265.1999.00598.x

Lindsay, 2003, Subcutaneous adipose 11β-hydroxysteroid dehydrogenase type 1 activity and messenger ribonucleic acid levels are associated with adiposity and insulinemia in Pima Indians and Caucasians., J Clin Endocrinol Metab, 88, 2738, 10.1210/jc.2002-030017

Wake, 2003, Local and systemic impact of transcriptional up-regulation of 11β-hydroxysteroid dehydrogenase type 1 in adipose tissue in human obesity., J Clin Endocrinol Metab, 88, 3983, 10.1210/jc.2003-030286

Whorwood, 2002, Increased glucocorticoid receptor expression in human skeletal muscle cells may contribute to the pathogenesis of the metabolic syndrome., Diabetes, 51, 1066, 10.2337/diabetes.51.4.1066

Gelding, 1998, The effect of growth hormone replacement therapy on cortisol-cortisone interconversion in hypopituitary adults: evidence for growth hormone modulation of extrarenal 11β-hydroxysteroid dehydrogenase activity., Clin Endocrinol (Oxf), 48, 153, 10.1046/j.1365-2265.1998.3641180.x

Moore, 1999, Modulation of 11β-hydroxysteroid dehydrogenase isozymes by growth hormone and insulin-like growth factor: in vivo and in vitro studies., J Clin Endocrinol Metab, 84, 4172

Liu, 1996, Effect of growth hormone, insulin and dexamethasone on 11β-hydroxysteroid dehydrogenase activity on a primary culture of rat hepatocytes., Life Sci, 59, 227, 10.1016/0024-3205(96)00288-3

Liu, 1997, Sex-specific effects of growth hormone on hepatic 11β-hydroxysteroid dehydrogenase activity and gene expression in hypothyroid rats., Life Sci, 61, 325, 10.1016/S0024-3205(97)00389-5

Brown, 1993, Human placental 11β-hydroxysteroid dehydrogenase: evidence for and partial purification of a distinct NAD-dependent isoform., Endocrinology, 132, 2614, 10.1210/endo.132.6.8504762

Krozowski, 1995, Immunohistochemical localization of the 11β-hydroxysteroid dehydrogenase type II enzyme in human kidney and placenta., J Clin Endocrinol Metab, 80, 2203

Lakshmi, 1993, Characterisation of 11β-hydroxysteroid dehydrogenase of human placenta: evidence for the existence of two species of 11β-hydroxysteroid dehydrogenase., J Steroid Biochem Mol Biol, 45, 391, 10.1016/0960-0760(93)90008-K

Pepe, 1996, Interconversion of cortisol and cortisone in the baboon placenta at midgestation: expression of 11β-hydroxysteroid dehydrogenase type 1 messenger RNA., J Steroid Biochem Mol Biol, 58, 403, 10.1016/0960-0760(96)00049-0

Yang, 1995, Co-expression of two distinct isoforms of 11β-hydroxysteroid dehydrogenase in the ovine placenta., J Steroid Biochem Mol Biol, 52, 337, 10.1016/0960-0760(94)00179-P

Lopez, 1981, Corticosteroid metabolism in vitro by human placenta, fetal membranes and decidua in early and late gestation., Placenta, 2, 279, 10.1016/S0143-4004(81)80025-2

Sun, 1997, Differential expression of 11β-hydroxysteroid dehydrogenase types 1 and 2 in human placenta and fetal membranes., J Clin Endocrinol Metab, 82, 300

Smith, 1997, The detection and confirmation of 11β-hydroxysteroid dehydrogenase type 1 transcripts in human luteinized granulosa cells using RT-PCR and plasmid pUC18., Mol Hum Reprod, 3, 651, 10.1093/molehr/3.8.651

Burton, 1998, Immunolocalization of 11β-hydroxysteroid dehydrogenase types 1 and 2 in rat uterus: variation across the estrous cycle and regulation by estrogen and progesterone., Endocrinology, 139, 376, 10.1210/endo.139.1.5692

Thompson, 2002, Spatial and temporal patterns of expression of 11β-hydroxysteroid dehydrogenase types 1 and 2 messenger RNA and glucocorticoid receptor protein in the murine placenta and uterus during late pregnancy., Biol Reprod, 67, 1708, 10.1095/biolreprod.102.005488

Yang, 1996, Pattern of 11β-hydroxysteroid dehydrogenase type 1 messenger ribonucleic acid expression in the ovine uterus during the estrous cycle and pregnancy., Biol Reprod, 55, 1231, 10.1095/biolreprod55.6.1231

Arcuri, 1996, Expression of 11β-hydroxysteroid dehydrogenase during decidualization of human endometrial stromal cells., Endocrinology, 137, 595, 10.1210/endo.137.2.8593807

Saruta, 1986, Multiple factors contribute to the pathogenesis of hypertension in Cushing’s syndrome., J Clin Endocrinol Metab, 62, 275, 10.1210/jcem-62-2-275

Whitworth, 1997, Glucocorticoids and hypertension in man., Clin Exp Hypertens, 19, 871, 10.3109/10641969709083192

Kornel, 1995, Study on the mechanisms of glucocorticoid-induced hypertension: glucocorticoids increase transmembrane Ca2+ influx in vascular smooth muscle in vivo., Endocr Res, 21, 203, 10.3109/07435809509030436

Funder, 1989, Vascular type I aldosterone binding sites are physiological mineralocorticoid receptors., Endocrinology, 125, 2224, 10.1210/endo-125-4-2224

Brem, 1997, Effects of licorice derivatives on vascular smooth muscle function., Life Sci, 60, 207, 10.1016/S0024-3205(96)00617-0

Souness, 2002, 11β-Hydroxysteroid dehydrogenase antisense affects vascular contractile response and glucocorticoid metabolism., Steroids, 67, 195, 10.1016/S0039-128X(01)00148-9

Ullian, 1996, Carbenoxolone damages endothelium and enhances vasoconstrictor action in aortic rings., Hypertension, 27, 1346, 10.1161/01.HYP.27.6.1346

Alzamora, 2000, Role of 11β-hydroxysteroid dehydrogenase in nongenomic aldosterone effects in human arteries., Hypertension, 35, 1099, 10.1161/01.HYP.35.5.1099

van Uum, 2002, Short-term cortisol infusion in the brachial artery, with and without inhibiting 11β-hydroxysteroid dehydrogenase, does not alter forearm vascular resistance in normotensive and hypertensive subjects., Eur J Clin Invest, 32, 874, 10.1046/j.1365-2362.2002.01097.x

Hadoke, 2001, Endothelial cell dysfunction in mice after transgenic knockout of type 2, but not type 1, 11β-hydroxysteroid dehydrogenase., Circulation, 104, 2832, 10.1161/hc4801.100077

Brem, 1998, Localization of 2 11β-OH steroid dehydrogenase isoforms in aortic endothelial cells., Hypertension, 31, 459, 10.1161/01.HYP.31.1.459

Walker, 1991, 11β-Hydroxysteroid dehydrogenase in vascular smooth muscle and heart: implications for cardiovascular responses to glucocorticoids., Endocrinology, 129, 3305, 10.1210/endo-129-6-3305

Brem, 1995, Bidirectional activity of 11β-hydroxysteroid dehydrogenase in vascular smooth muscle cells., Steroids, 60, 406, 10.1016/0039-128X(94)00074-M

Takeda, 1994, Decreased activity of 11β-hydroxysteroid dehydrogenase in mesenteric arteries of Dahl salt-sensitive rats., Life Sci, 54, 1343, 10.1016/0024-3205(94)00513-3

Sheppard, 2002, 11β-Hydroxysteroid dehydrogenase 1 transforms 11-dehydrocorticosterone into transcriptionally active glucocorticoid in neonatal rat heart., Endocrinology, 143, 198, 10.1210/endo.143.1.8583

Young, 2003, Early inflammatory responses in experimental cardiac hypertrophy and fibrosis: effects of 11β-hydroxysteroid dehydrogenase inactivation., Endocrinology, 144, 1121, 10.1210/en.2002-220926

Benediktsson, 1992, 11β-Hydroxysteroid dehydrogenase in the rat ovary: high expression in the oocyte., J Endocrinol, 135, 53, 10.1677/joe.0.1350053

Tetsuka, 1997, Differential expression of messenger ribonucleic acids encoding 11β-hydroxysteroid dehydrogenase types 1 and 2 in human granulosa cells., J Clin Endocrinol Metab, 82, 2006

Hsueh, 1978, Glucocorticoid inhibition of FSH-induced estrogen production in cultured rat granulosa cells., Steroids, 32, 639, 10.1016/0039-128X(78)90074-0

Harlow, 1987, Modulation of plasminogen activation by glucocorticoid hormones in the rat granulosa cell., J Endocrinol, 114, 207, 10.1677/joe.0.1140207

Wang, 1989, Glucocorticoids stimulate plasminogen activator production by rat granulosa cells., Endocrinology, 124, 1595, 10.1210/endo-124-4-1595

Michael, 1993, Direct inhibition of ovarian steroidogenesis by cortisol and the modulatory role of 11β-hydroxysteroid dehydrogenase., Clin Endocrinol (Oxf), 38, 641, 10.1111/j.1365-2265.1993.tb02147.x

Michael, 1997, Isoforms of 11β-hydroxysteroid dehydrogenase in human granulosa-lutein cells., Mol Cell Endocrinol, 132, 43, 10.1016/S0303-7207(97)00118-4

Tetsuka, 1999, Regulation of 11β-hydroxysteroid dehydrogenase type 1 gene expression by LH and interleukin-1β in cultured rat granulosa cells., J Endocrinol, 163, 417, 10.1677/joe.0.1630417

Yong, 2002, Regulation of 11β-hydroxysteroid dehydrogenase type 1 gene expression in human ovarian surface epithelial cells by interleukin-1., Hum Reprod, 17, 2300, 10.1093/humrep/17.9.2300

Hillier, 1998, An anti-inflammatory role for glucocorticoids in the ovaries?, J Reprod Immunol, 39, 21, 10.1016/S0165-0378(98)00011-4

Smith, 2000, Periovulatory human oocytes, cumulus cells, and ovarian leukocytes express type 1 but not type 2 11β-hydroxysteroid dehydrogenase RNA., Fertil Steril, 73, 825, 10.1016/S0015-0282(99)00607-X

Yong, 2000, Development-related increase in cortisol biosynthesis by human granulosa cells., J Clin Endocrinol Metab, 85, 4728

Tetsuka, 2003, Expression of 11β-hydroxysteroid dehydrogenases in bovine follicle and corpus luteum., J Endocrinol, 177, 445, 10.1677/joe.0.1770445

Thurston, 2002, Ovarian modulators of 11β-hydroxysteroid dehydrogenase (11βHSD) activity in follicular fluid from gonadotrophin-stimulated assisted conception cycles., Reproduction, 124, 801, 10.1530/rep.0.1240801

Michael, 1993, Ovarian 11β-hydroxysteroid dehydrogenase: potential predictor of conception by in-vitro fertilisation and embryo transfer., Lancet, 342, 711, 10.1016/0140-6736(93)91710-4

Michael, 1999, Relationship between ovarian cortisol:cortisone ratios and the clinical outcome of in vitro fertilization and embryo transfer (IVF-ET)., Clin Endocrinol (Oxf), 51, 535, 10.1046/j.1365-2265.1999.00892.x

Thomas, 1998, Corticosteroid metabolism in human granulosa-lutein cells., Clin Endocrinol (Oxf), 48, 509, 10.1046/j.1365-2265.1998.00457.x

Harlow, 1997, Increased follicular fluid total and free cortisol levels during the luteinizing hormone surge., Fertil Steril, 68, 48, 10.1016/S0015-0282(97)81474-4

Baillie, 1965, Histochemical demonstration of 11-β-hydroxysteroid dehydrogenase., J Endocrinol, 33, 119, 10.1677/joe.0.0330119

Brereton, 2001, Light and electron microscopy localization of the 11β-hydroxysteroid dehydrogenase type I enzyme in the rat., Endocrinology, 142, 1644, 10.1210/endo.142.4.8088

Waddell, 2003, Localization of 11β-hydroxysteroid dehydrogenase types 1 and 2 in the male reproductive tract., Endocrinology, 144, 3101, 10.1210/en.2003-0082

Hales, 1989, Glucocorticoid-mediated repression of P450scc mRNA and de novo synthesis in cultured Leydig cells., Endocrinology, 124, 2099, 10.1210/endo-124-5-2099

Payne, 1991, Multiple mechanisms for regulation of 3β-hydroxysteroid dehydrogenase/δ 5—δ4-isomerase, 17 α-hydroxylase/C17–20 lyase cytochrome P450, and cholesterol side-chain cleavage cytochrome P450 messenger ribonucleic acid levels in primary cultures of mouse Leydig cells., Endocrinology, 129, 1429, 10.1210/endo-129-3-1429

Monder, 1994, 11β-Hydroxysteroid dehydrogenase alleviates glucocorticoid-mediated inhibition of steroidogenesis in rat Leydig cells., Endocrinology, 134, 1199, 10.1210/endo.134.3.8119160

Monder, 1994, Reciprocal changes in plasma corticosterone and testosterone in stressed male rats maintained in a visible burrow system: evidence for a mediating role of testicular 11β-hydroxysteroid dehydrogenase., Endocrinology, 134, 1193, 10.1210/endo.134.3.8119159

Hardy, 2002, Trends of reproductive hormones in male rats during psychosocial stress: role of glucocorticoid metabolism in behavioral dominance., Biol Reprod, 67, 1750, 10.1095/biolreprod.102.006312

Ge, 1997, Developmental changes in glucocorticoid receptor and 11β-hydroxysteroid dehydrogenase oxidative and reductive activities in rat Leydig cells., Endocrinology, 138, 5089, 10.1210/endo.138.12.5614

Ferguson, 1999, The effects of different culture media, glucose, pyridine nucleotides and adenosine on the activity of 11β-hydroxysteroid dehydrogenase in rat Leydig cells., Mol Cell Endocrinol, 158, 37, 10.1016/S0303-7207(99)00186-0

Ge, 2002, Protein kinase C increases 11β-hydroxysteroid dehydrogenase oxidation and inhibits reduction in rat Leydig cells., J Androl, 23, 135, 10.1002/j.1939-4640.2002.tb02606.x

Ge, 1997, Identification of a kinetically distinct activity of 11β-hydroxysteroid dehydrogenase in rat Leydig cells., Endocrinology, 138, 2435, 10.1210/endo.138.6.5165

Gao, 2002, Glucocorticoid induces apoptosis in rat Leydig cells., Endocrinology, 143, 130, 10.1210/endo.143.1.8604

Gao, 2003, Mechanisms of glucocorticoid-induced Leydig cell apoptosis., Mol Cell Endocrinol, 199, 153, 10.1016/S0303-7207(02)00290-3

Lakshmi, 1991, Regional distribution of 11β-hydroxysteroid dehydrogenase in rat brain., Endocrinology, 128, 1741, 10.1210/endo-128-4-1741

Yau, 2001, Lack of tissue glucocorticoid reactivation in 11β-hydroxysteroid dehydrogenase type 1 knockout mice ameliorates age-related learning impairments., Proc Natl Acad Sci USA, 98, 4716, 10.1073/pnas.071562698

Seckl, 1993, The 11β-hydroxysteroid dehydrogenase inhibitor glycyrrhetinic acid affects corticosteroid feedback regulation of hypothalamic corticotrophin-releasing peptides in rats., J Endocrinol, 136, 471, 10.1677/joe.0.1360471

Seckl, 1991, Use of in situ hybridization to investigate the regulation of hippocampal corticosteroid receptors by monoamines., J Steroid Biochem Mol Biol, 40, 685, 10.1016/0960-0760(91)90291-C

Korbonits, 2001, Expression of 11β-hydroxysteroid dehydrogenase isoenzymes in the human pituitary: induction of the type 2 enzyme in corticotropinomas and other pituitary tumors., J Clin Endocrinol Metab, 86, 2728

Harris, 2001, Intracellular regeneration of glucocorticoids by 11β-hydroxysteroid dehydrogenase (11β-HSD)-1 plays a key role in regulation of the hypothalamic-pituitary-adrenal axis: analysis of 11β-HSD-1-deficient mice., Endocrinology, 142, 114, 10.1210/endo.142.1.7887

Rabbitt, 2003, Abnormal expression of 11β-hydroxysteroid dehydrogenase type 2 in human pituitary adenomas: a prereceptor determinant of pituitary cell proliferation., Oncogene, 22, 1663, 10.1038/sj.onc.1206293

Cooper, 1999, Glucocorticoid activity, inactivity and the osteoblast., J Endocrinol, 163, 159, 10.1677/joe.0.1630159

Manolagas, 2002, Sex steroids and bone., Recent Prog Horm Res, 57, 385, 10.1210/rp.57.1.385

Batista, 1986, Spironolactone-reversible rickets associated with 11 β-hydroxysteroid dehydrogenase deficiency syndrome., J Pediatr, 109, 989, 10.1016/S0022-3476(86)80282-7

Condon, 1998, Expression of type 2 11β-hydroxysteroid dehydrogenase and corticosteroid hormone receptors in early human fetal life., J Clin Endocrinol Metab, 83, 4490

Bland, 1999, Characterization of 11β-hydroxysteroid dehydrogenase activity and corticosteroid receptor expression in human osteosarcoma cell lines., J Endocrinol, 161, 455, 10.1677/joe.0.1610455

Eyre, 2001, Expression of 11β-hydroxysteroid dehydrogenase in rat osteoblastic cells: pre-receptor regulation of glucocorticoid responses in bone., J Cell Biochem, 81, 453, 10.1002/1097-4644(20010601)81:3<453::AID-JCB1059>3.0.CO;2-Z

Woitge, 2001, Cloning and in vitro characterization of α 1(I)-collagen 11β-hydroxysteroid dehydrogenase type 2 transgenes as models for osteoblast-selective inactivation of natural glucocorticoids., Endocrinology, 142, 1341, 10.1210/endo.142.3.8044

Rabbitt, 2003, 11β-Hydroxysteroid dehydrogenases, cell proliferation and malignancy., J Steroid Biochem Mol Biol, 85, 415, 10.1016/S0960-0760(03)00224-3

Bellows, 1998, Osteoprogenitor cells in cell populations derived from mouse and rat calvaria differ in their response to corticosterone, cortisol, and cortisone., Bone, 23, 119, 10.1016/S8756-3282(98)00084-2

Ishida, 2002, Expression of steroid-converting enzymes in osteoblasts derived from rat vertebrae., Osteoporos Int, 13, 235, 10.1007/s001980200020

Cooper, 2000, Expression and functional consequences of 11β-hydroxysteroid dehydrogenase activity in human bone., Bone, 27, 375, 10.1016/S8756-3282(00)00344-6

Cooper, 2002, Osteoblastic 11β-hydroxysteroid dehydrogenase type 1 activity increases with age and glucocorticoid exposure., J Bone Miner Res, 17, 979, 10.1359/jbmr.2002.17.6.979

Cooper, 2001, Modulation of 11β-hydroxysteroid dehydrogenase isozymes by proinflammatory cytokines in osteoblasts: an autocrine switch from glucocorticoid inactivation to activation., J Bone Miner Res, 16, 1037, 10.1359/jbmr.2001.16.6.1037

Canalis, 2002, 11β-Hydroxysteroid dehydrogenase, an amplifier of glucocorticoid action in osteoblasts., J Bone Miner Res, 17, 987, 10.1359/jbmr.2002.17.6.987

Cooper, 2003, 11β-Hydroxysteroid dehydrogenase type 1 activity predicts the effects of glucocorticoids on bone., J Clin Endocrinol Metab, 88, 3874, 10.1210/jc.2003-022025

Armaly, 1967, Dexamethasone ocular hypertension and eosinopenia, and glucose tolerance test., Arch Ophthalmol, 78, 193, 10.1001/archopht.1967.00980030195012

Mirshahi, 2001, Mineralocorticoid hormone signaling regulates the ’epithelial sodium channel’ in fibroblasts from human cornea., Ophthalmic Res, 33, 7, 10.1159/000055635

Weinreb, 1981, Detection of glucocorticoid receptors in cultured human trabecular cells., Invest Ophthalmol Vis Sci, 21, 403

Starka, 1976, Steroids and intraocular pressure., J Steroid Biochem, 7, 979, 10.1016/0022-4731(76)90021-2

Suzuki, 2001, Immunohistochemical distribution of 11β-hydroxysteroid dehydrogenase in human eye., Mol Cell Endocrinol, 173, 121, 10.1016/S0303-7207(00)00403-2

Stokes, 2000, Distribution of glucocorticoid and mineralocorticoid receptors and 11β-hydroxysteroid dehydrogenases in human and rat ocular tissues., Invest Ophthalmol Vis Sci, 41, 1629

Rauz, 2001, Expression and putative role of 11β-hydroxysteroid dehydrogenase isozymes within the human eye., Invest Ophthalmol Vis Sci, 42, 2037

Rauz, 2003, Inhibition of 11β-hydroxysteroid dehydrogenase type 1 lowers intraocular pressure in patients with ocular hypertension., QJM, 96, 481, 10.1093/qjmed/hcg085

Gronau, 2002, 11β-Hydroxysteroid dehydrogenase 1 expression in squamous cell carcinomas of the head and neck., Clin Otolaryngol, 27, 453, 10.1046/j.1365-2273.2002.00609.x

Mune, 2002, Role of local 11β-hydroxysteroid dehydrogenase type 2 (HSD11B2) expression in determining the phenotype of adrenal adenomas., Endocr Res, 28, 751, 10.1081/ERC-120017001

Wall, 1996, CAAT/enhancer-binding proteins are involved in β-globin gene expression and are differentially expressed in murine erythroleukemia and K562 cells., J Biol Chem, 271, 16477, 10.1074/jbc.271.28.16477

Koyama, 2001, Expression of the 11β-hydroxysteroid dehydrogenase type II enzyme in breast tumors and modulation of activity and cell growth in PMC42 cells., J Steroid Biochem Mol Biol, 76, 153, 10.1016/S0960-0760(00)00157-6

Sapolsky, 2000, How do glucocorticoids influence stress responses? Integrating permissive, suppressive, stimulatory, and preparative actions., Endocr Rev, 21, 55

Dougherty, 1960, 11 β-Hydroxy dehydrogenase system activity in thymi of mice following prolonged cortisol treatment., Endocrinology, 66, 550, 10.1210/endo-66-4-550

Hennebold, 1996, 11β-Hydroxysteroid dehydrogenase modulation of glucocorticoid activities in lymphoid organs, Am J Physiol, 270, R1296

Zhou, 1998, Prominent sex steroid metabolism in human lymphocytes., Mol Cell Endocrinol, 138, 61, 10.1016/S0303-7207(98)00052-5

Thieringer, 2001, 11β-Hydroxysteroid dehydrogenase type 1 is induced in human monocytes upon differentiation to macrophages., J Immunol, 167, 30, 10.4049/jimmunol.167.1.30

Marandici, 1993, Inhibition by glycyrrhetinic acid of rat tissue 11β-hydroxysteroid dehydrogenase in vivo., Steroids, 58, 153, 10.1016/0039-128X(93)90062-R

Hennebold, 1997, Active catabolism of glucocorticoids by 11β-hydroxysteroid dehydrogenase in vivo is a necessary requirement for natural resistance to infection with Listeria monocytogenes., Int Immunol, 9, 105, 10.1093/intimm/9.1.105

Rook, 1997, Pathogenetic role, in human and murine tuberculosis, of changes in the peripheral metabolism of glucocorticoids and antiglucocorticoids, Psychoneuroendocrinology, 22, S109, 10.1016/S0306-4530(97)00014-0

Baker, 2000, Increased cortisol: cortisone ratio in acute pulmonary tuberculosis., Am J Respir Crit Care Med, 162, 1641, 10.1164/ajrccm.162.5.9912119

Shimojo, 1996, 11 β-Hydroxysteroid dehydrogenase in the rat adrenal., J Mol Endocrinol, 17, 121, 10.1677/jme.0.0170121

Albertin, 2002, Human adrenal cortex and aldosterone secreting adenomas express both 11β-hydroxysteroid dehydrogenase type 1 and type 2 genes., Int J Mol Med, 9, 495

Hundertmark, 2002, 11β-Hydroxysteroid dehydrogenase type 1: a new regulator of fetal lung maturation., Horm Metab Res, 34, 537, 10.1055/s-2002-35424

Cole, 1995, Targeted disruption of the glucocorticoid receptor gene blocks adrenergic chromaffin cell development and severely retards lung maturation., Genes Dev, 9, 1608, 10.1101/gad.9.13.1608

Hundertmark, 2002, Foetal lung maturation in 11β-hydroxysteroid dehydrogenase type 1 knockout mice., Horm Metab Res, 34, 545, 10.1055/s-2002-35425

Davani, 2000, Type 1 11β-hydroxysteroid dehydrogenase mediates glucocorticoid activation and insulin release in pancreatic islets., J Biol Chem, 275, 34841, 10.1074/jbc.C000600200

Nikkila, 1993, Defects in the HSD11 gene encoding 11β-hydroxysteroid dehydrogenase are not found in patients with apparent mineralocorticoid excess or 11-oxoreductase deficiency., J Clin Endocrinol Metab, 77, 687

Jamieson, 1999, Apparent cortisone reductase deficiency: a functional defect in 11β-hydroxysteroid dehydrogenase type 1., J Clin Endocrinol Metab, 84, 3570

Nordenstrom, 1999, Failure of cortisone acetate treatment in congenital adrenal hyperplasia because of defective 11β-hydroxysteroid dehydrogenase reductase activity., J Clin Endocrinol Metab, 84, 1210

Biason-Lauber, 2000, Apparent cortisone reductase deficiency: a rare cause of hyperandrogenemia and hypercortisolism., Horm Res, 53, 260

Malunowicz, 2003, 11β-Hydroxysteroid dehydrogenase type 1 deficiency (’apparent cortisone reductase deficiency’) in a 6-year-old boy., Horm Res, 59, 205

Laing, 2002, Cortisone reductase deficiency (11β-hydroxysteroid dehydrogenase type 1) deficiency presenting with features of late onset congenital adrenal hyperplasia, Proc 21st Joint Meeting of the British Endocrine Societies, 264

Agarwal, 2001, Transcriptional influence of two poly purine-pyrimidine tracts located in the HSD11B2 (11β-hydroxysteroid dehydrogenase type 2) gene., Endocr Res, 27, 1, 10.1081/ERC-100107163

Kletzien, 1994, Glucose-6-phosphate dehydrogenase: a “housekeeping” enzyme subject to tissue-specific regulation by hormones, nutrients, and oxidant stress., FASEB J, 8, 174, 10.1096/fasebj.8.2.8119488

Mziaut, 1999, Targeting proteins to the lumen of endoplasmic reticulum using N-terminal domains of 11β-hydroxysteroid dehydrogenase and the 50-kDa esterase., J Biol Chem, 274, 14122, 10.1074/jbc.274.20.14122

Taylor, 1990, Corticosteroid 11-reductase deficiency: steroid studies in a further family, J Clin Invest, 13, P238

Kimura, 1979, Glucose dehydrogenase (hexose 6-phosphate dehydrogenase) and the microsomal electron transport system. Evidence supporting their possible functional relationship., J Biochem (Tokyo), 85, 319, 10.1093/oxfordjournals.jbchem.a132336

Stegeman, 1979, A possible role for microsomal hexose-6-phosphate dehydrogenase in microsomal electron transport and mixed-function oxygenase activity., Biochem Biophys Res Commun, 87, 410, 10.1016/0006-291X(79)91811-4

Oka, 1981, Differential effects of the NADPH/NADP+ ratio on the activities of hexose-6-phosphate dehydrogenase and glucose-6-phosphate dehydrogenase., Biochim Biophys Acta, 662, 318, 10.1016/0005-2744(81)90045-0

Kulkarni, 1982, Mouse liver microsomal hexose-6-phosphate dehydrogenase. NADPH generation and utilization in monooxygenation reactions., Biochem Pharmacol, 31, 1131, 10.1016/0006-2952(82)90353-7

Mason, 1999, Human hexose-6-phosphate dehydrogenase (glucose 1-dehydrogenase) encoded at 1p36: coding sequence and expression., Blood Cells Mol Dis, 25, 30, 10.1006/bcmd.1999.0224

Fabsitz, 1992, Evidence for independent genetic influences on obesity in middle age., Int J Obes Relat Metab Disord, 16, 657

Rankinen, 2002, The human obesity gene map: the 2001 update., Obes Res, 10, 196, 10.1038/oby.2002.30

Gelernter-Yaniv, 2003, Associations between a polymorphism in the 11β hydroxysteroid dehydrogenase type I gene and body composition., Int J Obes Relat Metab Disord, 27, 983, 10.1038/sj.ijo.0802327

Burtt, 2003, Haplotype-based association studies of HSD11B1 and abdominal obesity, Program of the 86th annual meeting of The Endocrine Society, 117

Hutton, 1984, Polycystic ovarian syndrome in identical twins., Postgrad Med J, 60, 64, 10.1136/pgmj.60.699.64

McDonough, 1972, Steroid, follicle-stimulating hormone, and luteinizing hormone profiles in identical twins with polycystic ovaries., Am J Obstet Gynecol, 113, 1072, 10.1016/0002-9378(72)90738-7

Legro, 1998, Evidence for a genetic basis for hyperandrogenemia in polycystic ovary syndrome., Proc Natl Acad Sci USA, 95, 14956, 10.1073/pnas.95.25.14956

Givens, 1988, Familial polycystic ovarian disease., Endocrinol Metab Clin North Am, 17, 771, 10.1016/S0889-8529(18)30409-2

Franks, 1999, Genetic abnormalities in polycystic ovary syndrome., Ann Endocrinol (Paris), 60, 131

Rodin, 1994, Hyperandrogenism in polycystic ovary syndrome: evidence of dysregulation of 11β-hydroxysteroid dehydrogenase., N Engl J Med, 330, 460, 10.1056/NEJM199402173300703

Lichter, 1994, Genetic clues to glaucoma’s secrets. The L Edward Jackson Memorial Lecture. Part 2., Am J Ophthalmol, 117, 706, 10.1016/S0002-9394(14)70314-9

Beckman, 1990, Heterozygosity effects in studies of genetic markers and disease., Hum Hered, 40, 322, 10.1159/000153955

Budde, 2000, Heredity in primary open-angle glaucoma., Curr Opin Ophthalmol, 11, 101, 10.1097/00055735-200004000-00006

Ralston, 2002, Genetic control of susceptibility to osteoporosis., J Clin Endocrinol Metab, 87, 2460, 10.1210/jcem.87.6.8621

Devoto, 1998, First-stage autosomal genome screen in extended pedigrees suggests genes predisposing to low bone mineral density on chromosomes 1p, 2p and 4q., Eur J Hum Genet, 6, 151, 10.1038/sj.ejhg.5200169

Devoto, 2001, Variance component linkage analysis indicates a QTL for femoral neck bone mineral density on chromosome 1p36., Hum Mol Genet, 10, 2447, 10.1093/hmg/10.21.2447

Quinkler, 1997, Evidence for isoforms of 11β-hydroxysteroid dehydrogenase in the liver and kidney of the guinea pig., J Endocrinol, 153, 291, 10.1677/joe.0.1530291

Hult, 1998, Selective inhibition of human type 1 11β-hydroxysteroid dehydrogenase by synthetic steroids and xenobiotics., FEBS Lett, 441, 25, 10.1016/S0014-5793(98)01515-4

Bujalska, 2000, Glucocorticoids, 11β-hydroxysteroid dehydrogenase and obesity, PhD thesis, 158

Whorwood, 1995, Detection of human 11β-hydroxysteroid dehydrogenase isoforms using reverse-transcriptase-polymerase chain reaction and localization of the type 2 isoform to renal collecting ducts, Mol Cell Endocrinol, 110, R7, 10.1016/0303-7207(95)03546-J

Cai, 2001, Induction of 11β-hydroxysteroid dehydrogenase type 1 but not -2 in human aortic smooth muscle cells by inflammatory stimuli., J Steroid Biochem Mol Biol, 77, 117, 10.1016/S0960-0760(01)00041-3

Driver, 2001, Expression of 11β-hydroxysteroid dehydrogenase isozymes and corticosteroid hormone receptors in primary cultures of human trophoblast and placental bed biopsies., Mol Hum Reprod, 7, 357, 10.1093/molehr/7.4.357

Pepe, 1999, Expression of the 11β-hydroxysteroid dehydrogenase types 1 and 2 proteins in human and baboon placental syncytiotrophoblast., Placenta, 20, 575, 10.1053/plac.1999.0416

Thurston, 2003, Expression of 11β-hydroxysteroid dehydrogenase (11βHSD) proteins in luteinizing human granulosa-lutein cells., J Endocrinol, 178, 127, 10.1677/joe.0.1780127

Sun, 2003, Enhancement of glucocorticoid-induced 11β-hydroxysteroid dehydrogenase type 1 expression by pro-inflammatory cytokines in cultured human amnion fibroblasts., Endocrinology, 144, 5568, 10.1210/en.2003-0780

Evagelatou, 1997, Leukocytes modulate 11β-hydroxysteroid dehydrogenase (11β-HSD) activity in human granulosa-lutein cell cultures., Mol Cell Endocrinol, 133, 81, 10.1016/S0303-7207(97)00146-9

Tomlinson, 2003, Low-dose growth hormone inhibits 11β-hydroxysteroid dehydrogenase type 1 but has no effect upon fat mass in patients with simple obesity., J Clin Endocrinol Metab, 88, 2113, 10.1210/jc.2002-021894

Sankar, 2000, Chronic corticosterone treatment impairs Leydig cell 11β-hydroxysteroid dehydrogenase activity and LH-stimulated testosterone production., Horm Metab Res, 32, 142, 10.1055/s-2007-978609

Gupta, 2003, Effects of cortisol and oestradiol on hepatic 11β-hydroxysteroid dehydrogenase type 1 and glucocorticoid receptor proteins in late-gestation sheep fetus., J Endocrinol, 176, 175, 10.1677/joe.0.1760175

Nwe, 2000, Differential regulation of the oxidative 11β-hydroxysteroid dehydrogenase activity in testis and liver., Steroids, 65, 40, 10.1016/S0039-128X(99)00078-1

Ma, 2003, Gestation-related and betamethasone-induced changes in 11β-hydroxysteroid dehydrogenase types 1 and 2 in the baboon placenta., Am J Obstet Gynecol, 188, 13, 10.1067/mob.2003.62

Sun, 2002, Intracrine induction of 11β-hydroxysteroid dehydrogenase type 1 expression by glucocorticoid potentiates prostaglandin production in the human chorionic trophoblast., Biol Reprod, 67, 1450, 10.1095/biolreprod.102.005892

Ho, 1999, Regulation of 11β-hydroxysteroid dehydrogenase isoforms and glucocorticoid receptor gene expression in the rat uterus., J Endocrinol, 163, 425, 10.1677/joe.0.1630425

Arcuri, 1997, Human endometrial decidual cell-associated 11 β-hydroxysteroid dehydrogenase expression: its potential role in implantation., Early Pregnancy, 3, 259

Gomez-Sanchez, 2003, Regulation of 11β-hydroxysteroid dehydrogenase enzymes in the rat kidney by estradiol, Am J Physiol Endocrinol Metab, 285, E272, 10.1152/ajpendo.00409.2002

Quinkler, 2003, Enhanced 11β-hydroxysteroid dehydrogenase type 1 activity in stress adaptation in the guinea pig., J Endocrinol, 176, 185, 10.1677/joe.0.1760185

Nwe, 2000, In vivo effects of stress, ACTH and corticosterone on testicular 11β-hydroxysteroid dehydrogenase oxidative activity in rats and the possible mechanism of actions., Exp Clin Endocrinol Diabetes, 108, 369, 10.1055/s-2000-8131

Gu, 2003, Dehydroepiandrosterone affects the expression of multiple genes in rat liver including 11 β-hydroxysteroid dehydrogenase type 1: a cDNA array analysis., Mol Pharmacol, 63, 722, 10.1124/mol.63.3.722

Sun, 1997, Differential regulation of 11β-hydroxysteroid dehydrogenase type 1 and 2 by nitric oxide in cultured human placental trophoblast and chorionic cell preparation., Endocrinology, 138, 4912, 10.1210/endo.138.11.5544

Alfaidy, 2001, Prostaglandin F2α potentiates cortisol production by stimulating 11β-hydroxysteroid dehydrogenase 1: a novel feedback loop that may contribute to human labor., J Clin Endocrinol Metab, 86, 5585

Laplante, 2003, PPAR-γ activation mediates adipose depot-specific effects on gene expression and lipoprotein lipase activity: mechanisms for modulation of postprandial lipemia and differential adipose accretion., Diabetes, 52, 291, 10.2337/diabetes.52.2.291

Hermanowski-Vosatka, 2000, PPARα agonists reduce 11β-hydroxysteroid dehydrogenase type 1 in the liver., Biochem Biophys Res Commun, 279, 330, 10.1006/bbrc.2000.3966

Taylor, 1984, Cortisone reductase deficiency: evidence for a new inborn error in metabolism of adrenal steroids, J Endocrinol, 102, 89

Phillipov, 1996, Apparent cortisone reductase deficiency: a unique form of hypercortisolism., J Clin Endocrinol Metab, 81, 3855

Savage, 1991, Increased metabolic clearance of cortisol in corticosteroid 11-reductase deficiency, J Endocrinol, 129, 219