Synthesis, enzyme localization, and regulation of neurosteroids

Neurosteroid Effects in the Central Nervous System: The Role of the GABA-A Receptor

Volumn , Issue , 2003, Pages 1-46

  Mellon, Synthia H ^a  

^a UNIVERSITY OF CALIFORNIA   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

EID: 84949516544     PISSN: None     EISSN: None     Source Type: Book    
DOI: None     Document Type: Chapter

References (245)

1. Corpechot, C., Robel, P., Axelson, M., Sjovall, J., and Baulieu, E.E., Characterization and measurement of dehydroepiandrosterone sulfate in rat brain, Proc. Natl. Acad. Sci. U.S.A., 78, 4704-4707, 1981.
2. Corpechot, C., Synguelakis, M., Talha, S., Axelson, M., Sjovall, J., Vihko, R., Baulieu, E.E., and Robel, P., Pregnenolone and its sulfate ester in the rat brain, Brain Res., 270, 119-125, 1983.
3. Miller, W.L., Molecular biology of steroid hormone synthesis, Endocr. Rev., 9, 295-318, 1988.
4. Penning, T.M., Molecular endocrinology of hydroxysteroid dehydrogenases, Endocr. Rev., 18, 281-305, 1997.
5. Pang, S., Yang, X., Wang, M., Tissot, R., Nino, M., Manaligod, J., Bullock, L.P., and Mason, J.I., Inherited congenital adrenal hyperplasia in the rabbit: absent cholesterol side-chain cleavage cytochrome P450 gene expression, Endocrinology, 131, 181-186, 1992.
6. Yang, X., Iwamoto, K., Wang, M., Artwohl, J., Mason, J.I., and Pang, S., Inherited congenital adrenal hyperplasia in the rabbit is caused by a deletion in the gene encoding cytochrome P450 cholesterol side-chain cleavage enzyme, Endocrinology, 132, 1977-1982, 1993.
7. Mellon, S.H. and Deschepper, C.F., Neurosteroid biosynthesis: genes for adrenal steroidogenic enzymes are expressed in the brain, Brain Res., 629, 283-292, 1993.
8. Mellon, S.H., Neurosteroids: biochemistry, modes of action, and clinical relevance, J. Clin. Endocrinol. Metab., 78, 1003-1008, 1994.
9. Tajima, T., Fujieda, K., Kouda, N., Nakae, J., and Miller, W.L., Heterozygous mutation in the cholesterol side chain cleavage enzyme (p450scc) gene in a patient with 46, XY sex reversal and adrenal insufficiency, J. Clin. Endocrinol. Metab., 86, 3820-3825, 2001.
10. Matteson, K.J., Chung, B.C., Urdea, M.S., and Miller, W.L., Study of cholesterol side-chain cleavage (20,22 desmolase) deficiency causing congenital lipoid adrenal hyperplasia using bovine-sequence P450scc oligodeoxyribonucleotide probes, Endocrinology, 118, 1296-1305, 1986.
11. Morohashi, K., Sogawa, K., Omura, T., and Fujii-Kuriyama, Y., Gene structure of human cytochrome P-450(SCC), cholesterol desmolase, J. Biochem. (Tokyo), 101, 879-887, 1987.
12. Oonk, R.B., Parker, K.L., Gibson, J.L., and Richards, J.S., Rat cholesterol side-chain cleavage cytochrome P-450 (P-450scc) gene: structure and regulation by cAMP in vitro, J. Biol. Chem., 265, 22392-22401, 1990.
13. Chung, B.C., Matteson, K.J., Voutilainen, R., Mohandas, T.K., and Miller, W.L., Human cholesterol side-chain cleavage enzyme, P450scc: cDNA cloning, assignment of the gene to chromosome 15, and expression in the placenta, Proc. Natl. Acad. Sci. U.S.A., 83, 8962-8966, 1986.
14. Omura, T., Sanders, S., Estabrook, R.W., Cooper, D.Y., and Rosenthal, O., Isolation from adrenal cortex of a non-heme iron protein and a flavoprotein functional as a reduced triphosphopyridine mucleotide-cytochrome P-450 reductase, Arch. Biochem. Biophys., 117, 660, 1966.
15. Nakamura, Y., Otsuka, H., and Tamaoki, B., Requirement of a new flavoprotein and a non-heme iron-containing protein in the steroid 11β - and 18-hydroxylase system, Biochim. Biophys. Acta, 122, 34-42, 1966.
16. Kimura, T. and Suzuki, K., Components of the electron transport system in adrenal steroid hydroxylase, J. Biol. Chem., 242, 485-491, 1967.
17. Coghlan, V.M. and Vickery, L.E., Expression of human ferredoxin and assembly of the [2Fe-2S] center in Escherichia coli, Proc. Natl. Acad. Sci. U.S.A., 86, 835-839, 1989.
18. Stocco, D.M. and Clark, B.J., Regulation of the acute production of steroids in steroidogenic cells, Endocr. Rev., 17, 221-244, 1996.
19. Clark, B.J., Wells, J., King, S.R., and Stocco, D.M., The purification, cloning, and expression of a novel luteinizing hormone-induced mitochondrial protein in MA-10 mouse Leydig tumor cells: characterization of the steroidogenic acute regulatory protein (StAR), J. Biol. Chem., 269, 28314-28322, 1994.
20. Lin, D., Sugawara, T., Strauss, J.F.I., Clark, B.J., Stocco, D.M., Saenger, P., Rogol, A., and Miller, W.L., Role of steroidogenic acute regulatory protein in adrenal and gonadal steroidogenesis, Science, 267, 1828-1831, 1995.
21. Lin, D., Black, S.M., Nagahama, Y., and Miller, W.L., Steroid 17 alpha-hydroxylase and 17,20-lyase activities of P450c17: contributions of serine106 and P450 reductase, Endocrinology, 132, 2498-2506, 1993.
22. Bose, H.S., Sugawara, T., Strauss, J.F.R., and Miller, W.L., The pathophysiology and genetics of congenital lipoid adrenal hyperplasia. International Congenital Lipoid Adrenal Hyperplasia Consortium, N. Engl. J. Med., 335, 1870-1878, 1996.
23. Papadopoulos, V., Peripheral-type benzodiazepine/diazepam binding inhibitor receptor: biological role in steroidogenic cell function, Endocr. Rev., 14, 222-240, 1993.
24. Saenger, P., Klonari, Z., Black, S.M., Compagnone, N., Mellon, S.H., Fleischer, A., Abrams, C.A., Shackelton, C.H., and Miller, W.L., Prenatal diagnosis of congenital lipoid adrenal hyperplasia, J. Clin. Endocrinol. Metab., 80, 200-205, 1995.
25. Furukawa, A., Miyatake, A., Ohnishi, T., and Ichikawa, Y., Steroidogenic acute regulatory protein (StAR) transcripts constitutively expressed in the adult rat central nervous system: colocalization of StAR, cytochrome P-450SCC (CYP XIA1), and 3beta-hydroxysteroid dehydrogenase in the rat brain, J. Neurochem., 71, 2231-2238, 1998.
26. King, S.R., Manna, P.R., Ishii, T., Syapin, P.J., Ginsberg, S.D., Wilson, K., Walsh, L.P., Parker, K.L., Stocco, D.M., Smith, R.G., and Lamb, D.J., An essential component in steroid synthesis, the steroidogenic acute regulatory protein, is expressed in discrete regions of the brain, J. Neurosci., 22, 10613-10620, 2002.
27. Luu-The, V., Lachance, Y., Labrie, C., Leblanc, G., Thomas, J.L., Strickler, R.C., and Labrie, F., Full length cDNA structure and deduced amino acid sequence of human 3 beta-hydroxy-5-ene steroid dehydrogenase, Mol. Endocrinol., 3, 1310-1312, 1989.
28. Thomas, J.L., Myers, R.P., and Strickler, R.C., Human placental 3 beta-hydroxy-5- ene-steroid dehydrogenase and steroid 5--4-ene-isomerase: purification from mitochondria and kinetic profiles, biophysical characterization of the purified mitochondrial and microsomal enzymes, J. Steroid Biochem., 33, 209-217, 1989.
29. Lorence, M.C., Murry, B.A., Trant, J.M., and Mason, J.I., Human 3 beta-hydroxysteroid dehydrogenase/delta 5--4isomerase from placenta: expression in nonsteroidogenic cells of a protein that catalyzes the dehydrogenation/isomerization of C21 and C19 steroids, Endocrinology, 126, 2493-2498, 1990.
30. Voutilainen, R., Ilvesmaki, V., and Miettinen, P.J., Low expression of 3 beta-hydroxy-5-ene steroid dehydrogenase gene in human fetal adrenals in vivo; adrenocorticotropin and protein kinase C-dependent regulation in adrenocortical cultures, J. Clin. Endocrinol. Metab., 72, 761-767, 1991.
31. Endoh, A., Kristiansen, S.B., Casson, P.R., Buster, J.E., and Hornsby, P.J., The zona reticularis is the site of biosynthesis of dehydroepiandrosterone and dehydroepiandrosterone sulfate in the adult human adrenal cortex resulting from its low expression of 3 beta-hydroxysteroid dehydrogenase, J. Clin. Endocrinol. Metab., 81, 3558-3565, 1996.
32. Coulter, C.L., Goldsmith, P.C., Mesiano, S., Voytek, C.C., Martin, M.C., Mason, J.I., and Jaffe, R.B., Functional maturation of the primate fetal adrenal in vivo. II. Ontogeny of corticosteroid synthesis is dependent upon specific zonal expression of 3 betahydroxysteroid dehydrogenase/isomerase, Endocrinology, 137, 4953-4959, 1996.
33. Yanase, T., Sasano, H., Yubisui, T., Sakai, Y., Takayanagi, R., and Nawata, H., Immunohistochemical study of cytochrome b5 in human adrenal gland and in adrenocortical adenomas from patients with Cushing’s syndrome, Endocr. J., 45, 89-95, 1998.
34. Zhang, L.H., Rodriguez, H., Ohno, S., and Miller, W.L., Serine phosphorylation of human P450c17 increases 17,20-lyase activity: implications for adrenarche and the polycystic ovary syndrome, Proc. Natl. Acad. Sci. U.S.A., 92, 10619-10623, 1995.
35. Pandey, A.V., Mellon, S.H., and Miller, W.L., Protein phosphatase 2A and phosphoprotein SET regulate androgen production by P450c17, J. Biol. Chem., 278, 2837-2844, 2003.
36. Matteson, K.J., Picado-Leonard, J., Chung, B.C., Mohandas, T.K., and Miller, W.L., Assignment of the gene for adrenal P450c17 (steroid 17 alpha-hydroxylase/17,20 lyase) to human chromosome 10, J. Clin. Endocrinol. Metab., 63, 789-791, 1986.
37. Casey, M.L. and MacDonald, P.C., Extraadrenal formation of a mineralocorticosteroid: deoxycorticosterone and deoxycorticosterone sulfate biosynthesis and metabolism, Endocr. Rev., 3, 396-403, 1982.
38. Casey, M.L., Winkel, C.A., and MacDonald, P.C., Conversion of progesterone to deoxycorticosterone in the human fetus: steroid 21-hydroxylase activity in fetal tissues, J. Steroid Biochem., 18, 449-452, 1983.
39. Mellon, S.H. and Miller, W.L., Extraadrenal steroid 21-hydroxylation is not mediated by P450c21, J. Clin. Invest., 84, 1497-1502, 1989.
40. Beyenburg, S., Watzka, M., Clusmann, H., Blumcke, I., Bidlingmaier, F., Elger, C.E., and Stoffel-Wagner, B., Messenger RNA of steroid 21-hydroxylase (CYP21) is expressed in the human hippocampus, Neurosci. Lett., 308, 111-114, 2001.
41. Yu, L., Romero, D.G., Gomez-Sanchez, C.E., and Gomez-Sanchez, E.P., Steroidogenic enzyme gene expression in the human brain, Mol. Cell. Endocrinol., 190, 9-17, 2002.
42. Zhou, Z., Agarwal, V.R., Dixit, N., White, P., and Speiser, P.W., Steroid 21-hydroxylase expression and activity in human lymphocytes, Mol. Cell. Endocrinol., 127, 11-18, 1997.
43. Hiroi, T., Kishimoto, W., Chow, T., Imaoka, S., Igarashi, T., and Funae, Y., Progesterone oxidation by cytochrome P450 2D isoforms in the brain, Endocrinology, 142, 3901-3908, 2001.
44. Yanagibashi, K. and Hall, P.F., Role of electron transport in the regulation of the lyase activity of C21 side-chain cleavage P-450 from porcine adrenal and testicular microsomes, J. Biol. Chem., 261, 8429-8433, 1986.
45. Yamano, S., Aoyama, T., McBride, O.W., Hardwick, J.P., Gelboin, H.V., and Gonzalez, F.J., Human NADPH-P450 oxidoreductase: complementary DNA cloning, sequence and vaccinia virus-mediated expression and localization of the CYPOR gene to chromosome 7, Mol. Pharmacol., 36, 83-88, 1989.
46. Onoda, M. and Hall, P.F., Cytochrome b5 stimulates purified testicular microsomal cytochrome P-450 (C21 side-chain cleavage), Biochem. Biophys. Res. Commun., 108, 454-460, 1982.
47. Kominami, S., Ogawa, N., Morimune, R., De-Ying, H., and Takemori, S., The role of cytochrome b5 in adrenal microsomal steroidogenesis, J. Steroid Biochem. Mol. Biol., 42, 57-64, 1992.
48. Auchus, R.J., Lee, T.C., and Miller, W.L., Cytochrome b5 augments the 17,20-lyase activity of human P450c17 without direct electron transfer, J. Biol. Chem., 273, 3158-3165, 1998.
49. Mapes, S., Corbin, C.J., Tarantal, A., and Conley, A., The primate adrenal zona reticularis is defined by expression of cytochrome b5, 17alpha-hydroxylase/17,20- lyase cytochrome P450 (P450c17) and NADPH-cytochrome P450 reductase (reductase) but not 3beta-hydroxysteroid dehydrogenase/delta5-4 isomerase (3beta-HSD), J. Clin. Endocrinol. Metab., 84, 3382-3385, 1999.
50. Mapes, S., Tarantal, A.F., Parker, C.R., Moran, F.M., Bahr, J.M., Pyter, L., and Conley, A.J., Adrenocortical cytochrome b5 expression during fetal development of the rhesus macaque, Endocrinology, 143, 1451-1458, 2002.
51. Mellon, S.H., Bair, S.R., and Monis, H., P450c11B3 mRNA, transcribed from a third P450c11 gene, is expressed in a tissue-specific, developmentally, and hormonally regulated fashion in the rodent adrenal and encodes a protein with both 11-hydroxylase and 18-hydroxylase activities, J. Biol. Chem., 270, 1643-1649, 1995.
52. Zhang, G. and Miller, W.L., The human genome contains only two CYP11B (P450c11) genes, J. Clin. Endocrinol. Metab., 81, 3254-3256, 1996.
53. Thompson, E.A. and Siiteri, P.K., Studies on the aromatization of C-19 androgens, Ann. N.Y. Acad. Sci., 212, 378-391, 1973.
54. Thompson, E.A., Jr. and Siiteri, P.K., The involvement of human placental microsomal cytochrome P-450 in aromatization, J. Biol. Chem., 249, 5373-5378, 1974.
55. Means, G.D., Mahendroo, M.S., Corbin, C.J., Mathis, J.M., Powell, F.E., Mendelson, C.R., and Simpson, E.R., Structural analysis of the gene encoding human aromatase cytochrome P-450, the enzyme responsible for estrogen biosynthesis, J. Biol. Chem., 264, 19385-19391, 1989.
56. Chen, S.A., Besman, M.J., Sparkes, R.S., Zollman, S., Klisak, I., Mohandas, T., Hall, P.F., and Shively, J.E., Human aromatase: cDNA cloning, Southern blot analysis, and assignment of the gene to chromosome 15, DNA, 7, 27-38, 1988.
57. Mahendroo, M.S., Means, G.D., Mendelson, C.R., and Simpson, E.R., Tissue-specific expression of human P-450AROM: the promoter responsible for expression in adipose tissue is different from that utilized in placenta, J. Biol. Chem., 266, 11276-11281, 1991.
58. Labrie, F., Luu-The, V., Lin, S.X., Labrie, C., Simard, J., Breton, R., and Belanger, A., The key role of 17 beta-hydroxysteroid dehydrogenases in sex steroid biology, Steroids, 62, 148-158, 1997.
59. Luu-The, V., Labrie, C., Zhao, H.F., Couet, J., Lachance, Y., Simard, J., Leblanc, G., Cote, J., Berube, D., Gagne, R., et al., Characterization of cDNAs for human estradiol 17 beta-dehydrogenase and assignment of the gene to chromosome 17: evidence of two mRNA species with distinct 5'-termini in human placenta, Mol. Endocrinol., 3, 1301-1309, 1989.
60. Peltoketo, H., Isomaa, V., Maentausta, O., and Vihko, R., Complete amino acid sequence of human placental 17 beta-hydroxysteroid dehydrogenase deduced from cDNA, FEBS Lett., 239, 73-77, 1988.
61. Tremblay, Y., Ringler, G.E., Morel, Y., Mohandas, T.K., Labrie, F., Strauss, J.F., III, and Miller, W.L., Regulation of the gene for estrogenic 17-ketosteroid reductase lying on chromosome 17cen--q25, J. Biol. Chem., 264, 20458-20462, 1989.
62. Carstensen, J.F., Tesdorpf, J.G., Kaufmann, M., Markus, M.M., Husen, B., Leenders, F., Jakob, F., de Launoit, Y., and Adamski, J., Characterization of 17 beta-hydroxysteroid dehydrogenase IV, J. Endocrinol., 150 (Suppl.), S3-S12, 1996.
63. Adamski, J., Carstensen, J., Husen, B., Kaufmann, M., de Launoit, Y., Leenders, F., Markus, M., and Jungblut, P.W., New 17 beta-hydroxysteroid dehydrogenases: molecular and cell biology of the type IV porcine and human enzymes, Ann. N.Y. Acad. Sci., 784, 124-136, 1996.
64. Leenders, F., Adamski, J., Husen, B., Thole, H.H., and Jungblut, P.W., Molecular cloning and amino acid sequence of the porcine 17 beta-estradiol dehydrogenase, Eur. J. Biochem., 222, 221-227, 1994.
65. Ghosh, D., Pletnev, V.Z., Zhu, D.W., Wawrzak, Z., Duax, W.L., Pangborn, W., Labrie, F., and Lin, S.X., Structure of human estrogenic 17 beta-hydroxysteroid dehydrogenase at 2.20 A resolution, Structure, 3, 503-513, 1995.
66. Sawicki, M.W., Erman, M., Puranen, T., Vihko, P., and Ghosh, D., Structure of the ternary complex of human 17beta-hydroxysteroid dehydrogenase type 1 with 3- hydroxyestra-1,3,5,7-tetraen-17-one (equilin) and NADP+, Proc. Natl. Acad. Sci. U.S.A., 96, 840-845, 1999.
67. Geissler, W.M., Davis, D.L., Wu, L., Bradshaw, K.D., Patel, S., Mendonca, B.B., Elliston, K.O., Wilson, J.D., Russell, D.W., and Andersson, S., Male pseudohermaphroditism caused by mutations of testicular 17 beta-hydroxysteroid dehydrogenase 3, Nat. Genet., 7, 34-39, 1994 (see Comments).
68. Leenders, F., Tesdorpf, J.G., Markus, M., Engel, T., Seedorf, U., and Adamski, J., Porcine 80-kDa protein reveals intrinsic 17 beta-hydroxysteroid dehydrogenase, fatty acyl-CoA-hydratase/dehydrogenase, and sterol transfer activities, J. Biol. Chem., 271, 5438-5442, 1996.
69. Dufort, I., Rheault, P., Huang, X.F., Soucy, P., and Luu-The, V., Characteristics of a highly labile human type 5 17beta-hydroxysteroid dehydrogenase, Endocrinology, 140, 568-574, 1999.
70. Bujalska, I., Shimojo, M., Howie, A., and Stewart, P.M., Human 11 beta-hydroxysteroid dehydrogenase: studies on the stably transfected isoforms and localization of the type 2 isozyme within renal tissue, Steroids, 62, 77-82, 1997.
71. Tannin, G.M., Agarwal, A.K., Monder, C., New, M.I., and White, P.C., The human gene for 11 beta-hydroxysteroid dehydrogenase: structure, tissue distribution, and chromosomal localization, J. Biol. Chem., 266, 16653-16658, 1991.
72. Moore, C.C., Mellon, S.H., Murai, J., Siiteri, P.K., and Miller, W.L., Structure and function of the hepatic form of 11 beta-hydroxysteroid dehydrogenase in the squirrel monkey, an animal model of glucocorticoid resistance, Endocrinology, 133, 368-375, 1993.
73. Stewart, P.M., Murry, B.A., and Mason, J.I., Human kidney 11 beta-hydroxysteroid dehydrogenase is a high affinity nicotinamide adenine dinucleotide-dependent enzyme and differs from the cloned type I isoform, J. Clin. Endocrinol. Metab., 79, 480-484, 1994.
74. Tannin, G.M., Agarwal, A.K., Monder, C., New, M.I., and White, P.C., The human gene for 11 beta-hydroxysteroid dehydrogenase: structure, tissue distribution, and chromosomal localization, J. Biol. Chem., 266, 16653-16658, 1991.
75. Whorwood, C.B., Mason, J.I., Ricketts, M.L., Howie, A.J., and Stewart, P.M., Detection of human 11 beta-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-R12, 1995.
76. Brown, R.W., Chapman, K.E., Edwards, C.R., and Seckl, J.R., Human placental 11 beta-hydroxysteroid dehydrogenase: evidence for and partial purification of a distinct NAD-dependent isoform, Endocrinology, 132, 2614-2621, 1993.
77. Brown, R.W., Diaz, R., Robson, A.C., Kotelevtsev, Y.V., Mullins, J.J., Kaufman, M.H., and Seckl, J.R., The ontogeny of 11 beta-hydroxysteroid dehydrogenase type 2 and mineralocorticoid receptor gene expression reveal intricate control of glucocorticoid action in development, Endocrinology, 137, 794-797, 1996.
78. Albiston, A.L., Obeyesekere, V.R., Smith, R.E., and Krozowski, Z.S., Cloning and tissue distribution of the human 11 beta-hydroxysteroid dehydrogenase type 2 enzyme, Mol. Cell. Endocrinol., 105, R11-R17, 1994.
79. Moore, C.C.D., Mellon, S.H., Murai, J., Siiteri, P.K., and Miller, W.L., Structure and function of the hepatic form of 11 beta-hydroxysteroid dehydrogenase in the squirrel monkey, an animal model of glucocorticoid resistance, Endocrinology, 133, 368-375, 1993.
80. Rusvai, E. and Naray-Fejes-Toth, A., A new isoform of 11 beta-hydroxysteroid dehydrogenase in aldosterone target cells, J. Biol. Chem., 268, 10717-10720, 1993.
81. Agarwal, A.K., Rogerson, F.M., Mune, T., and White, P.C., Gene structure and chromosomal localization of the human HSD11K gene encoding the kidney (type 2) isozyme of 11 beta-hydroxysteroid dehydrogenase, Genomics, 29, 195-199, 1995.
82. Andersson, S. and Russell, D.W., Structural and biochemical properties of cloned and expressed human and rat steroid 5 alpha-reductases, Proc. Natl. Acad. Sci. U.S.A., 87, 3640-3644, 1990.
83. Andersson, S., Berman, D.M., Jenkins, E.P., and Russell, D.W., Deletion of steroid 5 alpha-reductase 2 gene in male pseudohermaphroditism, Nature, 354, 159-161, 1991.
84. Jenkins, E.P., Andersson, S., Imperato-McGinley, J., Wilson, J.D., and Russell, D.W., Genetic and pharmacological evidence for more than one human steroid 5 alphareductase, J. Clin. Invest., 89, 293-300, 1992.
85. Thigpen, A.E., Davis, D.L., Gautier, T., Imperato-McGinley, J., and Russell, D.W., Brief report: the molecular basis of steroid 5 alpha-reductase deficiency in a large Dominican kindred, N. Engl. J. Med., 327, 1216-1219, 1992.
86. Russell, D.W. and Wilson, J.D., Steroid 5 alpha-reductase: two genes/two enzymes, Annu. Rev. Biochem., 63, 25-61, 1994.
87. Lephart, E., A review of brain 5α-reductase: cellular, enzymatic, and molecular perspectives and implications for biological function, Mol. Cell. Neurosci., 4, 473-484, 1993.
88. Thigpen, A.E., Davis, D.L., Milatovich, A., Mendonca, B.B., Imperato-McGinley, J., Griffin, J.E., Francke, U., Wilson, J.D., and Russell, D.W., Molecular genetics of steroid 5 alpha-reductase 2 deficiency, J. Clin. Invest., 90, 799-809, 1992.
89. Thigpen, A.E., Silver, R.I., Guileyardo, J.M., Casey, M.L., McConnell, J.D., and Russell, D.W., Tissue distribution and ontogeny of steroid 5 alpha-reductase isozyme expression, J. Clin. Invest., 92, 903-910, 1993.
90. Poletti, A., Coscarella, A., Negri-Cesi, P., Colciago, A., Celotti, F., and Martini, L., 5 Alpha-reductase isozymes in the central nervous system, Steroids, 63, 246-251, 1998.
91. Poletti, A., Negri-Cesi, P., Rabuffetti, M., Colciago, A., Celotti, F., and Martini, L., Transient expression of the 5alpha-reductase type 2 isozyme in the rat brain in late fetal and early postnatal life, Endocrinology, 139, 2171-2178, 1998.
92. Penning, T.M., Bennett, M.J., Smith-Hoog, S., Schlegel, B.P., Jez, J.M., and Lewis, M., Structure and function of 3 alpha-hydroxysteroid dehydrogenase, Steroids, 62, 101-111, 1997.
93. Hyndman, D., Bauman, D.R., Heredia, V.V., and Penning, T.M., The aldo-keto reductase superfamily homepage, Chem. Biol. Interact., 143/144, 621-631, 2003.
94. Pawlowski, J., Huizinga, M., and Penning, T.M., Isolation and partial characterization of a full-length cDNA clone for 3 alpha-hydroxysteroid dehydrogenase: a potential target enzyme for nonsteroidal anti-inflammatory drugs, Agents Actions, 34, 289-293, 1991.
95. Cheng, K.C., White, P.C., and Qin, K.N., Molecular cloning and expression of rat liver 3 alpha-hydroxysteroid dehydrogenase, Mol. Endocrinol., 5, 823-828, 1991.
96. Stolz, A., Rahimi-Kiani, M., Ameis, D., Chan, E., Ronk, M., and Shively, J.E., Molecular structure of rat hepatic 3 alpha-hydroxysteroid dehydrogenase: a member of the oxidoreductase gene family, J. Biol. Chem., 266, 15253-15257, 1991.
97. Usui, E., Okuda, K., Kato, Y., and Noshiro, M., Rat hepatic 3 alpha-hydroxysteroid dehydrogenase: expression of cDNA and physiological function in bile acid biosynthetic pathway, J. Biochem. (Tokyo), 115, 230-237, 1994.
98. Penning, T.M., Burczynski, M.E., Jez, J.M., Hung, C.F., Lin, H.K., Ma, H., Moore, M., Palackal, N., and Ratnam, K., Human 3alpha-hydroxysteroid dehydrogenase isoforms (AKR1C1-AKR1C4) of the aldo-keto reductase superfamily: functional plasticity and tissue distribution reveals roles in the inactivation and formation of male and female sex hormones, Biochem. J., 351 (Pt. 1), 67-77, 2000.
99. Deyashiki, Y., Ogasawara, A., Nakayama, T., Nakanishi, M., Miyabe, Y., Sato, K., and Hara, A., Molecular cloning of two human liver 3 alpha-hydroxysteroid/dihydrodiol dehydrogenase isoenzymes that are identical with chlordecone reductase and bile-acid binder, Biochem. J., 299 (Pt. 2), 545-552, 1994.
100. Hara, A., Matsuura, K., Tamada, Y., Sato, K., Miyabe, Y., Deyashiki, Y., and Ishida, N., Relationship of human liver dihydrodiol dehydrogenases to hepatic bile-acidbinding protein and an oxidoreductase of human colon cells, Biochem. J., 313 (Pt. 2), 373-376, 1996.
101. Khanna, M., Qin, K.N., Wang, R.W., and Cheng, K.C., Substrate specificity, gene structure, and tissue-specific distribution of multiple human 3 alpha-hydroxysteroid dehydrogenases, J. Biol. Chem., 270, 20162-20168, 1995.
102. Dufort, I., Soucy, P., Labrie, F., and Luu-The, V., Molecular cloning of human type 3 3 alpha-hydroxysteroid dehydrogenase that differs from 20 alpha-hydroxysteroid dehydrogenase by seven amino acids, Biochem. Biophys. Res. Commun., 228, 474-479, 1996.
103. Griffin, L.D. and Mellon, S.H., Selective serotonin reuptake inhibitors directly alter activity of neurosteroidogenic enzymes, Proc. Natl. Acad. Sci. U.S.A., 96, 13512-13517, 1999.
104. Usami, N., Yamamoto, T., Shintani, S., Ishikura, S., Higaki, Y., Katagiri, Y., and Hara, A., Substrate specificity of human 3(20)alpha-hydroxysteroid dehydrogenase for neurosteroids and its inhibition by benzodiazepines, Biol. Pharm. Bull., 25, 441-445, 2002.
105. Higaki, Y., Usami, N., Shintani, S., Ishikura, S., El-Kabbani, O., and Hara, A., Selective and potent inhibitors of human 20alpha-hydroxysteroid dehydrogenase (AKR1C1) that metabolizes neurosteroids derived from progesterone, Chem. Biol. Interact., 143/144, 503-513, 2003.
106. Jin, Y., Cooper, W.C., and Penning, T.M., Examination of the differences in structurefunction of human and rat 3alpha-hydroxysteroid dehydrogenase, Chem. Biol. Interact., 143/144, 383-392, 2003.
107. Griffin, L.D. and Mellon, S.H., Selective serotonin reuptake inhibitors directly alter activity of neurosteroidogenic enzymes, Proc. Natl. Acad. Sci. U.S.A., 96, 13512-13517, 1999.
108. Hobkirk, R., Steroid sulfotransferases and steroid sulfate sulfatases: characteristics and biological roles, Can. J. Biochem. Cell Biol., 63, 1127-1144, 1985.
109. Iwamori, M., Moser, H.W., and Kishimoto, Y., Steroid sulfatase in brain: comparison of sulfohydrolase activities for various steroid sulfates in normal and pathological brains, including the various forms of metachromatic leukodystrophy, J. Neurochem., 27, 1389-1395, 1976.
110. Parker, C.R., Jr., Falany, C.N., Stockard, C.R., Stankovic, A.K., and Grizzle, W.E., Immunohistochemical localization of dehydroepiandrosterone sulfotransferase in human fetal tissues, J. Clin. Endocrinol. Metab., 78, 234-236, 1994.
111. Barker, E.V., Hume, R., Hallas, A., and Coughtrie, W.H., Dehydroepiandrosterone Sulfotransferase in the Developing Human Fetus: Quantitative Biochemical and Immunological Characterization of the Hepatic, Renal, and Adrenal Enzymes, 1994.
112. Mulder, G., The Sulfatation of Drugs and Other Compounds, CRC Press, Boca Raton, FL, 1981.
113. De Meio, R., Sulfate activation and transfer, in Metabolism of Sulfur Compounds, Greenberg, D.M., Ed., Academic Press, New York, 1975, pp. 287-359.
114. Majewska, M., Neurosteroids: GABAA-agonistic and GABAA-antagonistic modulators of the GABAA receptor, in Neurosteroids and Brain Function, Costa, E. and Paul, S.M., Eds., Thieme, New York, 1991, pp. 109-117.
115. Majewska, M.D., Neurosteroids: endogenous bimodal modulators of the GABAA receptor: mechanism of action and physiological significance, Prog. Neurobiol., 38, 379-395, 1992.
116. Ogura, K., Kajita, J., Narihata, H., Watabe, T., Ozawa, S., Nagata, K., Yamazoe, Y., and Kato, R., Cloning and sequence analysis of a rat liver cDNA encoding hydroxysteroid sulfotransferase, Biochem. Biophys. Res. Commun., 165, 168-174, 1989.
117. Shapiro, L., Steroid Sulfatase, paper presented at International Symposium on DHEA Transformation into Androgens and Estrogens in Target Tissues: Intracrinology, Québec City, 1995.
118. Ballabio, A. and Shapiro, L.J., Steroid sulfatase deficiency and X-linked ichtyosis, in The Metabolic and Molecular Bases of Inherited Disease, Scriver, C.R., Beaudet, A.L., Sly, W.S., and Valle, D., Eds., McGraw-Hill, New York, 1995, pp. 2999-3022.
119. Mohandas, T., Geller, R.L., Yen, P.H., Rosendorff, J., Bernstein, R., Yoshida, A., and Shapiro, L.J., Cytogenetic and molecular studies on a recombinant human X chromosome: implications for the spreading of X chromosome inactivation, Proc. Natl. Acad. Sci. U.S.A., 84, 4954-4958, 1987.
120. Shapiro, L., Steroid sulfatase deficiency and x-linked ichthyosis, in The Metabolic Basis of Inherited Diseases, Stanbury, J.W., Fredrickson, D.S., Goldstein, J.L., Brown, M.S., Eds., McGraw Hill, New York, 1982, pp. 1027-1034.
121. Li, X.M., Salido, E.C., Gong, Y., Kitada, K., Serikawa, T., Yen, P.H., and Shapiro, L.J., Cloning of the rat steroid sulfatase gene (Sts), a non-pseudoautosomal X-linked gene that undergoes X inactivation, Mamm. Genome, 7, 420-424, 1996.
122. Salido, E.C., Li, X.M., Yen, P.H., Martin, N., Mohandas, T.K., and Shapiro, L.J., Cloning and expression of the mouse pseudoautosomal steroid sulphatase gene (Sts), Nat. Genet., 13, 83-86, 1996.
123. Rose, K.A., Stapleton, G., Dott, K., Kieny, M.P., Best, R., Schwarz, M., Russell, D.W., Bjorkhem, I., Seckl, J., and Lathe, R., Cyp7b, a novel brain cytochrome P450, catalyzes the synthesis of neurosteroids 7alpha-hydroxy dehydroepiandrosterone and 7alpha-hydroxy pregnenolone, Proc. Natl. Acad. Sci. U.S.A., 94, 4925-4930, 1997.
124. Stapleton, G., Steel, M., Richardson, M., Mason, J.O., Rose, K.A., Morris, R.G., and Lathe, R., A novel cytochrome P450 expressed primarily in brain, J. Biol. Chem., 270, 29739-29745, 1995.
125. Lund, E.G., Guileyardo, J.M., and Russell, D.W., cDNA cloning of cholesterol 24- hydroxylase, a mediator of cholesterol homeostasis in the brain, Proc. Natl. Acad. Sci. U.S.A., 96, 7238-7243, 1999.
126. Ray, W.J., Bain, G., Yao, M., and Gottlieb, D.I., CYP26, a novel mammalian cytochrome P450, is induced by retinoic acid and defines a new family, J. Biol. Chem., 272, 18702-18708, 1997.
127. White, J.A., Guo, Y.D., Baetz, K., Beckett-Jones, B., Bonasoro, J., Hsu, K.E., Dilworth, F.J., Jones, G., and Petkovich, M., Identification of the retinoic acid-inducible all-trans-retinoic acid 4-hydroxylase, J. Biol. Chem., 271, 29922-29927, 1996.
128. White, J.A., Beckett-Jones, B., Guo, Y.D., Dilworth, F.J., Bonasoro, J., Jones, G., and Petkovich, M., cDNA cloning of human retinoic acid-metabolizing enzyme (hP450RAI) identifies a novel family of cytochromes P450, J. Biol. Chem., 272, 18538-18541, 1997.
129. Morfin, R. and Courchay, G., Pregnenolone and dehydroepiandrosterone as precursors of native 7-hydroxylated metabolites which increase the immune response in mice, J. Steroid Biochem. Mol. Biol., 50, 91-100, 1994.
130. Doostzadeh, J., Cotillon, A.C., and Morfin, R., Hydroxylation of pregnenolone at the 7 alpha- and 7 beta-positions by mouse liver microsomes: effects of cytochrome p450 inhibitors and structure-specific inhibition by steroid hormones, Steroids, 63, 383-392, 1998.
131. Tzung, K.W., Ishimura-Oka, K., Kihara, S., Oka, K., and Chan, L., Structure of the mouse cholesterol 7 alpha-hydroxylase gene, Genomics, 21, 244-247, 1994.
132. Wood, A.W., Ryan, D.E., Thomas, P.E., and Levin, W., Regio- and stereoselective metabolism of two C19 steroids by five highly purified and reconstituted rat hepatic cytochrome P-450 isozymes, J. Biol. Chem., 258, 8839-8847, 1983.
133. Swinney, D.C., Ryan, D.E., Thomas, P.E., and Levin, W., Regioselective progesterone hydroxylation catalyzed by eleven rat hepatic cytochrome P-450 isozymes, Biochemistry, 26, 7073-7083, 1987.
134. Arlotto, M.P., Greenway, D.J., and Parkinson, A., Purification of two isozymes of rat liver microsomal cytochrome P450 with testosterone 7 alpha-hydroxylase activity, Arch. Biochem. Biophys., 270, 441-457, 1989.
135. Akwa, Y., Morfin, R.F., Robel, P., and Baulieu, E.E., Neurosteroid metabolism: 7 alpha-hydroxylation of dehydroepiandrosterone and pregnenolone by rat brain microsomes, Biochem. J., 288 (Pt. 3), 959-964, 1992.
136. Stoffel-Wagner, B., Neurosteroid metabolism in the human brain, Eur. J. Endocrinol., 145, 669-679, 2001.
137. Compagnone, N.A., Bulfone, A., Rubenstein, J.L., and Mellon, S.H., Steroidogenic enzyme P450c17 is expressed in the embryonic central nervous system, Endocrinology, 136, 5212-5223, 1995.
138. Compagnone, N.A., Bulfone, A., Rubenstein, J.L., and Mellon, S.H., Expression of the steroidogenic enzyme P450scc in the central and peripheral nervous systems during rodent embryogenesis, Endocrinology, 136, 2689-2696, 1995.
139. Compagnone, N.A., Salido, E., Shapiro, L.J., and Mellon, S.H., Expression of steroid sulfatase during embryogenesis, Endocrinology, 138, 4768-4773, 1997.
140. Lauber, M.E. and Lichtensteiger, W., Pre- and postnatal ontogeny of aromatase cytochrome P450 messenger ribonucleic acid expression in the male rat brain studied by in situ hybridization, Endocrinology, 135, 1661-1668, 1994.
141. Lauber, M.E. and Lichtensteiger, W., Ontogeny of 5 alpha-reductase (type 1) messenger ribonucleic acid expression in rat brain: early presence in germinal zones, Endocrinology, 137, 2718-2730, 1996.
142. Seckl, J.R. and Walker, B.R., Minireview: 11beta-hydroxysteroid dehydrogenase type 1: a tissue-specific amplifier of glucocorticoid action, Endocrinology, 142, 1371-1376, 2001.
143. Holmes, M.C., Kotelevtsev, Y., Mullins, J.J., and Seckl, J.R., Phenotypic analysis of mice bearing targeted deletions of 11beta-hydroxysteroid dehydrogenases 1 and 2 genes, Mol. Cell. Endocrinol., 171, 15-20, 2001.
144. Seckl, J.R., 11beta-Hydroxysteroid dehydrogenase in the brain: a novel regulator of glucocorticoid action? Front Neuroendocrinol., 18, 49-99, 1997.
145. Baulieu, E.E. and Robel, P., Neurosteroids: a new brain function? J. Steroid Biochem. Mol. Biol., 37, 395-403, 1990.
146. Ukena, K., Usui, M., Kohchi, C., and Tsutsui, K., Cytochrome P450 side-chain cleavage enzyme in the cerebellar Purkinje neuron and its neonatal change in rats, Endocrinology, 139, 137-147, 1998.
147. Tsutsui, K., Ukena, K., Usui, M., Sakamoto, H., and Takase, M., Novel brain function: biosynthesis and actions of neurosteroids in neurons, Neurosci. Res., 36, 261-273, 2000.
148. Sakamoto, H., Ukena, K., and Tsutsui, K., Effects of progesterone synthesized de novo in the developing Purkinje cell on its dendritic growth and synaptogenesis, J. Neurosci., 21, 6221-6232, 2001.
149. Mellon, S.H., Kushner, J.A., and Vaisse, C., Expression and regulation of adrenodoxin and P450scc mRNA in rodent tissues, DNA Cell Biol., 10, 339-347, 1991.
150. Oftebro, H., Stormer, F.C., and Pedersen, J.L., The presence of an adrenodoxin-like ferredoxin and cytochrome P-450 in brain mitochondria, J. Biol. Chem., 254, 4331-4334, 1979.
151. Lin, D., Sugawara, T., Strauss, J.F., III, Clark, B.J., Stocco, D.M., Saenger, P., Rogol, A., and Miller, W.L., Role of steroidogenic acute regulatory protein in adrenal and gonadal steroidogenesis, Science, 267, 1828-1831, 1995.
152. Caron, K.M., Soo, S.C., Wetsel, W.C., Stocco, D.M., Clark, B.J., and Parker, K.L., Targeted disruption of the mouse gene encoding steroidogenic acute regulatory protein provides insights into congenital lipoid adrenal hyperplasia, Proc. Natl. Acad. Sci. U.S.A., 94, 11540-11545, 1997.
153. Benmessahel, Y., Guennoun, R., Cadepond, F., Baulieu, E.E., Schumacher, M., and Groyer, G., Expression of steroidogenic acute regulatory protein in cultured Schwann cells and its regulation by cAMP, Ann. N.Y. Acad. Sci., 973, 83-87, 2002.
154. Mellon, S.H. and Deschepper, C.F., Neurosteroid biosynthesis: genes for adrenal steroidogenic enzymes are expressed in the brain, Brain Res., 629, 283-292, 1993.
155. Compagnone, N.A., Bulfone, A., Rubenstein, J.L.R., and Mellon, S.H., Steroidogenic enzyme P450c17 is expressed in the embryonic central nervous system, Endocrinology, 136, 5212-5223, 1995.
156. Rubenstein, J.L., Anderson, S., Shi, L., Miyashita-Lin, E., Bulfone, A., and Hevner, R., Genetic control of cortical regionalization and connectivity, Cereb. Cortex, 9, 524-532, 1999.
157. Miyashita-Lin, E.M., Hevner, R., Wassarman, K.M., Martinez, S., and Rubenstein, J.L., Early neocortical regionalization in the absence of thalamic innervation, Science, 285, 906-909, 1999.
158. Kohchi, C., Ukena, K., and Tsutsui, K., Age- and region-specific expressions of the messenger RNAs encoding for steroidogenic enzymes p450scc, P450c17 and 3beta-HSD in the postnatal rat brain, Brain Res., 801, 233-238, 1998.
159. Norris, P.J., Hardwick, J.P., and Emson, P.C., Localization of NADPH cytochrome P450 oxidoreductase in rat brain by immunohistochemistry and in situ hybridization and a comparison with the distribution of neuronal NADPH-diaphorase staining, Neuroscience, 61, 331-350, 1994.
160. Simmons, D.L. and Kasper, C.B., Quantitation of mRNAs specific for the mixedfunction oxidase system in rat liver and extrahepatic tissues during development, Arch. Biochem. Biophys., 271, 10-20, 1989.
161. Jung-Testas, I., Hu, Z.Y., Baulieu, E.E., and Robel, P., Neurosteroids: biosynthesis of pregnenolone and progesterone in primary cultures of rat glial cells, Endocrinology, 125, 2083-2091, 1989.
162. Hu, Z.Y., Bourreau, E., Jung-Testas, I., Robel, P., and Baulieu, E.E., Neurosteroids: oligodendrocyte mitochondria convert cholesterol to pregnenolone, Proc. Natl. Acad. Sci. U.S.A., 84, 8215-8219, 1987.
163. Kabbadj, K., el-Etr, M., Baulieu, E.E., and Robel, P., Pregnenolone metabolism in rodent embryonic neurons and astrocytes, Glia, 7, 170-175, 1993.
164. Bauer, H.C. and Bauer, H., Micromethod for the determination of 3-beta-HSD activity in cultured cells, J. Steroid Biochem., 33, 643-646, 1989.
165. Weidenfeld, J., Siegel, R.A., and Chowers, I., In vitro conversion of pregnenolone to progesterone by discrete brain areas of the male rat, J. Steroid Biochem., 13, 961-963, 1980.
166. Robel, P., Bourreau, E., Corpechot, C., Dang, D.C., Halberg, F., Clarke, C., Haug, M., Schlegel, M.L., Synguelakis, M., Vourch, C., et al., Neuro-steroids: 3 betahydroxy- delta 5-derivatives in rat and monkey brain, J. Steroid Biochem., 27, 649-655, 1987.
167. Guennoun, R., Fiddes, R.J., Gouezou, M., Lombes, M., and Baulieu, E.E., A key enzyme in the biosynthesis of neurosteroids, 3 beta-hydroxysteroid dehydrogenase/delta 5-delta 4-isomerase (3 beta-HSD), is expressed in rat brain, Brain Res. Mol. Brain Res., 30, 287-300, 1995.
168. Dupont, E., Labrie, F., Luu-The, V., and Pelletier, G., Immunocytochemical localization of 3 beta-hydroxysteroid dehydrogenase/delta 5-delta 4-isomerase in human ovary, J. Clin. Endocrinol. Metab., 74, 994-998, 1992.
169. Ukena, K., Kohchi, C., and Tsutsui, K., Expression and activity of 3beta-hydroxysteroid dehydrogenase/delta5-delta4-isomerase in the rat Purkinje neuron during neonatal life, Endocrinology, 140, 805-813, 1999.
170. Coirini, H., Gouezou, M., Liere, P., Delespierre, B., Pianos, A., Eychenne, B., Schumacher, M., and Guennoun, R., 3 Beta-hydroxysteroid dehydrogenase expression in rat spinal cord, Neuroscience, 113, 883-891, 2002.
171. Mensah-Nyagan, A.G., Feuilloley, M., Dupont, E., Do-Rego, J.L., Leboulenger, F., Pelletier, G., and Vaudry, H., Immunocytochemical localization and biological activity of 3 beta-hydroxysteroid dehydrogenase in the central nervous system of the frog, J. Neurosci., 14, 7306-7318, 1994.
172. Mathieu, M., Mensah-Nyagan, A.G., Vallarino, M., Do-Rego, J.L., Beaujean, D., Luu-The, V., Pelletier, G., and Vaudry, H., Immunohistochemical localization of 3β - hydroxyseroid dehydrogenase and 5α reductase in the brain of the African lungfish Protopterus amnectens, J. Comp. Neurol., 2000, submitted.
173. Sanne, J.L. and Krueger, K.E., Expression of cytochrome P450 side-chain cleavage enzyme and 3 beta-hydroxysteroid dehydrogenase in the rat central nervous system: a study by polymerase chain reaction and in situ hybridization, J. Neurochem., 65, 528-536, 1995.
174. Kabbadj, K., el-Etr, M., Baulieu, E.E., and Robel, P., Pregnenolone metabolism in rodent embryonic neurons and astrocytes, Glia, 7, 170-175, 1993.
175. Mensah-Nyagan, A.G., Feuilloley, M., Dupont, E., Do-Rego, J.L., Leboulenger, F., Pelletier, G., and Vaudry, H., Immunocytochemical localization and biological activity of 3 beta-hydroxysteroid dehydrogenase in the central nervous system of the frog, J. Neurosci., 14, 7306-7318, 1994.
176. Mensah-Nyagan, A.G., Beaujean, D., Luu-The, V., Pelletier, G., and Vaudry, H., Anatomical and biochemical evidence for the synthesis of unconjugated and sulfated neurosteroids in amphibians, Brain Res. Brain Res. Rev., 37, 13-24, 2001.
177. Robert, F., Guennoun, R., Desarnaud, F., Do-Thi, A., Benmessahel, Y., Baulieu, E.E., and Schumacher, M., Synthesis of progesterone in Schwann cells: regulation by sensory neurons, Eur. J. Neurosci., 13, 916-924, 2001.
178. Gago, N., Akwa, Y., Sananes, N., Guennoun, R., Baulieu, E.E., El-Etr, M., and Schumacher, M., Progesterone and the oligodendroglial lineage: stage-dependent biosynthesis and metabolism, Glia, 36, 295-308, 2001.
179. MacKenzie, S.M., Clark, C.J., Fraser, R., Gomez-Sanchez, C.E., Connell, J.M., and Davies, E., Expression of 11beta-hydroxylase and aldosterone synthase genes in the rat brain, J. Mol. Endocrinol., 24, 321-328, 2000.
180. Gomez-Sanchez, C.E., Zhou, M.Y., Cozza, E.N., Morita, H., Foecking, M.F., and Gomez-Sanchez, E.P., Aldosterone biosynthesis in the rat brain, Endocrinology, 138, 3369-3373, 1997.
181. Reddy, V.V., Estrogen metabolism in neural tissues of rabbits: 17 beta-hydroxysteroid oxidoreductase activity, Steroids, 34, 207-215, 1979.
182. Resko, J.A., Stadelman, H.L., and Norman, R.L., 17 beta-hydroxysteroid dehydrogenase activity in the pituitary gland and neural tissue of Rhesus monkeys, J. Steroid Biochem., 11, 1429-1434, 1979.
183. Pelletier, G., Luu-The, V., and Labrie, F., Immunocytochemical localization of type I 17 beta-hydroxysteroid dehydrogenase in the rat brain, Brain Res., 704, 233-239, 1995.
184. Mensah-Nyagan, A.M., Feuilloley, M., Do-Rego, J.L., Marcual, A., Lange, C., Tonon, M.C., Pelletier, G., and Vaudry, H., Localization of 17beta-hydroxysteroid dehydrogenase and characterization of testosterone in the brain of the male frog, Proc. Natl. Acad. Sci. U.S.A., 93, 1423-1428, 1996.
185. Mensah-Nyagan, A.G., Do-Rego, J.L., Feuilloley, M., Marcual, A., Lange, C., Pelletier, G., and Vaudry, H., In vivo and in vitro evidence for the biosynthesis of testosterone in the telencephalon of the female frog, J. Neurochem., 67, 413-422, 1996.
186. Steckelbroeck, S., Stoffel-Wagner, B., Reichelt, R., Schramm, J., Bidlingmaier, F., Siekmann, L., and Klingmuller, D., Characterization of 17beta-hydroxysteroid dehydrogenase activity in brain tissue: testosterone formation in the human temporal lobe, J. Neuroendocrinol., 11, 457-464, 1999.
187. Beyenburg, S., Watzka, M., Blumcke, I., Schramm, J., Bidlingmaier, F., Elger, C.E., and Stoffel-Wagner, B., Expression of mRNAs encoding for 17beta-hydroxisteroid dehydrogenase isozymes 1, 2, 3 and 4 in epileptic human hippocampus, Epilepsy Res., 41, 83-91, 2000.
188. Lephart, E.D., A review of brain aromatase cytochrome P450, Brain Res. Brain Res. Rev., 22, 1-26, 1996.
189. Hutchison, J.B., Wozniak, A., Beyer, C., and Hutchison, R.E., Regulation of sexspecific formation of oestrogen in brain development: endogenous inhibitors of aromatase, J. Steroid Biochem. Mol. Biol., 56, 201-207, 1996.
190. Roselli, C.E., Horton, L.E., and Resko, J.A., Distribution and regulation of aromatase activity in the rat hypothalamus and limbic system, Endocrinology, 117, 2471-2477, 1985.
191. George, F.W. and Ojeda, S.R., Changes in aromatase activity in the rat brain during embryonic, neonatal, and infantile development, Endocrinology, 111, 522-529, 1982.
192. MacLusky, N.J., Philip, A., Hurlburt, C., and Naftolin, F., Estrogen formation in the developing rat brain: sex differences in aromatase activity during early post-natal life, Psychoneuroendocrinology, 10, 355-361, 1985.
193. Lephart, E.D., Simpson, E.R., McPhaul, M.J., Kilgore, M.W., Wilson, J.D., and Ojeda, S.R., Brain aromatase cytochrome P-450 messenger RNA levels and enzyme activity during prenatal and perinatal development in the rat, Brain Res. Mol. Brain Res., 16, 187-192, 1992.
194. Honda, S., Harada, N., and Takagi, Y., Novel exon 1 of the aromatase gene specific for aromatase transcripts in human brain, Biochem. Biophys. Res. Commun., 198, 1153-1160, 1994.
195. Toda, K., Simpson, E.R., Mendelson, C.R., Shizuta, Y., and Kilgore, M.W., Expression of the gene encoding aromatase cytochrome P450 (CYP19) in fetal tissues, Mol. Endocrinol., 8, 210-217, 1994.
196. Kato, J., Yamada-Mouri, N., and Hirata, S., Structure of aromatase mRNA in the rat brain, J. Steroid Biochem. Mol. Biol., 61, 381-385, 1997.
197. Lephart, E.D., Molecular aspects of brain aromatase cytochrome P450, J. Steroid Biochem. Mol. Biol., 61, 375-380, 1997.
198. Wagner, C.K. and Morrell, J.I., Distribution and steroid hormone regulation of aromatase mRNA expression in the forebrain of adult male and female rats: a cellularlevel analysis using in situ hybridization, J. Comp. Neurol., 370, 71-84, 1996.
199. Sanghera, M.K., Simpson, E.R., McPhaul, M.J., Kozlowski, G., Conley, A.J., and Lephart, E.D., Immunocytochemical distribution of aromatase cytochrome P450 in the rat brain using peptide-generated polyclonal antibodies, Endocrinology, 129, 2834-2844, 1991.
200. Moisan, M.P., Seckl, J.R., and Edwards, C.R., 11 Beta-hydroxysteroid dehydrogenase bioactivity and messenger RNA expression in rat forebrain: localization in hypothalamus, hippocampus, and cortex, Endocrinology, 127, 1450-1455, 1990.
201. Lakshmi, V., Sakai, R.R., McEwen, B.S., and Monder, C., Regional distribution of 11 beta-hydroxysteroid dehydrogenase in rat brain, Endocrinology, 128, 1741-1748, 1991.
202. Roland, B.L., Li, K.X., and Funder, J.W., Hybridization histochemical localization of 11 beta-hydroxysteroid dehydrogenase type 2 in rat brain, Endocrinology, 136, 4697-4700, 1995.
203. Zhou, M.Y., Gomez-Sanchez, E.P., Cox, D.L., Cosby, D., and Gomez-Sanchez, C.E., Cloning, expression, and tissue distribution of the rat nicotinamide adenine dinucleotide- dependent 11 beta-hydroxysteroid dehydrogenase, Endocrinology, 136, 3729-3734, 1995.
204. Diaz, R., Brown, R.W., and Seckl, J.R., Distinct ontogeny of glucocorticoid and mineralocorticoid receptor and 11beta-hydroxysteroid dehydrogenase types I and II mRNAs in the fetal rat brain suggest a complex control of glucocorticoid actions, J. Neurosci., 18, 2570-2580, 1998.
205. Rajan, V., Edwards, C.R., and Seckl, J.R., 11 beta-Hydroxysteroid dehydrogenase in cultured hippocampal cells reactivates inert 11-dehydrocorticosterone, potentiating neurotoxicity, J. Neurosci., 16, 65-70, 1996.
206. Yau, J.L., Noble, J., Kenyon, C.J., Hibberd, C., Kotelevtsev, Y., Mullins, J.J., and Seckl, J.R., Lack of tissue glucocorticoid reactivation in 11beta-hydroxysteroid dehydrogenase type 1 knockout mice ameliorates age-related learning impairments, Proc. Natl. Acad. Sci. U.S.A., 98, 4716-4721, 2001.
207. Celotti, F., Melcangi, R.C., and Martini, L., The 5 alpha-reductase in the brain: molecular aspects and relation to brain function, Front Neuroendocrinol., 13, 163-215, 1992.
208. Melcangi, R.C., Celotti, F., Castano, P., and Martini, L., Differential localization of the 5 alpha-reductase and the 3 alpha-hydroxysteroid dehydrogenase in neuronal and glial cultures, Endocrinology, 132, 1252-1259, 1993.
209. Melcangi, R.C., Celotti, F., and Martini, L., Progesterone 5-alpha-reduction in neuronal and in different types of glial cell cultures: type 1 and 2 astrocytes and oligodendrocytes, Brain Res., 639, 202-206, 1994.
210. Pelletier, G., Luu-The, V., and Labrie, F., Immunocytochemical localization of 5 alpha-reductase in rat brain, Mol. Cell. Neurosci., 5, 394-399, 1994.
211. Normington, K. and Russell, D.W., Tissue distribution and kinetic characteristics of rat steroid 5 alpha-reductase isozymes: evidence for distinct physiological functions, J. Biol. Chem., 267, 19548-19554, 1992.
212. Stoffel-Wagner, B., Watzka, M., Steckelbroeck, S., Wickert, L., Schramm, J., Romalo, G., Klingmuller, D., and Schweikert, H.U., Expression of 5alpha-reductase in the human temporal lobe of children and adults, J. Clin. Endocrinol. Metab., 83, 3636-3642, 1998.
213. Stoffel-Wagner, B., Beyenburg, S., Watzka, M., Blumcke, I., Bauer, J., Schramm, J., Bidlingmaier, F., and Elger, C.E., Expression of 5alpha-reductase and 3alpha-hydroxisteroid oxidoreductase in the hippocampus of patients with chronic temporal lobe epilepsy, Epilepsia, 41, 140-147, 2000.
214. Pasmanik, M. and Callard, G.V., Aromatase and 5 alpha-reductase in the teleost brain, spinal cord, and pituitary gland, Gen. Comp. Endocrinol., 60, 244-251, 1985.
215. Pasmanik, M. and Callard, G.V., Changes in brain aromatase and 5 alpha-reductase activities correlate significantly with seasonal reproductive cycles in goldfish (Carassius auratus), Endocrinology, 122, 1349-1356, 1988.
216. Mahendroo, M.S., Cala, K.M., Landrum, D.P., and Russell, D.W., Fetal death in mice lacking 5alpha-reductase type 1 caused by estrogen excess, Mol. Endocrinol., 11, 917-927, 1997.
217. Compagnone, N.A. and Mellon, S.H., Neurosteroids: biosynthesis and function of these novel neuromodulators, Front Neuroendocrin., 21, 1-58, 2000.
218. Steckelbroeck, S., Watzka, M., Reichelt, R., Hans, V.H., Stoffel-Wagner, B., Heidrich, D.D., Schramm, J., Bidlingmaier, F., and Klingmuller, D., Characterization of the 5alpha-reductase-3alpha-hydroxysteroid dehydrogenase complex in the human brain, J. Clin. Endocrinol. Metab., 86, 1324-1331, 2001.
219. Karavolas, H.J. and Hodges, D.R., Neuroendocrine metabolism of progesterone and related progestins, Ciba Found. Symp., 153, 22-44, 1990.
220. Compagnone, N.A. and Mellon, S.H., Dehydroepiandrosterone: a potential signalling molecule for neocortical organization during development, Proc. Natl. Acad. Sci. U.S.A., 95, 4678-4683, 1998 (see Comments).
221. Beaujean, D., Mensah-Nyagan, A.G., Do-Rego, J.L., Luu-The, V., Pelletier, G., and Vaudry, H., Immunocytochemical localization and biological activity of hydroxysteroid sulfotransferase in the frog brain, J. Neurochem., 72, 848-857, 1999.
222. Sharp, S., Barker, E.V., Coughtrie, M.W., Lowenstein, P.R., and Hume, R., Immunochemical characterisation of a dehydroepiandrosterone sulfotransferase in rats and humans, Eur. J. Biochem., 211, 539-548, 1993.
223. Knapstein, P., David, A., Wu, C.H., Archer, D.F., FLickinger, G.L., and Touchstone, J.C., Metabolism of free and sulfoconjugated DHEA in brain tissue in vivo and in vitro, Steroids, 11, 885-896, 1968.
224. Rajkowski, K.M., Robel, P., and Baulieu, E.E., Hydroxysteroid sulfotransferase activity in the rat brain and liver as a function of age and sex, Steroids, 62, 427-436, 1997.
225. Price, D.J. and Lotto, R.B., Influences of the thalamus on the survival of subplate and cortical plate cells in cultured embryonic mouse brain, J. Neurosci., 16, 3247-3255, 1996.
226. Bixo, M., Andersson, A., Winblad, B., Purdy, R.H., and Backstrom, T., Progesterone, 5alpha-Pregnane-3,20-dione and 3alpha-hydroxy-5alpha-pregnane-20-one in specific regions of the human female brain in different endocrine states, Brain Res., 764, 173-178, 1997.
227. Korneyev, A., Guidotti, A., and Costa, E., Regional and interspecies differences in brain progesterone metabolism, J. Neurochem., 61, 2041-2047, 1993.
228. Milewich, L., Gomez-Sanchez, C., Crowley, G., Porter, J.C., Madden, J.D., and MacDonald, P.C., Progesterone and 5alpha-pregnane-3,20-dione in peripheral blood of normal young women: daily measurements throughout the menstrual cycle, J. Clin. Endocrinol. Metab., 45, 617-622, 1977.
229. Britton, K.T. and Koob, G.F., Neuropharmacology: premenstrual steroids? Nature, 392, 869-870, 1998.
230. Fry, J. and Honour, J., Understanding premenstrual syndrome, Lancet, 351, 1511, 1998 (author reply, Lancet, 351, 1512, 1998).
231. Lubbert, H., Understanding premenstrual syndrome, Lancet, 351, 1511, 1998 (author reply, Lancet, 351, 1512, 1998).
232. Rapkin, A.J., Morgan, M., Goldman, L., Brann, D.W., Simone, D., and Mahesh, V.B., Progesterone metabolite allopregnanolone in women with premenstrual syndrome, Obstet. Gynecol., 90, 709-714, 1997.
233. Schmidt, P.J., Purdy, R.H., Moore, P.H., Jr., Paul, S.M., and Rubinow, D.R., Circulating levels of anxiolytic steroids in the luteal phase in women with premenstrual syndrome and in control subjects, J. Clin. Endocrinol. Metab., 79, 1256-1260, 1994.
234. Monteleone, P., Luisi, S., Tonetti, A., Bernardi, F., Genazzani, A.D., Luisi, M., Petraglia, F., and Genazzani, A.R., Allopregnanolone concentrations and premenstrual syndrome, Eur. J. Endocrinol., 142, 269-273, 2000.
235. Wang, M., Seippel, L., Purdy, R.H., and Backstrom, T., Relationship between symptom severity and steroid variation in women with premenstrual syndrome: study on serum pregnenolone, pregnenolone sulfate, 5 alpha-pregnane-3,20-dione and 3 alphahydroxy-5 alpha-pregnan-20-one, J. Clin. Endocrinol. Metab., 81, 1076-1082, 1996.
236. Griffin, L.D., Conrad, S.C., and Mellon, S.H., Current perspectives on the role of neurosteroids in PMS and depression, Int. Rev. Neurobiol., 46, 479-492, 2001.
237. Dimmock, P.W., Wyatt, K.M., Jones, P.W., and O’Brien, P.M., Efficacy of selective serotonin-reuptake inhibitors in premenstrual syndrome: a systematic review, Lancet, 356, 1131-1136, 2000.
238. Uzunov, D.P., Cooper, T.B., Costa, E., and Guidotti, A., Fluoxetine-elicited changes in brain neurosteroid content measured by negative ion mass fragmentography, Proc. Natl. Acad. Sci. U.S.A., 93, 12599-12604, 1996.
239. Trauger, J.W., Jiang, A., Stearns, B.A., and LoGrasso, P.V., Kinetics of allopregnanolone formation catalyzed by human 3 alpha-hydroxysteroid dehydrogenase type III (AKR1C2), Biochemistry, 41, 13451-13459, 2002.
240. Hara, A., Matsuura, K., Tamada, Y., Sato, K., Miyabe, Y., Deyashiki, Y., and Ishida, N., Relationship of human liver dihydrodiol dehydrogenases to hepatic bile-acidbinding protein and an oxidoreductase of human colon cells, Biochem. J., 313 (Pt. 2), 373-376, 1996.
241. Mensah-Nyagan, A.G., Do-Rego, J.L., Beaujean, D., Luu-The, V., Pelletier, G., and Vaudry, H., Neurosteroids: expression of steroidogenic enzymes and regulation of steroid biosynthesis in the central nervous system, Pharmacol. Rev., 51, 63-81, 1999.
242. Do-Rego, J.L., Mensah-Nyagan, G.A., Beaujean, D., Vaudry, D., Sieghart, W., Luu-The, V., Pelletier, G., and Vaudry, H., gamma-Aminobutyric acid, acting through gamma-aminobutyric acid type A receptors, inhibits the biosynthesis of neurosteroids in the frog hypothalamus, Proc. Natl. Acad. Sci. U.S.A., 97, 13925-13930, 2000.
243. Suzdak, P.D., Schwartz, R.D., Skolnick, P., and Paul, S.M., Ethanol stimulates gamma-aminobutyric acid receptor-mediated chloride transport in rat brain synaptoneurosomes, Proc. Natl. Acad. Sci. U.S.A., 83, 4071-4075, 1986.
244. Grobin, A.C., Matthews, D.B., Devaud, L.L., and Morrow, A.L., The role of GABA(A) receptors in the acute and chronic effects of ethanol, Psychopharmacology (Berl.), 139, 2-19, 1998.
245. VanDoren, M.J., Matthews, D.B., Janis, G.C., Grobin, A.C., Devaud, L.L., and Morrow, A.L., Neuroactive steroid 3alpha-hydroxy-5alpha-pregnan-20-one modulates electrophysiological and behavioral actions of ethanol, J. Neurosci., 20, 1982-1989, 2000.