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Molecular Biology of Steroid Hormone Synthesis^*

Department of Pediatrics and The Metabolic Research Unit, University of California San Francisco, California 94143

Correspondence: Address requests for reprints to: Walter L. Miller, M.D., Room 677- S, Pediatrics, University of California, San Francisco, San Francisco, CA 94143.

Abstract

Introduction: STEROID hormones are familiar clinically and physiologically as regulators of physiological processes. Five groups of steroid hormones are generally recognized according to their physiological behavior: mineralocorticoids, which instruct the renal tubules to retain sodium; glucocorticoids, which are named for their carbohydratemobilizing properties but have many other effects as well; estrogens, which induce female secondary sexual characteristics; progestins, which are essential for reproduction; and androgens, which induce male secondary sexual characteristics. These classes of steroid hormones are structurally similar and arise from a common series of pathways. They are distinguished by their actions on one or more specific steroid hormone receptors. The hormone/receptor complexes function as tissue-specific transcriptional regulators of distinct domains of genes and, consequently, exert their broad array of physiological effects. (For reviews, see Refs. 1 and 2.) The pathways by which these steroid hormones are produced from cholesterol were initially studied by studying the structures of the steroids themselves. Over the past 30 yr, studies of enzyme kinetics and of steroidal precursorproduct relationships have led to the widely held belief that a very large number of distinct enzymes are involved in the conversion of cholesterol and its esters to active steroid hormones. The application of modern protein chemistry and molecular biology to these questions has radically changed this view.

Footnotes

^* Supported by NIH Grant DK-37922 and March of Dimes Grant 6-396.

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				P. R. Manna, M. T. Dyson, D. W. Eubank, B. J. Clark, E. Lalli, P. Sassone-Corsi, A. J. Zeleznik, and D. M. Stocco Regulation of Steroidogenesis and the Steroidogenic Acute Regulatory Protein by a Member of the cAMP Response-Element Binding Protein Family Mol. Endocrinol., January 1, 2002; 16(1): 184 - 199. [Abstract] [Full Text] [PDF]

				K. Tajima, K. Hosokawa, Y. Yoshida, A. Dantes, R. Sasson, F. Kotsuji, and A. Amsterdam Establishment of FSH-responsive cell lines by transfection of pre-ovulatory human granulosa cells with mutated p53 (p53val135) and Ha-ras genes Mol. Hum. Reprod., January 1, 2002; 8(1): 48 - 57. [Abstract] [Full Text] [PDF]

				K. Jia, P. S. Albert, and D. L. Riddle DAF-9, a cytochrome P450 regulating C. elegans larval development and adult longevity Development, January 1, 2002; 129(1): 221 - 231. [Abstract] [Full Text] [PDF]

				I. P. Artemenko, D. Zhao, D. B. Hales, K. H. Hales, and C. R. Jefcoate Mitochondrial Processing of Newly Synthesized Steroidogenic Acute Regulatory Protein (StAR), but Not Total StAR, Mediates Cholesterol Transfer to Cytochrome P450 Side Chain Cleavage Enzyme in Adrenal Cells J. Biol. Chem., November 30, 2001; 276(49): 46583 - 46596. [Abstract] [Full Text] [PDF]

				S. M. Baumgartner-Parzer, E. Schulze, W. Waldhausl, S. Pauschenwein, S. Rondot, P. Nowotny, K. Meyer, H. Frisch, F. Waldhauser, and H. Vierhapper Mutational Spectrum of the Steroid 21-Hydroxylase Gene in Austria: Identification of a Novel Missense Mutation J. Clin. Endocrinol. Metab., October 1, 2001; 86(10): 4771 - 4775. [Abstract] [Full Text] [PDF]

				T. Tajima, K. Fujieda, N. Kouda, J. Nakae, and W. L. Miller 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., August 1, 2001; 86(8): 3820 - 3825. [Abstract] [Full Text] [PDF]

				C. J. Lin, J. W. M. Martens, and W. L. Miller NF-1C, Sp1, and Sp3 Are Essential for Transcription of the Human Gene for P450c17 (Steroid 17{alpha}-hydroxylase/17,20 lyase) in Human Adrenal NCI-H295A Cells Mol. Endocrinol., August 1, 2001; 15(8): 1277 - 1293. [Abstract] [Full Text] [PDF]

				N. Sekar and J. D. Veldhuis Concerted Transcriptional Activation of the Low Density Lipoprotein Receptor Gene by Insulin and Luteinizing Hormone in Cultured Porcine Granulosa-Luteal Cells: Possible Convergence of Protein Kinase A, Phosphatidylinositol 3-Kinase, and Mitogen-Activated Protein Kinase Signaling Pathways Endocrinology, July 1, 2001; 142(7): 2921 - 2928. [Abstract] [Full Text] [PDF]

				M.-C. Huang and W. L. Miller Creation and Activity of COS-1 Cells Stably Expressing the F2 Fusion of the Human Cholesterol Side-Chain Cleavage Enzyme System Endocrinology, June 1, 2001; 142(6): 2569 - 2576. [Abstract] [Full Text] [PDF]

				M.-C. Hu, N.-C. Hsu, C.-I Pai, C.-K. L. Wang, and B.-c. Chung Functions of the Upstream and Proximal Steroidogenic Factor 1 (SF-1)-Binding Sites in the CYP11A1 Promoter in Basal Transcription and Hormonal Response Mol. Endocrinol., May 1, 2001; 15(5): 812 - 818. [Abstract] [Full Text]

				A. Lacroix, N. N'Diaye, J. Tremblay, and P. Hamet Ectopic and Abnormal Hormone Receptors in Adrenal Cushing's Syndrome Endocr. Rev., February 1, 2001; 22(1): 75 - 110. [Abstract] [Full Text]

				R. Sasson, K. Tajima, and A. Amsterdam Glucocorticoids Protect against Apoptosis Induced by Serum Deprivation, Cyclic Adenosine 3',5'-Monophosphate and p53 Activation in Immortalized Human Granulosa Cells: Involvement of Bcl-2 Endocrinology, February 1, 2001; 142(2): 802 - 811. [Abstract] [Full Text] [PDF]

				P. R. Manna, T. El-Hefnawy, J. Kero, and I. T. Huhtaniemi Biphasic Action of Prolactin in the Regulation of Murine Leydig Tumor Cell Functions Endocrinology, January 1, 2001; 142(1): 308 - 318. [Abstract] [Full Text] [PDF]

				P. R. Manna, J. Kero, M. Tena-Sempere, P. Pakarinen, D. M. Stocco, and I. T. Huhtaniemi Assessment of Mechanisms of Thyroid Hormone Action in Mouse Leydig Cells: Regulation of the Steroidogenic Acute Regulatory Protein, Steroidogenesis, and Luteinizing Hormone Receptor Function Endocrinology, January 1, 2001; 142(1): 319 - 331. [Abstract] [Full Text] [PDF]

				D. Boerboom and J. Sirois Equine P450 Cholesterol Side-Chain Cleavage and 3{beta}-Hydroxysteroid Dehydrogenase/{{Delta}}5-{{Delta}}4 Isomerase: Molecular Cloning and Regulation of Their Messenger Ribonucleic Acids in Equine Follicles During the Ovulatory Process Biol Reprod, January 1, 2001; 64(1): 206 - 215. [Abstract] [Full Text]

				L. Ibáñez, J. DiMartino-Nardi, N. Potau, and P. Saenger Premature Adrenarche--Normal Variant or Forerunner of Adult Disease? Endocr. Rev., December 1, 2000; 21(6): 671 - 696. [Abstract] [Full Text]

				B. Bao, N. Kumar, R. M. Karp, H. Allen Garverick, and K. Sundaram Estrogen Receptor-{beta} Expression in Relation to the Expression of Luteinizing Hormone Receptor and Cytochrome P450 Enzymes in Rat Ovarian Follicles Biol Reprod, December 1, 2000; 63(6): 1747 - 1755. [Abstract] [Full Text]

				H. S. Bose, S. Sato, J. Aisenberg, S. A. Shalev, N. Matsuo, and W. L. Miller Mutations in the Steroidogenic Acute Regulatory Protein (StAR) in Six Patients with Congenital Lipoid Adrenal Hyperplasia J. Clin. Endocrinol. Metab., October 1, 2000; 85(10): 3636 - 3639. [Abstract] [Full Text]

				G. W. Turner, J. Gershenzon, and R. B. Croteau Development of Peltate Glandular Trichomes of Peppermint Plant Physiology, October 1, 2000; 124(2): 665 - 680. [Abstract] [Full Text]

				G. Zhang, J. C. Garmey, and J. D. Veldhuis Interactive Stimulation by Luteinizing Hormone and Insulin of the Steroidogenic Acute Regulatory (StAR) Protein and 17{alpha}-Hydroxylase/17, 20-Lyase (CYP17) Genes in Porcine Theca Cells Endocrinology, August 1, 2000; 141(8): 2735 - 2742. [Abstract] [Full Text] [PDF]

				R. J. Urban, Y. Bodenburg, A. Kurosky, T. G. Wood, and S. Gasic Polypyrimidine Tract-Binding Protein-Associated Splicing Factor Is a Negative Regulator of Transcriptional Activity of the Porcine P450scc Insulin-Like Growth Factor Response Element Mol. Endocrinol., June 1, 2000; 14(6): 774 - 782. [Abstract] [Full Text]

				C. Le Roy, J. Y. Li, D. M. Stocco, D. Langlois, and J. M. Saez Regulation by Adrenocorticotropin (ACTH), Angiotensin II, Transforming Growth Factor-{beta}, and Insulin-Like Growth Factor I of Bovine Adrenal Cell Steroidogenic Capacity and Expression of ACTH Receptor, Steroidogenic Acute Regulatory Protein, Cytochrome P450c17, and 3{beta}-Hydroxysteroid Dehydrogenase Endocrinology, May 1, 2000; 141(5): 1599 - 1607. [Abstract] [Full Text] [PDF]

				L. S. Levine Congenital Adrenal Hyperplasia Pediatr. Rev., May 1, 2000; 21(5): 159 - 171. [Full Text]