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* Departments of Internal Medicine and Pharmacology, University of Texas Southwestern Medical Center, Dallas, Texas 75390
Banting and Best Department of Medical Research, University of Toronto, Toronto M5G 1L6 Canada
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ABSTRACT |
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 TOP ABSTRACT I. Initial Isolation of Steroidogenic...II. Developmental Profile of SF-1...III. The Roles of SF-1 in VivoIV. Regulation of SF-1 Expression and...V. Directions for Future ResearchACKNOWLEDGEMENTSREFERENCES |
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I. Initial Isolation of Steroidogenic Factor 1 |
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TOPABSTRACTI. Initial Isolation of Steroidogenic...II. Developmental Profile of SF-1...III. The Roles of SF-1 in VivoIV. Regulation of SF-1 Expression and...V. Directions for Future ResearchACKNOWLEDGEMENTSREFERENCES |
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30 kb of its 5'-flanking region (White et al., 1984a). The Schimmer laboratory had extensive experience with Y1 mouse adrenocortical tumor cells, which expressed several cytochrome P450 steroid hydroxylases - but not steroid 21-hydroxylase — in a hormonally responsive manner and were readily amenable to both stable and transient transfection (Schimmer, 1985). In a collaborative effort, the Schimmer and Seidman laboratories showed that Y1 cells stably transfected with the mouse 21-hydroxylase cosmid recovered hormonally regulated expression of 21-hydroxylase (Parker et al., 1985). Thereafter, 5'-deletion assays localized sequences essential for cell-selective and hormone-regulated expression of the 21-hydroxylase gene to the proximal 330 bp of 5'-flanking DNA (Parker et al., 1986; Handler et al., 1988). Using similar approaches, a number of groups analyzed the 5'-flanking regions of genes encoding the cytochrome P450 steroid hydroxylases (for reviews of these studies, see Omura and Morohashi, 1995; Parker and Schimmer, 1995). In particular, studies by two groups identified shared AGGTCA promoter elements in the proximal promoter regions of several of the steroid hydroxylases that interacted with the same DNA-binding protein (Rice et al., 1991; Morohashi et al., 1992). This protein, which initially was found only in steroidogenic cell lines, was designated steroidogenic factor 1 (SF-1) or adrenal 4-binding protein (Ad4BP). The selective expression of SF-1 in steroidogenic cells and its regulation of multiple genes encoding steroid hydroxylases provided the first clues that it was an important determinant of the cell-selective expression of the steroidogenic enzymes.
Based on evidence that SF-1 was a key determinant of the expression of the cytochrome P450 steroid hydroxylases, the Parker and Morohashi laboratories independently cloned cDNAs encoding SF-1. Morohashi and colleagues used an oligonucleotide affinity column to purify the protein from bovine adrenal glands, ultimately allowing them to obtain amino acid sequence and clone a bovine cDNA with an oligonucleotide probe (Honda et al., 1993). In contrast, Douglas Rice, a postdoctoral fellow in the Parker laboratory, reasoned that the AGGTCA DNA recognition motif represented a binding site for an atypical member of the nuclear hormone receptor family. Using a hybridization probe comprising the DNA-binding region of retinoid X receptor, the Parker laboratory isolated a cDNA clone that was expressed in adrenal gland, testes, and ovaries, but not in a variety of other tissues (Lala et al., 1992).
Subsequent studies established that the mouse and bovine cDNAs encoded orthologs of a protein that transactivated the steroid hydroxylase promoters in steroidogenic and nonsteroidogenic cells. As predicted from the cloning strategy used by the Parker group, the sequences of these cDNAs confirmed that SF-1 belonged to the nuclear hormone receptor family, with striking homology to the Drosophila nuclear receptor fushi tarazu factor 1 (Ftz-F1) and the mouse nuclear receptor embryonal long terminal repeat-binding protein (Tsukiyama et al., 1992). SF-1 homologs have been identified in a diverse group of species that includes humans, marmosets, cows, sheep, horses, mice, rats, pigs, tamarind wallabies, chickens, turtles, salmon, trout, zebrafish, flies, and worms.
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II. Developmental Profile of SF-1 Expression |
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TOPABSTRACTI. Initial Isolation of Steroidogenic...II. Developmental Profile of SF-1...III. The Roles of SF-1 in VivoIV. Regulation of SF-1 Expression and...V. Directions for Future ResearchACKNOWLEDGEMENTSREFERENCES |
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E12.5 to E13.5, SF-1 expression was restricted to the steroidogenic cells in the cortex. The initiation of SF-1 expression before the onset of steroidogenesis supported its key role in steroid hydroxylase expression and suggested additional roles in adrenal development.
In mice, gonadal development first becomes apparent at E9, when the intermediate mesoderm condenses into the urogenital ridge, which ultimately contributes cell lineages to the gonads, adrenal cortex, and kidneys. At this time, developing testes and ovaries are indistinguishable histologically and thus are termed indifferent or bipotential. By E12.5, the fetal testes have organized into the testicular cords, which contain fetal Sertoli cells and primordial germ cells, and the surrounding interstitial region, which contains the Leydig cells. Faint expression of SF-1 was seen in both male and female embryos from the inception of the indifferent stage (E9.0–E9.5), persisting thereafter throughout the indifferent gonad stage. Coincident with formation of the testicular cords at E12.5, SF-1 expression persisted in the testes but diminished in ovaries (Hatano et al., 1994; Ikeda et al., 1994). In addition, SF-1 transcripts were detected in both the interstitial region, where Leydig cells produce steroid hormones, and the testicular cords, where fetal Sertoli cells produce anti-Müllerian hormone. SF-1 transcripts also were detected in the embryonic diencephalon — the precursor to the endocrine hypothalamus - and the anterior pituitary gland (Ikeda et al., 1994). Taken together, these findings suggested roles for SF-1 in gonadal development that extended beyond its effects on the expression of the steroidogenic enzymes and actions to regulate multiple levels of the hypothalamic-pituitary-steroidogenic organ axis.
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III. The Roles of SF-1 in Vivo |
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TOPABSTRACTI. Initial Isolation of Steroidogenic...II. Developmental Profile of SF-1...III. The Roles of SF-1 in VivoIV. Regulation of SF-1 Expression and...V. Directions for Future ResearchACKNOWLEDGEMENTSREFERENCES |
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-subunit of glycoprotein hormones (GSU), and the receptor for gonadotropin-releasing hormone (Ingraham et al., 1994; Shinoda et al., 1995). As shown in Figure 2, these knockout mice also lacked the ventromedial hypothalamic nucleus (VMH), a hypothalamic region linked to feeding and appetite regulation and female reproductive behavior (Ikeda et al., 1995; Shinoda et al., 1995). Finally, although the functional consequences remain to be defined, the SF-1 knockout mice had defects in their splenic parenchyma (Morohashi et al., 1999).
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Because the original SF-1 knockout mice are globally deficient in SF-1, they cannot be used to delineate the roles of SF-1 at specific sites of expression. For example, the apparent defect in gonadotrope function could merely reflect the absence of gonadal steroids. Another limitation of the original SF-1 knockout mice is their early postnatal death from adrenocortical insufficiency, which precludes efforts to examine the roles of SF-1 after differentiation has occurred. Finally, the need to administer exogenous corticosteroids to keep SF-1 knockout mice alive complicates considerably our ability to use these mice to assess the role of the VMH in feeding and weight regulation. To obviate such limitations, the Parker laboratory has begun to use the Cre/loxP system to produce tissue-specific knockouts of SF-1. Marit Bakke initiated this process by modifying the SF-1 locus in embryonic stem cells to insert recognition sites for the bacteriophage Cre recombinase (termed loxP sites) around the last exon of SF-1, which encodes an essential domain for transcriptional activation and transcription termination sequences. The second essential step in the tissue-specific knockout was to generate a Cre transgene selectively active in a subset of SF-1-expressing cells. Lisa Cushman in the Camper laboratory generated a transgenic mouse line in which Cre expression was directed to the anterior pituitary gland by the 5'-flanking sequences of the subunit of glycoprotein hormones (Cushman et al., 2000). Liping Zhao bred the loxP-modified SF-1 line with the Cre transgenic mice, ultimately generating mice with pituitary-specific disruption of SF-1.
As shown in Figure 3, the GSU-Cre/loxP mice selectively lacked SF-1 immunoreactivity in the anterior pituitary (Zhao et al., 2001) but had normal levels at other sites, including the adrenal cortex and VMH. These mice had markedly diminished levels of pituitary gonadotropins and exhibited severe gonadal hypoplasia secondary to impaired gonadotropin stimulation. These pituitary-specific SF-1 knockout mice demonstrated that the local production of SF-1 in mice is essential for normal gonadotrope function, strongly supporting a direct role for SF-1 in gonadotropin gene expression.
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IV. Regulation of SF-1 Expression and Function |
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TOPABSTRACTI. Initial Isolation of Steroidogenic...II. Developmental Profile of SF-1...III. The Roles of SF-1 in VivoIV. Regulation of SF-1 Expression and...V. Directions for Future ResearchACKNOWLEDGEMENTSREFERENCES |
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T3 gonadotropes (Harris and Mellon, 1998). In the latter study, the transcription factor USF-1 was shown to regulate SF-1 expression. In view of the known developmental roles of basic helix-loop-helix proteins that bind these E-box motifs, these studies suggest an important role for basic helix-loop-helix proteins in regulating SF-1 expression in several cell types. Other promoter elements implicated in SF-1 promoter activity include a GC-rich sequence that may represent a binding site for Sp1 and a CCAAT-box motif (Woodson et al., 1997). The precise roles of these elements in different tissues remain to be defined. Moreover, the lack of success in the reported transgenic expression studies suggests that other elements also play important roles in vivo.
One might predict that distinct mechanisms regulate SF-1 expression in the adrenal cortex and gonads, which are believed to arise from the same embryonic lineage (Hatano et al., 1996), versus the anterior pituitary and hypothalamus, which are contiguous structures that interact reciprocally during development (Rosenfeld et al., 1996). In support of this, the pituitary transcripts in some species arise from a transcription initiation site distinct from that used in other sites (Ninomiya et al., 1995; Kimura et al., 2000). In an effort to explore the mechanisms that regulate SF-1 expression in vivo, the Parker laboratory recently used a 50-kb fragment derived from a bacterial artificial chromosome to direct expression of a green fluorescent protein (GFP) reporter gene in transgenic mice. As shown in Figure 4, GFP expression in the urogenital ridge was detected at E9.5, closely paralleling the onset of SF expression during gonadogenesis (Ikeda et al., 1994). Although GFP expression in the adrenal primordium and VMH also corresponded to the known expression profile of SF-1, GFP was not expressed in the anterior pituitary. These results document that 50 kb of 5'-flanking region suffice to direct regulated expression in most sites but also suggest that additional regulatory elements are required for activation of the pituitary-specific promoter.
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In the absence of a clear-cut ligand, others have examined the possibility that post-translational modifications alter SF-1 transcriptional activity. In their initial description of the bovine sequence, Morohashi and colleagues noted a potential phosphorylation site for cAMP-dependent protein kinase. Mellon and colleagues subsequently reported that recombinantly expressed SF-1 was phosphorylated in vitro by cAMP-dependent protein kinase (Zhang and Mellon, 1996). Mutation of the serine residue in the consensus motif did not impair SF-1 function in transfection assays (Lopez et al., 2001), suggesting that this site is not a key regulator of SF-1 activity.
In a separate line of investigations, the Ingraham laboratory used peptide mapping to define a site of SF-1 phosphorylation (Ser203) that mapped within a consensus motif for phosphorylation by mitogen-activated protein (MAP) kinase (Hammer et al., 1999). They further showed that activating the MAP kinase pathway increased SF-1 transcriptional activation, while mutation of Ser203 diminished SF-1 activity. Collectively, these findings raise the possibility that SF-1 function is modulated by posttranslational modification through extracellular signals that act via the MAP kinase pathway.
C. REGULATION THROUGH PROTEIN-PROTEIN INTERACTIONS
Analyses of human patients and knockout mouse models have identified a number of other genes that play key roles in the development of SF-1-expressing tissues (for a review, see Parker et al., 1999). It is almost certain that these genes interact with SF-1 - either in hierarchical cascades of gene regulation or via protein-protein interactions - to mediate endocrine development. Indeed, a number of factors have been shown to interact directly with SF-1, including the Wilm’s tumor related tumor suppressor gene WT-1 (Nachtigal et al., 1998), GATA-4 (Tremblay and Viger, 1999), Ptx1 (Tremblay et al., 1999), SOX9 (de Santa-Barbara et al., 1999), and EGR1 (Halvorson et al., 1998; Dorn et al., 1999). In contrast, SF-1 expression is markedly decreased in the gonads — but not the adrenal glands — of Lhx9 knockout mice, suggesting that Lhx9 regulates SF-1 expression in the gonads (Birk et al., 2000). Finally, DAX-1 and SF-1 apparently interact both hierarchically (i.e., SF-1 regulates DAX1 expression) (Yu et al., 1998; Kawabe et al., 1999) and by direct protein-protein interactions wherein DAX1 inhibits SF-1 transcriptional activation (Ito et al., 1997; Crawford et al., 1998).
Besides interactions with other tissue-specific transcription factors, it is apparent that coactivators and co-repressors are critical modulators of nuclear receptor transcriptional activity (for a review, see Xu et al., 1999). Predictably, a number of co-regulators have been reported to interact with SF-1, including CBP/P300 (Monte et al., 1998), GRIPI (Hammer et al., 1999), SRC-1 (Crawford et al., 1997a), MBP1 (Kabe et al., 1999), SMRT (Hammer et al., 1999), and N-CoR (Crawford et al., 1998; Nachtigal et al., 1998). It is possible that differential interactions with these co-regulators, as well as with the tissue-specific transcription factors described above, specify the differential expression of SF-1 target genes in various tissues. An increased understanding of the ways in which these genes interact to regulate the expression of specific target genes undoubtedly will provide important new insights into processes of endocrine development.
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V. Directions for Future Research |
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TOPABSTRACTI. Initial Isolation of Steroidogenic...II. Developmental Profile of SF-1...III. The Roles of SF-1 in VivoIV. Regulation of SF-1 Expression and...V. Directions for Future ResearchACKNOWLEDGEMENTSREFERENCES |
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-hydroxylase or Mullerian inhibiting substance (MIS) receptor promoters), and in Sertoli/granulosa cells (using the MIS or inhibin promoters). These experiments should provide novel insights into the specific functions of SF-1 at these sites. What is the relationship of SF-1 to other closely related members of the NR5 nuclear receptor family? SF-1 most closely resembles another orphan nuclear receptor, NR5A2, particularly within the DNA-binding domain, suggesting that these two transcription factors may regulate overlapping target genes. Indeed, both SF-1 and NR5A2 can activate promoter activity of the small heterodimerization partner (SHP) nuclear receptor (Lee et al., 1999), which is expressed in the adrenal cortex, liver, and other tissues. Interestingly, NR5A2 transcripts are expressed at high levels in the corpus luteum of the ovary (Boerboom et al., 2000), where it may replace SF-1 as a critical regulator of the cytochrome P450 steroid hydroxylases at certain stages of the ovulatory cycle.
While many laboratories have identified a diverse group of SF-1 target genes, as summarized in Table I, these analyses largely have focused on transient transfection assays using relatively limited stretches of promoter/regulatory DNA. These studies may overemphasize the importance of SF-1 in gene regulation and verification of important roles in vivo ultimately is needed. Among the SF-1 target genes in Table I, evidence supporting such in vivo roles has been provided for MIS (Giuili et al., 1997; Arango et al., 1999) and LHß (Keri and Nilson, 1996). Moreover, important questions about specific roles of SF-1 in development versus differentiated function may be too subtle to be addressed in either global or tissue-specific knockouts. Although it was anticipated that SF-1 knockout mice might provide in vivo evidence for the importance of SF-1 in gene expression, particularly for those genes involved in steroidogenesis, the failure of the knockout mice to develop the steroidogenic organs and the VMH precluded such analyses at these sites. Strong evidence for a developmental role of SF-1 came from studies in which forced expression of SF-1 in embryonic stem cells induced the expression of the cholesterol side-chain cleavage enzyme (Crawford et al., 1997b). In contrast, analyses of mutant Y1 cells with defects affecting SF-1 function provided evidence for multiple roles in differentiated function. The Schimmer laboratory showed that a SF-1 mutation was associated with decreased expression of the ACTH receptor, 11ß-hydroxylase, cholesterol side-chain cleavage enzyme, and steroidogenic acute regulator protein (StAR) (Frigeri et al., 2000). Interestingly, the SF-1 defect affected the expression of the ACTH receptor and 11ß-hydroxylase to a much greater degree than cholesterol side-chain cleavage enzyme or StAR, suggesting further subtleties among the target genes in their regulation by SF-1.
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ACKNOWLEDGEMENTS |
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TOPABSTRACTI. Initial Isolation of Steroidogenic...II. Developmental Profile of SF-1...III. The Roles of SF-1 in VivoIV. Regulation of SF-1 Expression and...V. Directions for Future ResearchACKNOWLEDGEMENTSREFERENCES |
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REFERENCES |
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TOPABSTRACTI. Initial Isolation of Steroidogenic...II. Developmental Profile of SF-1...III. The Roles of SF-1 in VivoIV. Regulation of SF-1 Expression and...V. Directions for Future ResearchACKNOWLEDGEMENTSREFERENCES |
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L. J. Martin, H. Taniguchi, N. M. Robert, J. Simard, J. J. Tremblay, and R. S. Viger GATA Factors and the Nuclear Receptors, Steroidogenic Factor 1/Liver Receptor Homolog 1, Are Key Mutual Partners in the Regulation of the Human 3{beta}-Hydroxysteroid Dehydrogenase Type 2 Promoter Mol. Endocrinol., September 1, 2005; 19(9): 2358 - 2370. [Abstract] [Full Text] [PDF]
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J. Seely, K. S. Amigh, T. Suzuki, B. Mayhew, H. Sasano, V. Giguere, J. Laganiere, B. R. Carr, and W. E. Rainey Transcriptional Regulation of Dehydroepiandrosterone Sulfotransferase (SULT2A1) by Estrogen-Related Receptor {alpha} Endocrinology, August 1, 2005; 146(8): 3605 - 3613. [Abstract] [Full Text] [PDF]
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Y. Li, M. Choi, K. Suino, A. Kovach, J. Daugherty, S. A. Kliewer, and H. E. Xu Structural and biochemical basis for selective repression of the orphan nuclear receptor liver receptor homolog 1 by small heterodimer partner PNAS, July 5, 2005; 102(27): 9505 - 9510. [Abstract] [Full Text] [PDF]
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 S. Y. Park, J. J. Meeks, G. Raverot, L. E. Pfaff, J. Weiss, G. D. Hammer, and J. L. Jameson Nuclear receptors Sf1 and Dax1 function cooperatively to mediate somatic cell differentiation during testis development Development, May 15, 2005; 132(10): 2415 - 2423. [Abstract] [Full Text] [PDF]
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 C. K. Cheng and P. C. K. Leung Molecular Biology of Gonadotropin-Releasing Hormone (GnRH)-I, GnRH-II, and Their Receptors in Humans Endocr. Rev., April 1, 2005; 26(2): 283 - 306. [Abstract] [Full Text] [PDF]
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G. D. Hammer, K. L. Parker, and B. P. Schimmer Minireview: Transcriptional Regulation of Adrenocortical Development Endocrinology, March 1, 2005; 146(3): 1018 - 1024. [Abstract] [Full Text] [PDF]
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L. J. Martin and J. J. Tremblay The Human 3{beta}-Hydroxysteroid Dehydrogenase/{Delta}5-{Delta}4 Isomerase Type 2 Promoter Is a Novel Target for the Immediate Early Orphan Nuclear Receptor Nur77 in Steroidogenic Cells Endocrinology, February 1, 2005; 146(2): 861 - 869. [Abstract] [Full Text] [PDF]
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J. L. Jameson Of Mice and Men: The Tale of Steroidogenic Factor-1 J. Clin. Endocrinol. Metab., December 1, 2004; 89(12): 5927 - 5929. [Full Text] [PDF]
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N. Ferrara Vascular Endothelial Growth Factor: Basic Science and Clinical Progress Endocr. Rev., August 1, 2004; 25(4): 581 - 611. [Abstract] [Full Text] [PDF]
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 L. A. Freeman, A. Kennedy, J. Wu, S. Bark, A. T. Remaley, S. Santamarina-Fojo, and H. B. Brewer Jr. The orphan nuclear receptor LRH-1 activates the ABCG5/ABCG8 intergenic promoter J. Lipid Res., July 1, 2004; 45(7): 1197 - 1206. [Abstract] [Full Text] [PDF]
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F. Hammer, N. A. Compagnone, J.-L. Vigne, S. R. Bair, and S. H. Mellon Transcriptional Regulation of P450scc Gene Expression in the Embryonic Rodent Nervous System Endocrinology, February 1, 2004; 145(2): 901 - 912. [Abstract] [Full Text] [PDF]
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 M. Maira, C. Couture, G. Le Martelot, A.-M. Pulichino, S. Bilodeau, and J. Drouin The T-box Factor Tpit Recruits SRC/p160 Co-activators and Mediates Hormone Action J. Biol. Chem., November 21, 2003; 278(47): 46523 - 46532. [Abstract] [Full Text] [PDF]
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A. E. Falender, R. Lanz, D. Malenfant, L. Belanger, and J. S. Richards Differential Expression of Steroidogenic Factor-1 and FTF/LRH-1 in the Rodent Ovary Endocrinology, August 1, 2003; 144(8): 3598 - 3610. [Abstract] [Full Text] [PDF]
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J. J. Tremblay and R. S. Viger Transcription Factor GATA-4 Is Activated by Phosphorylation of Serine 261 via the cAMP/Protein Kinase A Signaling Pathway in Gonadal Cells J. Biol. Chem., June 6, 2003; 278(24): 22128 - 22135. [Abstract] [Full Text] [PDF]
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J. LeCouter, R. Lin, G. Frantz, Z. Zhang, K. Hillan, and N. Ferrara Mouse Endocrine Gland-Derived Vascular Endothelial Growth Factor: A Distinct Expression Pattern from Its Human Ortholog Suggests Different Roles as a Regulator of Organ-Specific Angiogenesis Endocrinology, June 1, 2003; 144(6): 2606 - 2616. [Abstract] [Full Text] [PDF]
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A. Martinez, P. Val, I. Sahut-Barnola, C. Aigueperse, G. Veyssiere, and A.-M. Lefrancois-Martinez Steroidogenic Factor-1 Controls the Aldose Reductase akr1b7 Gene Promoter in Transgenic Mice through an Atypical Binding Site Endocrinology, May 1, 2003; 144(5): 2111 - 2120. [Abstract] [Full Text] [PDF]
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 Y. Yoshiura, B. Senthilkumaran, M. Watanabe, Y. Oba, T. Kobayashi, and Y. Nagahama Synergistic Expression of Ad4BP/SF-1 and Cytochrome P-450 Aromatase (Ovarian Type) in the Ovary of Nile Tilapia, Oreochromis niloticus, During Vitellogenesis Suggests Transcriptional Interaction Biol Reprod, May 1, 2003; 68(5): 1545 - 1553. [Abstract] [Full Text] [PDF]
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J. J. Meeks, S. E. Crawford, T. A. Russell, K.-i. Morohashi, J. Weiss, and J. L. Jameson Dax1 regulates testis cord organization during gonadal differentiation Development, March 1, 2003; 130(5): 1029 - 1036. [Abstract] [Full Text] [PDF]
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