Estrogen Receptor Null Mice - Endocrine Reviews

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June, 1999 ESTROGEN RECEPTOR NULL MICE 361 TABLE 1. Published reports involving the �ERKO and �ERKO models and the ER�-deficient human Author ER-� knockout mice: general Yr. Title Ref. Lubahn et al. 1993 Alteration of reproductive function but not prenatal sexual development after insertional disruption of the mouse estrogen receptor gene. 46 Couse et al. 1995 Analysis of transcription and estrogen insensitivity in the female mouse after targeted disruption of the estrogen receptor gene. 123 Couse et al. ER-� knockout mice: general 1997 Tissue distribution and quantitative analysis of estrogen receptor-� (ER�) and estrogen-� (ER�) messenger ribonucleic acid in the wild-type and ER�-knockout mouse. 93 Krege et al. Female reproductive tract 1998 Generation and reproductive phenotypes of mice lacking estrogen receptor-�. 47 Curtis et al. 1996 Physiological coupling of growth factor and steroid receptor-signaling pathways: estrogen receptor knockout mice lack estrogen-like response to epidermal growth factor. 170 Lindzey et al. 1996 Uterotropic effects of dihydrotestosterone in estrogen receptor knockout and wild-type mice. a 154 Das et al. 1997 Estrogenic responses in estrogen receptor-� deficient mice reveal a distinct signaling pathway. 173 Cooke et al. 1997 Stromal estrogen receptors mediate mitogenic effects of estradiol on uterine epithelium. 171 Buchanan et al. 1998 Role of stromal and epithelial estrogen receptors in vaginal epithelial proliferation, stratification, and cornification. 198 Ghosh et al. 1998 Methoxychlor acts in ER�-KO mice through an ER-� independent mechanism. a 174 Rosenfeld et al. 1998 Estrogen receptor-� knockout mice reveal a role for estrogen in mammalian female sexual development. a 494 Kurita et al. 1998 Estrogen induces progesterone receptors in uterine stroma of estrogen receptor � knockout mouse. a 495 Buchanan et al. 1999 Tissue compartment-specific estrogen receptor-� participation in the mouse uterine epithelial secretory response. 172 Curtis et al. 1999 Disruption of estrogen signaling does not prevent progesterone action in the estrogen receptor knockout mouse uterus. 183 Schomberg et al. Mammary gland 1999 Targeted disruption of the estrogen receptor-� (ER�) gene in mice: characterization of ovarian responses and phenotypes. 142 Bocchinfuso and Korach 1997 Mammary gland development and tumorigenesis in estrogen receptor knockout mice. 269 Cunha et al. 1997 Elucidation of a role of stromal steroid hormone receptors in mammary gland growth and development by tissue recombination experiments. 278 Kenney et al. 1997 The response of null estrogen receptor mammary parenchyma and mesenchyme in vivo. 496 Bocchinfuso et al. Male reproductive tract 1999 An MMTV-Wnt-1 transgene induces mammary gland hyperplasia and tumorigenesis in mice lacking estrogen receptor-�. 297 Eddy et al. 1996 Targeted disruption of the estrogen receptor gene in male mice causes alteration of spermatogenesis and infertility. 315 Donaldson et al. 1996 Morphometric study of the gubernaculum in male estrogen receptor mutant mice. 316 Hess et al. 1997 A role for oestrogens in the male reproductive system. 327 Rosenfeld et al. Neuroendocrine system 1998 Transcription and translation of estrogen receptor-beta in the male reproductive tract of estrogen knockout and wild-type mice. 121 Scully et al. 1997 Role of estrogen receptor � in the anterior pituitary gland. 282 Shughrue et al. 1997 Responses in the brain of estrogen receptor �-disrupted mice. 400 Shughrue et al. 1997 The disruption of estrogen receptor-� mRNA in forebrain regions of the estrogen receptor-� knockout mouse. 352 Simerly et al. 1997 Estrogen receptor-dependent sexual differentiation of dopaminergic neurons in the preoptic region of the mouse. 405 Young et al. 1998 Estrogen receptor � is essential in the induction of oxytocin receptor by estrogen. 497 De Vries et al. 1997 Steroid-responsive vasopressin innervation in the brain of estrogen receptor-�-minus mice. a 498 Lindzey et al. 1998 Effects of castration and chronic steroid treatments on hypothalamic gonadotropin-releasing hormone content and pituitary gonadotropins in male wild-type and estrogen receptor-� knockout mice. 317 Lindzey et al. 1998 Steroid regulation of gonadotrope function in female wild-type (WT) and estrogen receptor-� knockout (ERKO) mice. a 252 Ogawa et al. 1998 Forebrain steroid receptor immunoreactive cells in neonatal and prepubertal estrogen receptor- � gene deficient (ERKO) mice. a 426 Moffatt et al. 1998 Induction of progestin receptors by estradiol in the forebrain of estrogen receptor-� genedisrupted mice. 423 Wersinger et al. 1998 Estradiol decreases proGnRH immunoreactivity in female wild type but not estrogen receptor � knockout mice. a 499 B. Mechanism of ER action The ERs are classified as class I members of the superfamily of nuclear hormone receptors, defined as a ligandinducible transcription factor (30). Early studies indicated the ER was cytoplasmic and became localized to the nucleus only upon ligand binding, providing the basis of the initial “twostep mechanism” of hormone action (1, 3). However, it is now accepted that the ER is a predominantly nuclear protein regardless of whether or not it is complexed with ligand (74). The inactive ER exists in a complex consisting of several
362 COUSE AND KORACH Vol. 20, No. 3 TABLE 1. continued Author Yr. Title Ref. Gu et al. Behavior 1999 Rapid action of 17�-estradiol on kainate-induced currents in hippocampal neurons lacking intracellular estrogen receptors. 364 Ogawa et al. 1996 Reversal of sex roles in genetic female mice by disruption of estrogen receptor gene. 417 Ogawa et al. 1997 Behavioral effects of estrogen receptor gene disruption in male mice. 318 Rissman et al. 1997 Estrogen receptors are essential for female sexual receptivity. 419 Wersinger et al. 1997 Masculine sexual behavior is disrupted in male and female mice lacking a functional estrogen receptor � gene. 427 Fugger et al. 1998 Estrogen receptor � is not required for estradiol’s effects on inhibitory avoidance behavior in female mice. a 500 Fugger et al. 1998 Characterization of spatial behavior and hippocampal electrophysiology in male estrogen receptor-�-minus mice. a 501 Fugger et al. 1998 Sex differences in the activational effect of ER� on spatial learning. 502 Ogawa et al. 1998 Roles of estrogen receptor-� gene expression in reproduction-related behaviors in female mice. 418 Ogawa et al. Cardiovascular system 1998 Modifications of testosterone-dependent behaviors by estrogen receptor-� gene disruption in male mice. 428 Johns et al. 1996 Disruption of estrogen receptor gene prevents 17�-estradiol-induced angiogenesis in transgenic mice. 503 Iafrati et al. 1997 Estrogen inhibits the vascular injury response in estrogen receptor-� deficient mice. 464 Johnson et al. 1997 Increased expression of the cardiac L-type channel in estrogen receptor-deficient mice. 472 Rubanyi et al. 1997 Vascular estrogen receptors and endothelium-derived nitric oxide production in the mouse aorta: gender difference and effect of estrogen receptor gene disruption. 467 Srivastava et al. 1997 Estrogen up-regulates apolipoprotein E (Apo E) gene expression by increasing Apo E mRNA in the translating pool via the estrogen receptor �-mediated pathway. 460 Freay et al. 1997 Mechanism of vascular smooth muscle relaxation by estrogen in depolarized rat and mouse aorta. 504 Cross et al. 1998 Estrogen receptor knock-out mice exhibit higher myocardial contractility than wild-type mice but are equally susceptible to ischemic injury. a Bone 505 Roemmich et al. 1997 Bone strain independently augments bone mineral in pubertal mice with the disrupted estrogen receptor gene. a 506 Pan et al. 1997 Estrogen receptor-alpha knockout (ERKO) mice lose trabecular and cortical bone following ovariectomy. a 448 Korach et al. Other 1997 The effects of estrogen receptor gene disruption on bone. 447 Smithson et al. 1997 The role of estrogen receptors and androgen receptors in sex steroid regulation of B lymphopoiesis. 507 Hurn et al. 1998 Stroke in mice deficient in classical estrogen receptors (ERKO�). a 508 Couse et al. 1998 Use of the estrogen receptor-� to investigate the role of ER� in the developmental and carcinogenic actions of diethylstilbestrol. a 509 Taylor and Lubahn 1998 Impaired glucose tolerance in the ER�KO mouse. a 490 Yellayi et al. 1998 Role of estrogen receptor-� (ER�) in the development and function of the thymus in male and female mice. a ER-deficient humans 510 Smith et al. 1994 Estrogen resistance caused by a mutation in the estrogen receptor gene in a man. 116 Sudhir et al. 1997 Endothelial dysfunction in a man with disruptive mutation of the oestrogen-receptor gene. 492 Sudhir et al. 1997 Premature coronary artery disease associated with a disruptive mutation in the estrogen receptor gene in a man. 493 Dieudonne et al. 1998 Immortalization and characterization of bone marrow stromal fibroblasts from a patient with a loss of function mutation in the estrogen receptor-� gene. 511 a Abstract. heat-shock and other proteins that appear to disassociate upon ligand binding, resulting in a “transformation” of the receptor to an active state (75). With continued research, the two-step mechanism model has evolved to state that upon binding of estradiol, or an estrogenic ligand, the transformed receptors form dimers that tightly associate with specific consensus DNA sequences, consisting of 15-bp inverted palindromes in the regulatory regions of target genes (52, 74– 76). This complex then interacts with basal transcription factors, coregulator proteins, and other transcription factors to ultimately regulate transcription of the target gene (52, 55, 76). However, in recent years, pathways of gene activation by the steroid receptors that deviate from this classical model have been described. These include gene activation by ligand-bound steroid receptors without evidence of direct DNA binding, but rather via interaction with other DNAbound transcription factors, such as an AP-1 complex (77, 78). In addition, ligand-independent activation of the receptor through pathways that alter the activity of cellular kinases and phosphatases has been demonstrated both in vitro and in vivo (reviewed in Ref. 55). The discovery of these pathways strongly supports the great importance of the ER and its ability to possibly provide diverse physiological functions even in the absence of ligand.
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