Neuroprotective actions of brain aromatase

Abulrob, 2005, Protection by cholesterol-extracting cyclodextrins: a role for N-methyl-D-aspartate receptor redistribution, J. Neurochem., 92, 1477, 10.1111/j.1471-4159.2005.03001.x Adams, 2001, Different modes of hippocampal plasticity in response to estrogen in young and aged female rats, Proc. Natl. Acad. Sci. USA, 98, 8071, 10.1073/pnas.141215898 Adkins-Regan, 1990, Sexual dimorphism in the avian brain is not limited to the song system of songbirds: a morphometric analysis of the brain of the quail (Coturnix japonica), Brain Res., 514, 320, 10.1016/0006-8993(90)91427-I Aguirre, 2009, Progesterone reverses 17beta-estradiol-mediated neuroprotection and BDNF induction in cultured hippocampal slices, Eur. J. Neurosci., 10.1111/j.1460-9568.2008.06591.x Ahlbom, 1999, Androgen treatment of neonatal rats decreases susceptibility of cerebellar granule neurons to oxidative stress in vitro, Eur. J. Neurosci., 11, 1285, 10.1046/j.1460-9568.1999.00529.x Akwa, 1991, Neurosteroids: biosynthesis, metabolism and function of pregnenolone and dehydroepiandrosterone in the brain, J. Steroid Biochem. Mol. Biol., 40, 71, 10.1016/0960-0760(91)90169-6 Alvarez-Buylla, 1997, Birth, migration, incorporation, and death of vocal control neurons in adult songbirds, J. Neurobiol., 33, 585, 10.1002/(SICI)1097-4695(19971105)33:5<585::AID-NEU7>3.0.CO;2-0 Alvarez-Buylla, 2001, A unified hypothesis on the lineage of neural stem cells, Nat. Rev. Neurosci., 2, 287, 10.1038/35067582 Arai, 1994, Androgen enhances neuronal degeneration in the developing preoptic area: apoptosis in the anteroventral periventricular nucleus (AVPvN-POA), Horm. Behav., 28, 313, 10.1006/hbeh.1994.1027 Arnold, 1980, Effects of androgens on volumes of sexually dimorphic brain regions in the zebra finch, Brain Res., 185, 441, 10.1016/0006-8993(80)91083-5 Arvin, 1994, Effect of the non-NMDA receptor antagonist GYKI 52466 on the microdialysate and tissue concentrations of amino acids following transient forebrain ischaemia, J. Neurochem., 62, 1458, 10.1046/j.1471-4159.1994.62041458.x Azcoitia, 2001, Astroglia play a key role in the neuroprotective actions of estrogen, Prog. Brain. Res., 132, 469, 10.1016/S0079-6123(01)32096-4 Azcoitia, 2001, Brain aromatase is neuroprotective, J. Neurobiol., 47, 318, 10.1002/neu.1038 Azcoitia, 2003, Aromatase expression by reactive astroglia is neuroprotective, Ann. NY Acad. Sci., 1007, 298, 10.1196/annals.1286.028 Azcoitia, 2005, Brain steroidogenesis: emerging therapeutic strategies to prevent neurodegeneration, J. Neural Transm., 112, 171, 10.1007/s00702-004-0179-y Balthazart, 1996, Effects of testosterone and its metabolites on aromatase-immunoreactive cells in the quail brain: relationship with the activation of male reproductive behavior, J. Steroid Biochem. Mol. Biol., 56, 185, 10.1016/0960-0760(95)00236-7 Balthazart, 2006, Rapid control of brain aromatase activity by glutamatergic inputs, Endocrinology, 147, 359, 10.1210/en.2005-0845 Barreto, 2007, Testosterone decreases reactive astroglia and reactive microglia after brain injury in male rats: role of its metabolites, oestradiol and dihydrotestosterone, Eur. J. Neurosci., 25, 3039, 10.1111/j.1460-9568.2007.05563.x Barron, 2006, Reproductive hormones modulate oxidative stress in Alzheimer’s disease, Antioxid. Redox Signal., 8, 2047, 10.1089/ars.2006.8.2047 Bate, 2007, Squalestatin protects neurons and reduces the activation of cytoplasmic phospholipase A2 by Abeta(1-42), Neuropharmacology, 53, 222, 10.1016/j.neuropharm.2007.05.003 Bate, 2007, Cholesterol synthesis inhibitors protect against platelet-activating factor-induced neuronal damage, J. Neuroinflammation, 4, 5, 10.1186/1742-2094-4-5 Bate, 2008, Sequestration of free cholesterol in cell membranes by prions correlates with cytoplasmic phospholipase A2 activation, BMC Biol., 6, 8, 10.1186/1741-7007-6-8 Bate, 2008, Cholesterol esterification reduces the neurotoxicity of prions, Neuropharmacology, 54, 1247, 10.1016/j.neuropharm.2008.04.002 Behl, 2002, Antioxidant neuroprotection in Alzheimer’s disease as preventive and therapeutic approach, Free Radic. Biol. Med., 33, 182, 10.1016/S0891-5849(02)00883-3 Behl, 1997, Neuroprotection against oxidative stress by estrogens: structure–activity relationship, Mol. Pharmacol., 51, 535, 10.1124/mol.51.4.535 Belcredito, 2001, Estrogen neuroprotection: the involvement of the Bcl-2 binding protein BNIP2, Brain Res. Brain Res. Rev., 37, 335, 10.1016/S0165-0173(01)00138-2 Berger, 2008, Neuroprotection by pravastatin in acute ischemic stroke in rats, Brain. Res. Rev., 58, 48, 10.1016/j.brainresrev.2007.10.010 Bethea, 1996, Steroid regulation of estrogen and progestin receptor messenger ribonucleic acid in monkey hypothalamus and pituitary, Endocrinology, 137, 4372, 10.1210/en.137.10.4372 Bialek, 2004, Neuroprotective role of testosterone in the nervous system, Pol. J. Pharmacol., 56, 509 Bittman, 1990, Effects of day length on sheep neuroendocrine estrogen and progestin receptors, Am. J. Physiol., 258, R135 Black, 2005, Socially induced and rapid increases in aggression are inversely related to brain aromatase activity in a sex-changing fish, Lythrypnus dalli, Proc. Biol. Sci., 272, 2435, 10.1098/rspb.2005.3210 Bologa, 1987, Dehydroepiandrosterone and its sulfated derivative reduce neuronal death and enhance astrocytic differentiation in brain cell cultures, J. Neurosci. Res., 17, 225, 10.1002/jnr.490170305 Boruban, 2008, From endometrial hyperplasia to endometrial cancer: insight into the biology and possible medical preventive measures, Eur. J. Cancer Prev., 17, 133, 10.1097/CEJ.0b013e32811080ce Brailoiu, 2007, Distribution and characterization of estrogen receptor G protein-coupled receptor 30 in the rat central nervous system, J. Endocrinol., 193, 311, 10.1677/JOE-07-0017 Brann, 2007, Neurotrophic and neuroprotective actions of estrogen: basic mechanisms and clinical implications, Steroids, 72, 381, 10.1016/j.steroids.2007.02.003 Brannvall, 2002, Estrogen-receptor-dependent regulation of neural stem cell proliferation and differentiation, Mol. Cell. Neurosci., 21, 512, 10.1006/mcne.2002.1194 Brown, 1993, Neurogenesis in adult canary telencephalon is independent of gonadal hormone levels, J. Neurosci., 13, 2024, 10.1523/JNEUROSCI.13-05-02024.1993 Bryant, 2006, Multiple pathways transmit neuroprotective effects of gonadal steroids, Endocrine, 29, 199, 10.1385/ENDO:29:2:199 Bulun, 2008, Aromatase expression in women’s cancers, Adv. Exp. Med. Biol., 630, 112, 10.1007/978-0-387-78818-0_8 Cai, 2008, Two different molecular mechanisms underlying progesterone neuroprotection against ischemic brain damage, Neuropharmacology, 55, 127, 10.1016/j.neuropharm.2008.04.023 Callard, 1983, Identification of androgen metabolic pathways in the brain of little brown bats (Myotis lucifugus): sex and seasonal differences, Biol. Reprod., 28, 1155, 10.1095/biolreprod28.5.1155 Callard, 1990, Aromatization and estrogen action in brain, Prog. Clin. Biol. Res., 342, 105 Callard, 1990, Nonmammalian vertebrate models in studies of brain–steroid interactions, J. Exp. Zool., 4, 6, 10.1002/jez.1402560404 Callier, 2001, Neuroprotective properties of 17beta-estradiol, progesterone, and raloxifene in MPTP C57Bl/6 mice, Synapse, 41, 131, 10.1002/syn.1067 Canoine, 2006, Low sex steroids, high steroid receptors: increasing the sensitivity of the nonreproductive brain, J. Neurobiol., 67, 57, 10.1002/neu.20296 Cardona-Gomez, 2000, Estradiol and progesterone regulate the expression of insulin-like growth factor-I receptor and insulin-like growth factor binding protein-2 in the hypothalamus of adult female rats, J. Neurobiol., 43, 269, 10.1002/(SICI)1097-4695(20000605)43:3<269::AID-NEU5>3.0.CO;2-D Cardona-Gomez, 2001, Interactions of estrogens and insulin-like growth factor-I in the brain: implications for neuroprotection, Brain Res. Brain Res. Rev., 37, 320, 10.1016/S0165-0173(01)00137-0 Cardounel, 1999, Dehydroepiandrosterone protects hippocampal neurons against neurotoxin-induced cell death: mechanism of action, Proc. Soc. Exp. Biol. Med., 222, 145, 10.1046/j.1525-1373.1999.d01-124.x Carmeci, 1997, Identification of a gene (GPR30) with homology to the G-protein-coupled receptor superfamily associated with estrogen receptor expression in breast cancer, Genomics, 45, 607, 10.1006/geno.1997.4972 Carreau, 2007, Estrogens: a new player in spermatogenesis, Folia Histochem. Cytobiol., 45, S5 Carroll, 2008, Progesterone blocks estrogen neuroprotection from kainate in middle-aged female rats, Neurosci. Lett., 445, 229, 10.1016/j.neulet.2008.09.010 Carswell, 2005, Brain aromatase expression after experimental stroke: topography and time course, J. Steroid Biochem. Mol. Biol., 96, 89, 10.1016/j.jsbmb.2005.02.016 Cartier, 2005, Chemokine receptors in the central nervous system: role in brain inflammation and neurodegenerative diseases, Brain Res. Brain Res. Rev., 48, 16, 10.1016/j.brainresrev.2004.07.021 Cepa, 2008, Molecular mechanisms of aromatase inhibition by new A, D-ring modified steroids, Biol. Chem., 389, 1183, 10.1515/BC.2008.134 Cepa, 2008, Synthesis and biochemical studies of 17-substituted androst-3-enes and 3,4-epoxyandrostanes as aromatase inhibitors, Steroids, 73, 1409, 10.1016/j.steroids.2008.07.001 Charalampopoulos, 2006, G protein-associated specific membrane binding sites mediate the neuroprotective effect of dehydroepiandrosterone, Faseb J., 20, 577, 10.1096/fj.05-5078fje Charalampopoulos, 2008, Neurosteroids as modulators of neurogenesis and neuronal survival, Trends Endocrinol. Metab., 19, 300, 10.1016/j.tem.2008.07.004 Charlier, 2005, Inhibition of steroid receptor coactivator-1 blocks estrogen and androgen action on male sex behavior and associated brain plasticity, J. Neurosci., 25, 906, 10.1523/JNEUROSCI.3533-04.2005 Chisu, 2006, Effects of testosterone on differentiation and oxidative stress resistance in C1300 neuroblastoma cells, Neuro Endocrinol. Lett., 27, 807 Chiueh, 2003, Induction of antioxidative and antiapoptotic thioredoxin supports neuroprotective hypothesis of estrogen, Endocrine, 21, 27, 10.1385/ENDO:21:1:27 Choi, 2003, Estradiol increases pre- and post-synaptic proteins in the CA1 region of the hippocampus in female rhesus macaques (Macaca mulatta), Endocrinology, 144, 4734, 10.1210/en.2003-0216 Cimarosti, 2005, Neuroprotection and protein damage prevention by estradiol replacement in rat hippocampal slices exposed to oxygen-glucose deprivation, Neurochem. Res., 30, 583, 10.1007/s11064-005-2693-1 Clyne, 2004, Regulation of aromatase expression by the nuclear receptor LRH-1 in adipose tissue, Mol. Cell. Endocrinol., 215, 39, 10.1016/j.mce.2003.11.001 Combarros, 2005, Interaction between CYP19 aromatase and butyrylcholinesterase genes increases Alzheimer’s disease risk, Dement. Geriatr. Cogn. Disord., 20, 153, 10.1159/000087065 Combarros, 2008, Aromatase and interleukin-10 genetic variants interactively modulate Alzheimer’s disease risk, J. Neural. Transm., 115, 863, 10.1007/s00702-008-0028-5 Corasaniti, 2005, 17beta-estradiol reduces neuronal apoptosis induced by HIV-1 gp120 in the neocortex of rat, Neurotoxicology, 26, 893, 10.1016/j.neuro.2005.01.019 Culmsee, 1999, Neuroprotection by estrogens in a mouse model of focal cerebral ischemia and in cultured neurons: evidence for a receptor-independent antioxidative mechanism, J. Cereb. Blood Flow Metab., 19, 1263, 10.1097/00004647-199911000-00011 D’Astous, 2003, Dehydroepiandrosterone (DHEA) such as 17beta-estradiol prevents MPTP-induced dopamine depletion in mice, Synapse, 47, 10, 10.1002/syn.10145 D’Astous, 2006, Implication of the phosphatidylinositol-3 kinase/protein kinase B signaling pathway in the neuroprotective effect of estradiol in the striatum of 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine mice, Mol. Pharmacol., 69, 1492, 10.1124/mol.105.018671 De Nicola, 2006, Progesterone treatment of spinal cord injury: effects on receptors, neurotrophins, and myelination, J. Mol. Neurosci., 28, 3, 10.1385/JMN:28:1:3 Desjardins, 1995, Estrogen-induced hypothalamic beta-endorphin neuron loss: a possible model of hypothalamic aging, Exp. Gerontol., 30, 253, 10.1016/0531-5565(94)00040-A Dhandapani, 2002, Protective effects of estrogen and selective estrogen receptor modulators in the brain, Biol. Reprod., 67, 1379, 10.1095/biolreprod.102.003848 Dhandapani, 2005, Astrocyte-derived transforming growth factor-{beta} mediates the neuroprotective effects of 17{beta}-estradiol: involvement of nonclassical genomic signaling pathways, Endocrinology, 146, 2749, 10.1210/en.2005-0014 di Michele, 2000, Increased neurosteroids synthesis after brain and spinal cord injury in rats, Neurosci. Lett., 284, 65, 10.1016/S0304-3940(00)00965-4 Dieudonne, 2004, Evidence for functional estrogen receptors alpha and beta in human adipose cells: regional specificities and regulation by estrogens, Am. J. Physiol. Cell. Physiol., 286, C655, 10.1152/ajpcell.00321.2003 Dubal, 1999, Estradiol modulates bcl-2 in cerebral ischemia: a potential role for estrogen receptors, J. Neurosci., 19, 6385, 10.1523/JNEUROSCI.19-15-06385.1999 Dubal, 2001, Estrogen receptor alpha, not beta, is a critical link in estradiol-mediated protection against brain injury, Proc. Natl. Acad. Sci. USA, 98, 1952, 10.1073/pnas.041483198 Dubal, 2006, Differential modulation of estrogen receptors (ERs) in ischemic brain injury: a role for ERalpha in estradiol-mediated protection against delayed cell death, Endocrinology, 147, 3076, 10.1210/en.2005-1177 Fester, 2009, Cholesterol-promoted synaptogenesis requires the conversion of cholesterol to estradiol in the hippocampus, Hippocampus, 10.1002/hipo.20548 Filardo, 2000, Estrogen-induced activation of Erk-1 and Erk-2 requires the G protein-coupled receptor homolog, GPR30, and occurs via trans-activation of the epidermal growth factor receptor through release of HB-EGF, Mol. Endocrinol., 14, 1649, 10.1210/me.14.10.1649 Filardo, 2002, Estrogen action via the G protein-coupled receptor, GPR30: stimulation of adenylyl cyclase and cAMP-mediated attenuation of the epidermal growth factor receptor-to-MAPK signaling axis, Mol. Endocrinol., 16, 70, 10.1210/me.16.1.70 Fitzpatrick, 2002, Estrogen-mediated neuroprotection against beta-amyloid toxicity requires expression of estrogen receptor alpha or beta and activation of the MAPK pathway, J. Neurochem., 82, 674, 10.1046/j.1471-4159.2002.01000.x Folkestad, 2009, The effect of aromatase inhibitors on bone metabolism, Basic Clin. Pharmacol. Toxicol., 104, 3, 10.1111/j.1742-7843.2008.00337.x Forlano, 2001, Anatomical distribution and cellular basis for high levels of aromatase activity in the brain of teleost fish: aromatase enzyme and mRNA expression identify glia as source, J. Neurosci., 21, 8943, 10.1523/JNEUROSCI.21-22-08943.2001 Forlano, 2006, Brain aromatase: new lessons from non-mammalian model systems, Front. Neuroendocrinol., 10.1016/j.yfrne.2006.05.002 Fowler, 2003, Newly proliferated cells in the adult male amygdala are affected by gonadal steroid hormones, J. Neurobiol., 57, 257, 10.1002/neu.10273 Freking, 1998, Regulation of aromatase, 5 alpha- and 5 beta-reductase in primary cell cultures of developing zebra finch telencephalon, J. Neurobiol., 36, 30, 10.1002/(SICI)1097-4695(199807)36:1<30::AID-NEU3>3.0.CO;2-B Funakoshi, 2006, G protein-coupled receptor 30 is an estrogen receptor in the plasma membrane, Biochem. Biophys. Res. Commun., 346, 904, 10.1016/j.bbrc.2006.05.191 Fusani, 2000, Seasonal expression of androgen receptors, estrogen receptors, and aromatase in the canary brain in relation to circulating androgens and estrogens, J. Neurobiol., 43, 254, 10.1002/(SICI)1097-4695(20000605)43:3<254::AID-NEU4>3.0.CO;2-W Gahr, 1997, Distribution and dynamics in the expression of androgen and estrogen receptors in vocal control systems of songbirds, Brain Res. Bull., 44, 509, 10.1016/S0361-9230(97)00233-5 Gao, 2002, The roles of sex steroid receptor coregulators in cancer, Mol. Cancer, 1, 7, 10.1186/1476-4598-1-7 Garcia-Ovejero, 2002, Glial expression of estrogen and androgen receptors after rat brain injury, J. Comp. Neurol., 450, 256, 10.1002/cne.10325 Garcia-Ovejero, 2005, Glia-neuron crosstalk in the neuroprotective mechanisms of sex steroid hormones, Brain Res. Brain Res. Rev., 48, 273, 10.1016/j.brainresrev.2004.12.018 Garcia-Segura, 2006, Steroids and glial cell function, Glia, 54, 485, 10.1002/glia.20404 Garcia-Segura, 1999, Aromatase expression by astrocytes after brain injury: implications for local estrogen formation in brain repair, Neuroscience, 89, 567, 10.1016/S0306-4522(98)00340-6 Garcia-Segura, 2001, Neuroprotection by estradiol, Prog. Neurobiol., 63, 29, 10.1016/S0301-0082(00)00025-3 Garcia-Segura, 2006, Cross-talk between IGF-I and estradiol in the brain: focus on neuroprotection, Neuroendocrinology, 84, 275, 10.1159/000097485 Garcia-Segura, 2007, Estradiol insulin-like growth factor-I and brain aging, Psychoneuroendocrinology, 32, S57, 10.1016/j.psyneuen.2007.03.001 Gelinas, 1997, Immunolocalization of aromatase- and androgen receptor-positive neurons in the goldfish brain, Gen. Comp. Endocrinol., 106, 155, 10.1006/gcen.1997.6891 Gelinas, 2004, Alpha and beta estradiol protect neuronal but not native PC12 cells from paraquat-induced oxidative stress, Neurotox. Res., 6, 141, 10.1007/BF03033216 Gibson, 2005, Progesterone suppresses the inflammatory response and nitric oxide synthase-2 expression following cerebral ischemia, Exp. Neurol., 193, 522, 10.1016/j.expneurol.2005.01.009 Goldman, 1983, Neuronal production, migration and differentiation in a vocal control nucleus of the adult female canary brain, Proc. Natl. Acad. Sci. USA, 80, 2390, 10.1073/pnas.80.8.2390 Gorski, 1980, Evidence for the existence of a sexually dimorphic nucleus in the preoptic area of the rat, J. Comp. Neurol., 193, 529, 10.1002/cne.901930214 Gould, 1990, Gonadal steroids regulate dendritic spine density in hippocampal pyramidal cells in adulthood, J. Neurosci., 10, 1286, 10.1523/JNEUROSCI.10-04-01286.1990 Green, 1996, Estradiol protects against beta-amyloid (25-35)-induced toxicity in SK-N-SH human neuroblastoma cells, Neurosci. Lett., 218, 165, 10.1016/S0304-3940(96)13148-7 Green, 1997, 17 alpha-estradiol exerts neuroprotective effects on SK-N-SH cells, J. Neurosci., 17, 511, 10.1523/JNEUROSCI.17-02-00511.1997 Green, 2000, Neuroprotective effects of phenolic A ring oestrogens, Novart. Found. Symp., 230, 202, 10.1002/0470870818.ch15 Guerra, 2004, Plasma membrane oestrogen receptor mediates neuroprotection against beta-amyloid toxicity through activation of Raf-1/MEK/ERK cascade in septal-derived cholinergic SN56 cells, J. Neurochem., 91, 99, 10.1111/j.1471-4159.2004.02695.x Gursoy, 2001, Pregnenolone protects mouse hippocampal (HT-22) cells against glutamate and amyloid beta protein toxicity, Neurochem. Res., 26, 15, 10.1023/A:1007668213330 Hammond, 2001, Testosterone-mediated neuroprotection through the androgen receptor in human primary neurons, J. Neurochem., 77, 1319, 10.1046/j.1471-4159.2001.00345.x Hao, 2003, Estrogen increases the number of spinophilin-immunoreactive spines in the hippocampus of young and aged female rhesus monkeys, J. Comp. Neurol., 465, 540, 10.1002/cne.10837 Hatten, 1999, Central nervous system neuronal migration, Annu. Rev. Neurosci., 22, 511, 10.1146/annurev.neuro.22.1.511 Hatten, 1995, Mechanisms of neural patterning and specification in the developing cerebellum, Annu. Rev. Neurosci., 18, 385, 10.1146/annurev.ne.18.030195.002125 He, 2006, Lovastatin modulates increased cholesterol and oxysterol levels and has a neuroprotective effect on rat hippocampal neurons after kainate injury, J. Neuropathol. Exp. Neurol., 65, 652, 10.1097/01.jnen.0000225906.82428.69 Hill, 2009, Estrogen deficiency results in apoptosis in the frontal cortex of adult female aromatase knockout mice, Mol. Cell. Neurosci., 10.1016/j.mcn.2008.12.009 Honda, 1994, Novel exon 1 of the aromatase gene specific for aromatase transcripts in human brain, Biochem. Biophys. Res. Commun., 198, 1153, 10.1006/bbrc.1994.1163 Hsieh, 1996, Estradiol modulation of neuron loss in the medial division of medial preoptic nucleus in rats during aging, Gerontology, 42, 18, 10.1159/000213766 Hu, 2003, Decreased estrogen receptor-alpha expression in hippocampal neurons in relation to hyperphosphorylated tau in Alzheimer patients, Acta Neuropathol., 106, 213, 10.1007/s00401-003-0720-3 Huang, 2006, CYP19 haplotypes increase risk for Alzheimer’s disease, J. Med. Genet., 43, e42, 10.1136/jmg.2005.039461 Hurn, 2005, Adult or perinatal brain injury: does sex matter?, Stroke, 36, 193, 10.1161/01.STR.0000153064.41332.f6 Iivonen, 2004, Polymorphisms in the CYP19 gene confer increased risk for Alzheimer disease, Neurology, 62, 1170, 10.1212/01.WNL.0000118208.16939.60 Ing, 1997, Estradiol up-regulates estrogen receptor and progesterone receptor gene expression in specific ovine uterine cells, Biol. Reprod., 56, 1205, 10.1095/biolreprod56.5.1205 Ishunina, 2005, Diminished aromatase immunoreactivity in the hypothalamus, but not in the basal forebrain nuclei in Alzheimer’s disease, Neurobiol. Aging, 26, 173, 10.1016/j.neurobiolaging.2004.03.010 Jayaraman, 2008, Progesterone attenuates oestrogen neuroprotection via downregulation of oestrogen receptor expression in cultured neurones, J. Neuroendocrinol., 21, 77, 10.1111/j.1365-2826.2008.01801.x Jayaraman, 2009, Progesterone attenuates oestrogen neuroprotection via downregulation of oestrogen receptor expression in cultured neurones, J. Neuroendocrinol., 21, 77, 10.1111/j.1365-2826.2008.01801.x Jiang, 1996, Progesterone is neuroprotective after transient middle cerebral artery occlusion in male rats, Brain Res., 735, 101, 10.1016/0006-8993(96)00605-1 Kamada, 2003, Changes of free cholesterol and neutral lipids after transient focal brain ischemia in rats, Acta Neurochir., 86, 177, 10.1007/978-3-7091-0651-8_38 Kaslin, 2008, Proliferation, neurogenesis and regeneration in the non-mammalian vertebrate brain, Philos. Trans. R Soc. Lond B Biol. Sci., 363, 101, 10.1098/rstb.2006.2015 Kaur, 2007, Progesterone increases brain-derived neuroptrophic factor expression and protects against glutamate toxicity in a mitogen-activated protein kinase- and phosphoinositide-3 kinase-dependent manner in cerebral cortical explants, J. Neurosci. Res., 85, 2441, 10.1002/jnr.21370 Kawashima, 1994, Role of sex steroids on the survival, neuritic outgrowth of neurons, and dopamine neurons in cultured preoptic area and hypothalamus, Horm. Behav., 28, 305, 10.1006/hbeh.1994.1026 Kawata, 1995, Roles of steroid hormones and their receptors in structural organization in the nervous system, Neurosci. Res., 24, 1, 10.1016/0168-0102(96)81278-8 Kendall, 2005, Gonadal steroids differentially modulate neurotoxicity of HIV and cocaine: testosterone and ICI 182,780 sensitive mechanism, BMC Neurosci., 6, 40, 10.1186/1471-2202-6-40 Kim, 2008, Testosterone-induced matrix metalloproteinase activation is a checkpoint for neuronal addition to the adult songbird brain, J. Neurosci., 28, 208, 10.1523/JNEUROSCI.3674-07.2008 Koide, 1986, Circulating catecholamines modulate ischemic brain damage, J. Cereb. Blood Flow Metab., 6, 559, 10.1038/jcbfm.1986.102 Kretz, 2004, Hippocampal synapses depend on hippocampal estrogen synthesis, J. Neurosci., 24, 5913, 10.1523/JNEUROSCI.5186-03.2004 Le Page, 2008, Characterization of a cis-acting element involved in cell-specific expression of the zebrafish brain aromatase gene, Mol. Reprod. Dev., 75, 1549, 10.1002/mrd.20892 Lee, 2007, Estrogen mediation of injury-induced cell birth in neuroproliferative regions of the adult zebra finch brain, Dev. Neurobiol., 67, 1107, 10.1002/dneu.20399 Leranth, 1992, Transmitter content and afferent connections of estrogen-sensitive progestin receptor-containing neurons in the primate hypothalamus, Neuroendocrinology, 55, 667, 10.1159/000126187 Leranth, 2004, Androgens increase spine synapse density in the CA1 hippocampal subfield of ovariectomized female rats, J. Neurosci., 24, 495, 10.1523/JNEUROSCI.4516-03.2004 Li, 2006, Protection of androgen against hypoxic-ischemic brain damage in neonatal rats and possible mechanisms, Zhongguo Dang Dai Er Ke Za Zhi, 8, 441 Li, 2008, Effects of androgen on the expression of brain aromatase cytopigment and nerve growth factor in neonatal rats with hypoxic-ischemic brain damage, Zhongguo Dang Dai Er Ke Za Zhi, 10, 441 Lisztwan, 2008, The aromatase inhibitor letrozole and inhibitors of insulin-like growth factor I receptor synergistically induce apoptosis in in vitro models of estrogen-dependent breast cancer, Breast Cancer Res., 10, R56, 10.1186/bcr2113 Louissaint, 2002, Coordinated interaction of neurogenesis and angiogenesis in the adult songbird brain, Neuron, 34, 945, 10.1016/S0896-6273(02)00722-5 MacLusky, 1981, Sexual differentiation of the central nervous system, Science, 211, 1294, 10.1126/science.6163211 MacLusky, 1983, Anovulation in female rats induced by neonatal administration of the catechol estrogens, 2-hydroxy-estradiol and 4-hydroxy-estradiol, Neuroendocrinology, 37, 321, 10.1159/000123569 MacLusky, 2006, Androgen effects on hippocampal CA1 spine synapse numbers are retained in Tfm male rats with defective androgen receptors, Endocrinology, 147, 2392, 10.1210/en.2005-0673 MacLusky, 2006, Androgen modulation of hippocampal synaptic plasticity, Neuroscience, 138, 957, 10.1016/j.neuroscience.2005.12.054 Maggi, 2004, Estrogens in the nervous system: mechanisms and nonreproductive functions, Annu. Rev. Physiol., 66, 291, 10.1146/annurev.physiol.66.032802.154945 Mannella, 2006, Estrogen receptor protein interaction with phosphatidylinositol 3-kinase leads to activation of phosphorylated Akt and extracellular signal-regulated kinase 1/2 in the same population of cortical neurons: a unified mechanism of estrogen action, J. Neurosci., 26, 9439, 10.1523/JNEUROSCI.1443-06.2006 Martucci, 1993, P450 enzymes of estrogen metabolism, Pharmacol. Ther., 57, 237, 10.1016/0163-7258(93)90057-K Marz, 2007, Statins induce differentiation and cell death in neurons and astroglia, Glia, 55, 1, 10.1002/glia.20422 Mayo, 2001, Pregnenolone sulfate and aging of cognitive functions: behavioral, neurochemical, and morphological investigations, Horm. Behav., 40, 215, 10.1006/hbeh.2001.1677 McClean, 2008, 17alpha-estradiol is neuroprotective in male and female rats in a model of early brain injury, Exp. Neurol., 210, 41, 10.1016/j.expneurol.2007.09.027 McCullough, 2003, Aromatase cytochrome P450 and extragonadal estrogen play a role in ischemic neuroprotection, J. Neurosci., 23, 8701, 10.1523/JNEUROSCI.23-25-08701.2003 McEwen, 2001, Tracking the estrogen receptor in neurons: implications for estrogen-induced synapse formation, Proc. Natl. Acad. Sci. USA, 98, 7093, 10.1073/pnas.121146898 McKenna, 1999, Nuclear receptor coregulators: cellular and molecular biology, Endocr. Rev., 20, 321, 10.1210/er.20.3.321 Meffre, 2007, 3beta-Hydroxysteroid dehydrogenase/5-ene-4-ene isomerase mRNA expression in rat brain: effect of pseudopregnancy and traumatic brain injury, J. Steroid Biochem. Mol. Biol., 104, 293, 10.1016/j.jsbmb.2007.03.003 Meffre, 2007, Steroid profiling in brain and plasma of male and pseudopregnant female rats after traumatic brain injury: analysis by gas chromatography/mass spectrometry, Endocrinology, 148, 2505, 10.1210/en.2006-1678 Melcangi, 2008, Neurosteroids: measurement and pathophysiologic relevance, Neurochem. Int., 52, 503, 10.1016/j.neuint.2007.09.010 Melcangi, 2008, Neuroactive steroids: state of the art and new perspectives, Cell. Mol. Life Sci., 65, 777, 10.1007/s00018-007-7403-5 Mendelson, 2007, Mechanisms in the regulation of aromatase in developing ovary and placenta, J. Steroid Biochem. Mol. Biol., 106, 62, 10.1016/j.jsbmb.2007.05.001 Menuet, 2003, Distribution of aromatase mRNA and protein in the brain and pituitary of female rainbow trout: comparison with estrogen receptor alpha, J. Comp. Neurol., 462, 180, 10.1002/cne.10726 Menuet, 2005, Expression and estrogen-dependent regulation of the zebrafish brain aromatase gene, J. Comp. Neurol., 485, 304, 10.1002/cne.20497 Micevych, 2007, Estradiol stimulates progesterone synthesis in hypothalamic astrocyte cultures, Endocrinology, 148, 782, 10.1210/en.2006-0774 Miller, 2008, Steroidogenic enzymes, Endocr. Dev., 13, 1, 10.1159/000134751 Mohamed, 2000, Effect of long-term ovariectomy and estrogen replacement on the expression of estrogen receptor gene in female rats, Eur. J. Endocrinol., 142, 307, 10.1530/eje.0.1420307 Morale, 2008, Loss of aromatase cytochrome P450 function as a risk factor for Parkinson’s disease?, Brain Res. Rev., 57, 431, 10.1016/j.brainresrev.2007.10.011 Moreau, 2009, Aromatase expression in the normal human adult adrenal and in adrenocortical tumors: biochemical, immunohistochemical, and molecular studies, Eur. J. Endocrinol., 160, 93, 10.1530/EJE-08-0215 Morissette, 2008, Contribution of estrogen receptors alpha and beta to the effects of estradiol in the brain, J. Steroid Biochem. Mol. Biol., 108, 327, 10.1016/j.jsbmb.2007.09.011 Mouriec, 2008, Synthesis of estrogens in progenitor cells of adult fish brain: evolutive novelty or exaggeration of a more general mechanism implicating estrogens in neurogenesis?, Brain Res. Bull., 75, 274, 10.1016/j.brainresbull.2007.10.030 Naftolin, 1994, Brain aromatization of androgens, J. Reprod. Med., 39, 257 Naftolin, 2008, Sex steroids and the brain, Gynecol. Endocrinol., 24, 533, 10.1080/09513590802296104 Naftolin, 1971, Aromatization of androstenedione by the diencephalon, J. Clin. Endocrinol. Metab., 33, 368, 10.1210/jcem-33-2-368 Naftolin, 1996, Aromatase immunoreactivity in axon terminals of the vertebrate brain. An immunocytochemical study on quail, rat, monkey and human tissues, Neuroendocrinology, 63, 149, 10.1159/000126951 Nguyen, 2005, Androgens activate mitogen-activated protein kinase signaling: role in neuroprotection, J. Neurochem., 94, 1639, 10.1111/j.1471-4159.2005.03318.x Nichols, 1993, GFAP mRNA increases with age in rat and human brain, Neurobiol. Aging, 14, 421, 10.1016/0197-4580(93)90100-P Nilsson, 2001, Mechanisms of estrogen action, Physiol. Rev., 81, 1535, 10.1152/physrev.2001.81.4.1535 Obara, 2008, Regulation of pH in the mammalian central nervous system under normal and pathological conditions: facts and hypotheses, Neurochem. Int., 52, 905, 10.1016/j.neuint.2007.10.015 Ong, 2003, Increase in cholesterol and cholesterol oxidation products, and role of cholesterol oxidation products in kainate-induced neuronal injury, Brain Pathol., 13, 250, 10.1111/j.1750-3639.2003.tb00026.x Osawa, 1993, Multiple functions of aromatase and the active site structure; aromatase is the placental estrogen 2-hydroxylase, J. Steroid Biochem. Mol. Biol., 44, 469, 10.1016/0960-0760(93)90252-R Pekny, 2005, Astrocyte activation and reactive gliosis, Glia, 50, 427, 10.1002/glia.20207 Pellegrini, 2007, Identification of aromatase-positive radial glial cells as progenitor cells in the ventricular layer of the forebrain in zebrafish, J. Comp. Neurol., 501, 150, 10.1002/cne.21222 Peterson, 2001, Rapid upregulation of aromatase mRNA and protein following neural injury in the zebra finch (Taeniopygia guttata), J. Neuroendocrinol., 13, 317, 10.1046/j.1365-2826.2001.00647.x Peterson, 2004, Radial glia express aromatase in the injured zebra finch brain, J. Comp. Neurol., 475, 261, 10.1002/cne.20157 Phoenix, 1959, Organizing action of prenatally administered testosterone propionate on the tissues mediating mating behavior in the female guinea pig, Endocrinology, 65, 369, 10.1210/endo-65-3-369 Picazo, 2003, Neuroprotective and neurotoxic effects of estrogens, Brain Res., 990, 20, 10.1016/S0006-8993(03)03380-8 Prange-Kiel, 2006, Direct and indirect effects of estrogen on rat hippocampus, Neuroscience, 138, 765, 10.1016/j.neuroscience.2005.05.061 Prange-Kiel, 2006, Inhibition of hippocampal estrogen synthesis causes region-specific downregulation of synaptic protein expression in hippocampal neurons, Hippocampus, 16, 464, 10.1002/hipo.20173 Prokai, 2007, Structure-nongenomic neuroprotection relationship of estrogens and estrogen-derived compounds, Pharmacol. Ther., 114, 1, 10.1016/j.pharmthera.2007.01.006 Puglielli, 2003, Alzheimer’s disease: the cholesterol connection, Nat. Neurosci., 6, 345, 10.1038/nn0403-345 Quesada, 2004, Estrogen interacts with the IGF-1 system to protect nigrostriatal dopamine and maintain motoric behavior after 6-hydroxdopamine lesions, J. Neurosci. Res., 75, 107, 10.1002/jnr.10833 Quesada, 2008, PI3 kinase/Akt activation mediates estrogen and IGF-1 nigral DA neuronal neuroprotection against a unilateral rat model of Parkinson’s disease, Dev. Neurobiol., 68, 632, 10.1002/dneu.20609 Ramachandran, 1999, Zebra finch aromatase gene expression is regulated in the brain through an alternate promoter, Gene, 240, 209, 10.1016/S0378-1119(99)00399-6 Rasika, 1994, Testosterone increases the recruitment and/or survival of new high vocal center neurons in adult female canaries, Proc. Natl. Acad. Sci. USA, 91, 7854, 10.1073/pnas.91.17.7854 Rasika, 1999, BDNF mediates the effects of testosterone on the survival of new neurons in an adult brain, Neuron, 22, 53, 10.1016/S0896-6273(00)80678-9 Rau, 2003, Estradiol attenuates programmed cell death after stroke-like injury, J. Neurosci., 23, 11420, 10.1523/JNEUROSCI.23-36-11420.2003 Regan, 1997, Estrogens attenuate neuronal injury due to hemoglobin, chemical hypoxia, and excitatory amino acids in murine cortical cultures, Brain Res., 764, 133, 10.1016/S0006-8993(97)00437-X Reibel, 2000, Neuroprotective effects of chronic estradiol benzoate treatment on hippocampal cell loss induced by status epilepticus in the female rat, Neurosci. Lett., 281, 79, 10.1016/S0304-3940(00)00784-9 Reisert, 1987, Sex steroids promote neurite growth in mesencephalic tyrosine hydroxylase immunoreactive neurons in vitro, Int. J. Dev. Neurosci., 5, 91, 10.1016/0736-5748(87)90054-2 Repici, 2007, Neuronal death and neuroprotection: a review, Methods Mol. Biol., 399, 1, 10.1007/978-1-59745-504-6_1 Roberts, 1987, Effects of dehydroepiandrosterone and its sulfate on brain tissue in culture and on memory in mice, Brain Res., 406, 357, 10.1016/0006-8993(87)90807-9 Robertson, 2006, Physiologic progesterone reduces mitochondrial dysfunction and hippocampal cell loss after traumatic brain injury in female rats, Exp. Neurol., 197, 235, 10.1016/j.expneurol.2005.09.014 Roglio, 2007, Testosterone derivatives are neuroprotective agents in experimental diabetic neuropathy, Cell. Mol. Life Sci., 64, 1158, 10.1007/s00018-007-7002-5 Roglio, 2008, Neuroprotective effects of dihydroprogesterone and progesterone in an experimental model of nerve crush injury, Neuroscience, 155, 673, 10.1016/j.neuroscience.2008.06.034 Romero-Aleman, 2004, Radial glial cells, proliferating periventricular cells, and microglia might contribute to successful structural repair in the cerebral cortex of the lizard Gallotia galloti, Exp. Neurol., 188, 74, 10.1016/j.expneurol.2004.03.014 Roof, 2000, Gender differences in acute CNS trauma and stroke: neuroprotective effects of estrogen and progesterone, J. Neurotrauma, 17, 367, 10.1089/neu.2000.17.367 Roof, 1997, Progesterone protects against lipid peroxidation following traumatic brain injury in rats, Mol. Chem. Neuropathol., 31, 1, 10.1007/BF02815156 Rosario, 2006, Progestins inhibit the neuroprotective effects of estrogen in rat hippocampus, Brain Res., 1099, 206, 10.1016/j.brainres.2006.03.127 Roselli, 2007, Brain aromatase: roles in reproduction and neuroprotection, J. Steroid Biochem. Mol. Biol., 106, 143, 10.1016/j.jsbmb.2007.05.014 Rosen, 1994, Radial glia in the neocortex of adult rats: effects of neonatal brain injury, Brain Res. Dev. Brain Res., 82, 127, 10.1016/0165-3806(94)90155-4 Rune, 2005, Neurosteroid synthesis in the hippocampus: role in synaptic plasticity, Neuroscience, 136, 833, 10.1016/j.neuroscience.2005.03.056 Rune, 2002, Estrogen up-regulates estrogen receptor alpha and synaptophysin in slice cultures of rat hippocampus, Neuroscience, 113, 167, 10.1016/S0306-4522(02)00152-5 Russo, 2005, 17beta-estradiol protects SH-SY5Y Cells against HIV-1 gp120-induced cell death: evidence for a role of estrogen receptors, Neurotoxicology, 26, 905, 10.1016/j.neuro.2005.01.009 Saldanha, 1998, The passerine hippocampus is a site of high aromatase: inter- and intraspecies comparisons, Horm. Behav., 34, 85, 10.1006/hbeh.1998.1447 Saldanha, 2005, Estrogen provision by reactive glia decreases apoptosis in the zebra finch (Taeniopygia guttata), J. Neurobiol., 64, 192, 10.1002/neu.20147 Saleh, 2005, Estrogen synthesis in the central nucleus of the amygdala following middle cerebral artery occlusion: role in modulating neurotransmission, Neuroscience, 135, 1141, 10.1016/j.neuroscience.2005.06.061 Saleh, 2005, Estrogen in the parabrachial nucleus attenuates the sympathoexcitation following MCAO in male rats, Brain Res., 1066, 187, 10.1016/j.brainres.2005.10.062 Saredi, 2005, Effect of streptozotocin-induced diabetes on the gene expression and biological activity of 3beta-hydroxysteroid dehydrogenase in the rat spinal cord, Neuroscience, 135, 869, 10.1016/j.neuroscience.2005.06.033 Sasano, 1998, Aromatase in the human central nervous system, Clin. Endocrinol. (Oxf), 48, 325, 10.1046/j.1365-2265.1998.00390.x Sawada, 1998, Estradiol protects mesencephalic dopaminergic neurons from oxidative stress-induced neuronal death, J. Neurosci. Res., 54, 707, 10.1002/(SICI)1097-4547(19981201)54:5<707::AID-JNR16>3.0.CO;2-T Schlinger, 1991, Brain is the major site of estrogen synthesis in a male songbird, Proc. Natl. Acad. Sci. USA, 88, 4191, 10.1073/pnas.88.10.4191 Schlinger, 1992, Circulating estrogens in a male songbird originate in the brain, Proc. Natl. Acad. Sci. USA, 89, 7650, 10.1073/pnas.89.16.7650 Schlinger, 1994, Neuronal and non-neuronal aromatase in primary cultures of developing zebra finch telencephalon, J. Neurosci., 14, 7541, 10.1523/JNEUROSCI.14-12-07541.1994 Schlinger, 2001, Neurosteroids and brain sexual differentiation, Trends Neurosci., 24, 429, 10.1016/S0166-2236(00)01855-5 Schumacher, 2004, Local synthesis and dual actions of progesterone in the nervous system: neuroprotection and myelination, Growth Horm. IGF Res., 14, S18, 10.1016/j.ghir.2004.03.007 Schumacher, 2007, Progesterone: therapeutic opportunities for neuroprotection and myelin repair, Pharmacol. Ther., 116, 77, 10.1016/j.pharmthera.2007.06.001 Scott, 2000, The distribution of progesterone receptor immunoreactivity and mRNA in the preoptic area and hypothalamus of the ewe: upregulation of progesterone receptor mRNA in the mediobasal hypothalamus by oestrogen, J. Neuroendocrinol., 12, 565, 10.1046/j.1365-2826.2000.00490.x Shibata, 1997, Role of co-activators and co-repressors in the mechanism of steroid/thyroid receptor action, Recent Prog. Horm. Res., 52, 141 Shoulson, 1992, Neuroprotective clinical strategies for Parkinson’s disease, Ann. Neurol., 32, S143, 10.1002/ana.410320725 Shughrue, 1990, Developmental changes in estrogen receptors in mouse cerebral cortex between birth and postweaning: studied by autoradiography with 11 beta-methoxy-16 alpha-[125I]iodoestradiol, Endocrinology, 126, 1112, 10.1210/endo-126-2-1112 Shughrue, 1997, Responses in the brain of estrogen receptor alpha-disrupted mice, Proc. Natl. Acad. Sci. USA, 94, 11008, 10.1073/pnas.94.20.11008 Sierra, 2003, Steroidogenic acute regulatory protein in the rat brain: cellular distribution, developmental regulation and overexpression after injury, Eur. J. Neurosci., 18, 1458, 10.1046/j.1460-9568.2003.02872.x Simpkins, 2008, Mitochondrial mechanisms of estrogen neuroprotection, Brain Res. Rev., 57, 421, 10.1016/j.brainresrev.2007.04.007 Simpkins, 2005, Mitochondria play a central role in estrogen-induced neuroprotection, Curr. Drug. Targets CNS Neurol. Disord., 4, 69, 10.2174/1568007053005073 Simpson, 1994, Aromatase gene expression in adipose tissue: relationship to breast cancer, Int. J. Fertil. Menop. Stud., 39, 75 Simpson, 1994, Aromatase cytochrome P450 the enzyme responsible for estrogen biosynthesis, Endocr. Rev., 15, 342 Simpson, 2002, Aromatase – a brief overview, Annu. Rev. Physiol., 64, 93, 10.1146/annurev.physiol.64.081601.142703 Sinchak, 2003, Estrogen induces de novo progesterone synthesis in astrocytes, Dev. Neurosci., 25, 343, 10.1159/000073511 Singer, 1996, Estrogen protects primary cortical neurons from glutamate toxicity, Neurosci. Lett., 212, 13, 10.1016/0304-3940(96)12760-9 Singer, 1998, Modulation of Bcl-2 expression: a potential component of estrogen protection in NT2 neurons, Neuroreport, 9, 2565, 10.1097/00001756-199808030-00025 Soma, 1999, Androgen-metabolizing enzymes show region-specific changes across the breeding season in the brain of a wild songbird, J. Neurobiol., 41, 176, 10.1002/(SICI)1097-4695(19991105)41:2<176::AID-NEU2>3.0.CO;2-2 Soma, 2005, Neurosteroids and female reproduction: estrogen increases 3beta-HSD mRNA and activity in rat hypothalamus, Endocrinology, 146, 4386, 10.1210/en.2005-0569 Sponne, 2004, Membrane cholesterol interferes with neuronal apoptosis induced by soluble oligomers but not fibrils of amyloid-beta peptide, Faseb J., 18, 836, 10.1096/fj.03-0372fje Spritzer, 2007, Testosterone and dihydrotestosterone, but not estradiol, enhance survival of new hippocampal neurons in adult male rats, Dev. Neurobiol., 67, 1321, 10.1002/dneu.20457 Sribnick, 2004, 17beta-estradiol attenuates glutamate-induced apoptosis and preserves electrophysiologic function in primary cortical neurons, J. Neurosci. Res., 76, 688, 10.1002/jnr.20124 Stocco, 2008, Aromatase expression in the ovary: hormonal and molecular regulation, Steroids, 73, 473, 10.1016/j.steroids.2008.01.017 Sundholm-Peters, 2004, Radial glia-like cells at the base of the lateral ventricles in adult mice, J. Neurocytol., 33, 153, 10.1023/B:NEUR.0000029654.70632.3a Takagi, 1993, Culture of rat brain preoptic area neurons: effects of sex steroids, Int. J. Dev. Neurosci., 11, 63, 10.1016/0736-5748(93)90035-C Tapia-Gonzalez, 2008, Selective oestrogen receptor (ER) modulators reduce microglia reactivity in vivo after peripheral inflammation: potential role of microglial ERs, J. Endocrinol., 198, 219, 10.1677/JOE-07-0294 Teepker, 2003, 2-OH-estradiol, an endogenous hormone with neuroprotective functions, J. Psychiatr. Res., 37, 517, 10.1016/S0022-3956(03)00068-2 Tetel, 2000, Nuclear receptor coactivators in neuroendocrine function, J. Neuroendocrinol., 12, 927, 10.1046/j.1365-2826.2000.00557.x Thornton, 1989, Sex differences in cytosolic progestin receptors in microdissected regions of the hypothalamus/preoptic area of guinea pigs, Brain Res., 503, 253, 10.1016/0006-8993(89)91671-5 Toda, 1994, Expression of the gene encoding aromatase cytochrome P450 (CYP19) in fetal tissues, Mol. Endocrinol., 8, 210, 10.1210/me.8.2.210 Toran-Allerand, 2004, Minireview: a plethora of estrogen receptors in the brain: where will it end?, Endocrinology, 145, 1069, 10.1210/en.2003-1462 Toran-Allerand, 2005, Estrogen and the brain: beyond ER-alpha, ER-beta, and 17beta-estradiol, Ann. NY Acad. Sci., 1052, 136, 10.1196/annals.1347.009 Toran-Allerand, 2002, ER-X: a novel, plasma membrane-associated, putative estrogen receptor that is regulated during development and after ischemic brain injury, J. Neurosci., 22, 8391, 10.1523/JNEUROSCI.22-19-08391.2002 Toung, 1998, Estrogen-mediated neuroprotection after experimental stroke in male rats, Stroke, 29, 1666, 10.1161/01.STR.29.8.1666 VanLandingham, 2006, The enantiomer of progesterone acts as a molecular neuroprotectant after traumatic brain injury, Neuropharmacology, 51, 1078, 10.1016/j.neuropharm.2006.07.015 Veiga, 2003, Neuroprotection by the steroids pregnenolone and dehydroepiandrosterone is mediated by the enzyme aromatase, J. Neurobiol., 56, 398, 10.1002/neu.10249 Veiga, 2006, Neuroactive steroids prevent peripheral myelin alterations induced by diabetes, Neurosci. Lett., 402, 150, 10.1016/j.neulet.2006.03.058 Wakade, 2008, Tamoxifen neuroprotection in cerebral ischemia involves attenuation of kinase activation and superoxide production and potentiation of mitochondrial superoxide dismutase, Endocrinology, 149, 367, 10.1210/en.2007-0899 Wang, 2001, Morphological abnormalities in the brains of estrogen receptor beta knockout mice, Proc. Natl. Acad. Sci. USA, 98, 2792, 10.1073/pnas.041617498 Wise, 2003, Estradiol exerts neuroprotective actions against ischemic brain injury: insights derived from animal models, Endocrine, 21, 11, 10.1385/ENDO:21:1:11 Wojtal, 2006, Endogenous neuroprotective factors: neurosteroids, Pharmacol. Rep., 58, 335 Wright, 1991, Familial ALzheimer’s disease. Beta amyloid resurrected, Nature, 349, 653, 10.1038/349653a0 Wu, 2005, The orphan nuclear receptors NURR1 and NGFI-B modulate aromatase gene expression in ovarian granulosa cells: a possible mechanism for repression of aromatase expression upon luteinizing hormone surge, Endocrinology, 146, 237, 10.1210/en.2004-0889 Wu, 2008, Simvastatin-mediated upregulation of VEGF and BDNF, activation of the PI3K/Akt pathway, and increase of neurogenesis are associated with therapeutic improvement after traumatic brain injury, J. Neurotrauma, 25, 130, 10.1089/neu.2007.0369 Wynne, 2004, Glial aromatization decreases neural injury in the zebra finch (Taeniopygia guttata): influence on apoptosis, J. Neuroendocrinol., 16, 676, 10.1111/j.1365-2826.2004.01217.x Wynne, 2008, Inhibition of injury-induced glial aromatase reveals a wave of secondary degeneration in the songbird brain, Glia, 56, 97, 10.1002/glia.20594 Wynne, 2008, Molecular characterization of the injury-induced aromatase transcript in the adult zebra finch brain, J. Neurochem., 105, 1613, 10.1111/j.1471-4159.2008.05256.x Yague, 2006, Aromatase expression in the human temporal cortex, Neuroscience, 138, 389, 10.1016/j.neuroscience.2005.11.054 Yague, 2008, Selective transcriptional regulation of aromatase gene by vitamin D, dexamethasone, and mifepristone in human glioma cells, Endocrine Yanase, 2003, Aromatase in bone: roles of vitamin D3 and androgens, J. Steroid Biochem. Mol. Biol., 86, 393, 10.1016/S0960-0760(03)00349-2 Yang, 1997, A subpopulation of reactive astrocytes at the immediate site of cerebral cortical injury, Exp. Neurol., 146, 199, 10.1006/exnr.1997.6518 Yao, 2002, 22R-Hydroxycholesterol protects neuronal cells from beta-amyloid-induced cytotoxicity by binding to beta-amyloid peptide, J. Neurochem., 83, 1110, 10.1046/j.1471-4159.2002.01202.x Yokosuka, 1995, Transient expression of estrogen receptor-immunoreactivity (ER-IR) in the layer V of the developing rat cerebral cortex, Brain Res. Dev. Brain Res., 84, 99, 10.1016/0165-3806(94)00161-R Yue, 2005, Brain estrogen deficiency accelerates Abeta plaque formation in an Alzheimer’s disease animal model, Proc. Natl. Acad. Sci. USA, 102, 19198, 10.1073/pnas.0505203102 Zhang, 2004, Estrogen and androgen protection of human neurons against intracellular amyloid beta1-42 toxicity through heat shock protein 70, J. Neurosci., 24, 5315, 10.1523/JNEUROSCI.0913-04.2004 Zhao, 1997, Transcriptional regulation of CYP19 gene (aromatase) expression in adipose stromal cells in primary culture, J. Steroid Biochem. Mol. Biol., 61, 203, 10.1016/S0960-0760(97)80013-1 Zhou, 1998, Characterization of a silencer element in the human aromatase gene, Arch. Biochem. Biophys., 353, 213, 10.1006/abbi.1998.0641 Zhu, 1998, Functional role of estrogen metabolism in target cells: review and perspectives, Carcinogenesis, 19, 1, 10.1093/carcin/19.1.1 Zouboulis, 2007, Sexual hormones in human skin, Horm. Metab. Res., 39, 85, 10.1055/s-2007-961807 Zupanc, 2003, Potential role of radial glia in adult neurogenesis of teleost fish, Glia, 43, 77, 10.1002/glia.10236 Zwain, 1997, Astrocytes cultured in vitro produce estradiol-17beta and express aromatase cytochrome P-450 (P-450 AROM) mRNA, Biochim. Biophys. Acta, 1334, 338, 10.1016/S0304-4165(96)00115-8