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The FASEB Journal, Vol 6, 2311-2322, Copyright © 1992 by The Federation of American Societies for Experimental Biology

REVIEWS

Neuroactive steroids

SM Paul and RH Purdy
Section on Molecular Pharmacology, National Institute of Mental Health, Bethesda, Maryland 20892.

Neuroactive steroids are natural or synthetic steroids that rapidly alter the excitability of neurons by binding to membrane-bound receptors such as those for inhibitory and (or) excitatory neurotransmitters. The best-studied neuroactive steroids are a series of sedative-hypnotic 3 alpha-hydroxy ring A-reduced pregnane steroids that include the major metabolites of progesterone and deoxycorticosterone, 3 alpha-hydroxy-5 alpha-pregnan-20-one (allopregnanolone) and 3 alpha,21-dihydroxy-5 alpha-pregnan-20-one (allotetrahydroDOC), respectively. These 3 alpha-hydroxysteroids do not interact with classical intracellular steroid receptors but bind stereoselectively and with high affinity to receptors for the major inhibitory neurotransmitter in brain, gamma-amino-butyric acid (GABA). Biochemical and electrophysiological studies have shown that these steroids markedly augment GABA-activated chloride ion currents in a manner similar (but not identical) to that of anesthetic barbiturates. Several steroids have also been observed to have convulsant or proconvulsant properties, including the synthetic amidine 3 alpha- hydroxy-16-imino-5 beta-17-azaandrostan-11-one (RU5135) and the natural sulfate esters of pregnenolone and dehydroepiandrosterone. Several of these have been shown to be bicuculline or picrotoxin-like GABAA receptor antagonists. Examples of steroids that alter neuronal excitability rapidly by augmenting or inhibiting excitatory amino acid receptor-mediated responses have also been reported. Recently, allopregnanolone and allotetrahydroDOC have also been measured in brain and plasma where their levels have been shown to fluctuate in response to stress and during the estrous and menstrual cycles of rats and humans, respectively. Although the major fraction of allopregnanolone in tissue, including brain, is of adrenal and/or ovarian origin, appreciable levels of allopregnanolone can still be measured in the brains of adrenalectomized and/or oophorectomized animals. Receptor- active neurosteroids may represent an important class of neuromodulators that can rapidly alter central nervous system excitability via novel nongenomic mechanisms.


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The neurosteroid allopregnanolone modulates oxytocin expression in the hypothalamic paraventricular nucleus
Am J Physiol Regulatory Integrative Comp Physiol, March 1, 2000; 278(3): R684 - R691.
[Abstract] [Full Text] [PDF]

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R. A. Mulnard, C. W. Cotman, C. Kawas, C. H. van Dyck, M. Sano, R. Doody, E. Koss, E. Pfeiffer, S. Jin, A. Gamst, et al.
Estrogen Replacement Therapy for Treatment of Mild to Moderate Alzheimer Disease: A Randomized Controlled Trial
JAMA, February 23, 2000; 283(8): 1007 - 1015.
[Abstract] [Full Text] [PDF]

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D. A. Finn, E. J. Gallaher, and J. C. Crabbe
Differential Change in Neuroactive Steroid Sensitivity during Ethanol Withdrawal
J. Pharmacol. Exp. Ther., January 1, 2000; 292(1): 394 - 405.
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 StrokeHome page
P. Lyden, C. Shin, C. Jackson-Friedman, S. Hassid, A. Chong, and R. L. Macdonald
Effect of Ganaxolone in a Rodent Model of Cerebral Hematoma Editorial Comment
Stroke, January 1, 2000; 31(1): 169 - 175.
[Abstract] [Full Text] [PDF]

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 Endocr. Rev.Home page
G. A. Bray and F. L. Greenway
Current and Potential Drugs for Treatment of Obesity
Endocr. Rev., December 1, 1999; 20(6): 805 - 875.
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E. J Cooper, G. A R Johnston, and F. A Edwards
Effects of a naturally occurring neurosteroid on GABAA IPSCs during development in rat hippocampal or cerebellar slices
J. Physiol., December 1, 1999; 521(2): 437 - 449.
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 Exp. Biol. Med.Home page
A. S. Hazell and R. F. Butterworth
Hepatic Encephalopathy: An Update of Pathophysiologic Mechanisms
Experimental Biology and Medicine, November 1, 1999; 222(2): 99 - 112.
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 Mol. Pharmacol.Home page
K. D. W. Morris, C. N. Moorefield, and J. Amin
Differential Modulation of the gamma -Aminobutyric Acid Type C Receptor by Neuroactive Steroids
Mol. Pharmacol., October 1, 1999; 56(4): 752 - 759.
[Abstract] [Full Text]

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 Psychosom. Med.Home page
C. N. Epperson, K. L. Wisner, and B. Yamamoto
Gonadal Steroids in the Treatment of Mood Disorders
Psychosom Med, September 1, 1999; 61(5): 676 - 697.
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A. Y. Valeyev, J. C. Hackman, A. M. Holohean, P. M. Wood, J. L. Katz, and R. A. Davidoff
Alphaxalone Activates a Cl- Conductance Independent of GABAA Receptors in Cultured Embryonic Human Dorsal Root Ganglion Neurons
J Neurophysiol, July 1, 1999; 82(1): 10 - 15.
[Abstract] [Full Text] [PDF]

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R. A Wilke, P. J Lupardus, D. K Grandy, M. Rubinstein, M. J Low, and M. B Jackson
K+ channel modulation in rodent neurohypophysial nerve terminals by sigma receptors and not by dopamine receptors
J. Physiol., June 1, 1999; 517(2): 391 - 406.
[Abstract] [Full Text] [PDF]

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H. K. Gershenfeld, P. E. Neumann, X. Li, P. L. St. Jean, and S. M. Paul
Mapping Quantitative Trait Loci for Seizure Response to a GABAA Receptor Inverse Agonist in Mice
J. Neurosci., May 15, 1999; 19(10): 3731 - 3738.
[Abstract] [Full Text] [PDF]

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 PsychosomaticsHome page
A. G. Herzog
Psychoneuroendocrine Aspects of Temporolimbic Epilepsy: Part I. Brain, Reproductive Steroids, and Emotions
Psychosomatics, April 1, 1999; 40(2): 95 - 101.
[Abstract] [Full Text]

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C. A. Bowen, R. H. Purdy, and K. A. Grant
Ethanol-Like Discriminative Stimulus Effects of Endogenous Neuroactive Steroids: Effect of Ethanol Training Dose and Dosing Procedure
J. Pharmacol. Exp. Ther., April 1, 1999; 289(1): 405 - 411.
[Abstract] [Full Text]

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 Mol. Pharmacol.Home page
S. L. Hansen, B. Fjalland, and M. B. Jackson
Differential Blockade of gamma -Aminobutyric Acid Type A Receptors by the Neuroactive Steroid Dehydroepiandrosterone Sulfate in Posterior and Intermediate Pituitary
Mol. Pharmacol., March 1, 1999; 55(3): 489 - 496.
[Abstract] [Full Text]

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E. R Lumbers and Z.-Y. Yu
A method for determining baroreflex-mediated sympathetic and parasympathetic control of the heart in pregnant and non-pregnant sheep
J. Physiol., March 1, 1999; 515(2): 555 - 566.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
K. Ukena, C. Kohchi, and K. Tsutsui
Expression and Activity of 3{beta}-Hydroxysteroid Dehydrogenase/{Delta}5-{Delta}4-Isomerase in the Rat Purkinje Neuron during Neonatal Life
Endocrinology, February 1, 1999; 140(2): 805 - 813.
[Abstract] [Full Text]

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 J. Pharmacol. Exp. Ther.Home page
T. G. Kokate, M. K. Banks, T. Magee, S.-I. Yamaguchi, and M. A. Rogawski
Finasteride, a 5alpha -Reductase Inhibitor, Blocks the Anticonvulsant Activity of Progesterone in Mice
J. Pharmacol. Exp. Ther., February 1, 1999; 288(2): 679 - 684.
[Abstract] [Full Text]

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 DevelopmentHome page
M. Freeman, A Dobritsa, P Gaines, W. Segraves, and J. Carlson
The dare gene: steroid hormone production, olfactory behavior, and neural degeneration in Drosophila
Development, January 10, 1999; 126(20): 4591 - 4602.
[Abstract] [PDF]

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 Mol. Pharmacol.Home page
K. Sabey, K. Paradiso, J. Zhang, and J. H. Steinbach
Ligand Binding and Activation of Rat Nicotinic alpha 4beta 2 Receptors Stably Expressed in HEK293 Cells
Mol. Pharmacol., January 1, 1999; 55(1): 58 - 66.
[Abstract] [Full Text]

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 Am. J. Physiol. Regul. Integr. Comp. Physiol.Home page
Y. Deng and S. Kaufman
Pregnancy-induced changes in central response to atrial distension mimicked by progesterone metabolite
Am J Physiol Regulatory Integrative Comp Physiol, December 1, 1998; 275(6): R1875 - R1877.
[Abstract] [Full Text] [PDF]

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 J. Pharmacol. Exp. Ther.Home page
T. G. Kokate, S.-I. Yamaguchi, L. K. Pannell, U. Rajamani, D. M. Carroll, A. B. Grossman, and M. A. Rogawski
Lack of Anticonvulsant Tolerance to the Neuroactive Steroid Pregnanolone in Mice
J. Pharmacol. Exp. Ther., November 1, 1998; 287(2): 553 - 558.
[Abstract] [Full Text]

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 J. Pharmacol. Exp. Ther.Home page
M. J. Millan, A. Newman-Tancredi, M. Brocco, A. Gobert, F. Lejeune, V. Audinot, J.-M. Rivet, R. Schreiber, A. Dekeyne, M. Spedding, et al.
S 18126 ({2-[4-(2,3-dihydrobenzo[1,4]dioxin-6-yl)piperazin-1-yl methyl]indan-2-yl}), a Potent, Selective and Competitive Antagonist at Dopamine D4 Receptors: An In Vitro and In Vivo Comparison with L 745,870 (3-(4-[4-chlorophenyl]piperazin-1-yl)methyl-1H-pyrrolo[2,3b]pyridine) and Raclopride
J. Pharmacol. Exp. Ther., October 1, 1998; 287(1): 167 - 186.
[Abstract] [Full Text]

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 J. Clin. Endocrinol. Metab.Home page
B. Stoffel-Wagner, M. Watzka, S. Steckelbroeck, L. Wickert, J. Schramm, G. Romalo, D. Klingmüller, and H.-U. Schweikert
Expression of 5{alpha}-Reductase in the Human Temporal Lobe of Children and Adults
J. Clin. Endocrinol. Metab., October 1, 1998; 83(10): 3636 - 3642.
[Abstract] [Full Text]

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 Mol. Endocrinol.Home page
C. H. R. Wetzel, B. Hermann, C. Behl, E. Pestel, G. Rammes, W. Zieglgänsberger, F. Holsboer, and R. Rupprecht
Functional Antagonism of Gonadal Steroids at the 5-Hydroxytryptamine Type 3 Receptor
Mol. Endocrinol., September 1, 1998; 12(9): 1441 - 1451.
[Abstract] [Full Text]

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 Am. J. PsychiatryHome page
E. Romeo, A. Ströhle, G. Spalletta, F. d. Michele, B. Hermann, F. Holsboer, A. Pasini, and R. Rupprecht
Effects of Antidepressant Treatment on Neuroactive Steroids in Major Depression
Am J Psychiatry, July 1, 1998; 155(7): 910 - 913.
[Abstract] [Full Text]

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 J. Neurol. Neurosurg. PsychiatryHome page
A. G HERZOG, A. SOTREL, and M. RONTHAL
Reversible proximal myopathy in epilepsy related Cushing's syndrome
J. Neurol. Neurosurg. Psychiatry, July 1, 1998; 65(1): 134 - 134.
[Full Text]

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 J. Pharmacol. Exp. Ther.Home page
J. C. Crabbe
Provisional Mapping of Quantitative Trait Loci for Chronic Ethanol Withdrawal Severity in BXD Recombinant Inbred Mice
J. Pharmacol. Exp. Ther., July 1, 1998; 286(1): 263 - 271.
[Abstract] [Full Text]

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 Endocr. Rev.Home page
E. R. de Kloet, E. Vreugdenhil, M. S. Oitzl, and M. Joëls
Brain Corticosteroid Receptor Balance in Health and Disease
Endocr. Rev., June 1, 1998; 19(3): 269 - 301.
[Abstract] [Full Text]

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 J. Clin. Endocrinol. Metab.Home page
A. R. Genazzani, F. Petraglia, F. Bernardi, E. Casarosa, C. Salvestroni, A. Tonetti, R. E. Nappi, S. Luisi, M. Palumbo, R. H. Purdy, et al.
Circulating Levels of Allopregnanolone in Humans: Gender, Age, and Endocrine Influences
J. Clin. Endocrinol. Metab., June 1, 1998; 83(6): 2099 - 2103.
[Abstract] [Full Text]

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 J. Pharmacol. Exp. Ther.Home page
G. E. Duncan, D. J. Knapp, S. W. Carson, and G. R. Breese
Differential Effects of Chronic Antidepressant Treatment on Swim Stress- and Fluoxetine-Induced Secretion of Corticosterone and Progesterone
J. Pharmacol. Exp. Ther., May 1, 1998; 285(2): 579 - 587.
[Abstract] [Full Text]

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 EndocrinologyHome page
A. Poletti, P. Negri-Cesi, M. Rabuffetti, A. Colciago, F. Celotti, and L. Martini
Transient Expression of the 5{alpha}-Reductase Type 2 Isozyme in the Rat Brain in Late Fetal and Early Postnatal Life
Endocrinology, April 1, 1998; 139(4): 2171 - 2178.
[Abstract] [Full Text] [PDF]

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 Am. J. Physiol. Regul. Integr. Comp. Physiol.Home page
J. D. Laiprasert, R. C. Rogers, and C. M. Heesch
Neurosteroid modulation of arterial baroreflexsensitive neurons in rat rostral ventrolateral medulla
Am J Physiol Regulatory Integrative Comp Physiol, April 1, 1998; 274(4): R903 - R911.
[Abstract] [Full Text] [PDF]

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 Proc. Natl. Acad. Sci. USAHome page
C. Cascio, V. V. K. Prasad, Y. Y. Lin, S. Lieberman, and V. Papadopoulos
Detection of P450c17-independent pathways for dehydroepiandrosterone (DHEA) biosynthesis in brain glial tumor cells
PNAS, March 17, 1998; 95(6): 2862 - 2867.
[Abstract] [Full Text] [PDF]


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