Gunn et al. Carboni, E., Wieland, S., Lan, N. C., and Gee, K. W. (1996). Anxiolytic properties of endogenously occur- ring pregnanediols in two rodent models of anxiety. Psychopharmacol- ogy (Berl.) 126, 173–178. Carta, M., Mameli, M., and Valen- zuela, C. F. (2004). Alcohol enhances GABAergic transmission to cerebel- lar granule cells via an increase in golgi cell excitability. J. Neurosci. 24, 3746–3751. Chisari, M., Eisenman, L. N., Covey, D. F., Mennerick, S., and Zorumski, C. F. (2010). The sticky issue of neu- rosteroids and GABAA receptors. Trends Neurosci. 33, 299–306. Cole, R. L., and Sawchenko, P. E. (2002). Neurotransmitter regulation of cel- lular activation and neuropeptide gene expression in the paraventric- ular nucleus of the hypothalamus. J. Neurosci. 22, 959–969. Cooper, E. J., Johnston, G. A., and Edwards, F. A. (1999). Effects of a naturally occurring neurosteroid on GABAA IPSCs during develop- ment in rat hippocampal or cere- bellar slices. J. Physiol. 521(Pt 2), 437–449. Corpéchot, C., Young, J., Calvel, M., Wehrey, C., Veltz, J. N., Touyer, G., Mouren, M., Prasad, V. V., Banner, C., and Sjovall, J. (1993). Neuros- teroids: 3 alpha-hydroxy-5 alpha- pregnan-20-one and its precursors in the brain, plasma, and steroido- genic glands of male and female rats. Endocrinology 133, 1003–1009. Crawley, J. N., Glowa, J. R., Majew- ska, M. D., and Paul, S. M. (1986). Anxiolytic activity of an endoge- nous adrenal steroid. Brain Res. 398, 382–385. Crossley, K. J., Walker, D. W., Beart, P. M., and Hirst, J. J. (2000). Char- acterisation of GABAA receptors in fetal, neonatal and adult ovine brain: region and age related changes and the effects of allopregnanolone. Neu- ropharmacology 39, 1514–1522. Cullinan, W. E. (2000). GABAA receptor subunit expression within hypophysiotropic CRH neurons: a dual hybridization histochem- ical study. J. Comp. Neurol. 419, 344–351. Cullinan, W. E., and Wolfe, T. J. (2000). Chronic stress regulates levels of mRNA transcripts encoding beta subunits of the GABAA receptor in the rat stress axis. Brain Res. 887, 118–124. Cullinan, W. E., Ziegler, D. R., and Her- man, J. P. (2008). Functional role of local GABAergic inﬂuences on the HPA axis. Brain Struct. Funct. 213, 63–72. www.frontiersin.org GABAA receptors, neurosteroids, and stress Cunningham, E. T. Jr., and Sawchenko, P. E. (1988). Anatomical speciﬁcity of noradrenergic inputs to the par- aventricular and supraoptic nuclei of the rat hypothalamus. J. Comp. Neurol. 274, 60–76. Curley, J. P., Jensen, C. L., Mashoodh, R., and Champagne, F. A. (2011). Social inﬂuences on neurobiology and behavior: epigenetic effects dur- ing development. Psychoneuroen- docrinology 36, 352–371. Daftary, S. S., Boudaba, C., and Tasker, J. G. (2000). Noradrenergic reg- ulation of parvocellular neurons in the rat hypothalamic paraven- tricular nucleus. Neuroscience 96, 743–751. de Kloet, E. R., Joels, M., and Hols- boer, F. (2005). Stress and the brain: from adaptation to disease. Nat. Rev. Neurosci. 6, 463–475. Di, S., Malcher-Lopes, R., Halmos, K. C., and Tasker, J. G. (2003). Nongenomic glucocorticoid inhibi- tion via endocannabinoid release in the hypothalamus: a fast feed- back mechanism. J. Neurosci. 23, 4850–4857. Di, S., Malcher-Lopes, R., Marcheselli, V. L., Bazan, N. G., and Tasker, J. G. (2005). Rapid glucocorticoid- mediated endocannabinoid release and opposing regulation of gluta- mate and gamma-aminobutyric acid inputs to hypothalamic magnocel- lular neurons. Endocrinology 146, 4292–4301. Di, S., Maxson, M. M., Franco, A., and Tasker, J. G. (2009). Glucocorti- coids regulate glutamate and GABA synapse-speciﬁc retrograde trans- mission via divergent nongenomic signaling pathways. J. Neurosci. 29, 393–401. Do Rego, J. L., Seong, J. Y., Burel, D., Leprince, J., Luu-The, V., Tsut- sui, K., Tonon, M. C., Pelletier, G., and Vaudry, H. (2009). Neuros- teroid biosynthesis: enzymatic path- ways and neuroendocrine regula- tion by neurotransmitters and neu- ropeptides. Front. Neuroendocrinol. 30, 259–301. Dzhala, V. I., Talos, D. M., Sdrulla, D. A., Brumback, A. C., Mathews, G. C., Benke, T. A., Delpire, E., Jensen, F. E., and Staley, K. J. (2005). NKCC1 transporter facilitates seizures in the developing brain. Nat. Med. 11, 1205–1213. Eechaute, W. P., Dhooge, W. S., Gao, C. Q., Calders, P., Rubens, R., Weyne, J., and Kaufman, J. M. (1999). Progesterone-transforming enzyme activity in the hypothalamus of the male rat. J. Steroid Biochem. Mol. Biol. 70, 159–167. Ehrlich, I., Humeau, Y., Grenier, F., Cioc- chi, S., Herry, C., and Luthi, A. (2009). Amygdala inhibitory circuits and the control of fear memory. Neuron 62, 757–771. Engelmann, M., Landgraf, R., and Wot- jak, C. T. (2004). The hypothalamic- neurohypophysial system regulates the hypothalamic-pituitary-adrenal axis under stress: an old concept revisited. Front. Neuroendocrinol. 25, 132–149. Enoch, M. A., Hodgkinson, C. A., Yuan, Q., Shen, P. H., Goldman, D., and Roy, A. (2010). The inﬂuence of GABRA2, childhood trauma, and their interaction on alcohol, heroin, and cocaine dependence. Biol. Psy- chiatry 67, 20–27. Eser, D., di Michele, F., Zwanzger, P., Pasini, A., Baghai, T. C., Schüle, C., Rupprecht, R., and Romeo, E. (2005). Panic induction with cholecystokinin-tetrapeptide (CCK- 4) Increases plasma concentrations of the neuroactive steroid 3α,5α tetrahydrodeoxycorticosterone (3α, 5α-THDOC) in healthy volun- teers. Neuropsychopharmacology 30, 192–195. Eser, D., Romeo, E., Baghai, T. C., di Michele, F., Schule, C., Pasini, A., Zwanzger, P., Padberg, F., and Rupprecht, R. (2006). Neuroactive steroids as modulators of depres- sion and anxiety. Neuroscience 138, 1041–1048. Fancsik, A., Linn, D. M., and Tasker, J. G. (2000). Neurosteroid modu- lation of GABA IPSCs is phospho- rylation dependent. J. Neurosci. 20, 3067–3075. Farrant, M., and Nusser, Z. (2005). Vari- ations on an inhibitory theme: pha- sic and tonic activation of GABAA receptors. Nat. Rev. Neurosci. 6, 215–229. Fenoglio, K. A., Brunson, K. L., Avishai- Eliner, S., Stone, B. A., Kapadia, B. J., and Baram, T. Z. (2005). Endur- ing, handling-evoked enhancement of hippocampal memory func- tion and glucocorticoid receptor expression involves activation of the corticotropin-releasing factor type 1 receptor. Endocrinology 146, 4090–4096. Flak, J. N., Ostrander, M. M., Tasker, J. G., and Herman, J. P. (2009). Chronic stress-induced neu- rotransmitter plasticity in the PVN. J. Comp. Neurol. 517, 156–165. Ford, M. M., Nickel, J. D., Phillips, T. J., and Finn, D. A. (2005). Neuros- teroid modulators of GABAA recep- tors differentially modulate ethanol intake patterns in male C57BL/6J mice. Alcohol. Clin. Exp. Res. 29, 1630–1640. Francis, D., Diorio, J., Liu, D., and Meaney, M. J. (1999). Nonge- nomic transmission across genera- tions of maternal behavior and stress responses in the rat. Science 286, 1155–1158. Francis, D. D., and Meaney, M. J. (1999). Maternal care and the development of stress responses. Curr. Opin. Neu- robiol. 9, 128–134. Fritschy, J. M., and Brunig, I. (2003). Formation and plasticity of GABAergic synapses: physiological mechanisms and pathophysiological implications. Pharmacol. Ther. 98, 299–323. Fritschy, J. M., and Mohler, H. (1995). GABAA-receptor heterogeneity in the adult rat brain: differential regional and cellular distribution of seven major subunits. J. Comp. Neu- rol. 359, 154–194. Gallagher, J. P., Orozco-Cabal, L. F., Liu, J., and Shinnick-Gallagher, P. (2008). Synaptic physiology of cen- tral CRH system. Eur. J. Pharmacol. 583, 215–225. Gao, C. Q., Dhooge, W. S., Kaufman, J. M., Weyne, J. J., and Eechaute, W. P. (2002). Hypothalamic 5α-reductase and 3α-oxidoreductase activity in the male rat. J. Steroid Biochem. Mol. Biol. 80, 91–98. Gasior, M., Carter, R. B., and Witkin, J. M. (1999). Neuroactive steroids: potential therapeutic use in neu- rological and psychiatric disor- ders. Trends Pharmacol. Sci. 20, 107–112. Gee, K. W., Bolger, M. B., Brinton, R. E., Coirini, H., and McEwen, B. S. (1988). Steroid modula- tion of the chloride ionophore in rat brain: structure-activity require- ments, regional dependence and mechanism of action. J. Pharmacol. Exp. Ther. 246, 803–812. Gee, K. W., Chang, W. C., Brinton, R. E., and McEwen, B. S. (1987). GABA- dependent modulation of the Cl- ionophore by steroids in rat brain. Eur. J. Pharmacol. 136, 419–423. Goldwater, D. S., Pavlides, C., Hunter, R. G., Bloss, E. B., Hof, P. R., McEwen, B. S., and Morrison, J. H. (2009). Structural and functional alterations to rat medial prefrontal cortex following chronic restraint stress and recovery. Neuroscience 164, 798–808. Grifﬁn, L. D., and Mellon, S. H. (1999). Selective serotonin reup- take inhibitors directly alter activ- ity of neurosteroidogenic enzymes. Proc. Natl. Acad. Sci. U.S.A. 96, 13512–13517. December 2011 | Volume 5 | Article 131 | 15