The extra-adrenal effects of metyrapone and oxazepam on ongoing cocaine self-administration

Brain Research

Volumn 1575, Issue , 2014, Pages 45-54

  Guerin, Glenn F ^a   Schmoutz, Christopher D ^a   Goeders, Nicholas E ^a  

^a LOUISIANA STATE UNIVERSITY HEALTH SCIENCES CENTER   (United States)

Author keywords

Addiction; Adrenalectomy; Benzodiazepine; Cocaine; Metyrapone; Self administration

Indexed keywords

COCAINE; CORTICOSTERONE; CORTICOTROPIN; METYRAPONE; OXAZEPAM; ANXIOLYTIC AGENT;

ADRENAL GLAND; ADRENALECTOMY; ADULT; ANIMAL EXPERIMENT; ANIMAL MODEL; ARTICLE; COCAINE DEPENDENCE; CONTROLLED STUDY; CORTICOSTERONE BLOOD LEVEL; CORTICOTROPIN BLOOD LEVEL; MALE; NONHUMAN; PHYSIOLOGICAL STRESS; PRIORITY JOURNAL; RAT; REINFORCEMENT; ADRENAL MEDULLA; ANIMAL; BLOOD; DRUG EFFECTS; DRUG SEEKING BEHAVIOR; DRUG SELF ADMINISTRATION; INSTRUMENTAL CONDITIONING; PHYSIOLOGY; WISTAR RAT;

RATTUS; RATTUS NORVEGICUS; RODENTIA;

ADRENAL MEDULLA; ADRENALECTOMY; ANIMALS; ANTI-ANXIETY AGENTS; COCAINE; CONDITIONING, OPERANT; CORTICOSTERONE; DRUG-SEEKING BEHAVIOR; EXTINCTION, PSYCHOLOGICAL; MALE; METYRAPONE; OXAZEPAM; RATS; RATS, WISTAR; SELF ADMINISTRATION; STRESS, PHYSIOLOGICAL;

EID: 84925883590     PISSN: 00068993     EISSN: 18726240     Source Type: Journal    
DOI: 10.1016/j.brainres.2014.05.039     Document Type: Article

References (114)

1. Adinoff, B., 2004. Neurobiologic processes in drug reward and addiction. Harvard Rev. Psychiatry 12, 305-320.
2. Adinoff, B., Iranmanesh, A., Veldhuis, J., Fisher, L., 1998. Disturbances of the stress response: the role of the HPA axis during alcohol withdrawal and abstinence. Alcohol Health Res. World 22, 67-72.
3. Adinoff, B., Ruether, K., Krebaum, S., Iranmanesh, A., Williams, M.J., 2003. Increased salivary cortisol concentrations during chronic alcohol intoxication in a naturalistic clinical sample of men. Alcohol. Clin. Exp. Res. 27, 1420-1427.
4. Ambroggi, F., Turiault, M., Milet, A., Deroche-Gamonet, V., Parnaudeau, S., Balado, E., Barik, J., van der Veen, R., Maroteaux, G., Lemberger, T., Schütz, G., Lazar, M., Marinelli, M., Piazza, P.V., Tronche, F., 2009. Stress and addiction: glucocorticoid receptor in dopaminoceptive neurons facilitates cocaine seeking. Nat. Neurosci. 12, 247-249.
5. Barrett, A.C., Negus, S.S., Mello, N.K., Caine, S.B., 2005. Effect of GABA agonists and GABA-A receptor modulators on cocaine-and food-maintained responding and cocaine discrimination in rats. J. Pharmacol. Exp. Ther. 315, 858-871.
6. Barrot, M., Abrous, D.N., Marinelli, M., Rougé-Pont, F., Le Moal, M., Piazza, P.V., 2001. Influence of glucocorticoids on dopaminergic transmission in the rat dorsolateral striatum. Eur. J. Neurosci. (13), 812-818.
7. Barrot, M., Marinelli, M., Abrous, D.N., Rougé-Pont, F., Le Moal, M., Piazza, P.V., 2000. The dopaminergic hyper-responsiveness of the shell of the nucleus accumbens is hormone-dependent. Eur. J. Neurosci. (12), 973-979.
8. Belelli, D., Lambert, J.J., 2005. Neurosteroids: endogenous regulators of the GABA(A) receptor. Nat. Rev. Neurosci. 6, 565-575.
9. Brady, K.T., Sinha, R., 2005. Co-occurring mental and substance use disorders: the neurobiological effects of chronic stress. Am. J. Psychiatry 162, 1483-1493.
10. Brady, K.T., Sonne, S.C., 1999. The role of stress in alcohol use, alcoholism treatment, and relapse. Alcohol Res. Health 23, 263-271.
11. Brown, R.C., Papadopoulos, V., 2001. Role of the peripheral-type benzodiazepine receptor in adrenal and brain steroidogenesis. Int. Rev. Neurobiol. 46, 117-143.
12. Bruun-Meyer, S.E, 1987. The GABA/benzodiazepine receptorchloride ionophore complex: nature and modulation. Prog. Neuropsychopharmacol. Biol. Psychiatry 11, 365-387.
13. Buffalari, D.M., See, R.E., 2009. Footshock stress potentiates cue-induced cocaine-seeking in an animal model of relapse. Physiol. Behav. 98, 614-617.
14. Bureik, M., Lisurek, M., Bernhardt, R., 2002. The human steroid hydroxylases CYP1B1 and CYP11B2. Biol. Chem. 383, 1537-1551.
15. Chen, S.W., Rodriguez, L., Davies, M.F., Loew, G.H., 1996. The hyperphagic effect of 3 alpha-hydroxylated pregnane steroids in male rats. Pharmacol. Biochem. Behav. 53, 777-782.
16. Colby, H.D., Witorsch, R.J., Caffrey, J.L., Kitay, J.I., 1973. Effects of steroid suppression and gonadectomy on adrenal 5alpha-reductase activity and corticosterone production in rats. Acta Endocrinol. (Copenhagen) 74, 568-575.
17. Davies, E., MacKenzie, S.M., 2003. Extra-adrenal production of corticosteroids. Clin. Exp. Pharmacol. Physiol. 30, 437-445.
18. De Jong, I.E.M., de Kloet, E.R., 2009. Critical time-window for the actions of adrenal glucocorticoids in behavioural sensitisation to cocaine. Eur. J. Pharmacol. 604, 66-73.
19. De Jong, I.E.M., Oitzl, M.S., de Kloet, E.R., 2007. Adrenalectomy prevents behavioural sensitisation of mice to cocaine in a genotype-dependent manner. Behav. Brain Res. 177, 329-339.
20. De Jong, I.E.M., Steenbergen, P.J., de Kloet, E.R., 2008. Strain differences in the effects of adrenalectomy on the midbrain dopamine system: implication for behavioral sensitization to cocaine. Neuroscience 153, 594-604.
21. Deroche, V., Marinelli, M., Le Moal, M., Piazza, P.V., 1997. Glucocorticoids and behavioral effects of psychostimulants. II: Cocaine intravenous self-administration and reinstatement depend on glucocorticoid levels. J. Pharmacol. Exp. Ther. 281, 1401-1407.
22. Dixon, C.I., Halbout, B., King, S.L., Stephens, D.N., 2014. Deletion of the GABAA α2-subunit does not alter self-administration of cocaine or reinstatement of cocaine seeking. Psychopharmacology (Berlin).
23. Dixon, C.I., Morris, H.V., Breen, G., Desrivieres, S., Jugurnauth, S., Steiner, R.C., Vallada, H., Guindalini, C., Laranjeira, R., Messas, G., Rosahl, T.W., Atack, J.R., Peden, D.R., Belelli, D., Lambert, J.J., King, S.L., Schumann, G., Stephens, D.N., 2010. Cocaine effects on mouse incentive-learning and human addiction are linked to alpha2 subunit-containing GABAA receptors. Proc. Nat. Acad. Sci. U.S.A 107, 2289-2294.
24. Edwards, C.R., Stewart, P.M., Burt, D., Brett, L., McIntyre, M.A., Sutanto, W.S., de Kloet, E.R., Monder, C., 1988. Localisation of 11 beta-hydroxysteroid dehydrogenase - tissue specific protector of the mineralocorticoid receptor. Lancet 2, 986-989.
25. Engelhardt, D., Dörr, G., Jaspers, C., Knorr, D., 1985. Ketoconazole blocks cortisol secretion in man by inhibition of adrenal 11 beta-hydroxylase. Klin. Wochenschr. 63, 607-612.
26. Engelhardt, D., Weber, M.M., 1994. Therapy of Cushing's syndrome with steroid biosynthesis inhibitors. J. Steroid Biochem. Mol. Biol. 49, 261-267.
27. Engin, E., Bakhurin, K.I., Smith, K.S., Hines, R.M., Reynolds, L.M., Tang, W., Sprengel, R., Moss, S.J., Rudolph, U., 2014. Neural basis of benzodiazepine reward: requirement for α2 containing GABAA receptors in the nucleus accumbens. Neuropsychopharmacology.
28. Feltenstein, M.W., See, R.E., 2006. Potentiation of cue-induced reinstatement of cocaine-seeking in rats by the anxiogenic drug yohimbine. Behav. Brain Res. 174, 1-8.
29. Freel, E.M., Bernhardt, R., Bernhardt, M., Ingram, M., Ingram, R., Wallace, A.M., Fraser, R., Davies, E., Connell, J.M.C., 2007. Endogenous corticosteroid biosynthesis in subjects after bilateral adrenalectomy. Clin. Endocrinol. (Oxford) 66, 659-665.
30. Fryer, R.I., Gu, Z.Q., 1990. The rational design and synthesis of haptens having specific activity as full agonists or full antagonists at the benzodiazepine receptor. Life Sci. 47, 833-840.
31. Fudge, M.A., Kavaliers, M., Ossenkopp, K.-P., 2006. Allopregnanolone produces hyperphagia by reducing neophobia without altering food palatability. Eur. Neuropsychopharmacol. 16, 272-280.
32. Goeders, N.E., 2002. Stress and cocaine addiction. J. Pharmacol. Exp. Ther. 301, 785-789.
33. Goeders, N.E., 2004. Stress, motivation, and drug addiction. Curr. Dir. Psychol. Sci. 13, 33-35.
34. Goeders, N.E., 2010. Stress and drug craving. In: Koob Thompson, R.F., Le Moal, M.G. (Eds.), Encyclopaedia of Behavioral Neuroscience. Academic Press, Oxford, pp. 310-315.
35. Goeders, N.E., Guerin, G.F., 1994. Non-contingent electric footshock facilitates the acquisition of intravenous cocaine self-administration in rats. Psychopharmacology (Berlin) 114, 63-70.
36. Goeders, N.E., Guerin, G.F., 1996a. Effects of surgical and pharmacological adrenalectomy on the initiation and maintenance of intravenous cocaine self-administration in rats. Brain Res. 722, 145-152.
37. Goeders, N.E., Guerin, G.F., 1996b. Role of corticosterone in intravenous cocaine self-administration in rats. Neuroendocrinology 64, 337-348.
38. Goeders, N.E., Guerin, G.F., 2000. Effects of the CRH receptor antagonist CP-154,526 on intravenous cocaine self-administration in rats. Neuropsychopharmacology 23, 577-586.
39. Goeders, N.E., Guerin, G.F., 2008. Effects of the combination of metyrapone and oxazepam on cocaine and food self-administration in rats. Pharmacol. Biochem. Behav. 91, 181-189.
40. Goeders, N.E., Guerin, G.F., Schmoutz, C.D., 2014. The combination of metyrapone and oxazepam for the treatment of cocaine and other drug addictions. In: Dwoskin, L.P., Enna, S.J., LeCount, L. (Eds.), Advances in Pharmacology. Academic Press, San Diego, Calif, pp. 419-479.
41. Goeders, N.E., Irby, B.D., Shuster, C.C., Guerin, G.F., 1997. Tolerance and sensitization to the behavioral effects of cocaine in rats: relationship to benzodiazepine receptors. Pharmacol. Biochem. Behav. 57, 43-56.
42. Goeders, N.E., McNulty, M.A., Guerin, G.F., 1993. Effects of alprazolam on intravenous cocaine self-administration in rats. Pharmacol. Biochem. Behav. 44, 471-474.
43. Goeders, N.E., McNulty, M.A., Mirkis, S., McAllister, K.H., 1989. Chlordiazepoxide alters intravenous cocaine self-administration in rats. Pharmacol. Biochem. Behav. 33, 859-866.
44. Goeders, N.E., Peltier, R.L., Guerin, G.F., 1998. Ketoconazole reduces low dose cocaine self-administration in rats. Drug Alcohol Depend. 53, 67-77.
45. Gomez-Sanchez, C.E, Zhou, M.Y., Cozza, E.N., Morita, H., Eddleman, F.C., Gomez-Sanchez, E.P., 1996. Corticosteroid synthesis in the central nervous system. Endocr. Res. 22, 463-470.
46. Gomez-Sanchez, C.E, Zhou, M.Y., Cozza, E.N., Morita, H., Foecking, M.F., Gomez-Sanchez, E.P., 1997. Aldosterone biosynthesis in the rat brain. Endocrinology 138, 3369-3373.
47. Graf, E.N., Hoks, M.A., Baumgardner, J., Sierra, J., Vranjkovic, O., Bohr, C., Baker, D.A., Mantsch, J.R., 2011. Adrenal activity during repeated long-access cocaine self-administration is required for later CRF-Induced and CRF-dependent stressor-induced reinstatement in rats. Neuropsychopharmacology 36, 1444-1454.
48. Graf, E.N., Wheeler, R.A., Baker, D.A., Ebben, A.L., Hill, J.E., McReynolds, J.R., Robble, M.A., Vranjkovic, O., Wheeler, D.S., Mantsch, J.R., Gasser, P.J., 2013. Corticosterone acts in the nucleus accumbens to enhance dopamine signaling and potentiate reinstatement of cocaine seeking. J. Neurosci. 33, 11800-11810.
49. Greenblatt, D.J., 1981. Clinical pharmacokinetics of oxazepam and lorazepam. Clin. Pharmacokinet. 6, 89-105.
50. Greenblatt, D.J., Shader, R.I., Koch-Weser, J., 1975. Pharmacokinetics in clinical medicine: oxazepam versus other benzodiazepines. Dis. Nerv. Syst 36, 6-13.
51. Haleem, D.J., Kennett, G., Curzon, G., 1988. Adaptation of female rats to stress: shift to male pattern by inhibition of corticosterone synthesis. Brain Res. 458, 339-347.
52. Harris, J.J., Hoegel, C., Crane, M.G., 1967. Deoxycorticosterone secretion rates before and during metyrapone administration in normal subjects. J. Clin. Endocrinol. Metab. 27, 105-113.
53. Higgs, S., Cooper, S.J., 1998. Antineophobic effect of the neuroactive steroid 3alpha-hydroxy-5beta-pregnan-20-one in male rats. Pharmacol. Biochem. Behav. 60, 125-131.
54. Hirani, K., Khisti, R.T., Chopde, C.T., 2002. Behavioral action of ethanol in Porsolt's forced swim test: modulation by 3 alpha-hydroxy-5 alpha-pregnan-20-one. Neuropharmacology 43, 1339-1350.
55. Hirani, K., Sharma, A.N., Jain, N.S., Ugale, R.R., Chopde, C.T., 2005. Evaluation of GABAergic neuroactive steroid 3alpha-hydroxy-5alpha-pregnane-20-one as a neurobiological substrate for the anti-anxiety effect of ethanol in rats. Psychopharmacology (Berlin) 180, 267-278.
56. Holmberg, E., Bäckström, T., Johansson, M., Löfgren, M., Haage, D., 2013. Allopregnanolone induces a diurnally dependent hyperphagic effect and alters feeding latency and duration in male Wistar rats. Acta Physiol. (Oxford) 208, 400-409.
57. Holmes, M.C., Yau, J.L.W., Kotelevtsev, Y., Mullins, J.J., Seckl, J.R., 2003. 11 Beta-hydroxysteroid dehydrogenases in the brain: two enzymes two roles. Ann. N.Y. Acad. Sci. 1007, 357-366.
58. Kablinger, A.S., Lindner, M.A., Casso, S., Hefti, F., DeMuth, G., Fox, B.S., McNair, L.A., McCarthy, B.G., Goeders, N.E., 2012. Effects of the combination of metyrapone and oxazepam on cocaine craving and cocaine taking: a double-blind, randomized, placebo-controlled pilot study. J. Psychopharmacol. 26, 973-981.
59. Koob, G.F., 2010. The role of CRF and CRF-related peptides in the dark side of addiction. Brain Res. 1314, 3-14.
60. Koob, G.F., Goeders, N.E., 1989. Neuroanatomical substrates of drug self-administration, Oxford reviews in psychopharmacology. Neuropharmacological Basis of Reward, Vol. 1. Oxford University Press, London.
61. Koob, G.F., Zorrilla, E.P., 2010. Neurobiological mechanisms of addiction: focus on corticotropin-releasing factor. Curr. Opin. Invest. Drugs 11, 63-71.
62. Kreek, M.J., Nielsen, D.A., Butelman, E.R., LaForge, K.S., 2005. Genetic influences on impulsivity, risk taking, stress responsivity and vulnerability to drug abuse and addiction. Nat. Neurosci. 8, 1450-1457.
63. Krueger, K.E., Papadopoulos, V., 1990. Peripheral-type benzodiazepine receptors mediate translocation of cholesterol from outer to inner mitochondrial membranes in adrenocortical cells. J. Biol. Chem. 265, 15015-15022.
64. Lamberts, S.W., Bons, E.G., Bruining, H.A., de Jong, F.H., 1987. Differential effects of the imidazole derivatives etomidate, ketoconazole and miconazole and of metyrapone on the secretion of cortisol and its precursors by human adrenocortical cells. J. Pharmacol. Exp. Ther. 240, 259-264.
65. Li, H., Papadopoulos, V., 1998. Peripheral-type benzodiazepine receptor function in cholesterol transport. Identification of a putative cholesterol recognition/interaction amino acid sequence and consensus pattern. Endocrinology 139, 4991-4997.
66. Löfgren, M., Bengtsson, S.K., Johansson, M., Bäckström, T., 2013. Allopregnanolone promotes success in food competition in subordinate male rats. Neuropsychobiology 68, 15-23.
67. Lovallo, W.R., 2006. The hypothalamic-pituitary-adrenocortical axis in addiction. Int. J. Psychophysiol. 59, 193-194.
68. MacKenzie, S.M., Clark, C.J., Fraser, R., Gómez-Sanchez, C.E., Connell, J.M., Davies, E., 2000a. Expression of 11beta-hydroxylase and aldosterone synthase genes in the rat brain. J. Mol. Endocrinol. 24, 321-328.
69. MacKenzie, S.M., Clark, C.J., Ingram, M.C., Lai, M., Seckl, J., Gomez-Sanchez, CE., Fraser, R., Connell, J.M., Davies, E., 2000b. Corticosteroid production by fetal rat hippocampal neurons. Endocr. Res. 26, 531-535.
70. MacKenzie, S.M., Connell, J.M.C, Davies, E., 2012. Non-adrenal synthesis of aldosterone: a reality check. Mol. Cell. Endocrinol. 350, 163-167.
71. Maguire, E.P., Macpherson, T., Swinny, J.D., Dixon, C.I., Herd, M.B., Belelli, D., Stephens, D.N., King, S.L., Lambert, J.J., 2014. Tonic inhibition of accumbal spiny neurons by extrasynaptic α4β5 GABAA receptors modulates the actions of psychostimulants. J. Neurosci. 34, 823-838.
72. Mantsch, J.R., Baker, DA, Serge, J.P, Hoks, M.A., Francis, D.M., Katz, E.S., 2008. Surgical adrenalectomy with diurnal corticosterone replacement slows escalation and prevents the augmentation of cocaine-induced reinstatement in rats self-administering cocaine under long-access conditions. Neuropsychopharmacology 33, 814-826.
73. Mantsch, J.R., Vranjkovic, O., Twining, R.C, Gasser, P.J., McReynolds, J.R., Blacktop, J.M., 2014. Neurobiological mechanisms that contribute to stress-related cocaine use. Neuropharmacology 76, 383-394 (Pt B).
74. Marinelli, M., Rougé-Pont, F, De Jesus-Oliveira, C, Le Moal, M., Piazza, P.V., 1997. Acute blockade of corticosterone secretion decreases the psychomotor stimulant effects of cocaine. Neuropsychopharmacology (16), 156-161.
75. Mendelson, J.H., Sholar, M.B., Goletiani, N, Siegel, A.J., Mello, N.K., 2005. Effects of low- and high-nicotine cigarette smoking on mood states and the HPA axis in men. Neuropsychopharmacology 30, 1751-1763.
76. Nakhate, K.T., Subhedar, N.K., Bharne, A.P, Singru, P.S., Kokare, D.M., 2013. Involvement of cocaine- and amphetamine-regulated transcript peptide in the hyperphagic and body weight promoting effects of allopregnanolone in rats. Brain Res. 1532, 44-55.
77. Negus, S.S., Mello, N.K., Fivel, P.A., 2000. Effects of GABA agonists and GABA-A receptor modulators on cocaine discrimination in rhesus monkeys. Psychopharmacology (Berlin) 152, 398-407.
78. Papadopoulos, V., Nowzari, F.B., Krueger, K.E., 1991. Hormone-stimulated steroidogenesis is coupled to mitochondrial benzodiazepine receptors. Tropic hormone action on steroid biosynthesis is inhibited by flunitrazepam. J. Biol. Chem. 266, 3682-3687.
79. Papadopoulos, V., Amri, H., Boujrad, N., Cascio, C., Culty, M., Garnier, M., Hardwick, M., Li, H., Vidic, B., Brown, A.S., Reversa, J.L., Bernassau, J.M., Drieu, K., 1997. Peripheral benzodiazepine receptor in cholesterol transport and steroidogenesis. Steroids 62, 21-28.
80. Papadopoulos, V., Widmaier, E.P., Amri, H., Zilz, A., Li, H., Culty, M., Castello, R., Philip, G.H., Sridaran, R., Drieu, K., 1998. In vivo studies on the role of the peripheral benzodiazepine receptor (PBR) in steroidogenesis. Endocr. Res. 24, 479-487.
81. Papadopoulos, V., Baraldi, M., Guilarte, T.R., Knudsen, T.B., Lacapère, J.-J., Lindemann, P., Norenberg, M.D., Nutt, D., Weizman, A., Zhang, M.-R., Gavish, M., 2006a. Translocator protein (18 kDa): new nomenclature for the peripheral-type benzodiazepine receptor based on its structure and molecular function. Trends Pharmacol. Sci. 27, 402-409.
82. Papadopoulos, V., Lecanu, L., Brown, R.C., Han, Z., Yao, Z.-X., 2006b. Peripheral-type benzodiazepine receptor in neurosteroid biosynthesis, neuropathology and neurological disorders. Neuroscience 138, 749-756.
83. Peltier, R.L., Guerin, G.F., Dorairaj, N., Goeders, N.E., 2001. Effects of saline substitution on responding and plasma corticosterone in rats trained to self-administer different doses of cocaine. J. Pharmacol. Exp. Ther. 299, 114-120.
84. Piazza, P.V., Rougé-Pont, F., Deroche, V., Maccari, S., Simon, H., Le Moal, M., 1996. Glucocorticoids have state-dependent stimulant effects on the mesencephalic dopaminergic transmission. Proc. Nat. Acad. Sci. U.S.A 93, 8716-8720.
85. Pisu, M.G., Floris, I., Maciocco, E., Serra, M., Biggio, G., 2006. Anticipation and consumption of food each increase the concentration of neuroactive steroids in rat brain and plasma. Pharmacol. Biochem. Behav. 85, 76-81.
86. Prasad, B.M., Ulibarri, C., Kalivas, P.W., Sorg, B.A., 1996. Effect of adrenalectomy on the initiation and expression of cocaine-induced sensitization. Psychopharmacology (Berlin) 125, 265-273.
87. Prasad, B.M., Ulibarri, C., Sorg, B.A., 1998. Stress-induced cross-sensitization to cocaine: effect of adrenalectomy and corticosterone after short- and long-term withdrawal. Psychopharmacology (Berlin) 136, 24-33.
88. Preston, K.L., Epstein, D.H., 2011. Stress in the daily lives of cocaine and heroin users: relationship to mood, craving, relapse triggers, and cocaine use. Psychopharmacology (Berlin) 218, 29-37.
89. Przegaliński, E., Filip, M., Siwanowicz, J., Nowak, E., 2000. Effect of adrenalectomy and corticosterone on cocaine-induced sensitization in rats. J. Physiol. Pharmacol. 51, 193-204.
90. Raven, P.W., Checkley, S.A., Taylor, N.F., 1995. Extra-adrenal effects of metyrapone include inhibition of the 11-oxoreductase activity of 11 beta-hydroxysteroid dehydrogenase: a model for 11-HSD I deficiency. Clin. Endocrinol. 43, 637-644.
91. Raven, P.W., O'Dwyer, A.M., Taylor, N.F., Checkley, S.A., 1996. The relationship between the effects of metyrapone treatment on depressed mood and urinary steroid profiles. Psychoneuroendocrinology 21, 277-286.
92. Reddy, D.S., Kulkarni, S.K., 1998. The role of GABA-A and mitochondrial diazepam-binding inhibitor receptors on the effects of neurosteroids on food intake in mice. Psychopharmacology (Berlin) 137, 391-400.
93. Reddy, D.S., Kulkarni, S.K., 1999. Sex and estrous cycle-dependent changes in neurosteroid and benzodiazepine effects on food consumption and plus-maze learning behaviors in rats. Pharmacol. Biochem. Behav. 62, 53-60.
94. Reynolds, L.M., Engin, E., Tantillo, G., Lau, H.M., Muschamp, J.W., Carlezon, W.A., Rudolph, U., 2012. Differential roles of GABA (A) receptor subtypes in benzodiazepine-induced enhancement of brain-stimulation reward. Neuropsychopharmacology 37, 2531-2540.
95. Rupprecht, R., Ströhle, A., Hermann, B., di Michele, F., Spalletta, G., Pasini, A., Holsboer, F., Romeo, E., Strohle, A., 1998. Neuroactive steroid concentrations following metyrapone administration in depressed patients and healthy volunteers. Biol. Psychiatry 44, 912-914.
96. Sampath-Kumar, R., Yu, M., Khalil, M.W., Yang, K., 1997. Metyrapone is a competitive inhibitor of 11beta-hydroxysteroid dehydrogenase type 1 reductase. J. Steroid Biochem. Mol. Biol. 62, 195-199.
97. Schmoutz, C.D., 2013. The Intersection of Metyrapone, Inhibitory Neurosteroids and Cocaine Addiction. (Order no. 3572665, Louisiana State University Health Sciences Center - Shreveport). ProQuest Dissertations and Theses, 167.
98. Schmoutz, C.D., Runyon, S.P., Goeders, N.E., 2014. Effects of inhibitory GABA-active neurosteroids on cocaine seeking and cocaine taking in rats. Psychopharmacology (Berlin) http://dx.doi.org/10.1007/s00213-013-3404-2.
99. Shalev, U., Marinelli, M., Baumann, M.H., Piazza, P.-V., Shaham, Y., 2003. The role of corticosterone in food deprivation-induced reinstatement of cocaine seeking in the rat. Psychopharmacology (Berlin) 168, 170-176.
100. Sieghart, W., 1994. Pharmacology of benzodiazepine receptors: an update. J. Psychiatry Neurosci. 19, 24-29.
101. Sonino, N., 1987. The use of ketoconazole as an inhibitor of steroid production. N. Engl. J. Med. 317, 812-818.
102. Steptoe, A., Ussher, M., 2006. Smoking, cortisol and nicotine. Int. J. Psychophysiol. 59, 228-235.
103. Strömstedt, M., Waterman, M.R., Stromstedt, M., 1995. Messenger RNAs encoding steroidogenic enzymes are expressed in rodent brain. Brain Res. Mol. Brain Res. 34, 75-88.
104. Tan, K.R., Brown, M., Labouèbe, G., Yvon, C., Creton, C., Fritschy, J.-M., Rudolph, U., Lüscher, C., 2010. Neural bases for addictive properties of benzodiazepines. Nature 463, 769-774.
105. Tan, K.R., Rudolph, U., Lüscher, C., 2011. Hooked on benzodiazepines: GABAA receptor subtypes and addiction. Trends Neurosci. 34, 188-197.
106. Taves, M.D., Gomez-Sanchez, C.E., Soma, K.K., 2011. Extra-adrenal glucocorticoids and mineralocorticoids: evidence for local synthesis, regulation, and function. Am. J. Physiol. Endocrinol. Metab. 301, E11-E24.
107. Thienpont, D., Van Cutsem, J., Van Gerven, F., Heeres, J., Janssen, P.A., 1979. Ketoconazole - a new broad spectrum orally active antimycotic. Experientia 35, 606-607.
108. Ticku, M.K., 1983. Benzodiazepine-GABA receptor-ionophore complex. Current concepts. Neuropharmacology 22, 1459-1470.
109. Vashchinkina, E., Manner, A.K., Vekovischeva, O., Hollander, B. den, Uusi-Oukari, M., Aitta-Aho, T., Korpi, E.R., 2014. Neurosteroid agonist at GABAA receptor induces persistent neuroplasticity in VTA dopamine neurons. Neuropsychopharmacology 39, 727-737.
110. Vashchinkina, E., Panhelainen, A., Vekovischeva, O.Y., Aitta-aho, T., Ebert, B., Ator, N.A., Korpi, E.R., 2012. GABA site agonist gaboxadol induces addiction-predicting persistent changes in ventral tegmental area dopamine neurons but is not rewarding in mice or baboons. J. Neurosci. 32, 5310-5320.
111. Wang, J., Taniguchi, T., Spector, S., 1984a. Structural requirements for the binding of benzodiazepines to their peripheral-type sites. Mol. Pharmacol., 349-351.
112. Wang, J.K., Morgan, J.I., Spector, S., 1984b. Benzodiazepines that bind at peripheral sites inhibit cell proliferation. Proc. Nat. Acad. Sci. U.S.A 81, 753-756.
113. Ye, P., Kenyon, C.J., Mackenzie, S.M., Nichol, K., Seckl, J.R., Fraser, R., Connell, J.M.C., Davies, E., 2008. Effects of ACTH, dexamethasone, and adrenalectomy on 11beta-hydroxylase (CYP11B1) and aldosterone synthase (CYP11B2) gene expression in the rat central nervous system. J. Endocrinol. 196, 305-311.
114. Yin, L., Lucas, S., Maurer, F., Kazmaier, U., Hu, Q., Hartmann, R.W., 2012. Novel imidazol-1-ylmethyl substituted 1,2,5,6-tetrahydropyrrolo[3,2,1-ij]quinolin-4-ones as potent and selective CYP11B1 inhibitors for the treatment of Cushing's syndrome. J. Med. Chem. 55, 6629-6633.