Involvement of 5-HT 2A receptor and α 2-adrenoceptor blockade in the asenapine-induced elevation of prefrontal cortical monoamine outflow

Synapse

Volumn 66, Issue 7, 2012, Pages 650-660

  Frånberg, Olivia ^a   Marcus, Monica M ^a   Svensson, Torgny H ^a  

^a KAROLINSKA INSTITUTE   (Sweden)

Author keywords

Antipsychotic drug; Dopamine; Noradrenaline; Rats; Reverse microdialysis; Serotonin

Indexed keywords

4 IODO 2,5 DIMETHOXYAMPHETAMINE; ALPHA 2 ADRENERGIC RECEPTOR; ASENAPINE; CLONIDINE; DOPAMINE; IDAZOXAN; MONOAMINE; NORADRENALIN; SEROTONIN; SEROTONIN 2A RECEPTOR; VOLINANSERIN;

ANIMAL EXPERIMENT; ANIMAL MODEL; ARTICLE; BINDING AFFINITY; DRUG POTENCY; IN VIVO STUDY; MALE; MICRODIALYSIS; MONOAMINE RELEASE; MONOAMINERGIC SYSTEM; NONHUMAN; PREFRONTAL CORTEX; PRIORITY JOURNAL; RAT;

ADRENERGIC ALPHA-2 RECEPTOR ANTAGONISTS; AMPHETAMINES; ANIMALS; ANTIPSYCHOTIC AGENTS; BIOGENIC MONOAMINES; FLUOROBENZENES; HETEROCYCLIC COMPOUNDS WITH 4 OR MORE RINGS; MALE; MICRODIALYSIS; NOREPINEPHRINE; PIPERIDINES; PREFRONTAL CORTEX; RATS; RATS, WISTAR; RECEPTOR, SEROTONIN, 5-HT2A; RECEPTORS, ADRENERGIC, ALPHA-2; SEROTONIN; SEROTONIN 5-HT2 RECEPTOR ANTAGONISTS; UP-REGULATION;

EID: 84860883395     PISSN: 08874476     EISSN: 10982396     Source Type: Journal    
DOI: 10.1002/syn.21551     Document Type: Article

References (92)

1. Abi-Dargham A, Laruelle M, Aghajanian GK, Charney D, Krystal J. 1997. The role of serotonin in the pathophysiology and treatment of schizophrenia. J Neuropsychiatry Clin Neurosci 9: 1-17.
2. Alves Fda S, Figee M, Vamelsvoort T, Veltman D, de Haan L. 2008. The revised dopamine hypothesis of schizophrenia: Evidence from pharmacological MRI studies with atypical antipsychotic medication. Psychopharmacol Bull 41: 121-132.
3. Amargós-Bosch M, Bortolozzi A, Puig MV, Serrats J, Adell A, Celada P, Toth M, Mengod G, Artigas F. 2004. Coexpression and in vivo interaction of serotonin1A and serotonin2A receptors in pyramidal neurons of prefrontal cortex. Cereb Cortex 14: 281-299.
4. Andersson JL, Nomikos GG, Marcus M, Hertel P, Mathé JM, Svensson TH. 1995. Ritanserin potentiates the stimulatory effects of raclopride on neuronal activity and dopamine release selectivity in the mesolimbic dopaminergic system. Naunyn Schmiedebergs Arch Pharmacol 352: 374-385.
5. Berk M, Ichim C, Brook S. 2001. Efficacy of mirtazapine add on therapy to haloperidol in the treatment of the negative symptoms of schizophrenia: a double-blind randomized placebo-controlled study. Int Clin Psychopharmacol 16: 87-92.
6. Bortolozzi A, Diaz-Mataix L, Scorza MC, Celada P, Artigas F. 2005. The activation of 5-HT receptors in prefrontal cortex enhances dopaminergic activity. J Neurochem 95: 1597-1607.
7. Breier A. 1995. Serotonin, schizophrenia and antipsychotic drug action. Schizophr Res 14: 187-202.
8. Castner SA, Williams GV. 2007. Tuning the engine of cognition: A focus on NMDA/D1 receptor interactions in prefrontal cortex. Brain Cogn 63: 94-122.
9. Celada P, Puig MV, Díaz-Mataix L, Artigas F. 2008. The hallucinogen DOI reduces low-frequency oscillations in rat prefrontal cortex: Reversal by antipsychotic drugs. Biol Psychiatry 64: 392-400.
10. Chen G, Greengard P, Yan Z. 2004. Potentiation of NMDA receptor currents by dopamine D1 receptors in prefrontal cortex. Proc Natl Acad Sci USA 101: 2596-2600.
11. Chen L, Yang CR. 2002. Interaction of dopamine D1 and NMDA receptors mediates acute clozapine potentiation of glutamate EPSPs in rat prefrontal cortex. J Neurophysiol 87: 2324-2336.
12. Davis KL, Kahn RS, Ko G, Davidson M. 1991. Dopamine in schizophrenia: A review and reconceptualization. Am J Psychiatry 148: 1474-1486.
13. Davis JM, Chen N, Glick ID. 2003. A meta-analysis of the efficacy of second-generation antipsychotics. Arch Gen Psychiatry 60: 553-564.
14. Devoto P, Flore G, Pani L, Gessa GL. 2001. Evidence for co-release of noradrenaline and dopamine from noradrenergic neurons in the cerebral cortex. Mol Psychiatry 6: 657-664.
15. Devoto P, Flore G, Longu G, Pira L, Gessa GL. 2003. Origin of extracellular dopamine from dopamine and noradrenaline neurons in the medial prefrontal and occipital cortex. Synapse 50: 200-205.
16. Devoto P, Flore G, Pira L, Longu G, Gessa GL. 2004. Alpha2-adrenoceptor mediated co-release of dopamine and noradrenaline from noradrenergic neurons in the cerebral cortex. J Neurochem 88: 1003-1009.
17. Driesen NR, Leung HC, Calhoun VD, Constable RT, Gueorguieva R, Hoffman R, Skudlarski P, Goldman-Rakic PS, Krystal JH. 2008. Impairment of working memory maintenance and response in schizophrenia: Functional magnetic resonance imaging evidence. Biol Psychiatry 64: 1026-1034.
18. European Medicines Agency. Sycrest, asenapine. 2010. Available at:
19. Food and Drug Administration. Saphris, asenapine. 2009. Available at:
20. Fleming K, Potkin SG, Binneman B, Keller DS, Alphs L, Panagides J. 2007. Asenapine has cognitive function effects in acute schizophrenia: A placebo-controlled and risperidone-controlled trial. 62nd Annual Scientific Meeting of the Society of Biological Psychiatry. Biol Psychiatry 61: 241S-241S.
21. Frånberg O, Marcus MM, Ivanov V, Schilström B, Shahid M, Svensson TH. 2009. Asenapine elevates cortical dopamine, noradrenaline and serotonin release. Evidence for activation of cortical and subcortical dopamine systems by different mechanisms. Psychopharmacology (Berl) 204: 251-264.
22. Frånberg O, Wiker C, Marcus MM, Konradsson A, Jardemark K, Schilström B, Shahid M, Wong EH, Svensson TH. 2008. Asenapine, a novel psychopharmacologic agent: preclinical evidence for clinical effects in schizophrenia. Psychopharmacology (Berl) 196: 417-429.
23. Gelders Y, Vanden Bussche G, Reyntjens A, Janssen P. 1986. Serotonin-S2 receptor blockers in the treatment of chronic schizophrenia. Clin Neuropharmacol 9: 325-327.
24. Gelders YG. 1989. Thymosthenic agents, a novel approach in the treatment of schizophrenia. Br J Psychiatry Suppl 5: 33-36.
25. Ghanbari R, El Mansari M, Shahid M, Blier P. 2009. Electrophysiological characterization of the effects of asenapine at 5-HT(1A), 5-HT(2A), alpha(2)-adrenergic and D(2) receptors in the rat brain. Eur Neuropsychopharmacol 19: 177-187.
26. Gobert A, Rivet JM, Audinot V, Newman-Tancredi A, Cistarelli L, Millan MJ. 1998. Simultaneous quantification of serotonin, dopamine and noradrenaline levels in single frontal cortex dialysates of freely-moving rats reveals a complex pattern of reciprocal auto- and heteroreceptor-mediated control of release. Neuroscience 84: 413-429.
27. Goldman-Rakic PS, Castner SA, Svensson TH, Siever LJ, Williams GV. 2004. Targeting the dopamine D(1) receptor in schizophrenia: insights for cognitive dysfunction. Psychopharmacology (Berl) 174: 3-16.
28. Gresch PJ, Sved AF, Zigmond MJ, Finlay JM. 1995. Local influence of endogenous norepinephrine on extracellular dopamine in rat medial prefrontal cortex. J Neurochem 65: 111-116.
29. Grinshpoon A, Valevski A, Moskowitz M, Weizman A. 2000. Beneficial effect of the addition of the 5-HT 2A/2C and alpha2 antagonist mianserin to ongoing haloperidol treatment in drug-resistant chronically hospitalized schizophrenic patients. Eur Psychiatry 15: 388-390.
30. Gu H, Wall SC, Rudnick G. 1994. Stable expression of biogenic amine transporters reveals differences in inhibitor sensitivity, kinetics, and ion dependence. J Biol Chem 269: 7124-7130.
31. Hertel P, Nomikos GG, Iurlo M, Svensson TH. 1996. Risperidone: Regional effects in vivo on release and metabolism of dopamine and serotonin in the rat brain. Psychopharmacology 124(1-2): 74-86.
32. Hertel P, Nomikos GG, Schilström B, Arborelius L, Svensson TH. 1997. Risperidone dose-dependently increases extracellular concentrations of serotonin in the rat frontal cortex: Role of alpha 2-adrenoceptor antagonism. Neuropsychopharmacology 17: 44-55.
33. Hertel P, Fagerquist MV, Svensson TH. 1999a. Enhanced cortical dopamine output and antipsychotic-like effects of raclopride by alpha2 adrenoceptor blockade. Science 286: 105-107.
34. Hertel P, Nomikos GG, Svensson TH. 1999b. Idazoxan preferentially increases dopamine output in the rat medial prefrontal cortex at the nerve terminal level. Eur J Pharmacol 371: 153-158.
35. Hertel P, Nomikos GG, Svensson TH. 1999c. The antipsychotic drug risperidone interacts with auto- and hetero-receptors regulating serotonin output in the rat frontal cortex. Neuropharmacology 38: 1175-1184.
36. Huang M, Ichiwaka J, Li Z, Dai J, Meltzer HY. 2006. Augmentation by citalopram of risperidone-induced monoamine release in rat prefrontal cortex. Psychopharmacology (Berl) 185: 274-281.
37. Ichikawa J, Dai J, Meltzer HY. 2001a. DOI, a 5-HT2A/2C receptor agonist, attenuates clozapine-induced cortical dopamine release. Brain Res 907: 151-155.
38. Ichikawa J, Ishii H, Bonaccorso S, Fowler WL, O'Laughlin IA, Meltzer HY. 2001b. 5-HT(2A) and D(2) receptor blockade increases cortical DA release via 5-HT(1A) receptor activation: A possible mechanism of atypical antipsychotic-induced cortical dopamine release. J Neurochem 76: 1521-1531.
39. Jakab RL, Goldman-Rakic PS. 1998. 5-Hydroxytryptamine2A serotonin receptors in the primate cerebral cortex: Possible site of action of hallucinogenic and antipsychotic drugs in pyramidal cell apical dendrites. Proc Natl Acad Sci USA 95: 735-740.
40. Jardemark K, Marcus MM, Shahid M, Svensson TH. 2010. Effects of asenapine on prefrontal N-methyl-D-aspartate receptor-mediated transmission: Involvement of dopamine D1 receptors. Synapse 64: 870-874.
41. Javitt DC. 2010. Glutamatergic theories of schizophrenia. Isr J Psychiatry Relat Sci 47: 4-16.
42. Joffe G, Terevnikov V, Joffe M, Stenberg JH, Burkin M, Tiihonen J. 2009. Add-on mirtazapine enhances antipsychotic effect of first generation antipsychotics in schizophrenia: A double-blind, randomized, placebo-controlled trial. Schizophr Res 108: 245-251.
43. Kane JM, Cohen M, Zhao J, Alphs L, Panagides J. 2010. Efficacy and safety of asenapine in a placebo- and haloperidol-controlled trial in patients with acute exacerbation of schizophrenia. J Clin Psychopharmacol 30: 106-115.
44. Leucht S, Corves C, Arbter D, Engel RR, Li C, Davis JM. 2009. Second-generation versus first-generation antipsychotic drugs for schizophrenia: A meta-analysis. Lancet 373: 31-41.
45. Liégeois JF, Ichikawa J, Meltzer HY. 2002. 5-HT(2A) receptor antagonism potentiates haloperidol-induced dopamine release in rat medial prefrontal cortex and inhibits that in the nucleus accumbens in a dose-dependent manner. Brain Res 947: 157-165.
46. Litman RE, Hong WW, Weissman EM, Su TP, Potter WZ, Pickar D. 1993. Idazoxan, an alpha 2 antagonist, augments fluphenazine in schizophrenic patients : a pilot study. J Clin Psychopharmacol 13: 264-267.
47. Litman RE, Su TP, Potter WZ, Hong WW, Pickar D. 1996. Idazoxan and response to typical neuroleptics in treatment-resistant schizophrenia. Comparison with the atypical neuroleptic, clozapine. Br J Psychiatry 168: 571-579.
48. Marcus MM, Jardemark KE, Wadenberg ML, Langlois X, Hertel P, Svensson TH. 2005. Combined alpha2 and D2/3 receptor blockade enhances cortical glutamatergic transmission and reverses cognitive impairment in the rat. Int J Neuropsychopharmacol 8: 315-327.
49. Marcus MM, Wiker C, Frånberg O, Konradsson-Geuken A, Langlois X, Jardemark K, Svensson TH. 2010. Adjunctive alpha2-adrenoceptor blockade enhances the antipsychotic-like effect of risperidone and facilitates cortical dopaminergic and glutamatergic, NMDA receptor-mediated transmission. Int J Neuropsychopharmacol 13: 891-903.
50. Martin-Ruiz R, Puig MV, Celada P, Shapiro DA, Roth BL, Mengod G, Artigas F. 2001. Control of serotonergic function in medial prefrontal cortex by serotonin-2A receptors through a glutamate-dependent mechanism. J Neurosci 21: 9856-9866.
51. Masana M, Bortolozzi A, Artigas F. 2011. Selective enhacement of mesocortical dopaminergic transmission by noradrenergic drugs: therapeutic opportunities in schizophrenia. Int J Neuropsychopharmacol 14: 53-68.
52. Meltzer HY, Huang M. 2008. In vivo actions of atypical antipsychotic drug on serotonergic and dopaminergic systems. Prog Brain Res 172: 177-197.
53. Millan MJ. 2000. Improving the treatment of schizophrenia: Focus on serotonin (5-HT)(1A) receptors. J Pharmacol Exp Ther 295: 853-861.
54. Miner LH, Schroeter S, Blakely RD, Sesack SR. 2003. Ultrastructural localization of the norepinephrine transporter in superficial and deep layers of the rat prelimbic prefrontal cortex and its spatial relationship to probable dopamine terminals. J Comp Neurol 466: 478-494.
55. Mizuki Y, Kajimura N, Imai T, Suetsugi M, Kai S, Kaneyuki H, Yamada M. 1990. Effects of mianserin on negative symptoms in schizophrenia. Int Clin Psychopharmacol 5: 83-95.
56. Mizuki Y, Kajimura N, Kai S, Suetsugi M, Ushijima I, Yamada M. 1992. Effects of mianserin in chronic schizophrenia. Prog Neuropsychopharmacol Biol Psychiatry 16: 517-528.
57. Moron JA, Brockington A, Wise RA, Rocha BA, Hope BT. 2002. Dopamine uptake through the norepinephrine transporter in brain regions with low levels of the dopamine transporter: Evidence from knock-out mouse lines. J Neurosci 22: 389-395.
58. Newman-Tancredi A, Chaput C, Verriele L, Millan MJ. 1996. Clozapine is a partial agonist at cloned, human serotonin 5-HT1A receptors. Neuropharmacology 35: 119-121.
59. Newman-Tancredi A, Verrièle L, Touzard M, Millan MJ. 2001. Efficacy of antipsychotic agents at human 5-HT(1A) receptors determined by [3H]WAY100, 635 binding affinity ratios: Relationship to efficacy for G-protein activation. Eur J Pharmacol 428: 177-184.
60. Ninan I, Wang RY. 2003. Modulation of the ability of clozapine to facilitate NMDA- and electrically evoked responses in pyramidal cells of the rat medial prefrontal cortex by dopamine: Pharmacological evidence. Eur J Neurosci 17: 1306-1312.
61. Nomikos GG, Iurlo M, Andersson JL, Kimura K, Svensson TH. 1994. Systemic administration of amperozide, a new atypical antipsychotic drug, preferentially increases dopamine release in the rat medial prefrontal cortex. Psychopharmacology 115: 147-156.
62. Pehek EA, McFarlane HG, Maguschak K, Price B, Pluto CP. 2001. M100, 907, a selective 5-HT(2A) antagonist, attenuates dopamine release in the rat medial prefrontal cortex. Brain Res 888: 51-59.
63. Pehek EA, Nocjar C, Roth BL, Byrd TA, Mabrouk OS. 2006. Evidence for the preferential involvement of 5-HT2A serotonin receptors in stress- and drug-induced dopamine release in the rat medial prefrontal cortex. Neuropsychopharmacology 31: 265-277.
64. Potkin SG, Cohen M, Panagides J. 2007. Efficacy and tolerability of asenapine in acute schizophrenia: a placebo- and risperidone-controlled trial. J Clin Psychiatry 68: 1492-1500.
65. Poyurovsky M, Shardorodsky M, Fuchs C, Schneidman M, Weizman A. 1999. Treatment of neuroleptic-induced akathisia with the 5-HT2 antagonist mianserin. Double-blind, placebo-controlled study. Br J Psychiatry 174: 238-242.
66. Poyurovsky M, Koren D, Gonopolsky I, Schneidman M, Fuchs C, Weizman A, Weizman R. 2003. Effect of the 5-HT2 antagonist mianserin on cognitive dysfunction in chronic schizophrenia patients: An add-on, double-blind placebo-controlled study. Eur Neuropsychopharmacol 13: 123-128.
67. Pozzi L, Invernizzi R, Cervo L, Vallebuona F, Samanin R. 1994. Evidence that extracellular concentrations of dopamine are regulated by noradrenergic neurons in the frontal cortex of rats. J Neurochem 63: 195-200.
68. Reyntjens A, Gelders YG, Hoppenbrouwers MLJA, Vanden Bussche G. 1986. Thymosthenic effects of ritanserin (R-55667), a centrally acting serotonin-S2 receptor blocker. Drug Dev Res 8: 205-211.
69. Schotte A, Janssen PF, Gommeren W, Luyten WH, Van Gompel P, Lesage AS, De Loore K, Leysen JE. 1996. Risperidone compared with new and reference antipsychotic drugs: In vitro and in vivo receptor binding. Psychopharmacology (Berl) 124: 57-73.
70. Schreiber R, Brocco M, Millan MJ. 1994. Blockade of the discriminative stimulus effects of DOI by MDL 100907 and the "atypical" antipsychotics, clozapine and risperidone. Eur J Pharmacol 264: 99-102.
71. Schroeter S, Apparsundaram S, Wiley RG, Miner LH, Sesack SR, Blakely RD. 2000. Immunolocalization of the cocaine- and antidepressant-sensitive l-norepinephrine transporter. J Comp Neurol 420: 211-232.
72. Sesack SR, Hawrylak VA, Matus C, Guido MA, Levey AI. 1998. Dopamine axon varicosities in the prelimbic division of the rat prefrontal cortex exhibit sparse immunoreactivity for the dopamine transporter. J Neurosci 18: 2697-2708.
73. Shahid M, Walker GB, Zorn SH, Wong EH. 2009. Asenapine: A novel psychopharmacologic agent with a unique human receptor signature. J Psychopharmacol 23: 65-73.
74. Shiloh R, Zemishlany Z, Aizenberg D, Valevski A, Bodinger L, Munitz H, Weizman A. 2002. Mianserin or placebo as adjuncts to typical antipsychotics in resistant schizophrenia. Int Clin Psychopharmacol 17: 59-64.
75. Silver H. 2004. Selective serotonin re-uptake inhibitor augmentation in the treatment of negative symptoms of schizophrenia. Expert Opin Pharmacother 5: 2053-2058.
76. Snigdha S, Idris N, Grayson B, Shahid M, Neill JC. 2011. Asenapine improves phencyclidine-induced object recognition deficits in the rat: Evidence for engagement of a dopamine D(1) receptor mechanism. Psychopharmacology (Berl) 214: 843-853.
77. Stenberg JH, Terevnikov V, Joffe M, Tiihonen J, Tchoukhine E, Burkin M, Joffe G. 2010. Effects of add-on mirtazapine on neurocognition in schizophrenia: A double-blind, randomized, placebo-controlled study. Int J Neuropsychopharmacol 13: 433-441.
78. Svensson TH, Bunney BS, Aghajanian GK. 1975. Inhibition of both noradrenergic and serotonergic neurons in brain by the alpha-adrenergic agonist clonidine. Brain Res 92: 291-306.
79. Svensson TH. 2000. Dysfunctional brain dopamine systems induced by psychotomimetic NMDA-receptor antagonists and the effects of antipsychotic drugs. Brain Res Brain Res Rev 31: 320-329.
80. Svensson TH. 2003. Alpha-adrenoceptor modulation hypothesis of antipsychotic atypicality. Prog Neuropsychopharmacol Biol Psychiatry 27: 1145-1158.
81. Szegedi A, Zhao J, van Willigenburg A, Nations KR, Mackle M, Panagides J. 2011. Effects of asenapine on depressive symptoms in patients with bipolar I disorder experiencing acute manic or mixed episodes: A post hoc analysis of two 3-week clinical trials. BMC Psychiatry 11: 101.
82. Tan HY, Callicott JH, Weinberger DR. 2007. Dysfunctional and compensatory prefrontal cortical systems, genes and the pathogenesis of schizophrenia. Cereb Cortex 17( Suppl 1): i171-i181
83. Terevnikov V, Stenberg JH, Tiihonen J, Joffe M, Burkin M, Tchoukhine E, Joffe G. 2011. Add-on mirtazapine improves depressive symptoms in schizophrenia: A double-blind randomized placebo-controlled study with an open-label extension phase. Hum Psychopharmacol Clin Exp 26: 188-193.
84. Wadenberg ML, Hicks PB, Richter JT, Young KA. 1998. Enhancement of antipsychoticlike properties of raclopride in rats using the selective serotonin2A receptor antagonist MDL 100, 907. Biol Psychiatry 44: 508-515.
85. Wadenberg ML, Wiker C, Svensson TH. 2007. Enhanced efficacy of both typical and atypical antipsychotic drugs by adjunctive alpha2 adrenoceptor blockade: Experimental evidence. Int J Neuropsychopharmacol 10: 191-202.
86. Wang Y, Liu J, Gui ZH, Ali U, Fan LL, Hou C, Wang T, Chen L, Li Q. 2011. alpha(2)-Adrenoceptor regulates the spontaneous and the GABA/glutamate modulated firing activity of the rat medial prefrontal cortex pyramidal neurons. Neuroscience 182: 193-202.
87. Wedzony K, Chocyk A, Maćkowiak M. 2008. A search for colocalization of serotonin 5-HT2A and 5-HT1A receptors in the rat medial prefrontal and entorhinal cortices-Immunohistochemical studies. J Physiol Pharmacol 59: 229-238.
88. Westerink BH, Kawahara Y, De Boer P, Geels C, De Vries JB, Wikstrom HV, Van Kalkeren A, Van Vliet B, Kruse CG, Long SK. 2001. Antipsychotic drugs classified by their effects on the release of dopamine and noradrenaline in the prefrontal cortex and striatum. Eur J Pharmacol 412: 127-138.
89. Wiker C, Linner L, Wadenberg ML, Svensson TH. 2005. Adjunctive treatment with mianserin enhances effects of raclopride on cortical dopamine output and, in parallel, its antipsychotic-like effect. Neuropsychiatr Dis Treat 1: 253-260.
90. Willins DL, Deutch AY, Roth BL. 1997. Serotonin 5-HT2A receptors are expressed on pyramidal cells and interneurons in the rat cortex. Synapse 27: 79-82.
91. Wirkner K, Krause T, Koles L, Thummler S, Al-Khrasani M, Illes P. 2004. D(1) but not D(2) dopamine receptors or adrenoceptors mediate dopamine-induced potentiation of N-methyl-D-aspartate currents in the rat prefrontal cortex. Neurosci Lett 372: 89-93.
92. Yamamoto K, Hornykiewicz O. 2004. Proposal for a noradrenaline hypothesis of schizophrenia. Prog Neuropsychopharmacol Biol Psychiatry 28: 913-922.