Remodeling of neuronal networks by stress

Frontiers in Bioscience

Volumn 11, Issue SUPPL. 2, 2006, Pages 2746-2758

  Fuchs, Eberhard ^a,b,c   Flugge, Gabriele ^a,c   Czeh, Boldizsar ^a,c  

^a GERMAN PRIMATE CENTER   (Germany)

^b UNIVERSITY MEDICAL CENTER GÖTTINGEN   (Germany)

^c Deutsche Forschungsgemeinschaft   (Germany)

Author keywords

Animal Models; Antidepressants; CDC like kinase; Chronic Stress; Dendrites; GABAergic System; Hippocampus; HPA axis; Interneurons; M6A; Neurogenesis; Parvalbumin; Pyramidal Neurons; Review

Indexed keywords

ANIMALIA;

EID: 33744498085     PISSN: 27686701     EISSN: 27686698     Source Type: Journal    
DOI: 10.2741/2004     Document Type: Review

References (107)

1. de Kloet ER, E. Vreugdenhil, M. S. Oitzl, M. & M. Joels: Brain corticosteroid receptor balance in health and disease. Endocr Rev 19, 269-301 (1998)
2. Sapolsky RM, L. M. Romero & A. U. Munck: How do glucocorticoids influence stress responses? Integrating permissive, suppressive, stimulatory, and preparative actions. Endocr Rev 2l, 55-89 (2000)
3. Watanabe Y, E. Gould & B. S. McEwen: Stress induces atrophy of apical dendrites of hippocampal CA3 pyramidal neurons. Brain Res 588, 341-345 (1992)
4. Woolley CS, E. Gould & B. S. McEwen: Exposure to excess glucocorticoids alters dendritic morphology of adult hippocampal pyramidal neurons. Brain Res 531, 225-231 (1990)
5. Magariños AM, B. S. McEwen, G. Flugge & E. Fuchs: Chronic psychosocial stress causes apical dendritic atrophy of hippocampal CA3 pyramidal neurons in subordinate tree shrews. J Neurosci 16, 3534-3540 (1996)
6. Sousa N, N. V. Lukoyanov, M. D. Madeira, O. F. Almeida & M. M. Paula-Barbosa: Reorganization of the morphology of hippocampal neuntes and synapses after stress-induced damage correlates with behavioral improvement. Neuroscience 97, 253-266 (2000)
7. Vyas A, R. Mitra, B. S. Shankaranarayan Rao & S. Chattarji: Chronic stress induces contrasting patterns of dendritic remodeling in hippocampal and amygdaloid neurons. J Neurosci 22, 6810-6818 (2002)
8. Kole MH, B. Czeh & E. Fuchs: Homeostatic maintenance in excitability of tree shrew hippocampal CA3 pyramidal neurons after chronic stress. Hippocampus 14, 742-751 (2004)
9. Hume RI & D. Purves: Apportionment of the terminals from single preganglionic axons to target neurones in the rabbit ciliary ganglion. J Physiol 338, 259-275 (1983)
10. Magariños AM, J. M. Verdugo & B. S. McEwen: Chronic stress alters synaptic terminal structure in hippocampus. Proc Natl Acad Sci USA 94, 14002-14008 (1997)
11. Sandi C, H. A. Davies, M. I. Cordero, J. J. Rodriguez, V. I. Popov & M. G. Stewart: Rapid reversal of stress induced loss of synapses in CA3 of rat hippocampus following water maze training. Eur J Neurosci 17, 2447-256 (2003)
12. Kole MH, T. Costoli, J. M. Koolhaas & E. Fuchs: Bidirectional shift in the cornu ammonis 3 pyramidal dendritic organization following brief stress. Neuroscience 125, 337-347 (2004)
13. Wellman CL: Dendritic reorganization in pyramidal neurons in medial prefrontal cortex after chronic corticosterone administration. J Neurobiol 49, 245-253 (2001)
14. Cook SC & C. L. Wellman: Chronic stress alters dendritic morphology in rat medial prefrontal cortex. J Neurobiol 60, 236-248 (2004)
15. Radley JJ, A. B. Rocher, M. Miller, W. G. Janssen, C. Liston, P. R. Hof, B. S. McEwen & J. H. Morrison: Repeated Stress Induces Dendritic Spine Loss in the Rat Medial Prefrontal Cortex. Cereb Cortex 16, 313-320 (2006)
16. Radley JJ, H. M. Sisti, J. Hao, A. B. Rocher, T. McCall, P. R. Hof, B. S. McEwen & J. H. Morrison: Chronic behavioral stress induces apical dendritic reorganization in pyramidal neurons of the medial prefrontal cortex. Neuroscience 125, 1-6 (2004)
17. Brown SM, S. Henning & C. L. Wellman: Mild, short-term stress alters dendritic morphology in rat medial prefrontal cortex. Cereb Cortex 15, 1714-1722 (2005)
18. Ramon y Cajal S: Degeneration and regeneration of the nervous system (trans. Day RM, from the 1913 Spanish edn) Oxford Univ. Press, London. (1928)
19. Rakic P: DNA synthesis and cell division in the adult primate brain. Ann N Y Acad Sci 457, 193-211 (1985)
20. Altaian J & G. D. Das: Autoradiographic and histological evidence of postnatal hippocampal neurogenesis in rats. J Comp Neurol 124, 319-335 (1965)
21. Eriksson PS, E. Perfilieva, T. Bjork-Eriksson, A. M. Alborn, C. Nordborg, D. A. Peterson & F. H. Gage: Neurogenesis in the adult human hippocampus. Nat Med 4, 1313-1317 (1998)
22. Gould E, A. J. Reeves, M. Fallah, P. Tanapat, C. G. Gross & E. Fuchs: Hippocampal neurogenesis in adult Old World primates. Proc Natl Acad Sci USA 96, 5263-5267 (1999)
23. Kornack DR & P. Rakic: Continuation of neurogenesis in the hippocampus of the adult macaque monkey. Proc Natl Acad Sci USA 96, 5768-5773 (1999)
24. Altman J: Autoradiographic and histological studies of postnatal neurogenesis. IV. Cell proliferation and migration in the anterior forebrain, with special reference to persisting neurogenesis in the olfactory bulb. J Comp Neurol 137, 433-457 (1969)
25. Cameron HA, C. S. Woolley, B. S. McEwen & E. Gould: Differentiation of newly born neurons and glia in the dentate gyrus of the adult rat. Neuroscience 56, 337-344 (1993)
26. Kaplan MS & D. H. Bell: Mitotic neuroblasts in the 9-day-old and 11-month-old rodent hippocampus. J Neurosci 4, 1429-1441 (1984)
27. Stanfield BB & J. E. Trice: Evidence that granule cells generated in the dentate gyrus of adult rats extend axonal projections. Exp Brain Res 72, 399-406 (1988)
28. van Praag H, A. F. Schinder, B. R. Christie, N. Toni, T. D. Palmer & F. H. Gage: Functional neurogenesis in the adult hippocampus. Nature 415, 1030-1034 (2002)
29. Schmidt-Hieber C, P. Jonas & J. Bischofberger: Enhanced synaptic plasticity in newly generated granule cells of the adult hippocampus. Nature 429, 184-187 (2004)
30. Wang S, B. W. Scott & J. M. Wojtowicz: Heterogenous properties of dentate granule neurons in the adult rat. J Neurobiol 42, 248-257 (2000)
31. Miller MW & R. S. Nowakowski: Use of bromodeoxyuridine- immunohistochemistry to examine the proliferation, migration and time of origin of cells in the central nervous system. Brain Res 457, 44-52 (1988)
32. Cameron HA & R. D. McKay: Adult neurogenesis produces a large pool of new granule cells in the dentate gyrus. J Comp Neurol 435, 406-417 (2001)
33. Squire LR: Memory and the hippocampus: a synthesis from findings with rats, monkeys, and humans. Psychol. Rev 99, 195-231 (1992)
34. Shors TJ, G. Miesegaes, A. Beylin, M. Zhao, T. Rydel & E. Gould E: Neurogenesis in the adult is involved in the formation of trace memories. Nature 410, 372-376 (2001)
35. Bruel-Jungerman E, S. Laroche & C. Rampon: New neurons in the dentate gyrus are involved in the expression of enhanced long-term memory following environmental enrichment. Eur J Neurosci 21, 513-521 (2005)
36. Snyder JS, N. S. Hong, R. J. McDonald & J. M. Wojtowicz: A role for adult neurogenesis in spatial long-term memory. Neuroscience 130, 843-852 (2005)
37. Fuchs E & B. Czeh: Adult neurogenesis in rodents and primates: Functional Implications. In: Handbook of stress and the brain Volume 15 of Techniques in the Behavioral and Neural Sciences. Part 1: The neurobiology of stress. Eds: Steckler T, Kalin NH, Reul JMHM, Elsevier, Amsterdam, 711-727 (2005)
38. Gould E, B. S. McEwen, P. Tanapat, L. A. Galea & E. Fuchs: Neurogenesis in the dentate gyrus of the adult tree shrew is regulated by psychosocial stress and NMDA receptor activation. J Neurosci 17, 2492-2498 (1997)
39. Gould E, P. Tanapat, B. S. McEwen, G. Flugge & E. Fuchs. Proliferation of granule cell precursors in the dentate gyrus of adult monkeys is diminished by stress. Proc Natl Acad Sci USA 95, 3168-3171 (1998)
40. Tanapat P, L. A. Galea & E. Gould. Stress inhibits the proliferation of granule cell precursors in the developing dentate gyrus. Int J Dev Neurosci 16, 235-239 (1998)
41. Lemaire V, M. Koehl, M. Le Moal & D. N. Abrous: Prenatal stress produces learning deficits associated with an inhibition of neurogenesis in the hippocampus. Proc Natl Acad Sci USA 97, 11032-11037 (2000)
42. Czeh B, T. Michaelis, T. Watanabe, J. Frahm, G. de Biurrun, M. van Kampen, A. Bartolomucci & E. Fuchs: Stress-induced changes in cerebral metabolites, hippocampal volume, and cell proliferation are prevented by antidepressant treatment with tianeptine. Proc Natl Acad Sci USA 98, 12796-12801 (2001)
43. Czeh B, T. Welt, A. K. Fischer, A. Erhardt, W. Schmitt, M. B. Muller, N. Toschi, E. Fuchs & M. E. Keck: Chronic psychosocial stress and concomitant repetitive transcranial magnetic stimulation: effects on stress hormone levels and adult hippocampal neurogenesis. Biol Psychiatry 52, 1057-1065 (2003)
44. Coe CL, M. Kramer, B. Czeh, E. Gould, A. J. Reeves, C. Kirschbaum & E. Fuchs: Prenatal stress diminishes neurogenesis in the dentate gyrus of juvenile rhesus monkeys. Biol Psychiatry 54, 1025-1034 (2003)
45. Malberg JE & R. S. Duman: Cell proliferation in adult hippocampus is decreased by inescapable stress: reversal by fluoxetine treatment. Neuropsychopharmacology 28, 1562-1571 (2003)
46. Pham K, J. Nacher, P. R. Hof & B. S. McEwen: Repeated restraint stress suppresses neurogenesis and induces biphasic PSA-NCAM expression in the adult rat dentate gyrus. Eur J Neurosci 17, 879-886 (2003)
47. Vollmayr B, C. Simonis, S. Weber, P. Gass & F. Henn: Reduced cell proliferation in the dentate gyrus is not correlated with the development of learned helplessness. Biol Psychiatry 54, 1035-40 (2003)
48. Heine VM, S. Maslam, J. Zareno, M. Joels & P. J. Lucassen: Suppressed proliferation and apoptotic changes in the rat dentate gyrus after acute and chronic stress are reversible. Eur J Neurosci 19, 131-44 (2004)
49. Simon M, B. Czeh & E. Fuchs: Age-dependent susceptibility of adult hippocampal cell proliferation to chronic psychosocial stress. Brain Res 1049, 244-248 (2005)
50. Biebl M, C. M. Cooper, J. Winkler & H. G. Kuhn: Analysis of neurogenesis and programmed cell death reveals a self-renewing capacity in the adult rat brain. Neurosci Lett 291, 17-20 (2000)
51. Heine VM, S. Maslam, M. Joels & P. J. Lucassen: Prominent decline of newborn cell proliferation, differentiation, and apoptosis in the aging dentate gyrus, in absence of an age-related hypothalamus-pituitary-adrenal axis activation. Neurobiol Aging 25, 361-75 (2004)
52. Lucassen PJ, G. K. Vollmann-Honsdorf, M. Gleisberg, B. Czeh, E. R. De Kloet & E. Fuchs: Chronic psychosocial stress differentially affects apoptosis in hippocampal subregions and cortex of the adult tree shrew. Eur J Neurosci 14, 161-166 (2001)
53. Lucassen PJ, E. Fuchs & B. Czeh: Antidepressant treatment with tianeptine reduces apoptosis in the hippocampal dentate gyrus and temporal cortex. Biol Psychiatry 55, 789-796 (2004)
54. Alfarez DN, M. Joels & H. J. Krugers: Chronic unpredictable stress impairs long-term potentiation in rat hippocampal CA1 area and dentate gyrus in vitro. Eur J Neurosci 17, 1928-1934 (2003)
55. Karst H & M. Joels: Effect of chronic stress on synaptic currents in rat hippocampal dentate gyrus neurons. J Neurophysiol 89, 625-633 (2003)
56. McEwen BS: The neurobiology of stress: from serendipity to clinical relevance. Brain Res 886, 172-189 (2000)
57. Bowers G, W. E. Cullinan & J. P. Herman: Region-specific regulation of glutamic acid decarboxylase (GAD) mRNA expression in central stress circuits. J Neurosci 18, 5938-5947 (1998)
58. Stone DJ, J. P. Walsh, R. Sebro, R. Stevens, H. Pantazopolous & F. M Benes: Effects of pre- and postnatal corticosterone exposure on the rat hippocampal GABA system. Hippocampus 11, 492-507 (2001)
59. Orchinik M, S. S. Carroll, Y. H. Li, B. S. McEwen & N. G. Weiland: Heterogeneity of hippocampal GABA(A) receptors: regulation by corticosterone. J Neurosci 21, 330-339 (2001)
60. Czeh B, M. Simon, M. G. van der Hart, B. Schmelting, M. B. Hesselink & E. Fuchs: Chronic stress decreases the number of parvalbumin- immunoreactive intemeurons in the hippocampus: prevention by treatment with a substance P receptor (NK1) antagonist. Neuropsychopharmacology 30, 67-79 (2005)
61. Ishida H, Y. Shirayama, M. Iwata & R. Kawahara: Subchronic treatment with imipramine ameliorates the decreased number in neuropeptide Y-positive cells in the hippocampus of learned helplessness rats. Brain Res 1046, 239-243 (2005)
62. Sapolsky RM, L. C. Krey & B. S. McEwen: Prolonged glucocorticoid exposure reduces hippocampal neuron number: implications for aging. J Neurosci 5, 1222-1227 (1985)
63. Uno H, R. Tarara, J. G. Else, M. A. Suleman & R. M. Sapolsky: Hippocampal damage associated with prolonged and fatal stress in primates. J Neurosci 9, 1705-1711 (1989)
64. Lucassen PJ, M. B. Muller, F. Holsboer, J. Bauer, A. Holtrop & J. Wouda: Hippocampal apoptosis in major depression is a minor event and absent from subareas at risk for glucocorticoid overexposure. Am J Pathol 158, 453-468 (2001)
65. Muller MB, P. J. Lucassen, A. Yassouridis, W. J. Hoogendijk, F. Holsboer & D. F. Swaab: Neither major depression nor glucocorticoid treatment affects the cellular integrity of the human hippocampus. Eur J Neurosci 14, 1603-1612 (2001)
66. Leverenz JB, C. W. Wilkinson, M. Wamble, S. Corbin, J. E. Grabber & M. A. Raskind: Effect of chronic high-dose exogenous cortisol on hippocampal neuronal number in aged nonhuman primates. J Neurosci 19, 2356-2361 (1999)
67. Vollmann-Honsdorf GK, G. Flugge & E. Fuchs: Chronic psychosocial stress does not affect the number of pyramidal neurons in tree shrew hippocampus. Neurosci Lett 233, 121-124 (1997)
68. Vollmann-Honsdorf GK, M. J. West & E. Fuchs: Preserved neuron number in the hippocampus of chronic psychosocially stress tree shrews. Abstr Soc Neurosci 23, 239 (1997)
69. Vreugdenhil M, J. G. Jefferys, M. R. Celio & B. Schwaller: Parvalbumin-deficiency facilitates repetitive IPSCs and gamma oscillations in the hippocampus. J Neurophysiol 89, 1414-1422 (2003)
70. Scotti AL, O. Bollag, G. Kalt & C. Nitsch: Loss of perikaryal parvalbumin immunoreactivity from surviving GABAergic neurons in the CA1 field of epileptic gerbils. Hippocampus 7, 24-35 (1997)
71. McMillan PJ, C. W. Wilkinson, L. Greenup, M. A. Raskind, E. R. Peskind & J. B. Leverenz: Chronic cortisol exposure promotes the development of a GABAergic phenotype in the primate hippocampus. J Neurochem 91, 843-851 (2004)
72. Henze DA, N. N. Urban & G. Barrionuevo: The multifarious hippocampal mossy fiber pathway: a review. Neuroscience 98, 407-427 (2000)
73. Sloviter RS, M. A. Dichter, T. L. Rachinsky, E. Dean, J. H. Goodman, A. L. Sollas & D. L. Martin: Basal expression and induction of glutamate decarboxylase and GABA in excitatory granule cells of the rat and monkey hippocampal dentate gyrus. J Comp Neurol 373, 593-618 (1996)
74. Gomez-Lira G, E. Trillo, M. Ramirez, M. Asai, M. Sitges & R. Gutierrez: The expression of GABA in mossy fiber synaptosomes coincides with the seizure-induced expression of GABAergic transmission in the mossy fiber synapse. Exp Neurol 177, 276-283 (2002)
75. Freund TF & G. Buzsaki: Interneurons of the hippocampus. Hippocampus 6, 347-470 (1996)
76. Alfonso J, A. C. Frasch & G. Flugge: Chronic stress, depression and antidepressants: effects on gene transcription in the hippocampus. Rev Neurosci 16, 43-56 (2005)
77. Alfonso J, G. D. Pollevick, M. G. van der Hart, G. Flugge, E. Fuchs E & A. C. Frasch: Identification of genes regulated by chronic psychosocial stress and antidepressant treatment in the hippocampus. Eur J Neurosci 19, 659-666 (2004)
78. Heasley LE, B. Storey, G. R. Fanger, L. Butterfield, J. Zamarripa, D. Blumberg & R. A. Maue: GTPase-deficient G alpha 16 and G alpha q induce PC12 cell differentiation and persistent activation of cJun NH2-terminal kinases. Mal Cell Biol 16, 648-656 (1996)
79. Venero C, T. Tilling, I. Hermans-Borgmeyer, R. Schmidt, M. Schachner & C. Sandi: Chronic stress induces opposite changes in the mRNA expression of the cell adhesion molecules NCAM and L1. Neuroscience 115, 1211-1219 (2002)
80. Ronn LC, V. Berezin & E. Bock: The neural cell adhesion molecule in synaptic plasticity and ageing. Int J Dev Neurosci 18, 193-199 (2000)
81. Bianchi M, C. Heidbreder & F. Crespi: Cytoskeletal changes in the hippocampus following restraint stress: role of serotonin and microtubules. Synapse 49, 188-194 (2003)
82. Goldberg JL & B. A. Barres: The relationship between neuronal survival and regeneration. Annu Rev Neurosci 23, 579-612 (2000)
83. Smith MA, S. Makino, R. Kvetnansky & R. M. Post: Stress and glucocorticoids affect the expression of brain-derived neurotrophic factor and neurotrophin-3 mRNAs in the hippocampus. J Neurosci 15, 1768-1777 (1995)
84. Roceri M, F. Cirulli, C. Pessina, P. Peretto, G. Racagni & M. A. Riva: Postnatal repeated maternal deprivation produces age-dependent changes of brain-derived neurotrophic factor expression in selected rat brain regions. Biol Psychiatry 55, 708-714 (2004)
85. Nibuya M, M. Takahashi, D. S. Russell & R. S. Duman: Repeated stress increases catalytic TrkB mRNA in rat hippocampus. Neurosci Lett 267, 81-84 (1999)
86. Flugge G: Regulation of monoamine receptors in the brain: dynamic changes during stress. Int Rev Cytol 195, 145-213 (2000)
87. Flugge G, M. van Kampen & M. J. Mijnster: Perturbations in brain monoamine systems during stress. Cell Tissue Res 315, 1-14 (2004)
88. Datson NA, J. van der Perk, E. R. de Kloet & E. Vreugdenhil: Identification of corticosteroid-responsive genes in rat hippocampus using serial analysis of gene expression. Eur J Neurosci 14, 675-689 (2001)
89. Lopez JF, D. T. Chalmers, K. Y. Little & S. J. Watson: Regulation of serotonin1A, glucocorticoid, and mineralocorticoid receptor in rat and human hippocampus: implications for the neurobiology of depression. Biol Psychiatry 43, 547-573 (1998)
90. Meyer U, M. van Kampen, E. Isovich, G. Flugge & E. Fuchs: Chronic psychosocial stress regulates the expression of both GR and MR mRNA in the hippocampal formation of tree shrews. Hippocampus 11, 329-336 (2001)
91. Duman RS, G. R. Heninger & E. J. Nestler: A molecular and cellular theory of depression. Arch Gen Psychiatry 54, 597-606 (1997)
92. Rosa ML, F. S. Guimaraes, R. C. Pearson & E. A. Del Bel: Effects of single or repeated restraint stress on GluR1 and GluR2 flip and flop mRNA expression in the hippocampal formation. Brain Res Bull 59, 117-124 (2002)
93. Reagan LP, D. R. Rosell, G. E. Wood, M. Spedding, C. Munoz, J. Rothstein & B. S. McEwen: Chronic restraint stress up-regulates GLT-1 mRNA and protein expression in the rat hippocampus: reversal by tianeptine. Proc Natl Acad Sci USA 101, 2179-2184 (2004)
94. Cullinan WE & T. J. Wolfe: Chronic stress regulates levels of mRNA transcripts encoding beta subunits of the GABA(A) receptor in the rat stress axis. Brain Res 887, 118-124 (2000)
95. Janz R, T. C. Sudhof, R. E. Hammer, V. Unni, S. A. Siegelbaum & V. Y. Bolshakov: Essential roles in synaptic plasticity for synaptogyrin I and synaptophysin I. Neuron 24, 687-700 (1999)
96. Thome J, B. Pesold, M. Baader, M. Hu, J. C. Gewirtz, R. S. Duman & F. A. Henn: Stress differentially regulates synaptophysin and synaptotagmin expression in hippocampus. Biol Psychiatry 50, 809-812 (2001)
97. Andriamampandry C, C. Muller, C. Schmidt-Mutter, S. Gobaille, M. Spedding, D. Aunis & M. Maitre: Mss4 gene is up-regulated in rat brain after chronic treatment with antidepressant and down-regulated when rats are anhedonic. Mol Pharmacol 62, 1332-1338 (2002)
98. Bartolomucci A, P. Palanza, S. Parmigiani, T. Pederzani, E. Merlot, P. J. Neveu & R. Dantzer: Chronic psychosocial stress down-regulates central cytokines mRNA. Brain Res Bull 62, 173-178 (2003)
99. Drevets WC: Neuroplasticity in mood disorders. Dialogues Clin Neurosci 6, 199-216 (2004)
100. Sheline YI, M. H. Gado & H. C. Kraemer: Untreated depression and hippocampal volume loss. Am J Psychiatry 160, 1516-1518 (2003)
101. Manji HK & R. S. Duman: Impairments of neuroplasticity and cellular resilience in severe mood disorders: implications for the development of novel therapeutics. Psychopharmacol Bull 35, 5-49 (2001)
102. Kendler KS, L. M. Karkowski & C. A. Prescott: Causal relationship between stressful life events and the onset of major depression. Am J Psychiatry 156, 837-841 (1999)
103. Diamond DM, A. Campbell, C. R. Park & R. M. Vouimba: Preclinical research on stress, memory, and the brain in the development of pharmacotherapy for depression. Eur Neuropsychopharmacol 14, 491-495 (2004)
104. Sandi C: Stress, cognitive impairment and cell adhesion molecules. Nat Rev Neurosci 5, 917-930 (2004)
105. Bartolomucci A, G. de Biurrun, B. Czeh, M. van Kampen & E. Fuchs: Selective enhancement of spatial learning under chronic psychosocial stress. Eur J Neurosci 15, 1863-1866 (2002)
106. Luine V: Sex differences in chronic stress effects on memory in rats. Stress: The International Journal on the Biology of Stress 5, 205-216 (2002)
107. Sauro MD, R. S. Jorgensen & C. T. Pedlow: Stress, glucocorticoids, and memory: a meta-analytic review. Stress: Int J Biol of Stress 6, 235-245 (2003)