Threat processing: Models and mechanisms

Wiley Interdisciplinary Reviews: Cognitive Science

Volumn 6, Issue 5, 2015, Pages 427-439

  Bentz, Dorothée ^a,b   Schiller, Daniela ^c  

^a UNIVERSITY OF BASEL   (Switzerland)

^b UNIVERSITY OF BASEL   (Switzerland)

^c MOUNT SINAI SCHOOL OF MEDICINE   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

ANIMAL; BRAIN; CONDITIONED REFLEX; FEAR; HUMAN; LEARNING; PHYSIOLOGY; PSYCHOLOGICAL MODEL;

ANIMALS; BRAIN; CONDITIONING, CLASSICAL; FEAR; HUMANS; LEARNING; MODELS, PSYCHOLOGICAL;

EID: 84940959592     PISSN: 19395078     EISSN: 19395086     Source Type: Journal    
DOI: 10.1002/wcs.1353     Document Type: Review

References (134)

1. Johansen JP, Cain CK, Ostroff LE, LeDoux JE. Molecular mechanisms of fear learning and memory. Cell 2011, 147:509-524.
2. Duvarci S, Pare D. Amygdala microcircuits controlling learned fear. Neuron 2014, 82:966-980.
3. Herry C, Johansen JP. Encoding of fear learning and memory in distributed neuronal circuits. Nat Neurosci 2014, 17:1644-1654.
4. Ohman A, Mineka S. Fears, phobias, and preparedness: toward an evolved module of fear and fear learning. Psychol Rev 2001, 108:483-522.
5. Olsson A, Phelps EA. Social learning of fear. Nat Neurosci 2007, 10:1095-1102.
6. Pavlov IPC, Anrep GV. Conditioned Reflexes an Investigation of the Physiological Activity of the Cerebral Cortex. Oxford, England: Oxford University Press; 1927.
7. Rescorla RA, Wagner AR. A theory of Pavlovian conditioning: variations in the effectiveness of reinforcement and nonreinforcement. In: Black AH, Prokasy WF, eds. Classical Conditioning II. New York: Appleton-Century-Crofts; 1972, 64-99.
8. Boucsein W, Fowles DC, Grimnes S, Ben-Shakhar G, Roth WT, Dawson ME, Filion DL. Publication recommendations for electrodermal measurements. Psychophysiology 2012, 49:1017-1034.
9. Davis M, Falls WA, Campeau S, Kim M. Fear-potentiated startle: a neural and pharmacological analysis. Behav Brain Res 1993, 58:175-198.
10. Vurbic D, Bouton ME. Secondary extinction in Pavlovian fear conditioning. Learn Behav 2011, 39:202-211.
11. Kluver H, Bucy PC. Preliminary analysis of functions of the temporal lobes in monkeys. 1939. J Neuropsychiatry Clin Neurosci 1997, 9:606-620.
12. Weiskrantz L. Behavioral changes associated with ablation of the amygdaloid complex in monkeys. J Comp Physiol Psychol 1956, 49:381-391.
13. Blanchard DC, Blanchard RJ. Innate and conditioned reactions to threat in rats with amygdaloid lesions. J Comp Physiol Psychol 1972, 81:281-290.
14. LeDoux JE, Schiller D, Cain CK. Emotional Reaction and Action: From Threat Processing to Goal Directed Behavior. Gazzaniga, MS: MIT Press; 2009.
15. McNally GP, Johansen JP, Blair HT. Placing prediction into the fear circuit. Trends Neurosci 2011, 34:283-292.
16. Sierra-Mercado D, Padilla-Coreano N, Quirk GJ. Dissociable roles of prelimbic and infralimbic cortices, ventral hippocampus, and basolateral amygdala in the expression and extinction of conditioned fear. Neuropsychopharmacology 2011, 36:529-538.
17. Killcross S, Robbins TW, Everitt BJ. Different types of fear-conditioned behavior mediated by separate nuclei within amygdala. Nature 1997, 388:377-380.
18. Cardinal RN, Parkinson JA, Lachenal G, Halkerston KM, Rudarakanchana N, Hall J, Morrison CH, Howes SR, Robbins TW, Everitt BJ. Effects of selective excitotoxic lesions of the nucleus accumbens core, anterior cingulate cortex, and central nucleus of the amygdala on autoshaping performance in rats. Behav Neurosci 2002, 116:553-567.
19. Balleine BW, Killcross S. Parallel incentive processing: an integrated view of amygdala function. Trends Neurosci 2006, 29:272-279.
20. Maren S. Neurotoxic basolateral amygdala lesions impair learning and memory but not the performance of conditional fear in rats. J Neurosci 1999, 19:8696-8703.
21. Lee JL, Dickinson A, Everitt BJ. Conditioned suppression and freezing as measure of aversive Pavlovian conditioning: effects of discrete amygdala lesions and overtraining. Behav Brain Res 2005, 159:221-233.
22. Zimmerman JM, Rabinak CA, McLachlan IG, Maren S. The central nucleus of the amygdala is essential for acquiring and expressing conditional fear after overtraining. Learn Mem 2007, 14:634-644.
23. Phelps EA, LeDoux JE. Contributions of the amygdala to emotion processing: from animal models to human behavior. Neuron 2005, 48:175-187.
24. LaBar KS, LeDoux JE, Spencer DD, Phelps EA. Impaired fear conditioning following unilateral temporal lobectomy in humans. J Neurosci 1995, 15:6846-6855.
25. Adolphs R, Tranel D, Damasio H, Damasio A. Impaired recognition of emotion in facial expressions following bilateral damage to the human amygdala. Nature 1994, 372:669-672.
26. Adolphs R, Tranel D, Damasio AR. The human amygdala in social judgment. Nature 1998, 393:470-474.
27. Damsa C, Kosel M, Moussally J. Current status of brain imaging in anxiety disorders. Curr Opin Psychiatry 2009, 22:96-110.
28. Brohawn KH, Offringa R, Pfaff DL, Hughes KC, Shin LM. The neural correlates of emotional memory in posttraumatic stress disorder. Biol Psychiatry 2010, 68:1023-1030.
29. Parsons RG, Ressler KJ. Implications of memory modulation for post-traumatic stress and fear disorders. Nat Neurosci 2013, 16:146-153.
30. Graham BM, Milad MR. The study of fear extinction: implications for anxiety disorders. Am J Psychiatry 2011, 168:1255-1265.
31. Peri T, Ben-Shakhar G, Orr SP, Shalev AY. Psychophysiologic assessment of aversive conditioning in posttraumatic stress disorder. Biol Psychiatry 2000, 47:512-519.
32. Milad MR, Quirk GJ. Fear extinction as a model for translational neuroscience: ten years of progress. Annu Rev Psychol 2012, 63:129-151.
33. Sehlmeyer C, Schoning S, Zwitserlood P, Pfleiderer B, Kircher T, Arolt V, Konrad C. Human fear conditioning and extinction in neuroimaging: a systematic review. PLoS One 2009, 4:e5865.
34. Shin LM, Rauch SL, Pitman RK. Amygdala, medial prefrontal cortex, and hippocampal function in PTSD. Ann N Y Acad Sci 2006, 1071:67-79.
35. Schiller D, Levy I, Niv Y, LeDoux JE, Phelps EA. From fear to safety and back: reversal of fear in the human brain. J Neurosci 2008, 28:11517-11525.
36. Kalisch R, Korenfeld E, Stephan KE, Weiskopf N, Seymour B, Dolan RJ. Context-dependent human extinction memory is mediated by a ventromedial prefrontal and hippocampal network. J Neurosci 2006, 26:9503-9511.
37. Olsson A, Nearing KI, Phelps EA. Learning fears by observing others: the neural systems of social fear transmission. Soc Cogn Affect Neurosci 2007, 2:3-11.
38. Morris JS, Ohman A, Dolan RJ. Conscious and unconscious emotional learning in the human amygdala. Nature 1998, 393:467-470.
39. Raio CM, Carmel D, Carrasco M, Phelps EA. Nonconscious fear is quickly acquired but swiftly forgotten. Curr Biol 2012, 22:R477-R479.
40. Bach DR, Behrens TE, Garrido L, Weiskopf N, Dolan RJ. Deep and superficial amygdala nuclei projections revealed in vivo by probabilistic tractography. J Neurosci 2011, 31:618-623.
41. Bach DR, Weiskopf N, Dolan RJ. A stable sparse fear memory trace in human amygdala. J Neurosci 2011, 31:9383-9389.
42. Goosens KA, Maren S. NMDA receptors are essential for the acquisition, but not expression, of conditional fear and associative spike firing in the lateral amygdala. Eur J Neurosci 2004, 20:537-548.
43. Burgos-Robles A, Vidal-Gonzalez I, Quirk GJ. Sustained conditioned responses in prelimbic prefrontal neurons are correlated with fear expression and extinction failure. J Neurosci 2009, 29:8474-8482.
44. Milad MR, Quirk GJ, Pitman RK, Orr SP, Fischl B, Rauch SL. A role for the human dorsal anterior cingulate cortex in fear expression. Biol Psychiatry 2007, 62:1191-1194.
45. Hartley CA, Fischl B, Phelps EA. Brain structure correlates of individual differences in the acquisition and inhibition of conditioned fear. Cereb Cortex 2011, 21:1954-1962.
46. Schiller D, Delgado MR. Overlapping neural systems mediating extinction, reversal and regulation of fear. Trends Cogn Sci 2010, 14:268-276.
47. Sutton RS, Barto AG. Reinforcement Learning: An Introduction. Cambridge, MA: MIT Press; 1998.
48. Niv Y, Schoenbaum G. Dialogues on prediction errors. Trends Cogn Sci 2008, 12:265-272.
49. Schultz W, Dayan P, Montague PR. A neural substrate of prediction and reward. Science 1997, 275:1593-1599.
50. Li J, Schiller D, Schoenbaum G, Phelps EA, Daw ND. Differential roles of human striatum and amygdala in associative learning. Nat Neurosci 2011, 14:1250-1252.
51. Pearce JM, Hall G. A model for Pavlovian learning: variations in the effectiveness of conditioned but not of unconditioned stimuli. Psychol Rev 1980, 87:532-552.
52. Roesch MR, Calu DJ, Esber GR, Schoenbaum G. Neural correlates of variations in event processing during learning in basolateral amygdala. J Neurosci 2010, 30:2464-2471.
53. Klavir O, Genud-Gabai R, Paz R. Functional connectivity between amygdala and cingulate cortex for adaptive aversive learning. Neuron 2013, 80:1290-1300.
54. Zhang W, Schneider DM, Belova MA, Morrison SE, Paton JJ, Salzman CD. Functional circuits and anatomical distribution of response properties in the primate amygdala. J Neurosci 2013, 33:722-733.
55. Belova MA, Paton JJ, Morrison SE, Salzman CD. Expectation modulates neural responses to pleasant and aversive stimuli in primate amygdala. Neuron 2007, 55:970-984.
56. Antoniadis EA, Winslow JT, Davis M, Amaral DG. The nonhuman primate amygdala is necessary for the acquisition but not the retention of fear-potentiated startle. Biol Psychiatry 2009, 65:241-248.
57. Ohman A, Eriksson A, Olofsson C. One-trial learning and superior resistance to extinction of autonomic responses conditioned to potentially phobic stimuli. J Comp Physiol Psychol 1975, 88:619-627.
58. Hermans D, Craske MG, Mineka S, Lovibond PF. Extinction in human fear conditioning. Biol Psychiatry 2006, 60:361-368.
59. Quirk GJ, Mueller D. Neural mechanisms of extinction learning and retrieval. Neuropsychopharmacology 2008, 33:56-72.
60. Rescorla RA. Spontaneous recovery. Learn Mem 2004, 11:501-509.
61. Bouton ME. Context, ambiguity, and unlearning: sources of relapse after behavioral extinction. Biol Psychiatry 2002, 52:976-986.
62. Bouton ME. Context, time, and retrieval in the interference paradigms of Pavlovian learning. Psychol Bull 1993, 114:80-99.
63. Phelps EA, Delgado MR, Nearing KI, LeDoux JE. Extinction learning in humans: role of the amygdala and vmPFC. Neuron 2004, 43:897-905.
64. Milad MR, Pitman RK, Ellis CB, Gold AL, Shin LM, Lasko NB, Zeidan MA, Handwerger K, Orr SP, Rauch SL. Neurobiological basis of failure to recall extinction memory in posttraumatic stress disorder. Biol Psychiatry 2009, 66:1075-1082.
65. Milad MR, Rauch SL, Pitman RK, Quirk GJ. Fear extinction in rats: implications for human brain imaging and anxiety disorders. Biol Psychol 2006, 73:61-71.
66. Milad MR, Quirk GJ. Neurons in medial prefrontal cortex signal memory for fear extinction. Nature 2002, 420:70-74.
67. Sotres-Bayon F, Cain CK, LeDoux JE. Brain mechanisms of fear extinction: historical perspectives on the contribution of prefrontal cortex. Biol Psychiatry 2006, 60:329-336.
68. Bouton ME, Woods AM, Todd TP. Separation of time-based and trial-based accounts of the partial reinforcement extinction effect. Behav Process 2014, 101:23-31.
69. Ohman A, Fredrikson M, Hugdahl K, Rimmo PA. The premise of equipotentiality in human classical conditioning: conditioned electrodermal responses to potentially phobic stimuli. J Exp Psychol Gen 1976, 105:313-337.
70. Rescorla RA, Heth CD. Reinstatement of fear to an extinguished conditioned stimulus. J Exp Psychol Anim Behav Process 1975, 1:88-96.
71. Schiller D, Cain CK, Curley NG, Schwartz JS, Stern SA, Ledoux JE, Phelps EA. Evidence for recovery of fear following immediate extinction in rats and humans. Learn Mem 2008, 15:394-402.
72. Mystkowski JL, Mineka S, Vernon LL, Zinbarg RE. Changes in caffeine states enhance return of fear in spider phobia. J Consult Clin Psychol 2003, 71:243-250.
73. Bouton ME, Kenney FA, Rosengard C. State-dependent fear extinction with two benzodiazepine tranquilizers. Behav Neurosci 1990, 104:44-55.
74. Maren S, Phan KL, Liberzon I. The contextual brain: implications for fear conditioning, extinction and psychopathology. Nat Rev Neurosci 2013, 14:417-428.
75. Marschner A, Kalisch R, Vervliet B, Vansteenwegen D, Buchel C. Dissociable roles for the hippocampus and the amygdala in human cued versus context fear conditioning. J Neurosci 2008, 28:9030-9036.
76. McGaugh JL. Time-dependent processes in memory storage. Science 1966, 153:1351-1358.
77. Misanin JR, Miller RR, Lewis DJ. Retrograde amnesia produced by electroconvulsive shock after reactivation of a consolidated memory trace. Science 1968, 160:554-555.
78. Nader K, Schafe GE, Le Doux JE. Fear memories require protein synthesis in the amygdala for reconsolidation after retrieval. Nature 2000, 406:722-726.
79. Alberini CM. The Role of Reconsolidation and the Dynamic Process of Long-Term Memory Formation and Storage. Front Behav Neurosci 2011, 5:1-10.
80. Duvarci S, Nader K. Characterization of fear memory reconsolidation. J Neurosci 2004, 24:9269-9275.
81. Pedreira ME, Perez-Cuesta LM, Maldonado H. Mismatch between what is expected and what actually occurs triggers memory reconsolidation or extinction. Learn Mem 2004, 11:579-585.
82. Wang SH, de Oliveira AL, Nader K. Cellular and systems mechanisms of memory strength as a constraint on auditory fear reconsolidation. Nat Neurosci 2009, 12:905-912.
83. Exton-McGuinness MTJ, Lee JLC, Reichelt AC. Updating memories: the role of prediction errors in memory reconsolidation. Behav Brain Res 2014, 278:375-384.
84. Inda MC, Muravieva EV, Alberini CM. Memory retrieval and the passage of time: from reconsolidation and strengthening to extinction. J Neurosci 2011, 31:1635-1643.
85. Blake MG, Krawczyk MC, Baratti CM, Boccia MM. Neuropharmacology of memory consolidation and reconsolidation: insights on central cholinergic mechanisms. J Physiol Paris 2014, 108:286-291.
86. Dudai Y. The restless engram: consolidations never end. Annu Rev Neurosci 2012, 35:227-247.
87. Tronson NC, Taylor JR. Molecular mechanisms of memory reconsolidation. Nat Rev Neurosci 2007, 8:262-275.
88. Debiec J, Ledoux JE. Disruption of reconsolidation but not consolidation of auditory fear conditioning by noradrenergic blockade in the amygdala. Neuroscience 2004, 129:267-272.
89. Brunet A, Orr SP, Tremblay J, Robertson K, Nader K, Pitman RK. Effect of post-retrieval propranolol on psychophysiologic responding during subsequent script-driven traumatic imagery in post-traumatic stress disorder. J Psychiatr Res 2008, 42:503-506.
90. Kindt M, Soeter M, Vervliet B. Beyond extinction: erasing human fear responses and preventing the return of fear. Nat Neurosci 2009, 12:256-258.
91. Schiller D, Phelps EA. Does reconsolidation occur in humans? Front Behav Neurosci 2011, 5:24.
92. Agren T. Human reconsolidation: a reactivation and update. Brain Res Bull 2014, 105:70-82.
93. Soeter M, Kindt M. Disrupting reconsolidation: pharmacological and behavioral manipulations. Learn Mem 2011, 18:357-366.
94. Lonergan MH, Olivera-Figueroa LA, Pitman RK, Brunet A. Propranolol's effects on the consolidation and reconsolidation of long-term emotional memory in healthy participants: a meta-analysis. J Psychiatry Neurosci 2013, 38:222-231.
95. Soeter M, Kindt M. Dissociating response systems: erasing fear from memory. Neurobiol Learn Mem 2010, 94:30-41.
96. Spring JD, Wood NE, Mueller-Pfeiffer C, Milad MR, Pitman RK, Orr SP. Prereactivation propranolol fails to reduce skin conductance reactivity to prepared fear-conditioned stimuli. Psychophysiology 2014, 52:407-415. doi:10.1111/psyp.12326.
97. Bos MG, Beckers T, Kindt M. Noradrenergic blockade of memory reconsolidation: a failure to reduce conditioned fear responding. Front Behav Neurosci 2014, 8:412.
98. Gelinas JN, Nguyen PV. β-adrenergic receptor activation facilitates induction of a protein synthesis-dependent late phase of long-term potentiation. J Neurosci 2005, 25:3294-3303.
99. Wood NE, Rosasco ML, Suris AM, Spring JD, Marin MF, Lasko NB, Goetz JM, Fischer AM, Orr SP, Pitman RK. Pharmacological blockade of memory reconsolidation in posttraumatic stress disorder: three negative psychophysiological studies. Psychiatry Res 2015, 225:31-39.
100. Bustos SG, Giachero M, Maldonado H, Molina VA. Previous stress attenuates the susceptibility to Midazolam's disruptive effect on fear memory reconsolidation: influence of pre-reactivation D-cycloserine administration. Neuropsychopharmacology 2010, 35:1097-1108.
101. Glover EM, Ressler KJ, Davis M. Differing effects of systemically administered rapamycin on consolidation and reconsolidation of context vs. cued fear memories. Learn Mem 2010, 17:577-581.
102. Suris A, Smith J, Powell C, North CS. Interfering with the reconsolidation of traumatic memory: sirolimus as a novel agent for treating veterans with posttraumatic stress disorder. Ann Clin Psychiatry 2013, 25:33-40.
103. Choi JH, Kim JE, Kaang BK. Protein synthesis and degradation are required for the incorporation of modified information into the pre-existing object-location memory. Mol Brain 2010, 3:1.
104. Monfils MH, Cowansage KK, Klann E, LeDoux JE. Extinction-reconsolidation boundaries: key to persistent attenuation of fear memories. Science 2009, 324:951-955.
105. Schiller D, Monfils MH, Raio CM, Johnson DC, Ledoux JE, Phelps EA. Preventing the return of fear in humans using reconsolidation update mechanisms. Nature 2010, 463:49-53.
106. Clem RL, Huganir RL. Calcium-permeable AMPA receptor dynamics mediate fear memory erasure. Science 2010, 330:1108-1112.
107. Graff J, Joseph NF, Horn ME, Samiei A, Meng J, Seo J, Rei D, Bero AW, Phan TX, Wagner F. Epigenetic priming of memory updating during reconsolidation to attenuate remote fear memories. Cell 2014, 156:261-276.
108. Flavell CR, Barber DJ, Lee JL. Behavioural memory reconsolidation of food and fear memories. Nat Commun 2011, 2:504.
109. Tedesco V, Roquet RF, DeMis J, Chiamulera C, Monfils MH. Extinction, applied after retrieval of auditory fear memory selectively increases zinc-finger protein 268 and phosphorylated ribosomal protein S6 expression in prefrontal cortex and lateral amygdala. Neurobiol Learn Mem 2014, 115:78-85.
110. Olshavsky ME, Jones CE, Lee HJ, Monfils MH. Appetitive behavioral traits and stimulus intensity influence maintenance of conditioned fear. Front Behav Neurosci 2013, 7:179.
111. Agren T, Engman J, Frick A, Bjorkstrand J, Larsson EM, Furmark T, Fredrikson M. Disruption of reconsolidation erases a fear memory trace in the human amygdala. Science 2012, 337:1550-1552.
112. Agren T, Furmark T, Eriksson E, Fredrikson M. Human fear reconsolidation and allelic differences in serotonergic and dopaminergic genes. Transl Psychiatry 2012, 2:e76.
113. Oyarzún JP, Lopez-Barroso D, Fuentemilla L, Cucurell D, Pedraza C, Rodriguez-Fornells A, de Diego-Balaguer R. Updating fearful memories with extinction training during reconsolidation: a human study using auditory aversive stimuli. PLoS One 2012, 7:e38849.
114. Schiller D, Kanen JW, LeDoux JE, Monfils MH, Phelps EA. Extinction during reconsolidation of threat memory diminishes prefrontal cortex involvement. Proc Natl Acad Sci USA 2013, 110:20040-20045.
115. Steinfurth EC, Kanen JW, Raio CM, Clem RL, Huganir RL, Phelps EA. Young and old Pavlovian fear memories can be modified with extinction training during reconsolidation in humans. Learn Mem 2014, 21:338-341.
116. Xue YX, Luo YX, Shi HS, Chen C, Zhu WL, Ding ZB, Bao YP, Shi J, Epsteon DH, Shaham Y, Lu L. A memory retrieval-extinction procedure to prevent drug craving and relapse. Science 2012, 336:241-245.
117. Liu J, Zhao L, Xue Y, Shi J, Suo L, Luo Y, Chai B, Yang C, Fang Q, Zhang Y, et al. An unconditioned stimulus retrieval extinction procedure to prevent the return of fear memory. Biol Psychiatry 2014, 76:895-901.
118. Chan WY, Leung HT, Westbrook RF, McNally GP. Effects of recent exposure to a conditioned stimulus on extinction of Pavlovian fear conditioning. Learn Mem 2010, 17:512-521.
119. Costanzi M, Cannas S, Saraulli D, Rossi-Arnaud C, Cestari V. Extinction after retrieval: effects on the associative and nonassociative components of remote contextual fear memory. Learn Mem 2011, 18:508-518.
120. Perez-Cuesta LM, Maldonado H. Memory reconsolidation and extinction in the crab: mutual exclusion or coexistence? Learn Mem 2009, 16:714-721.
121. Ma X, Zhang JJ, Yu LC. Post-retrieval extinction training enhances or hinders the extinction of morphine-induced conditioned place preference in rats dependent on the retrieval-extinction interval. Psychopharmacology (Berl) 2012, 221:19-26.
122. Kindt M, Soeter M. Reconsolidation in a human fear conditioning study: a test of extinction as updating mechanism. Biol Psychol 2013, 92:43-50.
123. Auber A, Tedesco V, Jones CE, Monfils MH, Chiamulera C. Post-retrieval extinction as reconsolidation interference: methodological issues or boundary conditions? Psychopharmacology (Berl) 2013, 226:631-647.
124. Flavell CR, Lambert EA, Winters BD, Bredy TW. Mechanisms governing the reactivation-dependent destabilization of memories and their role in extinction. Front Behav Neurosci 2013, 7:214.
125. Eisenberg M, Kobilo T, Berman DE, Dudai Y. Stability of retrieved memory: inverse correlation with trace dominance. Science 2003, 301:1102-1104.
126. Merlo E, Milton AL, Goozée ZY, Theobald DE, Everitt BJ. Reconsolidation and extinction are dissociable and mutually exclusive processes: behavioral and molecular evidence. J Neurosci 2014, 34:2422-2431.
127. Pigott TA. Anxiety disorders in women. Psychiatr Clin North Am 2003, 26:621-672.
128. Bentz D, Michael T, de Quervain DJ, Wilhelm FH. Enhancing exposure therapy for anxiety disorders with glucocorticoids: from basic mechanisms of emotional learning to clinical applications. J Anxiety Disord 2010, 24:223-230.
129. Graham BM, Langton JM, Richardson R. Pharmacological enhancement of fear reduction: preclinical models. Br J Pharmacol 2011, 164:1230-1247.
130. Besnard A, Caboche J, Laroche S. Reconsolidation of memory: a decade of debate. Prog Neurobiol 2012, 99:61-80.
131. Delgado MR, Li J, Schiller D, Phelps EA. The role of the striatum in aversive learning and aversive prediction errors. Philos Trans R Soc Lond B Biol Sci 2008, 363:3787-3800.
132. Mineka S, Oehlberg K. The relevance of recent developments in classical conditioning to understanding the etiology and maintenance of anxiety disorders. Acta Psychol (Amst) 2008, 127:567-580.
133. Morrison FG, Ressler KJ. From the neurobiology of extinction to improved clinical treatments. Depress Anxiety 2014, 31:279-290.
134. Pare D, Duvarci S. Amygdala microcircuits mediating fear expression and extinction. Curr Opin Neurobiol 2012, 22:717-723.