条件性恐惧记忆消退的提取干预范式及其作用的神经机制
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摘要
基于记忆再巩固理论的恐惧记忆提取干预范式被证明可以有效消退恐惧记忆,能克服传统消退容易复发的缺点。该范式通过单独呈现条件刺激激活原有恐惧记忆,使记忆重返不稳定状态,随后在再巩固时间窗内实施干预则能改写原有记忆。目前该范式起作用的神经机制尚不明确,本文在现有的人类研究和动物研究基础上,总结了杏仁核、前额叶和海马三个脑区在提取干预过程中的作用,以及该领域研究的争议点,为之后的研究提供思路。
The memory reconsolidation theory holds that consolidated memories can be reactivated into an unstable state again, which is called the reconsolidation process. During this period, the activated memories are susceptive to new information and therefore, the interruption of it can update or eliminate original memories. Behavioral or pharmacological interventions after memory reactivation are proved to be an effective way to extinguish fear memories and prevent relapse. The behavioral intervention, which is the so-called retrieval-extinction paradigm can eliminate conditioned fear through applying extinction training during the reconsolidation time window. The neural mechanisms of this effect, however, are still largely unknown. Based on reviewing present studies in animals and humans, we concluded the critical brain areas which were proved to be playing an important part in retrieval extinction process and their effect mechanisms, including hippocampus, prefrontal cortex and amygdala. Furthermore, we summarized the controversial points in theories and manipulations in order to provide insights for future explorations.
The memory reconsolidation theory holds that consolidated memories can be reactivated into an unstable state again, which is called the reconsolidation process. During this period, the activated memories are susceptive to new information and therefore, the interruption of it can update or eliminate original memories. Behavioral or pharmacological interventions after memory reactivation are proved to be an effective way to extinguish fear memories and prevent relapse. The behavioral intervention, which is the so-called retrieval-extinction paradigm can eliminate conditioned fear through applying extinction training during the reconsolidation time window. The neural mechanisms of this effect, however, are still largely unknown. Based on reviewing present studies in animals and humans, we concluded the critical brain areas which were proved to be playing an important part in retrieval extinction process and their effect mechanisms, including hippocampus, prefrontal cortex and amygdala. Furthermore, we summarized the controversial points in theories and manipulations in order to provide insights for future explorations.
引文
安献丽,郑希耕.(2008).创伤后应激障碍的动物模型及其神经生物学机制.心理科学进展,16(3),371-377.
陈伟,李俊娇,曹杨婧文,杨勇,胡琰健,郑希付.(2018).预期错误在复合恐惧记忆提取消退中的作用.心理学报,50(7),739-749.
吴艳,李勇辉,隋南.(2009).记忆再巩固现象及其生物学机制.心理科学进展,17(4),699-705.
李平,王晓琴,孙振武,王功伍.(2018).药物成瘾记忆及其神经环路研究进展.生物学杂志,35(1),93-96+100.
曾祥星,杜娟,王凯欣,郑希付.(2015).记忆再巩固的时间动态性及其生物学机制.心理科学进展,23(4),582-590.
曾祥星,向燕辉,杜娟,郑希付.(2014).条件性恐惧记忆提取消退干预范式.心理科学进展,22(3),431-438.
庄楚群,王文清,胡静初,张蔚欣,王鹏贵,郑希付.(2017).提取-消退范式中复合刺激对恐惧消退的影响.心理学报,49(3),329-335.
Agren,T.,Engman,J.,Frick,A.,Bj?rkstrand,J.,Larsson,E-M.,Furmark,T.,&Fredrikson,M.(2012).Disruption of reconsolidation erases a fear memory trace in the human amygdala.Science,337(6101),1550-1552.
Asem,J.S.A.,Schiffino,F.L.,&Holland,P.C.(2015).Dorsolateral striatum is critical for the expression of surprise-induced enhancements in cue associability.European Journal of Neuroscience,42(5),2203-2213.
Beckers,T.,&Kindt,M.(2017).Memory reconsolidation interference as an emerging treatment for emotional disorders:Strengths,limitations,challenges,and opportunities.Annual Review of Clinical Psychology,13(1),99-121.
Besnard,A.,Caboche,J.,&Laroche,S.(2012).Reconsolidation of memory:A decade of debate.Progress in Neurobiology,99(1),61-80.
Bj?rkstrand,J.,Agren,T.,?hs,F.,Frick,A.,Larsson,E-M.,Hjorth,O.,...Fredrikson,M.(2016).Disrupting reconsolidation attenuates long-term fear memory in the human amygdala and facilitates approach behavior.Current Biology,26(19),2690-2695.
Bj?rkstrand,J.,Agren,T.,?hs,F.,Frick,A.,Larsson,E-M.,Hjorth,O.,...Fredrikson,M.(2017).Think twice,it's all right:Long lasting effects of disrupted reconsolidation on brain and behavior in human long-term fear.Behavioural Brain Research,324,125-129.
Bj?rkstrand,J.,Agren,T.,Frick,A.,Engman,J.,Larsson,E-M.,Furmark,T.,&Fredrikson,M.(2015).Disruption of memory reconsolidation erases a fear memory trace in the human amygdala:An 18-month follow-up.Plos One,10(7),e0129393.
Boes,A.D.,Grafft,A.H.,Joshi,C.,Chuang,N.A.,Nopoulos,P.,&Anderson,S.W.(2011).Behavioral effects of congenital ventromedial prefrontal cortex malformation.BMC Neurology,11,151.
Bouton,M.E.,García-gutiérrez,A.,Zilski,J.,&Moody,E.W.(2006).Extinction in multiple contexts does not necessarily make extinction less vulnerable to relapse.Behaviour Research and Therapy,44(7),983-994.
Chan,W.Y.M.,Leung,H.T.,Westbrook,R.F.,&Mcnally,G.P.(2010).Effects of recent exposure to a conditioned stimulus on extinction of Pavlovian fear conditioning.Learning and Memory,17(10),512-521.
Cheng,D.T.,Knight,D.C.,Smith,C.N.,Stein,E.A.,&Helmstetter,F.J.(2003).Functional MRI of human amygdala activity during Pavlovian fear conditioning:stimulus processing versus response expression.Behavioral Neuroscience,117(1),3-10.
Costanzi,M.,Cannas,S.,Saraulli,D.,Rossi-Arnaud,C.,&Cestari,V.(2011).Extinction after retrieval:Effects on the associative and nonassociative components of remote contextual fear memory.Learning and Memory,18(8),508-518.
Critchley,H.D.(2009).Psychophysiology of neural,cognitive and affective integration:FMRI and autonomic indicants.International Journal of Psychophysiology,73(2),88-94.
Debiec,J.,Ledoux,J.E.,&Nader,K.(2002).Cellular and systems reconsolidation in the hippocampus.Neuron,36(3),527-538.
Díaz-Mataix,L.,Tallot,L.,&Doyère,V.(2014).The amygdala:A potential player in timing CS-US intervals.Behavioural Processes,101,112-122.
Do-monte,F.H.,Qui?ones-laracuente,K.,&Quirk,G.J.(2015).A temporal shift in the circuits mediating retrieval of fear memory.Nature,519(7544),460-463.
Dudai,Y.(2004).The neurobiology of consolidations,or,how stable is the engram?Annual Review of Psychology,55(1),51-86.
Duvarci,S.,&Nader,K.(2004).Characterization of fear memory reconsolidation.Journal of Neuroscience the Official Journal of the Society for Neuroscience,24(42),9269-9275.
Elsey,J.W.B.,&Kindt,M.(2017a).Breaking boundaries:optimizing reconsolidation-based interventions for strong and old memories.Learning and Memory,24(9),472-479.
Elsey,J.W.B.,&Kindt,M.(2017b).Tackling maladaptive memories through reconsolidation:From neural to clinical science.Neurobiology of Learning and Memory,142,108-117.
Etkin,A.,Egner,T.,&Kalisch,R.(2011).Emotional processing in anterior cingulate and medial prefrontal cortex.Trends in Cognitiveences,15(2),85-93.
Ferrer Monti,R.I.,Alfei,J.M.,Mugnaini,M.,Bueno,A.M.,Beckers,T.,Urcelay,G.P.,&Molina,V.A.(2017).Acomparison of behavioral and pharmacological interventions to attenuate reactivated fear memories.Learning and Memory,24(8),369-374.
Foa,E.B.,&Kozak,M.J.(1986).Emotional processing of fear:Exposure to corrective information.Psychological Bulletin,99(1),20-35.
Fullana,M.A.,Albajes-eizagirre,A.,Soriano-mas,C.,Vervliet,B.,Cardoner,N.,Benet,O.,...Harrison,B.J.(2018).Fear extinction in the human brain:Ameta-analysis of fMRI studies in healthy participants.Neuroscience&Biobehavioral Reviews,88,16-25.
Fullana,M.A.,Harrison,B.J.,Soriano-mas,C.,Vervliet,B.,Cardoner,N.,àvila-parcet,A.,&Radua,J.(2016).Neural signatures of human fear conditioning:an updated and extended meta-analysis of fMRI studies.Mol Psychiatry,21(4),500-508.
Germeroth,L.J.,Carpenter,M.J.,Baker,N.L.,Froeliger,B.,Larowe,S.D.,&Saladin,M.E.(2017).Effect of a brief memory updating intervention on smoking behavior:Arandomized clinical trial.Jama Psychiatry,74(3),214-223.
Goode,T.D.,Holloway-Erickson,C.M.,&Maren,S.(2017).Extinction after fear memory reactivation fails to eliminate renewal in rats.Neurobiology of Learning and Memory,142(Pt A),41-47.
Gr?ff,J.,Joseph,N.F.,Horn,M.E.,Samiei,A.,Jia,M.,Seo,J.,...Tsai,L-H.(2014).Epigenetic priming of memory updating during reconsolidation to attenuate remote fear memories.Cell,156(1-2),261-276.
Herry,C.,Ciocchi,S.,Senn,V.,Demmou,L.,Müller,C.,&Lüthi,A.(2008).Switching on and off fear by distinct neuronal circuits.Nature,454(7204),600-606.
Hofmann,S.G.(2008).Cognitive processes during fear acquisition and extinction in animals and humans:Implications for exposure therapy of anxiety disorders.Clinical Psychology Review,28(2),199-210.
Hu,J.,Wang,W.,Homan,P.,Wang,P.,Zheng,X.,&Schiller,D.(2018).Reminder duration determines threat memory modification in humans.Sci Rep,8(1),8848.
Ishii,D.,Matsuzawa,D.,Matsuda,S.,Tomizawa,H.,Sutoh,C.,&Shimizu,E.(2012).No erasure effect of retrievalextinction trial on fear memory in the hippocampusindependent and dependent paradigms.Neuroscience Letters,523(1),76-81.
Ishikawa,A.,&Nakamura,S.(2003).Convergence and interaction of hippocampal and amygdalar projections within the prefrontal cortex in the rat.Journal of Neuroscience,23(31),9987-9995.
Kim,J.J.,&Jung,M.W.(2006).Neural circuits and mechanisms involved in Pavlovian fear conditioning:Acritical review.Neuroscience&Biobehavioral Reviews,30(2),188-202.
Kindt,M.,&Soeter,M.(2013).Reconsolidation in a human fear conditioning study:A test of extinction as updating mechanism.Biological Psychology,92(1),43-50.
Kindt,M.,Soeter,M.,&Vervliet,B.(2009).Beyond extinction:Erasing human fear responses and preventing the return of fear.Nature Neuroscience,12(3),256-258.
Knight,D.C.,Smith,C.N.,Cheng,D.T.,Stein,E.A.,&Helmstetter,F.J.(2004).Amygdala and hippocampal activity during acquisition and extinction of human fear conditioning.Cognitive Affective&Behavioral Neuroscience,4(3),317-325.
Koenigs,M.,Huey,E.D.,Raymont,V.,Cheon,B.,Solomon,J.,Wassermann,E.M.,&Grafman,J.(2008).Focal brain damage protects against post-traumatic stress disorder in combat veterans.Nature Neuroscience,11(2),232-237.
Kroes,M.C.,Tona,K-D.,den Ouden,H.E.M.,Vogel,S.,van Wingen,G.A.,&Fernández,G.(2016).How administration of the beta-blocker propranolol before extinction can prevent the return of fear.Neuropsychopharmacology Official Publication of the American College of Neuropsychopharmacology,41(6),1569-1578.
Laurent,V.,&Westbrook,R.F.(2009).Inactivation of the infralimbic but not the prelimbic cortex impairs consolidation and retrieval of fear extinction.Learning and Memory,16(9),520-529.
Li,J.,Chen,W.,Caoyang,J.,Wu,W.,Jie,J.,Xu,L.,&Zheng,X.(2017).Moderate partially reduplicated conditioned stimuli as retrieval cue can increase effect on preventing relapse of fear to compound stimuli.Frontiers in Human Neuroscience,11,575.
Li,S.S.Y.,&Mcnally,G.P.(2014).The conditions that promote fear learning:Prediction error and Pavlovian fear conditioning.Neurobiology of Learning and Memory,108(1),14-21.
Liu,J.,Zhao,L.,Xue,Y.,Shi,J.,Suo,L.,Luo,Y.,...Lu,L.(2014).An unconditioned stimulus retrieval extinction procedure to prevent the return of fear memory.Biological Psychiatry,76(11),895-901.
Medford,N.,&Critchley,H.D.(2010).Conjoint activity of anterior insular and anterior cingulate cortex:awareness and response.Brain Structure&Function,214(5-6),535-549.
Milad,M.R.,Pitman,R.K.,Ellis,C.B.,Gold,A.L.,Shin,L.M.,Lasko,N.B.,...Rauch,S.L.(2009).Neurobiological basis of failure to recall extinction memory in posttraumatic stress disorder.Biological Psychiatry,66(12),1075-1082.
Milad,M.R.,&Quirk,G.J.(2002).Neurons in medial prefrontal cortex signal memory for fear extinction.Nature,420(6911),70-74.
Milad,M.R.,Wright,C.I.,Orr,S.P.,Pitman,R.K.,Quirk,G.J.,&Rauch,S.L.(2007).Recall of fear extinction in humans activates the ventromedial prefrontal cortex and hippocampus in concert.Biological Psychiatry,62(5),446-454.
Monfils,M.H.,Cowansage,K.K.,Klann,E.,&Ledoux,J.E.(2009).Extinction-reconsolidation boundaries:Key to persistent attenuation of fear memories.Science,324(5929),951-955.
Morris,J.S.,Friston,K.J.,Büchel,C.,Frith,C.D.,Young,A.W.,Calder,A.J.,&Dolan,R.J.(1998).Aneuromodulatory role for the human amygdala in processing emotional facial expressions.Brain,121(1),47-57.
Myers,K.M.,&Davis,M.(2002).Behavioral and neural analysis of extinction.Neuron,36(4),567-584.
Nader,K.(2003).Memory traces unbound.Trends in Neurosciences,26(2),65-72.
Nader,K.,Schafe,G.E.,&Doux,J.E.(2000).Fear memories require protein synthesis in the amygdala for reconsolidation after retrieval.Nature,406(6797),722-726.
Orsini,C.A.,&Maren,S.(2012).Neural and cellular mechanisms of fear and extinction memory formation.Neuroscience&Biobehavioral Reviews,36(7),1773-1802.
Oyarzún,J.P.,Lopez-barroso,D.,Fuentemilla,L.,Cucurell,D.,Pedraza,C.,Rodriguez-fornells,A.,&Diego-balaguer,R.D.(2012).Updating fearful memories with extinction training during reconsolidation:A human study using auditory aversive stimuli.Plos One,7(6),e38849.
Penzo,M.A.,Robert,V.,Tucciarone,J.,De Bundel,D.,Wang,M.,Van Aelst,L.,...Li,B.(2015).The paraventricular thalamus controls a central amygdala fear circuit.Nature,519(7544),455-459.
Phelps,E.A.,Delgado,M.R.,Nearing,K.I.,&Ledoux,J.E.(2004).Extinction learning in humans:Role of the amygdala and vmPFC.Neuron,43(6),897-905.
Quirk,G.J.,Russo,G.K.,Barron,J.L.,&Lebron,K.(2000).The role of ventromedial prefrontal cortex in the recovery of extinguished fear.Journal of Neuroscience,20(16),6225-6231.
Reichelt,A.C.,&Lee,J.L.C.(2013).Memory reconsolidation in aversive and appetitive settings.Frontiers in Behavioral Neuroscience,7(7),118.
Rescorla,R.A.,&Wagner,A.R.(1972).A theory of Pavlovian conditioning:The effectiveness of reinforcement and non-reinforcement.Paper presented at the Classical Conditioning Ii:Current Theory&Research In:Black Ah,Prokasy Wf.
Sara,S.J.(2000).Retrieval and reconsolidation:Toward a neurobiology of remembering.Learning&Memory,7(2),73-84.
Schiller,D.,Kanen,J.W.,Ledoux,J.E.,Monfils,M-H.,&Phelps,E.A.(2013).Extinction during reconsolidation of threat memory diminishes prefrontal cortex involvement.Proceedings of the National Academy of Sciences of the United States of America,110(50),20040-20045.
Schiller,D.,Levy,I.,Niv,Y.,Ledoux,J.E.,&Phelps,E.A.(2008).From fear to safety and back:reversal of fear in the human brain.Journal of Neuroscience,28(45),11517.
Schiller,D.,Monfils,M.,Raio,C.M.,Johnson,D.C.,Ledoux,J.E.,&Phelps,E.A.(2010).Preventing the return of fear in humans using reconsolidation update mechanisms.Nature,463(7277),49.
Sehlmeyer,C.,Dannlowski,U.,Sch?ning,S.,Kugel,H.,Pyka,M.,Pfleiderer,B.,...Konrad,C.(2011).Neural correlates of trait anxiety in fear extinction.Psychological Medicine,41(4),789-798.
Sehlmeyer,C.,Sch?ning,S.,Zwitserlood,P.,Pfleiderer,B.,Kircher,T.,Arolt,V.,&Konrad,C.(2009).Human fear conditioning and extinction in neuroimaging:A systematic review.Plos One,4(6),e5865.
Sevenster,D.,Beckers,T.,&Kindt,M.(2013).Prediction error governs pharmacologically induced amnesia for learned fear.Science,339(6121),830-833.
Soeter,M.,&Kindt,M.(2015).An abrupt transformation of phobic behavior after a post-retrieval amnesic agent.Biological Psychiatry,78(12),880-886.
Tovote,P.,Fadok,J.P.,&Lüthi,A.(2015).Neuronal circuits for fear and anxiety.Nature Reviews Neuroscience,16(6),317-331.
Xue,Y-X.,Deng,J-H.,Chen,Y-Y.,Zhang,L-B.,Wu,P.,Huang,G-D.,...Lu,L(2017).Effect of selective inhibition of reactivated nicotine-associated memories with propranolol on nicotine craving.Jama Psychiatry,74(3),224-232.
Xue,Y-X.,Luo,Y-X.,Wu,P.,Shi,H-S.,Xue,L-F.,Chen,C.,...Lu,L.(2012).A memory retrieval-extinction procedure to prevent drug craving and relapse.Science,336(6078),241-245.
Zeng,X-X.,Du,J.,Zhuang,C-Q.,Zhang,J-H.,Jia,Y-L.,&Zheng,X-F.(2014).Unconditioned stimulus revaluation to promote conditioned fear extinction in the memory reconsolidation window.Plos One,9(7),e101589.
陈伟,李俊娇,曹杨婧文,杨勇,胡琰健,郑希付.(2018).预期错误在复合恐惧记忆提取消退中的作用.心理学报,50(7),739-749.
吴艳,李勇辉,隋南.(2009).记忆再巩固现象及其生物学机制.心理科学进展,17(4),699-705.
李平,王晓琴,孙振武,王功伍.(2018).药物成瘾记忆及其神经环路研究进展.生物学杂志,35(1),93-96+100.
曾祥星,杜娟,王凯欣,郑希付.(2015).记忆再巩固的时间动态性及其生物学机制.心理科学进展,23(4),582-590.
曾祥星,向燕辉,杜娟,郑希付.(2014).条件性恐惧记忆提取消退干预范式.心理科学进展,22(3),431-438.
庄楚群,王文清,胡静初,张蔚欣,王鹏贵,郑希付.(2017).提取-消退范式中复合刺激对恐惧消退的影响.心理学报,49(3),329-335.
Agren,T.,Engman,J.,Frick,A.,Bj?rkstrand,J.,Larsson,E-M.,Furmark,T.,&Fredrikson,M.(2012).Disruption of reconsolidation erases a fear memory trace in the human amygdala.Science,337(6101),1550-1552.
Asem,J.S.A.,Schiffino,F.L.,&Holland,P.C.(2015).Dorsolateral striatum is critical for the expression of surprise-induced enhancements in cue associability.European Journal of Neuroscience,42(5),2203-2213.
Beckers,T.,&Kindt,M.(2017).Memory reconsolidation interference as an emerging treatment for emotional disorders:Strengths,limitations,challenges,and opportunities.Annual Review of Clinical Psychology,13(1),99-121.
Besnard,A.,Caboche,J.,&Laroche,S.(2012).Reconsolidation of memory:A decade of debate.Progress in Neurobiology,99(1),61-80.
Bj?rkstrand,J.,Agren,T.,?hs,F.,Frick,A.,Larsson,E-M.,Hjorth,O.,...Fredrikson,M.(2016).Disrupting reconsolidation attenuates long-term fear memory in the human amygdala and facilitates approach behavior.Current Biology,26(19),2690-2695.
Bj?rkstrand,J.,Agren,T.,?hs,F.,Frick,A.,Larsson,E-M.,Hjorth,O.,...Fredrikson,M.(2017).Think twice,it's all right:Long lasting effects of disrupted reconsolidation on brain and behavior in human long-term fear.Behavioural Brain Research,324,125-129.
Bj?rkstrand,J.,Agren,T.,Frick,A.,Engman,J.,Larsson,E-M.,Furmark,T.,&Fredrikson,M.(2015).Disruption of memory reconsolidation erases a fear memory trace in the human amygdala:An 18-month follow-up.Plos One,10(7),e0129393.
Boes,A.D.,Grafft,A.H.,Joshi,C.,Chuang,N.A.,Nopoulos,P.,&Anderson,S.W.(2011).Behavioral effects of congenital ventromedial prefrontal cortex malformation.BMC Neurology,11,151.
Bouton,M.E.,García-gutiérrez,A.,Zilski,J.,&Moody,E.W.(2006).Extinction in multiple contexts does not necessarily make extinction less vulnerable to relapse.Behaviour Research and Therapy,44(7),983-994.
Chan,W.Y.M.,Leung,H.T.,Westbrook,R.F.,&Mcnally,G.P.(2010).Effects of recent exposure to a conditioned stimulus on extinction of Pavlovian fear conditioning.Learning and Memory,17(10),512-521.
Cheng,D.T.,Knight,D.C.,Smith,C.N.,Stein,E.A.,&Helmstetter,F.J.(2003).Functional MRI of human amygdala activity during Pavlovian fear conditioning:stimulus processing versus response expression.Behavioral Neuroscience,117(1),3-10.
Costanzi,M.,Cannas,S.,Saraulli,D.,Rossi-Arnaud,C.,&Cestari,V.(2011).Extinction after retrieval:Effects on the associative and nonassociative components of remote contextual fear memory.Learning and Memory,18(8),508-518.
Critchley,H.D.(2009).Psychophysiology of neural,cognitive and affective integration:FMRI and autonomic indicants.International Journal of Psychophysiology,73(2),88-94.
Debiec,J.,Ledoux,J.E.,&Nader,K.(2002).Cellular and systems reconsolidation in the hippocampus.Neuron,36(3),527-538.
Díaz-Mataix,L.,Tallot,L.,&Doyère,V.(2014).The amygdala:A potential player in timing CS-US intervals.Behavioural Processes,101,112-122.
Do-monte,F.H.,Qui?ones-laracuente,K.,&Quirk,G.J.(2015).A temporal shift in the circuits mediating retrieval of fear memory.Nature,519(7544),460-463.
Dudai,Y.(2004).The neurobiology of consolidations,or,how stable is the engram?Annual Review of Psychology,55(1),51-86.
Duvarci,S.,&Nader,K.(2004).Characterization of fear memory reconsolidation.Journal of Neuroscience the Official Journal of the Society for Neuroscience,24(42),9269-9275.
Elsey,J.W.B.,&Kindt,M.(2017a).Breaking boundaries:optimizing reconsolidation-based interventions for strong and old memories.Learning and Memory,24(9),472-479.
Elsey,J.W.B.,&Kindt,M.(2017b).Tackling maladaptive memories through reconsolidation:From neural to clinical science.Neurobiology of Learning and Memory,142,108-117.
Etkin,A.,Egner,T.,&Kalisch,R.(2011).Emotional processing in anterior cingulate and medial prefrontal cortex.Trends in Cognitiveences,15(2),85-93.
Ferrer Monti,R.I.,Alfei,J.M.,Mugnaini,M.,Bueno,A.M.,Beckers,T.,Urcelay,G.P.,&Molina,V.A.(2017).Acomparison of behavioral and pharmacological interventions to attenuate reactivated fear memories.Learning and Memory,24(8),369-374.
Foa,E.B.,&Kozak,M.J.(1986).Emotional processing of fear:Exposure to corrective information.Psychological Bulletin,99(1),20-35.
Fullana,M.A.,Albajes-eizagirre,A.,Soriano-mas,C.,Vervliet,B.,Cardoner,N.,Benet,O.,...Harrison,B.J.(2018).Fear extinction in the human brain:Ameta-analysis of fMRI studies in healthy participants.Neuroscience&Biobehavioral Reviews,88,16-25.
Fullana,M.A.,Harrison,B.J.,Soriano-mas,C.,Vervliet,B.,Cardoner,N.,àvila-parcet,A.,&Radua,J.(2016).Neural signatures of human fear conditioning:an updated and extended meta-analysis of fMRI studies.Mol Psychiatry,21(4),500-508.
Germeroth,L.J.,Carpenter,M.J.,Baker,N.L.,Froeliger,B.,Larowe,S.D.,&Saladin,M.E.(2017).Effect of a brief memory updating intervention on smoking behavior:Arandomized clinical trial.Jama Psychiatry,74(3),214-223.
Goode,T.D.,Holloway-Erickson,C.M.,&Maren,S.(2017).Extinction after fear memory reactivation fails to eliminate renewal in rats.Neurobiology of Learning and Memory,142(Pt A),41-47.
Gr?ff,J.,Joseph,N.F.,Horn,M.E.,Samiei,A.,Jia,M.,Seo,J.,...Tsai,L-H.(2014).Epigenetic priming of memory updating during reconsolidation to attenuate remote fear memories.Cell,156(1-2),261-276.
Herry,C.,Ciocchi,S.,Senn,V.,Demmou,L.,Müller,C.,&Lüthi,A.(2008).Switching on and off fear by distinct neuronal circuits.Nature,454(7204),600-606.
Hofmann,S.G.(2008).Cognitive processes during fear acquisition and extinction in animals and humans:Implications for exposure therapy of anxiety disorders.Clinical Psychology Review,28(2),199-210.
Hu,J.,Wang,W.,Homan,P.,Wang,P.,Zheng,X.,&Schiller,D.(2018).Reminder duration determines threat memory modification in humans.Sci Rep,8(1),8848.
Ishii,D.,Matsuzawa,D.,Matsuda,S.,Tomizawa,H.,Sutoh,C.,&Shimizu,E.(2012).No erasure effect of retrievalextinction trial on fear memory in the hippocampusindependent and dependent paradigms.Neuroscience Letters,523(1),76-81.
Ishikawa,A.,&Nakamura,S.(2003).Convergence and interaction of hippocampal and amygdalar projections within the prefrontal cortex in the rat.Journal of Neuroscience,23(31),9987-9995.
Kim,J.J.,&Jung,M.W.(2006).Neural circuits and mechanisms involved in Pavlovian fear conditioning:Acritical review.Neuroscience&Biobehavioral Reviews,30(2),188-202.
Kindt,M.,&Soeter,M.(2013).Reconsolidation in a human fear conditioning study:A test of extinction as updating mechanism.Biological Psychology,92(1),43-50.
Kindt,M.,Soeter,M.,&Vervliet,B.(2009).Beyond extinction:Erasing human fear responses and preventing the return of fear.Nature Neuroscience,12(3),256-258.
Knight,D.C.,Smith,C.N.,Cheng,D.T.,Stein,E.A.,&Helmstetter,F.J.(2004).Amygdala and hippocampal activity during acquisition and extinction of human fear conditioning.Cognitive Affective&Behavioral Neuroscience,4(3),317-325.
Koenigs,M.,Huey,E.D.,Raymont,V.,Cheon,B.,Solomon,J.,Wassermann,E.M.,&Grafman,J.(2008).Focal brain damage protects against post-traumatic stress disorder in combat veterans.Nature Neuroscience,11(2),232-237.
Kroes,M.C.,Tona,K-D.,den Ouden,H.E.M.,Vogel,S.,van Wingen,G.A.,&Fernández,G.(2016).How administration of the beta-blocker propranolol before extinction can prevent the return of fear.Neuropsychopharmacology Official Publication of the American College of Neuropsychopharmacology,41(6),1569-1578.
Laurent,V.,&Westbrook,R.F.(2009).Inactivation of the infralimbic but not the prelimbic cortex impairs consolidation and retrieval of fear extinction.Learning and Memory,16(9),520-529.
Li,J.,Chen,W.,Caoyang,J.,Wu,W.,Jie,J.,Xu,L.,&Zheng,X.(2017).Moderate partially reduplicated conditioned stimuli as retrieval cue can increase effect on preventing relapse of fear to compound stimuli.Frontiers in Human Neuroscience,11,575.
Li,S.S.Y.,&Mcnally,G.P.(2014).The conditions that promote fear learning:Prediction error and Pavlovian fear conditioning.Neurobiology of Learning and Memory,108(1),14-21.
Liu,J.,Zhao,L.,Xue,Y.,Shi,J.,Suo,L.,Luo,Y.,...Lu,L.(2014).An unconditioned stimulus retrieval extinction procedure to prevent the return of fear memory.Biological Psychiatry,76(11),895-901.
Medford,N.,&Critchley,H.D.(2010).Conjoint activity of anterior insular and anterior cingulate cortex:awareness and response.Brain Structure&Function,214(5-6),535-549.
Milad,M.R.,Pitman,R.K.,Ellis,C.B.,Gold,A.L.,Shin,L.M.,Lasko,N.B.,...Rauch,S.L.(2009).Neurobiological basis of failure to recall extinction memory in posttraumatic stress disorder.Biological Psychiatry,66(12),1075-1082.
Milad,M.R.,&Quirk,G.J.(2002).Neurons in medial prefrontal cortex signal memory for fear extinction.Nature,420(6911),70-74.
Milad,M.R.,Wright,C.I.,Orr,S.P.,Pitman,R.K.,Quirk,G.J.,&Rauch,S.L.(2007).Recall of fear extinction in humans activates the ventromedial prefrontal cortex and hippocampus in concert.Biological Psychiatry,62(5),446-454.
Monfils,M.H.,Cowansage,K.K.,Klann,E.,&Ledoux,J.E.(2009).Extinction-reconsolidation boundaries:Key to persistent attenuation of fear memories.Science,324(5929),951-955.
Morris,J.S.,Friston,K.J.,Büchel,C.,Frith,C.D.,Young,A.W.,Calder,A.J.,&Dolan,R.J.(1998).Aneuromodulatory role for the human amygdala in processing emotional facial expressions.Brain,121(1),47-57.
Myers,K.M.,&Davis,M.(2002).Behavioral and neural analysis of extinction.Neuron,36(4),567-584.
Nader,K.(2003).Memory traces unbound.Trends in Neurosciences,26(2),65-72.
Nader,K.,Schafe,G.E.,&Doux,J.E.(2000).Fear memories require protein synthesis in the amygdala for reconsolidation after retrieval.Nature,406(6797),722-726.
Orsini,C.A.,&Maren,S.(2012).Neural and cellular mechanisms of fear and extinction memory formation.Neuroscience&Biobehavioral Reviews,36(7),1773-1802.
Oyarzún,J.P.,Lopez-barroso,D.,Fuentemilla,L.,Cucurell,D.,Pedraza,C.,Rodriguez-fornells,A.,&Diego-balaguer,R.D.(2012).Updating fearful memories with extinction training during reconsolidation:A human study using auditory aversive stimuli.Plos One,7(6),e38849.
Penzo,M.A.,Robert,V.,Tucciarone,J.,De Bundel,D.,Wang,M.,Van Aelst,L.,...Li,B.(2015).The paraventricular thalamus controls a central amygdala fear circuit.Nature,519(7544),455-459.
Phelps,E.A.,Delgado,M.R.,Nearing,K.I.,&Ledoux,J.E.(2004).Extinction learning in humans:Role of the amygdala and vmPFC.Neuron,43(6),897-905.
Quirk,G.J.,Russo,G.K.,Barron,J.L.,&Lebron,K.(2000).The role of ventromedial prefrontal cortex in the recovery of extinguished fear.Journal of Neuroscience,20(16),6225-6231.
Reichelt,A.C.,&Lee,J.L.C.(2013).Memory reconsolidation in aversive and appetitive settings.Frontiers in Behavioral Neuroscience,7(7),118.
Rescorla,R.A.,&Wagner,A.R.(1972).A theory of Pavlovian conditioning:The effectiveness of reinforcement and non-reinforcement.Paper presented at the Classical Conditioning Ii:Current Theory&Research In:Black Ah,Prokasy Wf.
Sara,S.J.(2000).Retrieval and reconsolidation:Toward a neurobiology of remembering.Learning&Memory,7(2),73-84.
Schiller,D.,Kanen,J.W.,Ledoux,J.E.,Monfils,M-H.,&Phelps,E.A.(2013).Extinction during reconsolidation of threat memory diminishes prefrontal cortex involvement.Proceedings of the National Academy of Sciences of the United States of America,110(50),20040-20045.
Schiller,D.,Levy,I.,Niv,Y.,Ledoux,J.E.,&Phelps,E.A.(2008).From fear to safety and back:reversal of fear in the human brain.Journal of Neuroscience,28(45),11517.
Schiller,D.,Monfils,M.,Raio,C.M.,Johnson,D.C.,Ledoux,J.E.,&Phelps,E.A.(2010).Preventing the return of fear in humans using reconsolidation update mechanisms.Nature,463(7277),49.
Sehlmeyer,C.,Dannlowski,U.,Sch?ning,S.,Kugel,H.,Pyka,M.,Pfleiderer,B.,...Konrad,C.(2011).Neural correlates of trait anxiety in fear extinction.Psychological Medicine,41(4),789-798.
Sehlmeyer,C.,Sch?ning,S.,Zwitserlood,P.,Pfleiderer,B.,Kircher,T.,Arolt,V.,&Konrad,C.(2009).Human fear conditioning and extinction in neuroimaging:A systematic review.Plos One,4(6),e5865.
Sevenster,D.,Beckers,T.,&Kindt,M.(2013).Prediction error governs pharmacologically induced amnesia for learned fear.Science,339(6121),830-833.
Soeter,M.,&Kindt,M.(2015).An abrupt transformation of phobic behavior after a post-retrieval amnesic agent.Biological Psychiatry,78(12),880-886.
Tovote,P.,Fadok,J.P.,&Lüthi,A.(2015).Neuronal circuits for fear and anxiety.Nature Reviews Neuroscience,16(6),317-331.
Xue,Y-X.,Deng,J-H.,Chen,Y-Y.,Zhang,L-B.,Wu,P.,Huang,G-D.,...Lu,L(2017).Effect of selective inhibition of reactivated nicotine-associated memories with propranolol on nicotine craving.Jama Psychiatry,74(3),224-232.
Xue,Y-X.,Luo,Y-X.,Wu,P.,Shi,H-S.,Xue,L-F.,Chen,C.,...Lu,L.(2012).A memory retrieval-extinction procedure to prevent drug craving and relapse.Science,336(6078),241-245.
Zeng,X-X.,Du,J.,Zhuang,C-Q.,Zhang,J-H.,Jia,Y-L.,&Zheng,X-F.(2014).Unconditioned stimulus revaluation to promote conditioned fear extinction in the memory reconsolidation window.Plos One,9(7),e101589.