{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T08:57:27Z","timestamp":1785574647935,"version":"3.56.0"},"reference-count":75,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"Deutsche Gesellschaft f\u00fcr Psychologie"},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["WE 4801\/6-1"],"award-info":[{"award-number":["WE 4801\/6-1"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Transl Psychiatry"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Laboratory research suggests that the attenuation of defensive responses during threat exposure relies on the inhibition of subcortical defense circuits orchestrated by the prefrontal cortex. This process may be reflected by increased vagal efferent activity on the autonomic nervous system. Here, we tested whether cardiac vagal efferent activity, indicated by vagally-mediated heart rate variability (vmHRV), is associated with reduced defensive responses during in-vivo threat exposure and whether it may serve as a physiological index of multi-level exposure responding. Thirty-three women with excessive fear of spiders underwent standardized exposure, where they first approached a living tarantula (approach phase) and subsequently progressed through stages of increasingly direct interactions with the spider (threat imminence task). Indices of cardiac vagal activity (vmHRV) and overall cardiac autonomic arousal (heart rate) were measured alongside self-reported fear and avoidance behavior (distance to spider, performance in threat imminence task). During approach, self-reported fear increased until it reached a stable ceiling. Concurrent vmHRV leveled accordingly, with greater activity being related to lower fear and reduced avoidance during the subsequent threat imminence task. From the very first minute, cardiac vagal activity was associated with the participants\u2019 ability to physically engage with the fear cue during threat imminence task, irrespective of covariations in self-reported fear. Our results suggest that cardiac vagal activity may represent inhibition of fear and avoidance during exposure to perceived threat. Cardiac vagal activity, indexed by vmHRV, could therefore represent an index of within-session multi-level responding to exposure and a promising target for adjuncts to promote fear reduction.<\/jats:p>","DOI":"10.1038\/s41398-026-04333-7","type":"journal-article","created":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T05:53:56Z","timestamp":1785563636000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Cardiac vagal activity during in-vivo threat exposure is associated with within-session inhibition of fear and avoidance"],"prefix":"10.1038","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2819-8536","authenticated-orcid":false,"given":"Christoph","family":"Szeska","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kai","family":"Klepzig","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alfons O.","family":"Hamm","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3968-9656","authenticated-orcid":false,"given":"Mathias","family":"Weymar","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,8,1]]},"reference":[{"key":"4333_CR1","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1002\/da.22353","volume":"32","author":"P Duits","year":"2015","unstructured":"Duits P, Cath DC, Lissek S, Hox JJ, Hamm AO, Engelhard IM, et al. Updated meta-analysis of classical fear conditioning in the anxiety disorders. Depress Anxiety. 2015;32:239\u201353.","journal-title":"Depress Anxiety"},{"key":"4333_CR2","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1016\/j.brat.2014.08.006","volume":"62","author":"MR Milad","year":"2014","unstructured":"Milad MR, Rosenbaum BL, Simon NM. Neuroscience of fear extinction: Implications for assessment and treatment of fear-based and anxiety related disorders. Behav Res Ther. 2014;62:17\u201323.","journal-title":"Behav Res Ther"},{"key":"4333_CR3","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1146\/annurev-clinpsy-050212-185542","volume":"9","author":"B Vervliet","year":"2013","unstructured":"Vervliet B, Craske MG, Hermans D. Fear extinction and relapse: state of the art. Annu Rev Clin Psychol. 2013;9:215\u201348.","journal-title":"Annu Rev Clin Psychol"},{"key":"4333_CR4","doi-asserted-by":"publisher","first-page":"502","DOI":"10.1002\/da.22728","volume":"35","author":"JK Carpenter","year":"2018","unstructured":"Carpenter JK, Andrews LA, Witcraft SM, Powers MB, Smits JAJ, Hofmann SG. Cognitive behavioral therapy for anxiety and related disorders: A meta-analysis of randomized placebo-controlled trials. Depress Anxiety. 2018;35:502\u201314.","journal-title":"Depress Anxiety"},{"key":"4333_CR5","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1016\/j.brat.2014.04.006","volume":"58","author":"MG Craske","year":"2014","unstructured":"Craske MG, Treanor M, Conway CC, Zbozinek T, Vervliet B. Maximizing exposure therapy: An inhibitory learning approach. Behav Res Ther. 2014;58:10\u201323.","journal-title":"Behav Res Ther"},{"key":"4333_CR6","doi-asserted-by":"publisher","first-page":"17024","DOI":"10.1038\/nrdp.2017.24","volume":"3","author":"MG Craske","year":"2017","unstructured":"Craske MG, Stein MB, Eley TC, Milad MR, Holmes A, Rapee RM, et al. Anxiety disorders. Nat Rev Dis Primer. 2017;3:17024.","journal-title":"Nat Rev Dis Primer"},{"key":"4333_CR7","doi-asserted-by":"publisher","first-page":"102407","DOI":"10.1016\/j.janxdis.2021.102407","volume":"81","author":"HC Levy","year":"2021","unstructured":"Levy HC, O\u2019Bryan EM, Tolin DF. A meta-analysis of relapse rates in cognitive-behavioral therapy for anxiety disorders. J Anxiety Disord. 2021;81:102407.","journal-title":"J Anxiety Disord"},{"key":"4333_CR8","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1016\/j.cpr.2015.08.004","volume":"42","author":"AG Loerinc","year":"2015","unstructured":"Loerinc AG, Meuret AE, Twohig MP, Rosenfield D, Bluett EJ, Craske MG. Response rates for CBT for anxiety disorders: need for standardized criteria. Clin Psychol Rev. 2015;42:72\u201382.","journal-title":"Clin Psychol Rev"},{"key":"4333_CR9","doi-asserted-by":"publisher","first-page":"212","DOI":"10.1177\/23727322231196789","volume":"10","author":"S Laborde","year":"2023","unstructured":"Laborde S, Ackermann S, Borges U, D\u2019Agostini M, Giraudier M, Iskra M, et al. Leveraging vagally mediated heart rate variability as an actionable, noninvasive biomarker for self-regulation: assessment, intervention, and evaluation. Policy Insights Behav Brain Sci. 2023;10:212\u201320.","journal-title":"Policy Insights Behav Brain Sci"},{"key":"4333_CR10","volume-title":"Behavior therapy techniques: A guide to the treatment of neuroses","author":"J Wolpe","year":"1966","unstructured":"Wolpe J, Lazarus AA Behavior therapy techniques: A guide to the treatment of neuroses. Elmsford, NY, US: Pergamon Press; 1966."},{"key":"4333_CR11","doi-asserted-by":"publisher","first-page":"123","DOI":"10.3390\/clinpract15070123","volume":"15","author":"E Mattera","year":"2025","unstructured":"Mattera E, Zaboski B. Rethinking the subjective units of distress scale: validity and clinical utility of the SUDS. Clin Pract. 2025;15:123.","journal-title":"Clin Pract"},{"key":"4333_CR12","doi-asserted-by":"publisher","first-page":"101833","DOI":"10.1016\/j.jbtep.2022.101833","volume":"79","author":"AL McGlade","year":"2023","unstructured":"McGlade AL, Treanor M, Kim R, Craske MG. Does fear reduction predict treatment response to exposure for social anxiety disorder?. J Behav Ther Exp Psychiatry. 2023;79:101833.","journal-title":"J Behav Ther Exp Psychiatry"},{"key":"4333_CR13","doi-asserted-by":"publisher","first-page":"494","DOI":"10.1016\/j.janxdis.2010.03.006","volume":"24","author":"CP McLean","year":"2010","unstructured":"McLean CP, Hope DA. Subjective anxiety and behavioral avoidance: Gender, gender role, and perceived confirmability of self-report. J Anxiety Disord. 2010;24:494\u2013502.","journal-title":"J Anxiety Disord"},{"key":"4333_CR14","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1016\/j.biopsych.2005.10.006","volume":"60","author":"D Hermans","year":"2006","unstructured":"Hermans D, Craske MG, Mineka S, Lovibond PF. Extinction in human fear conditioning. Biol Psychiatry. 2006;60:361\u20138.","journal-title":"Biol Psychiatry"},{"key":"4333_CR15","doi-asserted-by":"publisher","first-page":"428","DOI":"10.1016\/j.neuron.2013.11.006","volume":"81","author":"V Senn","year":"2014","unstructured":"Senn V, Wolff SBE, Herry C, Grenier F, Ehrlich I, Gr\u00fcndemann J, et al. Long-range connectivity defines behavioral specificity of amygdala neurons. Neuron. 2014;81:428\u201337.","journal-title":"Neuron"},{"key":"4333_CR16","doi-asserted-by":"publisher","DOI":"10.1038\/s41398-022-02174-8","volume":"12","author":"C Szeska","year":"2022","unstructured":"Szeska C, P\u00fcnjer H, Riemann S, Meinzer M, Hamm AO. Stimulation of the ventromedial prefrontal cortex blocks the return of subcortically mediated fear responses. Transl Psychiatry. 2022;12:394.","journal-title":"Transl Psychiatry"},{"key":"4333_CR17","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1038\/nrn3945","volume":"16","author":"P Tovote","year":"2015","unstructured":"Tovote P, Fadok JP, L\u00fcthi A. Neuronal circuits for fear and anxiety. Nat Rev Neurosci. 2015;16:317\u201331.","journal-title":"Nat Rev Neurosci"},{"key":"4333_CR18","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/S0079-6123(06)56001-7","volume":"156","author":"PJ Lang","year":"2006","unstructured":"Lang PJ, Davis M. Emotion, motivation, and the brain: Reflex foundations in animal and human research. Prog Brain Res. 2006;156:3\u201329.","journal-title":"Prog Brain Res"},{"key":"4333_CR19","doi-asserted-by":"crossref","unstructured":"Thayer JF, Lane RD A model of neurovisceral integration in emotion regulation and dysregulation. 2000.","DOI":"10.1016\/S0165-0327(00)00338-4"},{"key":"4333_CR20","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1016\/j.neubiorev.2008.08.004","volume":"33","author":"JF Thayer","year":"2009","unstructured":"Thayer JF, Lane RD. Claude bernard and the heart\u2013brain connection: further elaboration of a model of neurovisceral integration. Neurosci Biobehav Rev. 2009;33:81\u201388.","journal-title":"Neurosci Biobehav Rev"},{"key":"4333_CR21","doi-asserted-by":"publisher","first-page":"747","DOI":"10.1016\/j.neubiorev.2011.11.009","volume":"36","author":"JF Thayer","year":"2012","unstructured":"Thayer JF, \u00c5hs F, Fredrikson M, Sollers JJ, Wager TD. A meta-analysis of heart rate variability and neuroimaging studies: Implications for heart rate variability as a marker of stress and health. Neurosci Biobehav Rev. 2012;36:747\u201356.","journal-title":"Neurosci Biobehav Rev"},{"key":"4333_CR22","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1016\/j.biopsycho.2005.11.014","volume":"74","author":"P Grossman","year":"2007","unstructured":"Grossman P, Taylor EW. Toward understanding respiratory sinus arrhythmia: relations to cardiac vagal tone, evolution and biobehavioral functions. Biol Psychol. 2007;74:263\u201385.","journal-title":"Biol Psychol"},{"key":"4333_CR23","doi-asserted-by":"publisher","first-page":"e13688","DOI":"10.1111\/psyp.13688","volume":"58","author":"J Koenig","year":"2021","unstructured":"Koenig J, Abler B, Agartz I, \u00c5kerstedt T, Andreassen OA, Anthony M, et al. Cortical thickness and resting-state cardiac function across the lifespan: A cross-sectional pooled mega-analysis. Psychophysiology. 2021;58:e13688.","journal-title":"Psychophysiology"},{"key":"4333_CR24","doi-asserted-by":"publisher","first-page":"446","DOI":"10.1523\/JNEUROSCI.2815-16.2016","volume":"37","author":"SU Maier","year":"2017","unstructured":"Maier SU, Hare TA. Higher heart-rate variability is associated with ventromedial prefrontal cortex activity and increased resistance to temptation in dietary self-control challenges. J Neurosci. 2017;37:446\u201355.","journal-title":"J Neurosci"},{"key":"4333_CR25","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1016\/j.neuroimage.2016.05.076","volume":"139","author":"M Sakaki","year":"2016","unstructured":"Sakaki M, Yoo HJ, Nga L, Lee T-H, Thayer JF, Mather M. Heart rate variability is associated with amygdala functional connectivity with MPFC across younger and older adults. Neuroimage. 2016;139:44\u201352.","journal-title":"Neuroimage"},{"key":"4333_CR26","doi-asserted-by":"publisher","first-page":"e105054","DOI":"10.1371\/journal.pone.0105054","volume":"9","author":"M Pappens","year":"2014","unstructured":"Pappens M, Schroijen M, S\u00fctterlin S, Smets E, Bergh OV, den, Thayer JF, et al. Resting heart rate variability predicts safety learning and fear extinction in an interoceptive fear conditioning paradigm. PLoS ONE. 2014;9:e105054.","journal-title":"PLoS ONE"},{"key":"4333_CR27","doi-asserted-by":"publisher","first-page":"1161","DOI":"10.1111\/psyp.12456","volume":"52","author":"J Wendt","year":"2015","unstructured":"Wendt J, Neubert J, Koenig J, Thayer JF, Hamm AO. Resting heart rate variability is associated with inhibition of conditioned fear. Psychophysiology. 2015;52:1161\u20136.","journal-title":"Psychophysiology"},{"key":"4333_CR28","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1007\/s10484-025-09686-1","volume":"50","author":"S Fagioli","year":"2025","unstructured":"Fagioli S, Watanabe DK, Koenig J, Free M, Fazio RH, Vasey MW, et al. Heart rate variability predicts therapy outcome in anxiety disorders: the role of inhibitory learning. Appl Psychophysiol Biofeedback. 2025;50:235\u201345.","journal-title":"Appl Psychophysiol Biofeedback"},{"key":"4333_CR29","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1159\/000487599","volume":"87","author":"J Wendt","year":"2018","unstructured":"Wendt J, Hamm AO, Pan\u00e9-Farr\u00e9 CA, Thayer JF, Gerlach A, Gloster AT, et al. Pretreatment cardiac vagal tone predicts dropout from and residual symptoms after exposure therapy in patients with panic disorder and agoraphobia. Psychother Psychosom. 2018;87:187\u20139.","journal-title":"Psychother Psychosom"},{"key":"4333_CR30","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-021-86867-y","volume":"11","author":"J Richter","year":"2021","unstructured":"Richter J, Pietzner A, Koenig J, Thayer JF, Pan\u00e9-Farr\u00e9 CA, Gerlach AL, et al. Vagal control of the heart decreases during increasing imminence of interoceptive threat in patients with panic disorder and agoraphobia. Sci Rep. 2021;11:1\u201312.","journal-title":"Sci Rep"},{"key":"4333_CR31","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1016\/S0005-7894(74)80008-0","volume":"5","author":"R Klorman","year":"1974","unstructured":"Klorman R, Weerts TC, Hastings JE, Mela BG, Lang PJ. Psychometric description of some specific-fear questionnaires 1 all rights of reproduction in any form reserved. Behav Ther. 1974;5:401\u20139.","journal-title":"Behav Ther"},{"key":"4333_CR32","doi-asserted-by":"publisher","first-page":"114","DOI":"10.1016\/j.beth.2008.04.002","volume":"40","author":"BO Olatunji","year":"2009","unstructured":"Olatunji BO, Woods CM, de Jong PJ, Teachman BA, Sawchuk CN, David B. Development and initial validation of an abbreviated spider phobia questionnaire using item response theory. Behav Ther. 2009;40:114\u201330.","journal-title":"Behav Ther"},{"key":"4333_CR33","doi-asserted-by":"crossref","unstructured":"Margraf J, Cwik JC, Brachel R von, Suppiger A, Schneider S DIPS Open Access 1.2: Diagnostisches Interview bei psychischen St\u00f6rungen. 2021. http:\/\/hss-opus.ub.ruhr-unibochum.de\/opus4\/frontdoor\/index\/index\/docId\/14101 (accessed 5 Feb2026).","DOI":"10.46586\/rub.172.149"},{"key":"4333_CR34","doi-asserted-by":"publisher","unstructured":"American Psychiatric Association. Diagnostic and statistical manual of mental disorders: DSM-5. American Psychiatric Publishing Inc.: Washington, DC, 2013 https:\/\/doi.org\/10.1176\/appi.books.9780890425596.","DOI":"10.1176\/appi.books.9780890425596"},{"key":"4333_CR35","doi-asserted-by":"publisher","first-page":"399","DOI":"10.1016\/S0005-7894(02)80035-1","volume":"33","author":"JL Mystkowski","year":"2002","unstructured":"Mystkowski JL, Craske MG, Echiverri AM. Treatment context and return of fear in spider phobia. Behav Ther. 2002;33:399\u2013416.","journal-title":"Behav Ther"},{"key":"4333_CR36","doi-asserted-by":"publisher","first-page":"6536","DOI":"10.3390\/s22176536","volume":"22","author":"M Schaffarczyk","year":"2022","unstructured":"Schaffarczyk M, Rogers B, Reer R, Gronwald T. Validity of the polar H10 sensor for heart rate variability analysis during resting state and incremental exercise in recreational men and women. Sensors. 2022;22:6536.","journal-title":"Sensors"},{"key":"4333_CR37","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1016\/j.cmpb.2013.07.024","volume":"113","author":"MP Tarvainen","year":"2014","unstructured":"Tarvainen MP, Niskanen J-P, Lipponen JA, Ranta-aho PO, Karjalainen PA. Kubios HRV \u2013 Heart rate variability analysis software. Comput Methods Programs Biomed. 2014;113:210\u201320.","journal-title":"Comput Methods Programs Biomed"},{"key":"4333_CR38","doi-asserted-by":"publisher","first-page":"172","DOI":"10.1109\/10.979357","volume":"49","author":"MP Tarvainen","year":"2002","unstructured":"Tarvainen MP, Ranta-aho PO, Karjalainen PA. An advanced detrending method with application to HRV analysis. IEEE Trans Biomed Eng. 2002;49:172\u20135.","journal-title":"IEEE Trans Biomed Eng"},{"key":"4333_CR39","first-page":"71","volume":"45","author":"L Hill","year":"2009","unstructured":"Hill L, Siebenbrock A. All are measures created equal? Heart rate variability and respiration. Biomed Sci Instrum. 2009;45:71\u20136.","journal-title":"Biomed Sci Instrum"},{"key":"4333_CR40","doi-asserted-by":"publisher","first-page":"365","DOI":"10.1046\/j.1365-2281.2001.00337.x","volume":"21","author":"J Penttil\u00e4","year":"2001","unstructured":"Penttil\u00e4 J, Helminen A, Jartti T, Kuusela T, Huikuri HV, Tulppo MP, et al. Time domain, geometrical and frequency domain analysis of cardiac vagal outflow: effects of various respiratory patterns. Clin Physiol. 2001;21:365\u201376.","journal-title":"Clin Physiol"},{"key":"4333_CR41","doi-asserted-by":"publisher","first-page":"e14604","DOI":"10.1111\/psyp.14604","volume":"61","author":"KS Quigley","year":"2024","unstructured":"Quigley KS, Gianaros PJ, Norman GJ, Jennings JR, Berntson GG, de Geus EJC. Publication guidelines for human heart rate and heart rate variability studies in psychophysiology\u2014Part 1: Physiological underpinnings and foundations of measurement. Psychophysiology. 2024;61:e14604.","journal-title":"Psychophysiology"},{"key":"4333_CR42","doi-asserted-by":"publisher","first-page":"1706","DOI":"10.1177\/0956797615597332","volume":"26","author":"A L\u00f6w","year":"2015","unstructured":"L\u00f6w A, Weymar M, Hamm AO. When threat is near, get out of here: dynamics of defensive behavior during freezing and active avoidance. Psychol Sci. 2015;26:1706\u201316.","journal-title":"Psychol Sci"},{"key":"4333_CR43","unstructured":"R Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing. 2022. https:\/\/www.r-project.org\/."},{"key":"4333_CR44","first-page":"1","volume":"3","author":"J Pinheiro","year":"2022","unstructured":"Pinheiro J, Bates D, Core Team R. nlme: Linear and Nonlinear Mixed Effects Models. R package version. 2022;3:1\u2013158.","journal-title":"R package version"},{"key":"4333_CR45","doi-asserted-by":"publisher","first-page":"175","DOI":"10.3758\/BF03193146","volume":"39","author":"F Faul","year":"2007","unstructured":"Faul F, Erdfelder E, Lang A-G, Buchner A. G*Power 3: A flexible statistical power analysis program for the social, behavioral, and biomedical sciences. Behav Res Methods. 2007;39:175\u201391.","journal-title":"Behav Res Methods"},{"key":"4333_CR46","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-020-58412-w","volume":"10","author":"C Szeska","year":"2020","unstructured":"Szeska C, Richter J, Wendt J, Weymar M, Hamm AO. Promoting long-term inhibition of human fear responses by non-invasive transcutaneous vagus nerve stimulation during extinction training. Sci Rep. 2020;10:1529. https:\/\/doi.org\/10.1038\/s41598-020-58412-w.","journal-title":"Sci Rep"},{"key":"4333_CR47","doi-asserted-by":"publisher","DOI":"10.1038\/s41398-020-0786-x","volume":"10","author":"M Hollandt","year":"2020","unstructured":"Hollandt M, Wroblewski A, Yang Y, Ridderbusch IC, Kircher T, Hamm AO, et al. Facilitating translational science in anxiety disorders by adjusting extinction training in the laboratory to exposure-based therapy procedures. Transl Psychiatry. 2020;10:110.","journal-title":"Transl Psychiatry"},{"key":"4333_CR48","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/j.neuron.2015.09.028","volume":"88","author":"JE Dunsmoor","year":"2015","unstructured":"Dunsmoor JE, Niv Y, Daw N, Phelps EA. Rethinking extinction. Neuron. 2015;88:47\u201363.","journal-title":"Neuron"},{"key":"4333_CR49","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fpubh.2017.00258","volume":"5","author":"F Shaffer","year":"2017","unstructured":"Shaffer F, Ginsberg JP. An overview of heart rate variability metrics and norms. Front Public Health. 2017;5:1\u201317.","journal-title":"Front Public Health"},{"key":"4333_CR50","doi-asserted-by":"publisher","first-page":"108728","DOI":"10.1016\/j.biopsycho.2023.108728","volume":"185","author":"T Ritz","year":"2024","unstructured":"Ritz T. Putting back respiration into respiratory sinus arrhythmia or high-frequency heart rate variability: Implications for interpretation, respiratory rhythmicity, and health. Biol Psychol. 2024;185:108728.","journal-title":"Biol Psychol"},{"key":"4333_CR51","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1111\/psyp.13812","volume":"58","author":"C Szeska","year":"2021","unstructured":"Szeska C, Richter J, Wendt J, Weymar M, Hamm AO. Attentive immobility in the face of inevitable distal threat\u2014Startle potentiation and fear bradycardia as an index of emotion and attention. Psychophysiology. 2021;58:1\u201317.","journal-title":"Psychophysiology"},{"key":"4333_CR52","doi-asserted-by":"publisher","first-page":"568","DOI":"10.1038\/s41583-022-00608-2","volume":"23","author":"K Roelofs","year":"2022","unstructured":"Roelofs K, Dayan P. Freezing revisited: coordinated autonomic and central optimization of threat coping. Nat Rev Neurosci. 2022;23:568\u201380.","journal-title":"Nat Rev Neurosci"},{"key":"4333_CR53","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1111\/psyp.13474","volume":"57","author":"AO Hamm","year":"2020","unstructured":"Hamm AO. Fear, anxiety, and their disorders from the perspective of psychophysiology. Psychophysiology. 2020;57:1\u201314.","journal-title":"Psychophysiology"},{"key":"4333_CR54","doi-asserted-by":"publisher","first-page":"429","DOI":"10.3758\/BF03210947","volume":"1","author":"MS Fanselow","year":"1994","unstructured":"Fanselow MS. Neural organization of the defensive behavior system responsible for fear. Psychon Bull Rev. 1994;1:429\u201338.","journal-title":"Psychon Bull Rev"},{"key":"4333_CR55","unstructured":"Lang PJ, Bradley MM, Cuthbert BN Motivated attention: affect, activation and action. In: Lang PJ, Simons RF, Balaban MT (eds). Attention and Orienting: Sensory and Motivational Processes. Hillsdale, NJ: Lawrence Erlbaum Associates, Inc., 1997, pp 97\u2013135."},{"key":"4333_CR56","unstructured":"Campbell BA, Wood G, McBride T Origins of orienting and defensive responses: An evolutionary perspective. In: Lang PJ, Simons RF, Balaban MT (eds). Attention and orienting: Sensory and motivational processes. Erlbaum: Hillsdale, NJ, 1997, pp 41\u201367."},{"key":"4333_CR57","doi-asserted-by":"publisher","first-page":"20170025","DOI":"10.1098\/rstb.2017.0025","volume":"373","author":"MG Craske","year":"2018","unstructured":"Craske MG, Hermans D, Vervliet B. State-of-the-art and future directions for extinction as a translational model for fear and anxiety. Philos Trans R Soc B Biol Sci. 2018;373:20170025.","journal-title":"Philos Trans R Soc B Biol Sci"},{"key":"4333_CR58","doi-asserted-by":"publisher","first-page":"143","DOI":"10.5127\/jep.048615","volume":"7","author":"TJ Barry","year":"2016","unstructured":"Barry TJ, Griffith JW, Vervliet B, Hermans D. The role of stimulus specificity and attention in the generalization of extinction. J Exp Psychopathol. 2016;7:143\u201352.","journal-title":"J Exp Psychopathol"},{"key":"4333_CR59","doi-asserted-by":"publisher","first-page":"76","DOI":"10.5127\/jep.051115","volume":"8","author":"TJ Barry","year":"2017","unstructured":"Barry TJ, Vervliet B, Hermans D. Feature specific attention and return of fear after extinction. J Exp Psychopathol. 2017;8:76\u201387.","journal-title":"J Exp Psychopathol"},{"key":"4333_CR60","doi-asserted-by":"publisher","first-page":"101640","DOI":"10.1016\/j.jbtep.2021.101640","volume":"72","author":"Z Klein","year":"2021","unstructured":"Klein Z, Ginat-Frolich R, Barry TJ, Shechner T. Effects of increased attention allocation to threat and safety stimuli on fear extinction and its recall. J Behav Ther Exp Psychiatry. 2021;72:101640.","journal-title":"J Behav Ther Exp Psychiatry"},{"key":"4333_CR61","doi-asserted-by":"publisher","first-page":"102790","DOI":"10.1016\/j.janxdis.2023.102790","volume":"100","author":"K Roesmann","year":"2023","unstructured":"Roesmann K, Leehr EJ, B\u00f6hnlein J, Gathmann B, Herrmann MJ, Jungh\u00f6fer M, et al. Mechanisms of action underlying virtual reality exposure treatment in spider phobia: Pivotal role of within-session fear reduction. J Anxiety Disord. 2023;100:102790.","journal-title":"J Anxiety Disord"},{"key":"4333_CR62","doi-asserted-by":"publisher","first-page":"256","DOI":"10.1002\/da.22720","volume":"35","author":"MG Wheaton","year":"2018","unstructured":"Wheaton MG, Gershkovich M, Gallagher T, Foa EB, Simpson HB. Behavioral avoidance predicts treatment outcome with exposure and response prevention for obsessive\u2013compulsive disorder. Depress Anxiety. 2018;35:256\u201363.","journal-title":"Depress Anxiety"},{"key":"4333_CR63","doi-asserted-by":"publisher","first-page":"748","DOI":"10.1176\/appi.ajp.2010.09091379","volume":"167","author":"T Insel","year":"2010","unstructured":"Insel T, Cuthbert BN, Garvie M, Heinssen R, Pine DS, Quinn K, et al. Research domain criteria (RDoC): toward a new classification framework for research on mental disorders. Am J Psychiatry. 2010;167:748\u201351.","journal-title":"Am J Psychiatry"},{"key":"4333_CR64","doi-asserted-by":"publisher","DOI":"10.1186\/1741-7015-11-126","volume":"11","author":"BN Cuthbert","year":"2013","unstructured":"Cuthbert BN, Insel TR. Toward the future of psychiatric diagnosis: the seven pillars of RDoC. BMC Med. 2013;11:126.","journal-title":"BMC Med"},{"key":"4333_CR65","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1038\/nature17996","volume":"534","author":"P Tovote","year":"2016","unstructured":"Tovote P, Esposito MS, Botta P, Chaudun F, Fadok JP, Markovic M, et al. Midbrain circuits for defensive behaviour. Nature. 2016;534:206\u201312.","journal-title":"Nature"},{"key":"4333_CR66","doi-asserted-by":"publisher","first-page":"489","DOI":"10.1038\/nn.2499","volume":"13","author":"T Amano","year":"2010","unstructured":"Amano T, Unal CT, Par\u00e9 D. Synaptic correlates of fear extinction in the amygdala. Nat Neurosci. 2010;13:489\u201394.","journal-title":"Nat Neurosci"},{"key":"4333_CR67","doi-asserted-by":"publisher","first-page":"999","DOI":"10.1038\/s41593-022-01130-5","volume":"25","author":"JM Moscarello","year":"2022","unstructured":"Moscarello JM, Penzo MA. The central nucleus of the amygdala and the construction of defensive modes across the threat-imminence continuum. Nat Neurosci. 2022;25:999\u20131008.","journal-title":"Nat Neurosci"},{"key":"4333_CR68","doi-asserted-by":"publisher","first-page":"236","DOI":"10.1002\/wps.21078","volume":"22","author":"A Abi-Dargham","year":"2023","unstructured":"Abi-Dargham A, Moeller SJ, Ali F, DeLorenzo C, Domschke K, Horga G, et al. Candidate biomarkers in psychiatric disorders: state of the field. World Psychiatry. 2023;22:236\u201362.","journal-title":"World Psychiatry"},{"key":"4333_CR69","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1016\/j.neubiorev.2016.04.005","volume":"66","author":"U Lueken","year":"2016","unstructured":"Lueken U, Zierhut KC, Hahn T, Straube B, Kircher T, Reif A, et al. Neurobiological markers predicting treatment response in anxiety disorders: A systematic review and implications for clinical application. Neurosci Biobehav Rev. 2016;66:143\u201362.","journal-title":"Neurosci Biobehav Rev"},{"key":"4333_CR70","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1097\/00002508-199509000-00005","volume":"11","author":"TL Deshields","year":"1995","unstructured":"Deshields TL, Tait RC, Gfeller JD, Chibnall JT. Relationship between social desirability and self-report in chronic pain patients. Clin J Pain. 1995;11:189\u201393.","journal-title":"Clin J Pain"},{"key":"4333_CR71","doi-asserted-by":"publisher","DOI":"10.1038\/s41398-019-0481-y","volume":"9","author":"C Rayner","year":"2019","unstructured":"Rayner C, Coleman JRI, Purves KL, Hodsoll J, Goldsmith K, Alpers GW, et al. A genome-wide association meta-analysis of prognostic outcomes following cognitive behavioural therapy in individuals with anxiety and depressive disorders. Transl Psychiatry. 2019;9:150.","journal-title":"Transl Psychiatry"},{"key":"4333_CR72","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1016\/j.euroneuro.2021.01.004","volume":"44","author":"MA Schiele","year":"2021","unstructured":"Schiele MA, Reif A, Lin J, Alpers GW, Andersson E, Andersson G, et al. Therapygenetic effects of 5-HTTLPR on cognitive-behavioral therapy in anxiety disorders: A meta-analysis. Eur Neuropsychopharmacol. 2021;44:105\u201320.","journal-title":"Eur Neuropsychopharmacol"},{"key":"4333_CR73","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41398-025-03352-0","volume":"15","author":"C Szeska","year":"2025","unstructured":"Szeska C, Klepzig K, Hamm AO, Weymar M. Ready for translation: non-invasive auricular vagus nerve stimulation inhibits psychophysiological indices of stimulus-specific fear and facilitates responding to repeated exposure in phobic individuals. Transl Psychiatry. 2025;15:1\u201313.","journal-title":"Transl Psychiatry"},{"key":"4333_CR74","doi-asserted-by":"publisher","first-page":"112374","DOI":"10.1016\/j.ijpsycho.2024.112374","volume":"204","author":"BL Blaser","year":"2024","unstructured":"Blaser BL, Weymar M, Wendt J. Premenstrual syndrome is associated with differences in heart rate variability and attentional control throughout the menstrual cycle: A pilot study. Int J Psychophysiol. 2024;204:112374.","journal-title":"Int J Psychophysiol"},{"key":"4333_CR75","doi-asserted-by":"publisher","first-page":"448","DOI":"10.1186\/s12905-024-03273-y","volume":"24","author":"KM Schmalenberger","year":"2024","unstructured":"Schmalenberger KM, Eisenlohr-Moul TA, Jarczok MN, Schneider E, Barone JC, Thayer JF, et al. Associations of luteal phase changes in vagally mediated heart rate variability with premenstrual emotional changes. BMC Womens Health. 2024;24:448.","journal-title":"BMC Womens Health"}],"container-title":["Translational Psychiatry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41398-026-04333-7","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41398-026-04333-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41398-026-04333-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T08:02:18Z","timestamp":1785571338000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41398-026-04333-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,1]]},"references-count":75,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["4333"],"URL":"https:\/\/doi.org\/10.1038\/s41398-026-04333-7","relation":{},"ISSN":["2158-3188"],"issn-type":[{"value":"2158-3188","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,1]]},"assertion":[{"value":"15 October 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 July 2026","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 July 2026","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 August 2026","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Ethics"}},{"value":"All methods were performed in accordance with relevant guidelines and regulations. The study was approved by the ethical committee of the University Medicine Greifswald (date of approval: 08.11.2020; reference number: BB 128\/20). Informed consent was obtained from all participants.","order":2,"name":"Ethics","label":"Ethics approval and consent to participate","group":{"name":"EthicsHeading","label":"Ethics"}}],"article-number":"388"}}