{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,10]],"date-time":"2026-11-10T12:01:54Z","timestamp":1794312114150,"version":"4.9.0"},"reference-count":115,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2021,3,31]],"date-time":"2021-03-31T00:00:00Z","timestamp":1617148800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000781","name":"European Research Council","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000781","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Neurosci."],"abstract":"<jats:p>Acutely challenging or threatening situations frequently require approach-avoidance decisions. Acute threat triggers fast autonomic changes that prepare the body to freeze, fight or flee. However, such autonomic changes may also influence subsequent instrumental approach-avoidance decisions. Since defensive bodily states are often not considered in value-based decision-making models, it remains unclear how they influence the decision-making process. Here, we aim to bridge this gap by discussing the existing literature on the potential role of threat-induced bodily states on decision making and provide a new neurocomputational framework explaining how these effects can facilitate or bias approach-avoid decisions under threat. Theoretical accounts have stated that threat-induced parasympathetic activity is involved in information gathering and decision making. Parasympathetic dominance over sympathetic activity is particularly seen during threat-anticipatory freezing, an evolutionarily conserved response to threat demonstrated across species and characterized by immobility and bradycardia. Although this state of freezing has been linked to altered information processing and action preparation, a full theoretical treatment of the interactions with value-based decision making has not yet been achieved. Our neural framework, which we term the Threat State\/Value Integration (TSI) Model, will illustrate how threat-induced bodily states may impact valuation of competing incentives at three stages of the decision-making process, namely at threat evaluation, integration of rewards and threats, and action initiation. Additionally, because altered parasympathetic activity and decision biases have been shown in anxious populations, we will end with discussing how biases in this system can lead to characteristic patterns of avoidance seen in anxiety-related disorders, motivating future pre-clinical and clinical research.<\/jats:p>","DOI":"10.3389\/fnins.2021.621517","type":"journal-article","created":{"date-parts":[[2021,3,31]],"date-time":"2021-03-31T00:48:42Z","timestamp":1617151722000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":66,"title":["Approach-Avoidance Decisions Under Threat: The Role of Autonomic Psychophysiological States"],"prefix":"10.3389","volume":"15","author":[{"given":"James J. A.","family":"Livermore","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Felix H.","family":"Klaassen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bob","family":"Bramson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anneloes M.","family":"Hulsman","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sjoerd W.","family":"Meijer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leslie","family":"Held","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Floris","family":"Klumpers","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lycia D.","family":"de Voogd","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Karin","family":"Roelofs","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2021,3,31]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"449","DOI":"10.1002\/hbm.22639","article-title":"Neural substrates of approach-avoidance conflict decision-making.","volume":"36","author":"Aupperle","year":"2015","journal-title":"Hum. Brain Mapp."},{"key":"B2","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1111\/j.1469-8986.2005.00287.x","article-title":"A freezing-like posture to pictures of mutilation.","volume":"42","author":"Azevedo","year":"2005","journal-title":"Psychophysiology"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1004646","article-title":"Anxiety-like behavioural inhibition is normative under environmental threat-reward correlations.","volume":"11","author":"Bach","year":"2015","journal-title":"PLoS Comput. Biol."},{"key":"B4","doi-asserted-by":"publisher","first-page":"311","DOI":"10.1038\/nrn.2017.35","article-title":"Algorithms for survival: a comparative perspective on emotions.","volume":"18","author":"Bach","year":"2017","journal-title":"Nat. Rev. Neurosci."},{"key":"B5","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1016\/j.cub.2014.01.046","article-title":"Human hippocampus arbitrates approach-avoidance conflict.","volume":"24","author":"Bach","year":"2014","journal-title":"Curr. Biol."},{"key":"B6","doi-asserted-by":"publisher","first-page":"1805","DOI":"10.1016\/j.brs.2020.10.007","article-title":"Sonication of the anterior thalamus with MRI-Guided transcranial focused ultrasound (tFUS) alters pain thresholds in healthy adults: a double-blind, sham-controlled study.","volume":"13","author":"Badran","year":"2020","journal-title":"Brain Stimul."},{"key":"B7","doi-asserted-by":"crossref","first-page":"327","DOI":"10.31887\/DCNS.2015.17.3\/bbandelow","article-title":"Epidemiology of anxiety disorders in the 21st century.","volume":"17","author":"Bandelow","year":"2015","journal-title":"Dialogues Clin. Neurosci."},{"key":"B8","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/B978-0-12-386525-0.00002-0","article-title":"Chapter 2 \u2013 central autonomic control","author":"Benarroch","year":"2012","journal-title":"Primer on the Autonomic Nervous System"},{"key":"B9","doi-asserted-by":"publisher","first-page":"991","DOI":"10.1016\/j.neubiorev.2010.10.016","article-title":"Risk assessment as an evolved threat detection and analysis process.","volume":"35","author":"Blanchard","year":"2011","journal-title":"Neurosci. Biobehav. Rev."},{"key":"B10","doi-asserted-by":"publisher","first-page":"758","DOI":"10.1111\/ejn.12094","article-title":"Separate amygdala subregions signal surprise and predictiveness during associative fear learning in humans.","volume":"37","author":"Boll","year":"2013","journal-title":"Eur. J. Neurosci."},{"key":"B11","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1016\/j.tics.2004.10.003","article-title":"Conflict monitoring and anterior cingulate cortex: an update.","volume":"8","author":"Botvinick","year":"2004","journal-title":"Trends Cogn. Sci."},{"key":"B12","doi-asserted-by":"publisher","first-page":"276","DOI":"10.1037\/1528-3542.1.3.276","article-title":"Emotion and motivation I: defensive and appetitive reactions in picture processing.","volume":"1","author":"Bradley","year":"2001","journal-title":"Emotion"},{"key":"B13","doi-asserted-by":"publisher","first-page":"602","DOI":"10.1111\/j.1469-8986.2008.00654.x","article-title":"The pupil as a measure of emotional arousal and autonomic activation.","volume":"45","author":"Bradley","year":"2008","journal-title":"Psychophysiology"},{"key":"B14","doi-asserted-by":"publisher","first-page":"468","DOI":"10.1111\/j.0956-7976.2005.01558.x","article-title":"When good things go bad: the reflex physiology of defense.","volume":"16","author":"Bradley","year":"2005","journal-title":"Psychol. Sci."},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.3389\/fpsyg.2021.586553","article-title":"Breathing biofeedback for police officers in a stressful virtual environment: challenges and opportunities.","volume":"12","author":"Brammer","year":"2021","journal-title":"Front. Psychol."},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1101\/2020.06.04.129569","article-title":"Improving emotional-action control by targeting long-range phase-amplitude neuronal coupling.","author":"Bramson","year":"2020","journal-title":"bioRxiv"},{"key":"B17","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.bbr.2007.10.018","article-title":"Different patterns of freezing behavior organized in the periaqueductal gray of rats: association with different types of anxiety.","volume":"188","author":"Brand\u00e3o","year":"2008","journal-title":"Behav. Brain Res."},{"key":"B18","first-page":"16","article-title":"Biological basis of generalized anxiety disorder.","volume":"58","author":"Brawman-Mintzer","year":"1997","journal-title":"J. Clin. Psychiatry"},{"key":"B19","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1016\/j.janxdis.2016.04.012","article-title":"The default response to uncertainty and the importance of perceived safety in anxiety and stress: an evolution-theoretical perspective.","volume":"41","author":"Brosschot","year":"2016","journal-title":"J. Anxiety Disord."},{"key":"B20","doi-asserted-by":"publisher","first-page":"590","DOI":"10.1038\/nn.3961","article-title":"Anxious individuals have difficulty learning the causal statistics of aversive environments.","volume":"18","author":"Browning","year":"2015","journal-title":"Nat. Neurosci."},{"key":"B21","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/j.brat.2017.01.003","article-title":"From avoidance to approach: the influence of threat-of-shock on reward-based decision making.","volume":"96","author":"Bublatzky","year":"2017","journal-title":"Behav. Res. Ther."},{"key":"B22","doi-asserted-by":"publisher","first-page":"2595","DOI":"10.1016\/j.cortex.2013.07.008","article-title":"Functional and structural amygdala \u2013 anterior cingulate connectivity correlates with attentional bias to masked fearful faces.","volume":"49","author":"Carlson","year":"2013","journal-title":"Cortex"},{"key":"B23","doi-asserted-by":"publisher","first-page":"21773","DOI":"10.1073\/pnas.1010079108","article-title":"Amygdala regulates risk of predation in rats foraging in a dynamic fear environment.","volume":"107","author":"Choi","year":"2010","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B24","doi-asserted-by":"publisher","first-page":"885","DOI":"10.1016\/j.neuroimage.2005.05.047","article-title":"Anterior cingulate activity during error and autonomic response.","volume":"27","author":"Critchley","year":"2005","journal-title":"NeuroImage"},{"key":"B25","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1111\/j.1469-8986.2004.00197.x","article-title":"Heart rate and skin conductance analysis of antecendents and consequences of decision making.","volume":"41","author":"Crone","year":"2004","journal-title":"Psychophysiology"},{"key":"B26","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1037\/a0022619","article-title":"The skin conductance response, anticipation, and decision-making.","volume":"4","author":"Dawson","year":"2011","journal-title":"J. Neurosci. Psychol. Econ."},{"key":"B27","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/ncomms10996","article-title":"Computations of uncertainty mediate acute stress responses in humans.","volume":"7","author":"de Berker","year":"2016","journal-title":"Nat. Commun."},{"key":"B28","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1177\/026988119100500414","article-title":"5-HT and mechanisms of defence.","volume":"5","author":"Deakin","year":"1991","journal-title":"J. Psychopharmacol."},{"key":"B29","doi-asserted-by":"publisher","first-page":"414","DOI":"10.1080\/13803395.2015.1023265","article-title":"Reaction time, cerebral blood flow, and heart rate responses in fibromyalgia: Evidence of alterations in attentional control.","volume":"37","author":"del Paso","year":"2015","journal-title":"J. Clin. Exp. Neuropsychol."},{"key":"B30","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1093\/brain\/118.1.279","article-title":"Contributions of anterior cingulate cortex to behaviour.","volume":"118","author":"Devinsky","year":"1995","journal-title":"Brain"},{"key":"B31","doi-asserted-by":"publisher","first-page":"1181","DOI":"10.1016\/j.neubiorev.2005.03.027","article-title":"Die hard: a blend of freezing and fleeing as a dynamic defense\u2014implications for the control of defensive behavior.","volume":"29","author":"Eilam","year":"2005","journal-title":"Neurosci. Biobehav. Rev."},{"key":"B32","doi-asserted-by":"publisher","first-page":"4812","DOI":"10.1073\/pnas.1519829113","article-title":"Striatal structure and function predict individual biases in learning to avoid pain.","volume":"113","author":"Eldar","year":"2016","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B33","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1038\/nature21047","article-title":"A competitive inhibitory circuit for selection of active and passive fear responses.","volume":"542","author":"Fadok","year":"2017","journal-title":"Nature"},{"key":"B34","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1016\/j.neubiorev.2020.05.002","article-title":"Context and trade-offs characterize real-world threat detection systems: A review and comprehensive framework to improve research practice and resolve the translational crisis.","volume":"115","author":"Fendt","year":"2020","journal-title":"Neurosci. Biobehav. Rev."},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1101\/2020.11.25.20234401","article-title":"Transcranial focused ultrasound alters conflict and emotional processing, physiology, and performance I: dorsal anterior cingulate cortex targeting.","author":"Fini","year":"2020","journal-title":"medRxiv"},{"key":"B36","doi-asserted-by":"publisher","first-page":"1109","DOI":"10.1016\/j.neuron.2019.01.019","article-title":"Manipulation of subcortical and deep cortical activity in the primate brain using transcranial focused ultrasound stimulation.","volume":"101","author":"Folloni","year":"2019","journal-title":"Neuron"},{"key":"B37","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1016\/j.biopsycho.2005.08.009","article-title":"An autonomic flexibility\u2013neurovisceral integration model of anxiety and cardiac vagal tone.","volume":"74","author":"Friedman","year":"2007","journal-title":"Biol. Psychol."},{"key":"B38","doi-asserted-by":"publisher","first-page":"1428","DOI":"10.1162\/jocn_a_00425","article-title":"Aversive pavlovian control of instrumental behavior in humans.","volume":"25","author":"Geurts","year":"2013","journal-title":"J. Cogn. Neurosci."},{"key":"B39","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1016\/j.neulet.2016.03.027","article-title":"Ready and waiting: freezing as active action preparation under threat.","volume":"619","author":"Gladwin","year":"2016","journal-title":"Neurosci. Lett."},{"key":"B40","doi-asserted-by":"publisher","first-page":"394","DOI":"10.1016\/j.biopsych.2014.03.030","article-title":"Amygdala\u2013prefrontal cortex functional connectivity during threat-induced anxiety and goal distraction.","volume":"77","author":"Gold","year":"2015","journal-title":"Biol. Psychiatry"},{"key":"B41","doi-asserted-by":"publisher","first-page":"656","DOI":"10.1016\/j.neuron.2010.07.008","article-title":"A neural switch for active and passive fear.","volume":"67","author":"Gozzi","year":"2010","journal-title":"Neuron"},{"key":"B42","doi-asserted-by":"publisher","first-page":"1029","DOI":"10.1016\/j.cell.2017.05.024","article-title":"Noninvasive deep brain stimulation via temporally interfering electric fields.","volume":"169","author":"Grossman","year":"2017","journal-title":"Cell"},{"key":"B43","doi-asserted-by":"publisher","first-page":"194","DOI":"10.1016\/j.tics.2014.01.003","article-title":"Action versus valence in decision making.","volume":"18","author":"Guitart-Masip","year":"2014","journal-title":"Trends Cogn. Sci."},{"key":"B44","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/j.neubiorev.2014.07.021","article-title":"Updating freeze: aligning animal and human research.","volume":"47","author":"Hagenaars","year":"2014","journal-title":"Neurosci. Biobehav. Rev."},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.18422","article-title":"Perceptual decisions are biased by the cost to act.","volume":"6","author":"Hagura","year":"2017","journal-title":"eLife"},{"key":"B46","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1016\/j.biopsych.2011.12.027","article-title":"Anxiety and decision-making.","volume":"72","author":"Hartley","year":"2012","journal-title":"Biol. Psychiatry"},{"key":"B47","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-019-40917-8","article-title":"Neural dynamics of shooting decisions and the switch from freeze to fight.","volume":"9","author":"Hashemi","year":"","journal-title":"Sci. Rep."},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1101\/554840","article-title":"Human defensive freezing is associated with acute threat coping, long term hair cortisol levels and trait anxiety.","author":"Hashemi","year":"","journal-title":"bioRxiv"},{"key":"B49","doi-asserted-by":"publisher","first-page":"22918","DOI":"10.1073\/pnas.1915504116","article-title":"Inexperienced preys know when to flee or to freeze in front of a threat.","volume":"116","author":"H\u00e9bert","year":"2019","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B50","doi-asserted-by":"publisher","first-page":"278","DOI":"10.1016\/j.neuroimage.2012.10.063","article-title":"Fear bradycardia and activation of the human periaqueductal grey.","volume":"66","author":"Hermans","year":"2013","journal-title":"NeuroImage"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2020.00147","article-title":"Linking pain sensation to the autonomic nervous system: the role of the anterior cingulate and periaqueductal gray resting-state networks.","volume":"14","author":"Hohenschurz-Schmidt","year":"2020","journal-title":"Front. Neurosci."},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1016\/j.brat.2020.103788","article-title":"Individual differences in costly fearful avoidance and the relation to psychophysiology.","volume":"137","author":"Hulsman","year":"2021","journal-title":"Behav. Res. Ther."},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1002028","article-title":"Disentangling the roles of approach, activation and valence in instrumental and pavlovian responding.","volume":"7","author":"Huys","year":"2011","journal-title":"PLoS Comput. Biol."},{"key":"B54","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/0165-1838(92)90121-V","article-title":"Specialized functional pathways are the building blocks of the autonomic nervous system.","volume":"41","author":"J\u00e4nig","year":"1992","journal-title":"J. Auton. Nerv. Syst."},{"key":"B55","doi-asserted-by":"publisher","first-page":"434","DOI":"10.1037\/0033-2909.131.3.434","article-title":"Preparation for speeded action as a psychophysiological concept.","volume":"131","author":"Jennings","year":"2005","journal-title":"Psychol. Bull."},{"key":"B56","doi-asserted-by":"publisher","first-page":"746","DOI":"10.1073\/pnas.1524272113","article-title":"Sympathetic innervation controls homeostasis of neuromuscular junctions in health and disease.","volume":"113","author":"Khan","year":"2016","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B57","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1016\/j.neulet.2018.05.028","article-title":"Afferent connections of the dorsal, perigenual, and subgenual anterior cingulate cortices of the monkey: amygdalar inputs and intrinsic connections.","volume":"681","author":"Kim","year":"2018","journal-title":"Neurosci. Lett."},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1101\/2021.01.29.428809","article-title":"Defensive freezing and its relation to approach-avoidance decision-making under threat.","author":"Klaassen","year":"2021","journal-title":"BioRXiv."},{"key":"B59","doi-asserted-by":"publisher","first-page":"186","DOI":"10.1007\/s11065-019-09401-y","article-title":"Roles of the amygdala and basal forebrain in defense: a reply to luyck et al. and implications for defensive action.","volume":"29","author":"Klumpers","year":"2019","journal-title":"Neuropsychol. Rev."},{"key":"B60","doi-asserted-by":"publisher","first-page":"9645","DOI":"10.1523\/JNEUROSCI.3830-16.2017","article-title":"how human amygdala and bed nucleus of the stria terminalis may drive distinct defensive responses.","volume":"37","author":"Klumpers","year":"2017","journal-title":"J. Neurosci."},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.14814\/phy2.12831","article-title":"Role played by periaqueductal gray neurons in parasympathetically mediated fear bradycardia in conscious rats.","volume":"4","author":"Koba","year":"2016","journal-title":"Physiol. Rep."},{"key":"B62","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1097\/HRP.0000000000000065","article-title":"Fear and the defense cascade: clinical implications and management.","volume":"23","author":"Kozlowska","year":"2015","journal-title":"Harvard Rev. Psychiatry"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2015.00189","article-title":"Avoidance learning: a review of theoretical models and recent developments.","volume":"9","author":"Krypotos","year":"2015","journal-title":"Front. Behav. Neurosci."},{"key":"B64","doi-asserted-by":"publisher","first-page":"1120","DOI":"10.1016\/j.biopsych.2009.12.002","article-title":"Sidman instrumental avoidance initially depends on lateral and basal amygdala and is constrained by central amygdala-mediated pavlovian processes.","volume":"67","author":"L\u00e1zaro-Mu\u00f1oz","year":"2010","journal-title":"Biol. Psychiatry"},{"key":"B65","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1038\/mp.2016.1","article-title":"Overshadowed by the amygdala: the bed nucleus of the stria terminalis emerges as key to psychiatric disorders.","volume":"21","author":"Lebow","year":"2016","journal-title":"Mol. Psychiatry"},{"key":"B66","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1038\/nrn.2018.22","article-title":"Surviving threats: neural circuit and computational implications of a new taxonomy of defensive behaviour.","volume":"19","author":"LeDoux","year":"2018","journal-title":"Nat. Rev. Neurosci."},{"key":"B67","doi-asserted-by":"crossref","first-page":"1995","DOI":"10.1002\/hbm.23981","article-title":"Neuromodulation with single-element transcranial focused ultrasound in human thalamus.","volume":"39","author":"Legon","year":"2018","journal-title":"Hum. Brain Mapp."},{"key":"B68","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1016\/j.neuroimage.2012.04.059","article-title":"Avoidance of harm and anxiety: a role for the nucleus accumbens.","volume":"62","author":"Levita","year":"2012","journal-title":"NeuroImage"},{"key":"B69","doi-asserted-by":"publisher","first-page":"1080","DOI":"10.1037\/xge0000117","article-title":"Freezing promotes perception of coarse visual features.","volume":"144","author":"Lojowska","year":"2015","journal-title":"J. Exp. Psychol. Gen."},{"key":"B70","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1016\/j.neuroimage.2018.06.013","article-title":"Visuocortical changes during a freezing-like state in humans.","volume":"179","author":"Lojowska","year":"2018","journal-title":"NeuroImage"},{"key":"B71","doi-asserted-by":"publisher","first-page":"1706","DOI":"10.1177\/0956797615597332","article-title":"When threat is near, get out of here: dynamics of defensive behavior during freezing and active avoidance.","volume":"26","author":"L\u00f6w","year":"2015","journal-title":"Psychol. Sci."},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2014.00237","article-title":"Individual differences in bodily freezing predict emotional biases in decision making.","volume":"8","author":"Ly","year":"2014","journal-title":"Front. Behav. Neurosci."},{"key":"B73","doi-asserted-by":"publisher","first-page":"446","DOI":"10.1101\/lm.031047.113","article-title":"Active vs. reactive threat responding is associated with differential c-Fos expression in specific regions of amygdala and prefrontal cortex.","volume":"20","author":"Martinez","year":"2013","journal-title":"Learn. Mem."},{"key":"B74","doi-asserted-by":"publisher","first-page":"1151","DOI":"10.1016\/j.neuroimage.2004.03.005","article-title":"Functional subdivisions within anterior cingulate cortex and their relationship to autonomic nervous system function.","volume":"22","author":"Matthews","year":"2004","journal-title":"NeuroImage"},{"key":"B75","doi-asserted-by":"publisher","first-page":"439","DOI":"10.1002\/cphy.c140014","article-title":"Autonomic control of the eye.","volume":"5","author":"McDougal","year":"2014","journal-title":"Compr. Physiol."},{"key":"B76","doi-asserted-by":"publisher","first-page":"9024","DOI":"10.1523\/JNEUROSCI.4465-13.2014","article-title":"Aversive prediction error signals in the amygdala.","volume":"34","author":"McHugh","year":"2014","journal-title":"J. Neurosci."},{"key":"B77","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1016\/j.neubiorev.2004.03.005","article-title":"A two-dimensional neuropsychology of defense: fear\/anxiety and defensive distance.","volume":"28","author":"McNaughton","year":"2004","journal-title":"Neurosci. Biobehav. Rev."},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2015.00055","article-title":"The ecology of human fear: survival optimization and the nervous system.","volume":"9","author":"Mobbs","year":"2015","journal-title":"Front. Neurosci."},{"key":"B79","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1016\/j.cobeha.2015.06.005","article-title":"Neuroethological studies of fear, anxiety, and risky decision-making in rodents and humans.","volume":"5","author":"Mobbs","year":"2015","journal-title":"Curr. Opin. Behav. Sci."},{"key":"B80","doi-asserted-by":"publisher","first-page":"1079","DOI":"10.1126\/science.1144298","article-title":"When fear is near: threat imminence elicits prefrontal-periaqueductal gray shifts in humans.","volume":"317","author":"Mobbs","year":"2007","journal-title":"Science"},{"key":"B81","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1016\/j.brainresbull.2011.12.005","article-title":"Cytoarchitecture and cortical connections of the anterior cingulate and adjacent somatomotor fields in the rhesus monkey.","volume":"87","author":"Morecraft","year":"2012","journal-title":"Brain Res. Bull."},{"key":"B82","doi-asserted-by":"publisher","first-page":"905","DOI":"10.1016\/S0306-4522(00)00513-3","article-title":"Activation of the ventrolateral periaqueductal gray reduces locomotion but not mean arterial pressure in awake, freely moving rats.","volume":"102","author":"Morgan","year":"2001","journal-title":"Neuroscience"},{"key":"B83","doi-asserted-by":"publisher","first-page":"3815","DOI":"10.1523\/JNEUROSCI.2596-12.2013","article-title":"Active avoidance learning requires prefrontal suppression of amygdala-mediated defensive reactions.","volume":"33","author":"Moscarello","year":"2013","journal-title":"J. Neurosci."},{"key":"B84","doi-asserted-by":"crossref","DOI":"10.1037\/10802-000","author":"Mowrer","year":"1960","journal-title":"Learning Theory and Behavior."},{"key":"B85","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1016\/j.psyneuen.2017.05.001","article-title":"Defensive freezing links Hypothalamic-Pituitary-Adrenal-axis activity and internalizing symptoms in humans.","volume":"82","author":"Niermann","year":"2017","journal-title":"Psychoneuroendocrinology"},{"key":"B86","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1016\/S0014-2999(98)00261-1","article-title":"Conditioned fear-induced tachycardia in the rat; vagal involvement.","volume":"350","author":"Nijsen","year":"1998","journal-title":"Eur. J. Pharmacol."},{"key":"B87","doi-asserted-by":"publisher","first-page":"769","DOI":"10.1016\/j.conb.2004.10.016","article-title":"Reward representations and reward-related learning in the human brain: insights from neuroimaging.","volume":"14","author":"O\u2019Doherty","year":"2004","journal-title":"Curr. Opin. Neurobiol."},{"key":"B88","doi-asserted-by":"publisher","first-page":"9307","DOI":"10.1523\/JNEUROSCI.4973-10.2011","article-title":"Neurobiology of value integration: when value impacts valuation.","volume":"31","author":"Park","year":"2011","journal-title":"J. Neurosci."},{"key":"B89","doi-asserted-by":"publisher","first-page":"555","DOI":"10.1016\/j.brainresrev.2005.02.005","article-title":"The yin and yang of cardiac autonomic control: vago-sympathetic interactions revisited.","volume":"49","author":"Paton","year":"2005","journal-title":"Brain Res. Rev."},{"key":"B90","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1007\/s11055-019-00878-x","article-title":"Behavior of rats with high and low levels of freezing in defensive situations and on selection of food reinforcement.","volume":"50","author":"Pavlova","year":"2020","journal-title":"Neurosci. Behav. Phys."},{"key":"B91","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1016\/S0166-2236(97)01125-9","article-title":"Organization of intra-amygdaloid circuitries in the rat: an emerging framework for understanding functions of the amygdala.","volume":"20","author":"Pitk\u00e4nen","year":"1997","journal-title":"Trends Neurosci."},{"key":"B92","doi-asserted-by":"publisher","first-page":"116","DOI":"10.1016\/j.biopsycho.2006.06.009","article-title":"The polyvagal perspective.","volume":"74","author":"Porges","year":"2007","journal-title":"Biol. Psychol."},{"key":"B93","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1016\/B978-0-323-03354-1.50022-5","article-title":"Evolutionary aspects of anxiety disorders.","volume":"5","author":"Price","year":"2003","journal-title":"Dialogues Clin. Neurosci."},{"key":"B94","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1016\/j.neuroimage.2019.06.004","article-title":"Neural correlates of anticipatory cardiac deceleration and its association with the speed of perceptual decision-making, in young and older adults.","volume":"199","author":"Ribeiro","year":"2019","journal-title":"NeuroImage"},{"key":"B95","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2016.0206","article-title":"Freeze for action: neurobiological mechanisms in animal and human freezing.","volume":"372","author":"Roelofs","year":"2017","journal-title":"Philos. Trans. R. Soc. B Biol. Sci."},{"key":"B96","doi-asserted-by":"publisher","first-page":"1575","DOI":"10.1177\/0956797610384746","article-title":"Facing freeze: social threat induces bodily freeze in humans.","volume":"21","author":"Roelofs","year":"2010","journal-title":"Psychol. Sci."},{"key":"B97","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-019-53683-4","article-title":"Freezing of gaze during action preparation under threat imminence.","volume":"9","author":"R\u00f6sler","year":"2019","journal-title":"Sci. Rep."},{"key":"B98","doi-asserted-by":"publisher","first-page":"1607","DOI":"10.1038\/nn.3832","article-title":"Representation of aversive prediction errors in the human periaqueductal gray.","volume":"17","author":"Roy","year":"2014","journal-title":"Nat. Neurosci."},{"key":"B99","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1111\/j.1440-1681.2005.04210.x","article-title":"Role of the central nucleus of the amygdala in the control of blood pressure: descending pathways to medullary cardiovascular nuclei.","volume":"32","author":"Saha","year":"2005","journal-title":"Clin. Exp. Pharmacol. Physiol."},{"key":"B100","doi-asserted-by":"publisher","first-page":"25941","DOI":"10.1073\/pnas.1904843116","article-title":"The association between serotonin transporter availability and the neural correlates of fear bradycardia.","volume":"116","author":"Schipper","year":"2019","journal-title":"Proc. Natl. Acad. Sci. U.S.A"},{"key":"B101","doi-asserted-by":"publisher","first-page":"94","DOI":"10.1016\/j.neuroimage.2016.04.070","article-title":"The tipping point: value differences and parallel dorsal\u2013ventral frontal circuits gating human approach\u2013avoidance behavior.","volume":"136","author":"Schlund","year":"2016","journal-title":"NeuroImage"},{"key":"B102","doi-asserted-by":"publisher","first-page":"1234","DOI":"10.1038\/nn1527","article-title":"Opponent appetitive-aversive neural processes underlie predictive learning of pain relief.","volume":"8","author":"Seymour","year":"2005","journal-title":"Nat. Neurosci."},{"key":"B103","doi-asserted-by":"publisher","first-page":"8050","DOI":"10.1523\/JNEUROSCI.0982-16.2016","article-title":"Contributions of the central extended amygdala to fear and anxietycontributions of the central extended amygdala to fear and anxiety.","volume":"36","author":"Shackman","year":"2016","journal-title":"J. Neurosci."},{"key":"B104","doi-asserted-by":"publisher","first-page":"1868","DOI":"10.1016\/j.neuroimage.2009.09.016","article-title":"Neural reward processing is modulated by approach- and avoidance-related personality traits.","volume":"49","author":"Simon","year":"2010","journal-title":"NeuroImage"},{"key":"B105","first-page":"331","article-title":"Innervation of eccrine sweat glands.","volume":"7","author":"Sokolov","year":"1980","journal-title":"Biol. Bull. Acad. Sci. USSR"},{"key":"B106","doi-asserted-by":"publisher","DOI":"10.3389\/fnhum.2013.00284","article-title":"Hierarchical brain networks active in approach and avoidance goal pursuit.","volume":"7","author":"Spielberg","year":"2013","journal-title":"Front. Hum. Neurosci."},{"key":"B107","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2016.0010","article-title":"Interoception, homeostatic emotions and sympathovagal balance.","volume":"371","author":"Strigo","year":"2016","journal-title":"Philos. Trans. R. Soc. B Biol. Sci."},{"key":"B108","doi-asserted-by":"publisher","first-page":"14617","DOI":"10.1523\/JNEUROSCI.2026-09.2009","article-title":"How humans integrate the prospects of pain and reward during choice.","volume":"29","author":"Talmi","year":"2009","journal-title":"J. Neurosci."},{"key":"B109","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1038\/nature17996","article-title":"Midbrain circuits for defensive behaviour.","volume":"534","author":"Tovote","year":"2016","journal-title":"Nature"},{"key":"B110","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1016\/j.ijpsycho.2007.07.004","article-title":"cardiac defense: from attention to action.","volume":"66","author":"Vila","year":"2007","journal-title":"Int. J. Psychophysiol."},{"key":"B111","doi-asserted-by":"publisher","first-page":"1766","DOI":"10.1016\/j.cub.2011.08.050","article-title":"Anterior prefrontal cortex inhibition impairs control over social emotional actions.","volume":"21","author":"Volman","year":"2011","journal-title":"Curr. Biol."},{"key":"B112","doi-asserted-by":"publisher","first-page":"897","DOI":"10.1016\/S0306-4522(02)00744-3","article-title":"Role of ventrolateral periaqueductal gray neurons in the behavioral and cardiovascular responses to contextual conditioned fear and poststress recovery.","volume":"116","author":"Walker","year":"2003","journal-title":"Neuroscience"},{"key":"B113","doi-asserted-by":"publisher","first-page":"196","DOI":"10.1016\/j.neuroimage.2017.06.054","article-title":"Active avoidance and attentive freezing in the face of approaching threat.","volume":"158","author":"Wendt","year":"2017","journal-title":"NeuroImage"},{"key":"B114","doi-asserted-by":"publisher","first-page":"110","DOI":"10.1176\/appi.ajp.2016.15111410","article-title":"Prediction error representation in individuals with generalized anxiety disorder during passive avoidance.","volume":"174","author":"White","year":"2016","journal-title":"Am. J. Psychiatry"},{"key":"B115","doi-asserted-by":"publisher","first-page":"6488","DOI":"10.1523\/JNEUROSCI.4419-15.2016","article-title":"Central amygdala somatostatin neurons gate passive and active defensive behaviors.","volume":"36","author":"Yu","year":"2016","journal-title":"J. Neurosci."}],"container-title":["Frontiers in Neuroscience"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnins.2021.621517\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,3,31]],"date-time":"2021-03-31T00:48:48Z","timestamp":1617151728000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnins.2021.621517\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,3,31]]},"references-count":115,"alternative-id":["10.3389\/fnins.2021.621517"],"URL":"https:\/\/doi.org\/10.3389\/fnins.2021.621517","relation":{},"ISSN":["1662-453X"],"issn-type":[{"value":"1662-453X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,3,31]]},"article-number":"621517"}}