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Ragozzino","year":"2007","journal-title":"Ann N Y Acad Sci"},{"key":"pcbi.1007475.ref089","doi-asserted-by":"crossref","first-page":"1587","DOI":"10.1162\/jocn.2010.21548","article-title":"Pupil diameter predicts changes in the exploration\u2013exploitation trade-off: Evidence for the adaptive gain theory","volume":"23","author":"M Jepma","year":"2011","journal-title":"J Cogn Neurosci"},{"key":"pcbi.1007475.ref090","doi-asserted-by":"crossref","first-page":"628","DOI":"10.1016\/j.neuron.2014.12.053","article-title":"Neuronal activity in primate dorsal anterior cingulate cortex signals task conflict and predicts adjustments in pupil-linked arousal","volume":"85","author":"RB Ebitz","year":"2015","journal-title":"Neuron"},{"key":"pcbi.1007475.ref091","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1177\/1745691611400234","article-title":"Arousal-biased competition in perception and memory","volume":"6","author":"M Mather","year":"2011","journal-title":"Perspect Psychol Sci J 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Stuss","year":"2000","journal-title":"Neuropsychologia"},{"key":"pcbi.1007475.ref102","doi-asserted-by":"crossref","first-page":"927","DOI":"10.1016\/j.neuron.2010.02.027","article-title":"Separable learning systems in the macaque brain and the role of orbitofrontal cortex in contingent learning","volume":"65","author":"ME Walton","year":"2010","journal-title":"Neuron"},{"key":"pcbi.1007475.ref103","doi-asserted-by":"crossref","first-page":"1140","DOI":"10.1038\/nn.3440","article-title":"Prefrontal mechanisms of behavioral flexibility, emotion regulation and value updating","volume":"16","author":"PH Rudebeck","year":"2013","journal-title":"Nat Neurosci"},{"key":"pcbi.1007475.ref104","doi-asserted-by":"crossref","first-page":"1278","DOI":"10.1038\/nn.4387","article-title":"Dorsal anterior cingulate: A Rorschach test for cognitive neuroscience","volume":"19","author":"RB Ebitz","year":"2016","journal-title":"Nat 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