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Anterior cingulate cortex
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==== Evidence from electrical studies ==== Evidence for ACC as having an error detection function comes from observations of [[error-related negativity]] (ERN) uniquely generated within the ACC upon error occurrences.<ref name=Bush00/><ref>{{cite journal |vauthors=Gehring WJ, Goss B, Coles MG, Meyer DE, Donchin E |title=A neural system for error-detection and compensation |journal=Psychological Science |volume=4 |issue=6 |pages=385β90 |date=November 1993 |doi=10.1111/j.1467-9280.1993.tb00586.x |s2cid=17422146 }}</ref><ref name="Holroyd04">{{cite book |vauthors=Holroyd CB, Nieuwenhuis S, Mars RB, Coles MG |chapter=Anterior cingulate cortex, selection for action, and error processing |editor=Posner MI |title=Cognitive neuroscience of attention |publisher=Guilford Press |location=New York |year=2004 |pages=[https://archive.org/details/cognitiveneurosc0000unse_q4e4/page/219 219β31] |isbn=1-59385-048-4 |chapter-url=https://archive.org/details/cognitiveneurosc0000unse_q4e4/page/219 }}</ref><ref name="Luu04">{{cite book |vauthors=Luu P, Pederson SM |chapter=The anterior cingulate cortex: Regulating actions in context |editor=Posner MI |title=Cognitive neuroscience of attention |publisher=Guilford Press |location=New York |year=2004 |isbn=1-59385-048-4 |url-access=registration |url=https://archive.org/details/cognitiveneurosc0000unse_q4e4 }}</ref> A distinction has been made between an [[Event-related potential|ERP]] following incorrect responses (response ERN) and a signal after subjects receive feedback after erroneous responses (feedback ERN). Patients with lateral prefrontal cingulate (PFC) damage show reduced ERNs.<ref>{{cite journal | vauthors = Gehring WJ, Knight RT | title = Prefrontal-cingulate interactions in action monitoring | journal = Nature Neuroscience | volume = 3 | issue = 5 | pages = 516β20 | date = May 2000 | pmid = 10769394 | doi = 10.1038/74899 | s2cid = 11136447 }}</ref> Reinforcement learning ERN theory poses that there is a mismatch between actual response execution and appropriate response execution, which results in an ERN discharge.<ref name=Bush00/><ref name=Holroyd04/> Furthermore, this theory predicts that, when the ACC receives conflicting input from control areas in the brain, it determines and allocates which area should be given control over the motor system. Varying levels of dopamine are believed to influence the optimization of this filter system by providing expectations about the outcomes of an event. The ERN, then, serves as a beacon to highlight the violation of an expectation.<ref name=Luu04/> Research on the occurrence of the feedback ERN shows evidence that this potential has larger amplitudes when violations of expectancy are large. In other words, if an event is not likely to happen, the feedback ERN will be larger if no error is detected. Other studies have examined whether the ERN is elicited by varying the cost of an error and the evaluation of a response.<ref name=Holroyd04/> In these trials, feedback is given about whether the participant has gained or lost money after a response. Amplitudes of ERN responses with small gains and small losses were similar. No ERN was elicited for any losses as opposed to an ERN for no wins, even though both outcomes are the same. The finding in this paradigm suggests that monitoring for wins and losses is based on the relative expected gains and losses. If you get a different outcome than expected, the ERN will be larger than for expected outcomes. ERN studies have also localized specific functions of the ACC.<ref name=Luu04/> The rostral ACC seems to be active after an error commission, indicating an error response function, whereas the dorsal ACC is active after both an error and feedback, suggesting a more evaluative function (for fMRI evidence, see also<ref name="Bush02">{{cite journal | vauthors = Bush G, Vogt BA, Holmes J, Dale AM, Greve D, Jenike MA, Rosen BR | title = Dorsal anterior cingulate cortex: a role in reward-based decision making | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 99 | issue = 1 | pages = 523β8 | date = January 2002 | pmid = 11756669 | pmc = 117593 | doi = 10.1073/pnas.012470999 | bibcode = 2002PNAS...99..523B | doi-access = free }}</ref><ref name="Polli">{{cite journal | vauthors = Polli FE, Barton JJ, Cain MS, Thakkar KN, Rauch SL, Manoach DS | title = Rostral and dorsal anterior cingulate cortex make dissociable contributions during antisaccade error commission | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 102 | issue = 43 | pages = 15700β5 | date = October 2005 | pmid = 16227444 | pmc = 1255733 | doi = 10.1073/pnas.0503657102 | bibcode = 2005PNAS..10215700P | doi-access = free }}</ref><ref name="Taylor06">{{cite journal | vauthors = Taylor SF, Martis B, Fitzgerald KD, Welsh RC, Abelson JL, Liberzon I, Himle JA, Gehring WJ | title = Medial frontal cortex activity and loss-related responses to errors | journal = The Journal of Neuroscience | volume = 26 | issue = 15 | pages = 4063β70 | date = April 2006 | pmid = 16611823 | pmc = 6673891 | doi = 10.1523/JNEUROSCI.4709-05.2006 }}</ref> ). This evaluation is emotional in nature and highlights the amount of distress associated with a certain error.<ref name=Bush00/> Summarizing the evidence found by ERN studies, it appears to be the case that ACC receives information about a stimulus, selects an appropriate response, monitors the action, and adapts behavior if there is a violation of expectancy.<ref name=Luu04/>
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