Editing Insular cortex (section)

==Function==
===Multimodal sensory processing, sensory binding===
Functional imaging studies show activation of the insula during audio-visual integration tasks.<ref>{{cite journal|last1=Bushara|first1=KO|last2=Grafman|first2=J|last3=Hallett|first3=M|title=Neural correlates of auditory-visual stimulus onset asynchrony detection.|journal=The Journal of Neuroscience|date=1 January 2001|volume=21|issue=1|pages=300–4|pmid=11150347|pmc=6762435|doi=10.1523/JNEUROSCI.21-01-00300.2001}}</ref><ref>{{cite journal|last1=Bushara|first1=KO|last2=Hanakawa|first2=T|last3=Immisch|first3=I|last4=Toma|first4=K|last5=Kansaku|first5=K|last6=Hallett|first6=M|title=Neural correlates of cross-modal binding.|journal=Nature Neuroscience|date=February 2003|volume=6|issue=2|pages=190–5|pmid=12496761|doi=10.1038/nn993|s2cid=1098979}}</ref><ref>{{Cite journal |last=Kostanyan |first=Daria |last2=Vartanov |first2=Alexander |last3=Kiselnikov |first3=Andrey |last4=Kozlovskiy |first4=Stanislav |author-link4=Stanislav Kozlovsky |date=2018 |title=The role of insula in processes of audio- visual integration |url=https://istina.msu.ru/publications/article/144166858/ |journal=[[Psychophysiology (journal)|Psychophysiology]] |publisher=Blackwell Publishing Inc. |volume=55 |issue=1 |pages=133-134}}</ref>

=== Taste ===
The anterior insula is part of the primary [[gustatory cortex]].<ref>{{Cite book   | last1 = Marieb | first1 = Elaine N.  | first2 = Katja | last2 = Hoehn  | title = Anatomy & Physiology, Third Edition  | location = Boston  | publisher = Benjamin Cummings/Pearson  | year= 2008  | pages = 391–395  | isbn = 978-0-8053-0094-9  }}</ref><ref>{{cite journal|last=Pritchard|first=TC|author2=Macaluso, DA |author3=Eslinger, PJ |title=Taste perception in patients with insular cortex lesions.|journal=Behavioral Neuroscience|date=August 1999|volume=113|issue=4|pages=663–71|pmid=10495075|doi=10.1037/0735-7044.113.4.663}}</ref> Research in rhesus monkeys has also reported that apart from numerous taste-sensitive neurons, the insular cortex also responds to non-taste properties of oral stimuli related to the texture (viscosity, grittiness) or temperature of food.<ref>{{Cite journal |last1=Verhagen |first1=Justus V. |last2=Kadohisa |first2=Mikiko |last3=Rolls |first3=Edmund T. |date=September 2004 |title=Primate Insular/Opercular Taste Cortex: Neuronal Representations of the Viscosity, Fat Texture, Grittiness, Temperature, and Taste of Foods |url=https://www.physiology.org/doi/10.1152/jn.00321.2004 |journal=Journal of Neurophysiology |language=en |volume=92 |issue=3 |pages=1685–1699 |doi=10.1152/jn.00321.2004 |pmid=15331650 |issn=0022-3077|url-access=subscription }}</ref>

=== Speech ===
The sensory speech region, Wernicke’s area, and the motor speech region, Broca’s area, are interconnected by a large axonal fiber system known as the arcuate fasciculus which passes directly beneath the insular cortex. On account of this anatomical architecture, ischemic strokes in the insular region can disrupt the arcuate fasciculus.<ref>{{Cite journal |url=https://academic.oup.com/brain/article-abstract/103/2/337/378338 |access-date=2023-12-11 |doi=10.1093/brain/103.2.337 |title=The Anatomical Basis of Conduction Aphasia |date=1980 |last1=Damasio |first1=Hanna |last2=Damasio |first2=Antonio R. |journal=Brain |volume=103 |issue=2 |pages=337–350 |pmid=7397481 |url-access=subscription }}</ref> Functional imaging studies on the cerebral correlates of language production also suggest that the anterior insula forms part of the brain network of speech motor control.<ref>{{Cite journal |last1=Bohland |first1=Jason W. |last2=Guenther |first2=Frank H. |date=2006-08-15 |title=An fMRI investigation of syllable sequence production |url=https://www.sciencedirect.com/science/article/pii/S1053811906004472 |journal=NeuroImage |volume=32 |issue=2 |pages=821–841 |doi=10.1016/j.neuroimage.2006.04.173 |pmid=16730195 |s2cid=9909543 |issn=1053-8119|url-access=subscription }}</ref> Moreover, electrical stimulation of the posterior insular can evoke speech disturbances such as speech arrest and reduced voice intensity.<ref>{{Cite journal |last1=Afif |first1=Afif |last2=Minotti |first2=Lorella |last3=Kahane |first3=Philippe |last4=Hoffmann |first4=Dominique |date=November 2010 |title=Anatomofunctional organization of the insular cortex: A study using intracerebral electrical stimulation in epileptic patients |journal=Epilepsia |language=en |volume=51 |issue=11 |pages=2305–2315 |doi=10.1111/j.1528-1167.2010.02755.x |pmid=20946128 |issn=0013-9580|doi-access=free }}</ref>

Lesion of the pre-central gyrus of the insula can also cause “pure speech apraxia” (i.e. the inability to speak with no apparent aphasic or orofacial motor impairments).<ref name=":1">{{Cite journal |url=https://academic.oup.com/cercor/article/31/8/3723/6213947 |access-date=2023-12-11 |doi=10.1093/cercor/bhab043 |title=The Precentral Insular Cortical Network for Speech Articulation |date=2021 |last1=Tomaiuolo |first1=Francesco |last2=Campana |first2=Serena |last3=Voci |first3=Loredana |last4=Lasaponara |first4=Stefano |last5=Doricchi |first5=Fabrizio |last6=Petrides |first6=Michael |journal=Cerebral Cortex |volume=31 |issue=8 |pages=3723–3731 |pmid=33825880 |url-access=subscription }}</ref> This demonstrates that the insular cortex forms part of a critical circuit for the coordination of complex articulatory movements prior to and during the execution of the motor speech plans.<ref name=":1" /> Importantly, this specific cortical circuit is different from those that relate to the cognitive aspects of language production (e.g., Broca’s area on the inferior frontal gyrus).<ref name=":1" /> Subvocal, or silent, speech has also been shown to activate right insular cortex, further supporting the theory that the motor control of speech proceeds from the insula.<ref>{{Cite journal |last1=Kato |first1=Yutaka |last2=Muramatsu |first2=Taro |last3=Kato |first3=Motoichiro |last4=Shintani |first4=Masuro |last5=Kashima |first5=Haruo |date=2007-03-26 |title=Activation of right insular cortex during imaginary speech articulation |url=https://journals.lww.com/neuroreport/abstract/2007/03260/activation_of_right_insular_cortex_during.21.aspx |journal=NeuroReport |language=en-US |volume=18 |issue=5 |pages=505–509 |doi=10.1097/WNR.0b013e3280586862 |pmid=17496812 |s2cid=2040545 |issn=0959-4965|url-access=subscription }}</ref>

===Interoceptive awareness===
{{Further |Interoception}}
There is evidence that, in addition to its base functions, the insula may play a role in certain higher-level functions that operate only in humans and other [[Hominidae|great apes]]. The spindle neurons found at a higher density in the right frontal insular cortex are also found in the [[anterior cingulate cortex]], which is another region that has reached a high level of specialization in great apes. It has been speculated that these neurons are involved in [[Cognition|cognitive]]-[[emotion]]al processes that are specific to primates including great apes, such as [[empathy]] and [[metacognition|metacognitive]] emotional feelings. This is supported by functional imaging results showing that the structure and function of the right frontal insula is correlated with the ability to feel one's own heartbeat, or to empathize with the pain of others. It is thought that these functions are not distinct from the lower-level functions of the insula but rather arise as a consequence of the role of the insula in conveying homeostatic information to [[consciousness]].<ref>{{cite journal |author1=Benedetto De Martino |author2=Dharshan Kumaran |author3=Ben Seymour |author4=Raymond J. Dolan |title=Frames, Biases, and Rational Decision-Making in the Human Brain |journal=Science |volume=313 |issue=6|date=August 2006 |pages= 684–687 |doi=10.1126/science.1128356 |pmid=16888142 |pmc=2631940|bibcode=2006Sci...313..684D }}</ref><ref>{{cite journal |author1=Gui Xue |author2=Zhonglin Lu |author3=Irwin P. Levin d |author4=Antoine Bechara |title=The impact of prior risk experiences on subsequent risky decision-making: The role of the insula|journal=NeuroImage|volume=50 |year=2010 |pages= 709–716 |doi=10.1016/j.neuroimage.2009.12.097 |pmid=20045470 |issue=2 |pmc=2828040}}</ref> The right anterior insula is engaged in [[Interoception|interoceptive]] awareness of homeostatic emotions such as thirst, pain and fatigue,<ref>{{cite journal|title=Gut feelings: the emerging biology of gut–brain communication|author=Emeran A. Mayer|journal=Nature Reviews Neuroscience|volume=12|issue=8|pages=453–466|date=August 2011|doi=10.1038/nrn3071|url= |pmid=21750565|pmc=3845678}}</ref> and the ability to time one's own [[heart rate|heartbeat]]. Moreover, greater right anterior insular [[gray matter]] volume correlates with increased accuracy in this subjective sense of the inner body, and with [[negative emotion]]al experience.<ref>{{cite journal |vauthors=Critchley HD, Wiens S, Rotshtein P, Ohman A, Dolan RJ |title=Neural systems supporting interoceptive awareness |journal=Nat. Neurosci. |volume=7 |issue=2 |pages=189–95 |date=February 2004 |pmid=14730305 |doi=10.1038/nn1176|hdl=21.11116/0000-0001-A2FB-D |s2cid=13344271 |hdl-access=free }}</ref> It is also involved in the control of [[blood pressure]],<ref name="Lamb">{{cite journal |vauthors=Lamb K, Gallagher K, McColl R, Mathews D, Querry R, Williamson JW |title=Exercise-induced decrease in insular cortex rCBF during postexercise hypotension |journal=Med Sci Sports Exerc |volume=39 |issue=4 |pages=672–9 |date=April 2007 |pmid=17414805 |doi=10.1249/mss.0b013e31802f04e0 |doi-access=free }}</ref> in particular during and after exercise,<ref name="Lamb"/> and its activity varies with the amount of effort a person believes he/she is exerting.<ref>{{cite journal |vauthors=Williamson JW, McColl R, Mathews D, Mitchell JH, Raven PB, Morgan WP |title=Hypnotic manipulation of effort sense during dynamic exercise: cardiovascular responses and brain activation |journal=J. Appl. Physiol. |volume=90 |issue=4 |pages=1392–9 |date=April 2001 |pmid=11247939 |doi=10.1152/jappl.2001.90.4.1392 |s2cid=8653997 }}</ref><ref>{{cite journal |vauthors=Williamson JW, McColl R, Mathews D, Ginsburg M, Mitchell JH |title=Activation of the insular cortex is affected by the intensity of exercise |journal=J. Appl. Physiol. |volume=87 |issue=3 |pages=1213–9 |date=September 1999 |pmid=10484598 |doi=10.1152/jappl.1999.87.3.1213|citeseerx=10.1.1.492.2730 |s2cid=1078691 }}</ref>

The insular cortex also is where the sensation of [[pain]] is judged as to its degree.<ref>{{cite journal |vauthors=Baliki MN, Geha PY, Apkarian AV |title=Parsing pain perception between nociceptive representation and magnitude estimation |journal=J. Neurophysiol. |volume=101 |issue=2 |pages=875–87 |date=February 2009 |pmid=19073802 |pmc=3815214 |doi=10.1152/jn.91100.2008 }}</ref> Lesion of the insula is associated with dramatic loss of pain perception and isolated insular infarction can  lead to contralateral elimination of pinprick perception.<ref>{{Cite journal |last1=Birklein |first1=Frank |last2=Rolke |first2=Roman |last3=Müller-Forell |first3=Wibke |date=2005-11-08 |title=Isolated insular infarction eliminates contralateral cold, cold pain, and pinprick perception |journal=Neurology |language=en |volume=65 |issue=9 |pages=1381 |doi=10.1212/01.wnl.0000181351.82772.b3 |pmid=16275823 |issn=0028-3878|doi-access=free }}</ref> Further, the insula is where a person imagines pain when looking at images of painful events while thinking about their happening to one's own body.<ref>{{cite journal |vauthors=Ogino Y, Nemoto H, Inui K, Saito S, Kakigi R, Goto F |title=Inner experience of pain: imagination of pain while viewing images showing painful events forms subjective pain representation in human brain |journal=Cereb. Cortex |volume=17 |issue=5 |pages=1139–46 |date=May 2007 |pmid=16855007 |doi=10.1093/cercor/bhl023 |doi-access=free }}</ref>  Those with [[irritable bowel syndrome]] have abnormal processing of [[Viscus|visceral]] pain in the insular cortex related to dysfunctional inhibition of pain within the brain.<ref>{{cite journal |vauthors=Song GH, Venkatraman V, Ho KY, Chee MW, Yeoh KG, Wilder-Smith CH |title=Cortical effects of anticipation and endogenous modulation of visceral pain assessed by functional brain MRI in irritable bowel syndrome patients and healthy controls |journal=Pain |volume=126 |issue=1–3 |pages=79–90 |date=December 2006 |pmid=16846694 |doi=10.1016/j.pain.2006.06.017 |s2cid=21437784 }}</ref>

Physiological studies in rhesus monkeys have shown that neurons in the insula respond to skin stimulation.<ref>{{Cite journal |last1=Schneider |first1=Richard J. |last2=Friedman |first2=David P. |last3=Mishkin |first3=Mortimer |date=1993-09-03 |title=A modality-specific somatosensory area within the insula of the rhesus monkey |url=https://dx.doi.org/10.1016/0006-8993%2893%2990305-7 |journal=Brain Research |volume=621 |issue=1 |pages=116–120 |doi=10.1016/0006-8993(93)90305-7 |pmid=8221062 |s2cid=20207990 |issn=0006-8993|url-access=subscription }}</ref> PET studies have also revealed that the human insula can also be activated by vibrational stimulation to the skin.<ref>{{Cite journal |last1=Burton |first1=H. |last2=Videen |first2=T. O. |last3=Raichle |first3=M. E. |date=January 1993 |title=Tactile-Vibration-Activated Foci in Insular and Parietal-Opercular Cortex Studied with Positron Emission Tomography: Mapping the Second Somatosensory Area in Humans |url=http://www.tandfonline.com/doi/full/10.3109/08990229309028839 |journal=Somatosensory & Motor Research |language=en |volume=10 |issue=3 |pages=297–308 |doi=10.3109/08990229309028839 |pmid=8237217 |issn=0899-0220|url-access=subscription }}</ref>

Another perception of the right anterior insula is the degree of nonpainful [[Sense#Temperature|warmth]]<ref>{{cite journal |vauthors=Olausson H, Charron J, Marchand S, Villemure C, Strigo IA, Bushnell MC |title=Feelings of warmth correlate with neural activity in right anterior insular cortex |journal=Neurosci. Lett. |volume=389 |issue=1 |pages=1–5 |date=November 2005 |pmid=16051437 |doi=10.1016/j.neulet.2005.06.065 |s2cid=20068852 }}</ref> or nonpainful coldness<ref>{{cite journal |vauthors=Craig AD, Chen K, Bandy D, Reiman EM |title=Thermosensory activation of insular cortex |journal=Nat. Neurosci. |volume=3 |issue=2 |pages=184–90 |date=February 2000 |pmid=10649575 |doi=10.1038/72131|s2cid=7077496 }}</ref> of a skin sensation. Other internal sensations processed by the insula include stomach or [[abdominal distension]].<ref>{{cite journal |vauthors=Ladabaum U, Minoshima S, Hasler WL, Cross D, Chey WD, Owyang C |title=Gastric distention correlates with activation of multiple cortical and subcortical regions |journal=Gastroenterology |volume=120 |issue=2 |pages=369–76 |date=February 2001 |pmid=11159877 |doi=10.1053/gast.2001.21201|doi-access=free }}</ref><ref>{{cite journal  |vauthors=Hamaguchi T, Kano M, Rikimaru H, etal |title=Brain activity during distention of the descending colon in humans |journal=Neurogastroenterol. Motil. |volume=16 |issue=3 |pages=299–309 |date=June 2004 |pmid=15198652 |doi=10.1111/j.1365-2982.2004.00498.x |s2cid=20437580 }}{{dead link|date=February 2019|bot=medic}}{{cbignore|bot=medic}}</ref> A full [[Urinary bladder|bladder]] also activates the insular cortex.<ref>{{cite journal |vauthors=Matsuura S, Kakizaki H, Mitsui T, Shiga T, Tamaki N, Koyanagi T |title=Human brain region response to distention or cold stimulation of the bladder: a positron emission tomography study |journal=J. Urol. |volume=168 |issue=5 |pages=2035–9 |date=November 2002 |pmid=12394703 |doi=10.1016/s0022-5347(05)64290-5}}</ref>

One brain imaging study suggests that the unpleasantness of subjectively perceived [[dyspnea]] is processed in the right human anterior insula and [[amygdala]].<ref>{{cite journal|last=von Leupoldt|first=A.|author2=Sommer, T.|author3=Kegat, S.|author4=Baumann, H. J.|author5=Klose, H.|author6=Dahme, B.|author7=Buchel, C.|title=The Unpleasantness of Perceived Dyspnea Is Processed in the Anterior Insula and Amygdala|journal=American Journal of Respiratory and Critical Care Medicine|date=24 January 2008|volume=177|issue=9|pages=1026–1032|doi=10.1164/rccm.200712-1821OC|pmid=18263796|url=http://171.66.122.149/content/177/9/1026.full.pdf+html|url-access=subscription}}{{Dead link|date=April 2019 |bot=InternetArchiveBot |fix-attempted=yes }}</ref>

The cerebral cortex processing [[Vestibular system|vestibular]] sensations extends into the insula,<ref>{{cite journal  |vauthors=Kikuchi M, Naito Y, Senda M, etal |title=Cortical activation during optokinetic stimulation — an fMRI study |journal=Acta Otolaryngol. |volume=129 |issue=4 |pages=440–3 |date=April 2009 |pmid=19116795 |doi=10.1080/00016480802610226 |s2cid=42990194 }}
</ref> with small lesions in the anterior insular cortex being able to cause loss of [[Equilibrioception|balance]] and [[Vertigo (medical)|vertigo]].<ref>{{cite journal |vauthors=Papathanasiou ES, Papacostas SS, Charalambous M, Eracleous E, Thodi C, Pantzaris M |title=Vertigo and imbalance caused by a small lesion in the anterior insula |journal=Electromyogr Clin Neurophysiol |volume=46 |issue=3 |pages=185–92 |year=2006 |pmid=16918202 }}</ref>

Other noninteroceptive perceptions include passive listening to music,<ref>{{cite journal |vauthors=Brown S, Martinez MJ, Parsons LM |title=Passive music listening spontaneously engages limbic and paralimbic systems |journal=NeuroReport |volume=15 |issue=13 |pages=2033–7 |date=September 2004 |pmid=15486477 |doi=10.1097/00001756-200409150-00008|s2cid=12308683 }}</ref> laughter and crying,<ref>{{cite journal |vauthors=Sander K, Scheich H |title=Left auditory cortex and amygdala, but right insula dominance for human laughing and crying |journal=J Cogn Neurosci |volume=17 |issue=10 |pages=1519–31 |date=October 2005 |pmid=16269094 |doi=10.1162/089892905774597227 |s2cid=9509954 }}</ref> empathy and compassion,<ref>{{Cite web |url=http://ccare.stanford.edu/node/89 |title=Interview with Tania Singer &#124; the Center for Compassion and Altruism Research and Education |access-date=2010-07-04 |archive-url=https://web.archive.org/web/20100714152432/http://ccare.stanford.edu/node/89 |archive-date=2010-07-14 |url-status=dead }}</ref> and language.<ref>{{cite journal |vauthors=Bamiou DE, Musiek FE, Luxon LM |title=The insula (Island of Reil) and its role in auditory processing. Literature review |journal=Brain Res. Brain Res. Rev. |volume=42 |issue=2 |pages=143–54 |date=May 2003 |pmid=12738055 |doi=10.1016/S0165-0173(03)00172-3|s2cid=22339177 }}</ref>

===Motor control===
In motor control, it contributes to hand-and-eye motor movement,<ref>{{cite journal |vauthors=Anderson TJ, Jenkins IH, Brooks DJ, Hawken MB, Frackowiak RS, Kennard C |title=Cortical control of saccades and fixation in man. A PET study |journal=Brain |volume=117 |issue=Pt 5 |pages=1073–84 |date=October 1994 |pmid=7953589 |doi=10.1093/brain/117.5.1073}}</ref><ref>{{cite journal |vauthors=Fink GR, Frackowiak RS, Pietrzyk U, Passingham RE |title=Multiple nonprimary motor areas in the human cortex |journal=J. Neurophysiol. |volume=77 |issue=4 |pages=2164–74 |date=April 1997 |pmid=9114263 |doi=10.1152/jn.1997.77.4.2164 |s2cid=15881491 }}</ref> swallowing,<ref>{{cite journal |vauthors=Sörös P, Inamoto Y, Martin RE |title=Functional brain imaging of swallowing: an activation likelihood estimation meta-analysis |journal=Hum Brain Mapp |volume=30 |issue=8 |pages=2426–39 |date=August 2009 |pmid=19107749 |doi=10.1002/hbm.20680 |pmc=6871071 |s2cid=15438676 }}</ref> gastric motility,<ref>{{cite journal |vauthors=Penfield W, Faulk ME |title=The insula; further observations on its function |journal=Brain |volume=78 |issue=4 |pages=445–70 |year=1955 |pmid=13293263 |doi=10.1093/brain/78.4.445}}</ref> and speech articulation.<ref>{{cite journal |author=Dronkers NF |title=A new brain region for coordinating speech articulation |journal=Nature |volume=384 |issue=6605 |pages=159–61 |date=November 1996 |pmid=8906789 |doi=10.1038/384159a0|bibcode=1996Natur.384..159D |s2cid=4305696 }}</ref><ref>{{cite journal |vauthors=Ackermann H, Riecker A |title=The contribution of the insula to motor aspects of speech production: a review and a hypothesis |journal=Brain Lang |volume=89 |issue=2 |pages=320–8 |date=May 2004 |pmid=15068914 |doi=10.1016/S0093-934X(03)00347-X |s2cid=36867434 }}</ref> It has been identified as a "central command” centre that ensures that [[heart rate]] and [[blood pressure]] increase at the onset of [[exercise physiology|exercise]].<ref>{{cite journal |vauthors=Nowak M, Holm S, Biering-Sørensen F, Secher NH, Friberg L |title="Central command" and insular activation during attempted foot lifting in paraplegic humans |journal=Hum Brain Mapp |volume=25 |issue=2 |pages=259–65 |date=June 2005 |pmid=15849712 |doi=10.1002/hbm.20097|pmc=6871668 }}</ref> Research upon conversation links it to the capacity for long and complex spoken sentences.<ref>{{cite journal |vauthors=Borovsky A, Saygin AP, Bates E, Dronkers N |title=Lesion correlates of conversational speech production deficits |journal=Neuropsychologia |volume=45 |issue=11 |pages=2525–33 |date=June 2007 |pmid=17499317 |pmc=5610916 |doi=10.1016/j.neuropsychologia.2007.03.023 }}</ref> It is also involved in motor learning<ref>{{cite journal |vauthors=Mutschler I, Schulze-Bonhage A, Glauche V, Demandt E, Speck O, Ball T  |editor1-last=Fitch |editor1-first=Tecumseh |title=A rapid sound-action association effect in human insular cortex |journal=PLOS ONE |volume=2 |issue=2 |pages=e259 |year=2007 |pmid=17327919 |pmc=1800344 |doi=10.1371/journal.pone.0000259 |bibcode=2007PLoSO...2..259M |doi-access=free }}</ref> and has been identified as playing a role in the motor recovery from stroke.<ref>{{cite journal |vauthors=Weiller C, Ramsay SC, Wise RJ, Friston KJ, Frackowiak RS |title=Individual patterns of functional reorganization in the human cerebral cortex after capsular infarction |journal=Annals of Neurology |volume=33 |issue=2 |pages=181–9 |date=February 1993 |pmid=8434880 |doi=10.1002/ana.410330208 |s2cid=25131597 }}</ref>

===Homeostasis===
It plays a role in a variety of homeostatic functions related to basic survival needs, such as taste, visceral sensation, and autonomic control. The insula controls autonomic functions through the regulation of the sympathetic and parasympathetic systems.<ref>{{cite journal |vauthors=Oppenheimer SM, Gelb A, Girvin JP, Hachinski VC |title=Cardiovascular effects of human insular cortex stimulation |journal=Neurology |volume=42 |issue=9 |pages=1727–32 |date=September 1992 |pmid=1513461 |doi=10.1212/wnl.42.9.1727|s2cid=32371468 }}</ref><ref name="Critchley">{{cite journal |author=Critchley HD |title=Neural mechanisms of autonomic, affective, and cognitive integration |journal=J. Comp. Neurol. |volume=493 |issue=1 |pages=154–66 |date=December 2005 |pmid=16254997 |doi=10.1002/cne.20749|s2cid=32616395 }}</ref> It has a role in regulating the immune system.<ref>{{cite journal |vauthors=Pacheco-López G, Niemi MB, Kou W, Härting M, Fandrey J, Schedlowski M |title=Neural substrates for behaviorally conditioned immunosuppression in the rat |journal=J. Neurosci. |volume=25 |issue=9 |pages=2330–7 |date=March 2005 |pmid=15745959 |pmc=6726099 |doi=10.1523/JNEUROSCI.4230-04.2005 }}</ref><ref>{{cite journal |vauthors=Ramírez-Amaya V, Alvarez-Borda B, Ormsby CE, Martínez RD, Pérez-Montfort R, Bermúdez-Rattoni F |title=Insular cortex lesions impair the acquisition of conditioned immunosuppression |journal=Brain Behav. Immun. |volume=10 |issue=2 |pages=103–14 |date=June 1996 |pmid=8811934 |doi=10.1006/brbi.1996.0011|s2cid=24813018 }}</ref><ref>{{cite journal |vauthors=Ramírez-Amaya V, Bermúdez-Rattoni F |title=Conditioned enhancement of antibody production is disrupted by insular cortex and amygdala but not hippocampal lesions |journal=Brain Behav. Immun. |volume=13 |issue=1 |pages=46–60 |date=March 1999 |pmid=10371677 |doi=10.1006/brbi.1998.0547 |s2cid=20527835 }}</ref>

===Self===

The insula has been identified as playing a role in the experience of bodily self-awareness,<ref>{{cite journal |vauthors=Karnath HO, Baier B, Nägele T |title=Awareness of the functioning of one's own limbs mediated by the insular cortex? |journal=J. Neurosci. |volume=25 |issue=31 |pages=7134–8 |date=August 2005 |pmid=16079395 |pmc=6725240 |doi=10.1523/JNEUROSCI.1590-05.2005 }}</ref><ref>{{cite journal |author=Craig AD |title=How do you feel—now? The anterior insula and human awareness |journal=Nature Reviews Neuroscience |volume=10 |issue=1 |pages=59–70 |date=January 2009 |pmid=19096369 |doi=10.1038/nrn2555|s2cid=2340032 }}</ref> sense of agency,<ref>{{cite journal |vauthors=Farrer C, Frith CD |title=Experiencing oneself vs another person as being the cause of an action: the neural correlates of the experience of agency |journal=NeuroImage |volume=15 |issue=3 |pages=596–603 |date=March 2002 |pmid=11848702 |doi=10.1006/nimg.2001.1009 |s2cid=768408 }}</ref> and sense of body ownership.<ref>{{cite journal |vauthors=Tsakiris M, Hesse MD, Boy C, Haggard P, Fink GR |title=Neural signatures of body ownership: a sensory network for bodily self-consciousness |journal=Cereb. Cortex |volume=17 |issue=10 |pages=2235–44 |date=October 2007 |pmid=17138596 |doi=10.1093/cercor/bhl131 |doi-access=free }}</ref>

===Social emotions===

The anterior insula processes a person's sense of [[disgust]] both to smells<ref name="Wicker">{{cite journal |vauthors=Wicker B, Keysers C, Plailly J, Royet JP, Gallese V, Rizzolatti G |title=Both of us disgusted in My insula: the common neural basis of seeing and feeling disgust |journal=Neuron |volume=40 |issue=3 |pages=655–64 |date=October 2003 |pmid=14642287 |doi=10.1016/S0896-6273(03)00679-2|s2cid=766157 |doi-access=free }}</ref> and to the sight of contamination and mutilation<ref>{{cite journal |vauthors=Wright P, He G, Shapira NA, Goodman WK, Liu Y |title=Disgust and the insula: fMRI responses to pictures of mutilation and contamination |journal=NeuroReport |volume=15 |issue=15 |pages=2347–51 |date=October 2004 |pmid=15640753 |doi=10.1097/00001756-200410250-00009|s2cid=6864309 }}</ref> — even when just imagining the experience.<ref name="Jabbi">{{cite journal |vauthors=Jabbi M, Bastiaansen J, Keysers C |editor1-last=Lauwereyns |editor1-first=Jan |title=A common anterior insula representation of disgust observation, experience and imagination shows divergent functional connectivity pathways |journal=PLOS ONE |volume=3 |issue=8 |pages=e2939 |year=2008 |pmid=18698355 |pmc=2491556 |doi=10.1371/journal.pone.0002939 |bibcode=2008PLoSO...3.2939J |doi-access=free }}</ref>  This associates with a [[mirror neuron]]-like link between external and internal experiences.

In social experience, it is involved in the processing of norm violations,<ref>{{cite journal |vauthors=Sanfey AG, Rilling JK, Aronson JA, Nystrom LE, Cohen JD |title=The neural basis of economic decision-making in the Ultimatum Game |journal=Science |volume=300 |issue=5626 |pages=1755–8 |date=June 2003 |pmid=12805551 |doi=10.1126/science.1082976 |bibcode=2003Sci...300.1755S |s2cid=7111382 }}</ref>  emotional processing,<ref>{{cite journal |vauthors=Phan KL, Wager T, Taylor SF, Liberzon I |title=Functional neuroanatomy of emotion: a meta-analysis of emotion activation studies in PET and fMRI |journal=NeuroImage |volume=16 |issue=2 |pages=331–48 |date=June 2002 |pmid=12030820 |doi=10.1006/nimg.2002.1087 |s2cid=7150871 }}</ref> empathy,<ref>{{cite journal |author=Singer T |title=The neuronal basis and ontogeny of empathy and mind reading: review of literature and implications for future research |journal=Neurosci Biobehav Rev |volume=30 |issue=6 |pages=855–63 |year=2006 |pmid=16904182 |doi=10.1016/j.neubiorev.2006.06.011 |s2cid=15411628 }}</ref> and orgasms.<ref>{{cite journal |vauthors=Ortigue S, Grafton ST, Bianchi-Demicheli F |title=Correlation between insula activation and self-reported quality of orgasm in women |journal=NeuroImage |volume=37 |issue=2 |pages=551–60 |date=August 2007 |pmid=17601749 |doi=10.1016/j.neuroimage.2007.05.026 |s2cid=3377994 |url=https://archive-ouverte.unige.ch/unige:29406 }}</ref>

The insula is active during social decision making. Tiziana Quarto et al. measured [[emotional intelligence]] (EI) (the ability to identify, regulate, and process emotions of themselves and of others) of sixty-three healthy subjects. Using [[functional magnetic resonance imaging|fMRI]] EI was measured in correlation with left insular activity. The subjects were shown various pictures of [[facial expression]]s and tasked with deciding to approach or avoid the person in the picture. The results of the social decision task yielded that individuals with high EI scores had left insular activation when processing fearful faces. Individuals with low EI scores had left insular activation when processing angry faces.<ref>{{Cite journal|last1=Quarto|first1=Tiziana|last2=Blasi|first2=Giuseppe|last3=Maddalena|first3=Chiara|last4=Viscanti|first4=Giovanna|last5=Lanciano|first5=Tiziana|last6=Soleti|first6=Emanuela|last7=Mangiulli|first7=Ivan|last8=Taurisano|first8=Paolo|last9=Fazio|first9=Leonardo|date=2016-02-09|title=Association between Ability Emotional Intelligence and Left Insula during Social Judgment of Facial Emotions|journal=PLOS ONE|volume=11|issue=2|pages=e0148621|doi=10.1371/journal.pone.0148621|issn=1932-6203|pmc=4747486|pmid=26859495|bibcode=2016PLoSO..1148621Q|doi-access=free}}</ref>

=== Emotions ===
The insular cortex, in particular its most anterior portion, is considered a [[limbic]]-related cortex. The insula has increasingly become the focus of attention for its role in body representation and subjective emotional experience. In particular, [[Antonio Damasio]] has proposed that this region plays a role in mapping visceral states that are associated with emotional experience, giving rise to conscious feelings. This is in essence a neurobiological formulation of the ideas of [[William James]], who first proposed that subjective emotional experience (i.e., feelings) arise from our brain's interpretation of bodily states that are elicited by emotional events. This is an example of [[embodied cognition]].{{Citation needed|date=February 2025}}

In terms of function, the insula is believed to process convergent information to produce an ''emotionally relevant context for sensory experience''. To be specific, the anterior insula is related more to ''olfactory, gustatory, viscero-autonomic, and limbic function'', whereas the posterior insula is related more to ''auditory-somesthetic-skeletomotor'' function. [[fMRI|Functional imaging experiments]] have revealed that the insula has an important role in [[pain]] experience and the experience of a number of basic [[emotions]], including [[anger]], [[fear]], [[disgust]], [[happiness]], and [[sadness]].<ref>{{cite journal|last=Wager|first=Tor|title=Functional Neuroanatomy of Emotion: A Meta-Analysis of Emotion Activation Studies in PET and fMRI|doi=10.1006/nimg.2002.1087|pmid=12030820|volume=16|issue=2|date=June 2002|journal=NeuroImage|pages=331–48|s2cid=7150871}}</ref>

The anterior insular cortex (AIC) is believed to be correlated to emotional sensations, including maternal and romantic love, anger, fear, sadness, happiness, sexual arousal, disgust, aversion, unfairness, inequity, indignation, uncertainty,<ref>{{cite journal |vauthors=Vilares I, Howard JD, Fernandes HL, Gottfried JA, Kording KP |title=Differential Representations of Prior and Likelihood Uncertainty in the Human Brain |journal=Current Biology |volume=22 |issue=18 |pages=1641–1648 |date=2012 |doi=10.1016/j.cub.2012.07.010 |pmid=22840519 |pmc=3461114 |bibcode=2012CBio...22.1641V }}</ref>{{Dubious|date=February 2025}} disbelief, [[social exclusion]], trust, empathy, sculptural beauty, a ‘state of union with God’, and hallucinogenic states.<ref>{{cite journal |last=Craig |first=A. D. (Bud) |year=2009 |title=How do you feel — now? The anterior insula and human awareness |journal=Nature Reviews Neuroscience |volume=10 |pages=59–70 |url=http://www.allmanlab.caltech.edu/biCNS217/PDFs/Craig2009.pdf |archive-url=https://wayback.archive-it.org/all/20130107223506/http://www.allmanlab.caltech.edu/biCNS217/PDFs/Craig2009.pdf |url-status=dead |archive-date=2013-01-07 |doi=10.1038/nrn2555 |issue=1 |pmid=19096369 |s2cid=2340032 }}</ref>

Functional imaging studies have also implicated the insula in conscious desires, such as food craving and drug craving. What is common to all of these emotional states is that they each change the body in some way and are associated with highly salient subjective qualities. The insula is well-situated for the integration of information relating to bodily states into higher-order cognitive and emotional processes. The insula receives information from "homeostatic afferent" sensory pathways via the thalamus and sends output to a number of other limbic-related structures, such as the [[amygdala]], the [[ventral striatum]], and the [[orbitofrontal cortex]], as well as to [[Motor cortex|motor cortices]].<ref>{{cite journal |last=Craig |first=A. D. (Bud) |year=2002 |title=A new view of pain as a homeostatic emotion |journal=Trends in Neurosciences |volume=26 |issue=6 |pages=303–307 |url=http://meagherlab.tamu.edu/M-Meagher/%20Health%20Psyc%20630/Readings%20630/Pain%20mech%20read/Craig%2003%20pain%20emotion.pdf |doi=10.1016/s0166-2236(03)00123-1 |pmid=12798599 |s2cid=19794544 |access-date=2009-09-03 |archive-url=https://web.archive.org/web/20100622001631/http://meagherlab.tamu.edu/M-Meagher/%20Health%20Psyc%20630/Readings%20630/Pain%20mech%20read/Craig%2003%20pain%20emotion.pdf |archive-date=2010-06-22 |url-status=dead }}</ref>

A study using [[magnetic resonance imaging]] found that the right anterior insula is significantly thicker in people that [[meditate]].<ref name="pmid16272874">{{cite journal |vauthors=Lazar SW, Kerr CE, Wasserman RH, Gray JR, Greve DN, Treadway MT, McGarvey M, Quinn BT, Dusek JA, Benson H, Rauch SL, Moore CI, Fischl B |title=Meditation experience is associated with increased cortical thickness |journal=NeuroReport |volume=16 |issue=17 |pages=1893–7 |year=2005 |pmid=16272874 |pmc=1361002 |doi=10.1097/01.wnr.0000186598.66243.19}}</ref> Other research into [[brain activity and meditation]] has shown an increase in grey matter in areas of the brain including the insular cortex.<ref name="Fox 48–73">{{cite journal|last=Fox|first=Kieran C.R.|author2=Nijeboer, Savannah |author3=Dixon, Matthew L. |author4=Floman, James L. |author5=Ellamil, Melissa |author6=Rumak, Samuel P. |author7=Sedlmeier, Peter |author8= Christoff, Kalina |title=Is meditation associated with altered brain structure? A systematic review and meta-analysis of morphometric neuroimaging in meditation practitioners|journal=Neuroscience & Biobehavioral Reviews|date=June 2014|volume=43|pages=48–73|doi=10.1016/j.neubiorev.2014.03.016 |pmid=24705269|s2cid=207090878}}</ref>

Another study using voxel-based morphometry and MRI on experienced [[Vipassana meditation|Vipassana meditators]] was done to extend the findings of Lazar et al., which found increased grey matter concentrations in this and other areas of the brain in experienced meditators.<ref>{{Cite journal | url= | doi=10.1093/scan/nsm038| pmid=19015095| title=Investigation of mindfulness meditation practitioners with voxel-based morphometry| journal=Social Cognitive and Affective Neuroscience| volume=3| issue=1| pages=55–61| year=2008| last1=Hölzel| first1=Britta K.| last2=Ott| first2=Ulrich| last3=Gard| first3=Tim| last4=Hempel| first4=Hannes| last5=Weygandt| first5=Martin| last6=Morgen| first6=Katrin| last7=Vaitl| first7=Dieter| pmc=2569815}}</ref>

The strongest evidence against a causative role for the insula cortex in emotion comes from Damasio et al. (2012) <ref>{{Cite journal | doi=10.1093/cercor/bhs077| pmid=22473895| title=Persistence of Feelings and Sentience after Bilateral Damage of the Insula| journal=Cerebral Cortex| volume=23| issue=4| pages=833–846| year=2013| last1=Damasio| first1=A.| last2=Damasio| first2=H.| last3=Tranel| first3=D.| pmc=3657385}}</ref> which showed that a patient who suffered bilateral lesions of the insula cortex expressed the full complement of human emotions, and was fully capable of emotional learning.

===Salience===
[[Functional neuroimaging]] research suggests the insula is involved in two types of [[Salience (neuroscience)|salience]]. Interoceptive information processing that links interoception with emotional salience to generate a subjective representation of the body. This involves, first, the anterior insular cortex with the [[Anterior cingulate cortex|pregenual anterior cingulate cortex]] ([[Brodmann area 33]]) and the anterior and [[posterior cingulate cortex|posterior mid-cingulate cortices]], and, second, a general [[salience network]] concerned with environmental monitoring, response selection, and skeletomotor body orientation that involves all of the insular cortex and the mid-cingulate cortex.<ref>{{cite journal |vauthors=Taylor KS, Seminowicz DA, Davis KD |title=Two systems of resting state connectivity between the insula and cingulate cortex |journal=Hum Brain Mapp |volume=30 |issue=9 |pages=2731–45 |date=September 2009 |pmid=19072897 |doi=10.1002/hbm.20705|pmc=6871122 |s2cid=12917288 }}</ref> A related idea is that the anterior insula, as part of the salience network, interacts with the mid-posterior insula to combine salient stimuli with autonomic information, leading to a high state of physiological awareness of salient stimuli.<ref>{{Cite journal|last1=Menon|first1=Vinod|last2=Uddin|first2=Lucina Q.|date=2010-05-29|title=Saliency, switching, attention and control: a network model of insula function|url=http://dx.doi.org/10.1007/s00429-010-0262-0|journal=Brain Structure and Function|volume=214|issue=5–6|pages=655–667|doi=10.1007/s00429-010-0262-0|pmid=20512370 |pmc=2899886 |issn=1863-2653}}</ref>

An alternative or perhaps complementary proposal is that the right anterior insular regulates the interaction between the salience of the [[selective attention]] created to achieve a task (the dorsal attention system) and the salience of [[arousal]] created to keep focused upon the relevant part of the environment (ventral attention system).<ref name="Eckert">{{cite journal |vauthors=Eckert MA, Menon V, Walczak A, Ahlstrom J, Denslow S, Horwitz A, Dubno JR | year = 2009 | title = At the heart of the ventral attention system: the right anterior insula | journal = Hum. Brain Mapp. | volume = 30 | issue = 8| pages = 2530–41 | doi = 10.1002/hbm.20688 | pmid = 19072895 | pmc=2712290}}</ref> This regulation of salience might be particularly important during challenging tasks where attention might [[Fatigue (safety)|fatigue]] and so cause careless mistakes but if there is too much arousal it risks creating poor performance by turning into [[anxiety]].<ref name="Eckert"/>

=== Decision making ===
Studies have shown that damage or dysfunction in the insular cortex can impair decision-making, emotional regulation, and social behavior. The insula is considered a key brain structure in the neural circuitry underlying complex decision-making processes.<ref>{{cite journal |last1=Billeke |first1=Pablo |last2=Ossandon |first2=Tomas |last3=Perrone-Bertolotti |first3=Marcela |last4=Kahane |first4=Philippe |last5=Bastin |first5=Julien |last6=Jerbi |first6=Karim |last7=Lachaux |first7=Jean-Philippe |last8=Fuentealba |first8=Pablo |title=Human Anterior Insula Encodes Performance Feedback and Relays Prediction Error to the Medial Prefrontal Cortex |journal=Cerebral Cortex |date=1 June 2020 |volume=30 |issue=7 |pages=4011–4025 |doi=10.1093/cercor/bhaa017|pmid=32108230 }}</ref> It plays a significant role in integrating internal and external cues to facilitate adaptive choices.

=== Auditory perception ===
Research indicates that the insular cortex is involved in [[Hearing|auditory perception]]. Responses to sound stimuli were obtained using [[Stereoelectroencephalography|intracranial EEG]] recordings acquired from patients with epilepsy. The posterior part of the insula showed auditory responses that resemble those observed in [[Heschl's gyrus]], whereas the anterior part responded to the emotional contents of the auditory stimuli.<ref name=":0">{{Cite journal|last1=Zhang|first1=Yang|last2=Zhou|first2=Wenjing|last3=Wang|first3=Siyu|last4=Zhou|first4=Qin|last5=Wang|first5=Haixiang|last6=Zhang|first6=Bingqing|last7=Huang|first7=Juan|last8=Hong|first8=Bo|last9=Wang|first9=Xiaoqin|date=2019-02-01|title=The Roles of Subdivisions of Human Insula in Emotion Perception and Auditory Processing|url=https://academic.oup.com/cercor/article/29/2/517/4802217|journal=Cerebral Cortex|language=en|volume=29|issue=2|pages=517–528|doi=10.1093/cercor/bhx334|pmid=29342237|s2cid=36927038|issn=1047-3211|url-access=subscription}}</ref> Clinical data additionally shows that bilateral damage to the insula after ischemic injury or trauma can lead to auditory agnosia.<ref>{{Citation |last=Nieuwenhuys |first=Rudolf |title=Chapter 7 - The insular cortex: A review |date=2012-01-01 |url=https://www.sciencedirect.com/science/article/pii/B9780444538604000076 |journal=Progress in Brain Research |volume=195 |pages=123–163 |editor-last=Hofman |editor-first=Michel A. |access-date=2023-12-11 |series=Evolution of the Primate Brain |publisher=Elsevier |doi=10.1016/B978-0-444-53860-4.00007-6 |pmid=22230626 |isbn=978-0-444-53860-4 |editor2-last=Falk |editor2-first=Dean|url-access=subscription }}</ref> Functional magnetic resonance studies have also demonstrated that the insular cortex participates in many key auditory processes such as tuning into novel auditory stimuli and allocating auditory attention.<ref >{{Cite journal |last1=Bamiou |first1=Doris-Eva |last2=Musiek |first2=Frank E |last3=Luxon |first3=Linda M |date=2003-05-01 |title=The insula (Island of Reil) and its role in auditory processing: Literature review |url=https://www.sciencedirect.com/science/article/pii/S0165017303001723 |journal=Brain Research Reviews |volume=42 |issue=2 |pages=143–154 |doi=10.1016/S0165-0173(03)00172-3 |pmid=12738055 |s2cid=22339177 |issn=0165-0173|url-access=subscription }}</ref>

Direct recordings from the posterior part of the insula showed responses to unexpected sounds within regular auditory streams, a process known as [[Mismatch negativity|auditory deviance detection]]. Researchers observed a [[mismatch negativity]] (MMN) potential, a well known [[Event-related potential|event related potential]], as well as the high frequency activity signals originating from local neurons.<ref>{{Cite journal|last1=Blenkmann|first1=Alejandro O.|last2=Collavini|first2=Santiago|last3=Lubell|first3=James|last4=Llorens|first4=Anaïs|last5=Funderud|first5=Ingrid|last6=Ivanovic|first6=Jugoslav|last7=Larsson|first7=Pål G.|last8=Meling|first8=Torstein R.|last9=Bekinschtein|first9=Tristan|last10=Kochen|first10=Silvia|last11=Endestad|first11=Tor|date=December 2019|title=Auditory deviance detection in the human insula: An intracranial EEG study|journal=Cortex|language=en|volume=121|pages=189–200|doi=10.1016/j.cortex.2019.09.002|pmid=31629197|s2cid=202749677|url=https://www.biorxiv.org/content/10.1101/487306v1|hdl=10852/75077|hdl-access=free}}</ref>

Simple auditory illusions and hallucinations were elicited by electrical functional mapping.<ref>{{Cite journal|last1=Afif|first1=Afif|last2=Minotti|first2=Lorella|last3=Kahane|first3=Philippe|last4=Hoffmann|first4=Dominique|date=November 2010|title=Anatomofunctional organization of the insular cortex: A study using intracerebral electrical stimulation in epileptic patients: Functional Organization of the Insula|journal=Epilepsia|language=en|volume=51|issue=11|pages=2305–2315|doi=10.1111/j.1528-1167.2010.02755.x|pmid=20946128|s2cid=2506125|doi-access=free}}</ref><ref name=":0" />