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Insular cortex
Right insula, exposed by removing theopercula
3D view of the insular cortex
Details
Part ofCerebral cortexofbrain
ArteryMiddle cerebral
Identifiers
Latincortex insularis
MeSHD000087623
NeuroNames111
NeuroLexIDbirnlex_1117
TA98A14.1.09.149
A12.2.07.053
TA25502
FMA67329
Anatomical terms of neuroanatomy

Theinsular cortex(alsoinsulaandinsular lobe) is a portion of thecerebral cortexfolded deep within thelateral sulcus(the fissure separating thetemporal lobefrom theparietalandfrontal lobes) within eachhemisphereof themammalianbrain.

The insulae are believed to be involved inconsciousnessand play a role in diverse functions usually linked toemotionor the regulation of the body'shomeostasis.These functions includecompassion,empathy,taste,perception,motor control,self-awareness,cognitive functioning,interpersonal relationships,and awareness ofhomeostatic emotionssuch ashunger,painandfatigue.In relation to these, it is involved inpsychopathology.

The insular cortex is divided into two parts: the anterior insula and the posterior insula in which more than a dozen field areas have been identified. The cortical area overlying the insula toward the lateral surface of the brain is theoperculum(meaninglid). The opercula are formed from parts of the enclosing frontal, temporal, and parietal lobes.

Structure

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Connections

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The anterior part of the insula is subdivided by shallow sulci into three or fourshort gyri.

The anterior insula receives a direct projection from the basal part of theventral medial nucleusof the thalamus and a particularly large input from thecentral nucleus of the amygdala.In addition, the anterior insula itself projects to theamygdala.

One study on rhesus monkeys revealed widespread reciprocal connections between the insular cortex and almost all subnuclei of the amygdaloid complex. The posterior insula projects predominantly to the dorsal aspect of the lateral and to the central amygdaloid nuclei. In contrast, the anterior insula projects to the anterior amygdaloid area as well as the medial, the cortical, the accessory basal magnocellular, the medial basal, and the lateral amygdaloid nuclei.[1]

The posterior part of the insula is formed by along gyrus.

The posterior insula connects reciprocally with thesecondary somatosensory cortexand receives input fromspinothalamicallyactivatedventral posterior inferiorthalamic nuclei. It has also been shown that this region receives inputs from the ventromedial nucleus (posterior part) of the thalamus that are highly specialized to convey homeostatic information such as pain, temperature, itch, local oxygen status, and sensual touch.[2]

A human neuroimaging study usingdiffusion tensor imagingrevealed that the anterior insula is interconnected to regions in the temporal and occipital lobe, opercular and orbitofrontal cortex, triangular and opercular parts of the inferior frontal gyrus. The same study revealed differences in the anatomical connection patterns between the left and right hemisphere.[3]

The 'circular sulcus of insula'(orsulcus of Reil[4]) is a semicircularsulcusorfissure[4]that separates the insula from the neighboring gyri of theoperculum[5]in the front, above, and behind.[4]

Cytoarchitecture

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The insular cortex has regions of variable cell structure orcytoarchitecture,changing fromgranularin theposteriorportion to agranular in theanteriorportion. The insula also receives differentialcorticalandthalamicinput along its length. The anterior insular cortex contains a population ofspindle neurons(also calledvon Economo neurons), identified as characterising a distinctive subregion as the agranular frontal insula.[6]

Development

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The insular cortex is considered a separatelobeof thetelencephalonby some authorities.[7]Other sources see the insula as a part of thetemporal lobe.[8]It is also sometimes grouped with limbic structures deep in the brain into alimbic lobe.[citation needed]As a paralimbic cortex, the insular cortex is considered to be a relatively old structure.

Function

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Multimodal sensory processing, sensory binding

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Functional imaging studies show activation of the insula during audio-visual integration tasks.[9][10]

Taste

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The anterior insula is part of the primarygustatory cortex.[11][12]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.[13]

Speech

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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.[14]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.[15]Moreover, electrical stimulation of the posterior insular can evoke speech disturbances such as speech arrest and reduced voice intensity.[16]

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).[17]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.[17]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).[17]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.[18]

Interoceptive awareness

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Further information: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 othergreat apes.The spindle neurons found at a higher density in the right frontal insular cortex are also found in theanterior 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 incognitive-emotionalprocesses that are specific to primates including great apes, such asempathyandmetacognitiveemotional 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 toconsciousness.[19][20]The right anterior insula is engaged ininteroceptiveawareness of homeostatic emotions such as thirst, pain and fatigue,[21]and the ability to time one's ownheartbeat.Moreover, greater right anterior insulargray mattervolume correlates with increased accuracy in this subjective sense of the inner body, and withnegative emotionalexperience.[22]It is also involved in the control ofblood pressure,[23]in particular during and after exercise,[23]and its activity varies with the amount of effort a person believes he/she is exerting.[24][25]

The insular cortex also is where the sensation ofpainis judged as to its degree.[26]Lesion of the insula is associated with dramatic loss of pain perception and isolated insular infarction can lead to contralateral elimination of pinprick perception.[27]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.[28]Those withirritable bowel syndromehave abnormal processing ofvisceralpain in the insular cortex related to dysfunctional inhibition of pain within the brain.[29]

Physiological studies in rhesus monkeys have shown that neurons in the insula respond to skin stimulation.[30]PET studies have also revealed that the human insula can also be activated by vibrational stimulation to the skin.[31]

Another perception of the right anterior insula is the degree of nonpainfulwarmth[32]or nonpainful coldness[33]of a skin sensation. Other internal sensations processed by the insula include stomach orabdominal distension.[34][35]A fullbladderalso activates the insular cortex.[36]

One brain imaging study suggests that the unpleasantness of subjectively perceiveddyspneais processed in the right human anterior insula andamygdala.[37]

The cerebral cortex processingvestibularsensations extends into the insula,[38]with small lesions in the anterior insular cortex being able to cause loss ofbalanceandvertigo.[39]

Other noninteroceptive perceptions include passive listening to music,[40]laughter and crying,[41]empathy and compassion,[42]and language.[43]

Motor control

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In motor control, it contributes to hand-and-eye motor movement,[44][45]swallowing,[46]gastric motility,[47]and speech articulation.[48][49]It has been identified as a "central command” centre that ensures thatheart rateandblood pressureincrease at the onset ofexercise.[50]Research upon conversation links it to the capacity for long and complex spoken sentences.[51]It is also involved in motor learning[52]and has been identified as playing a role in the motor recovery from stroke.[53]

Homeostasis

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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.[54][55]It has a role in regulating the immune system.[56][57][58]

Self

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The insula has been identified as playing a role in the experience of bodily self-awareness,[59][60]sense of agency,[61]and sense of body ownership.[62]

Social emotions

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The anterior insula processes a person's sense ofdisgustboth to smells[63]and to the sight of contamination and mutilation[64]— even when just imagining the experience.[65]This associates with amirror neuron-like link between external and internal experiences.

In social experience, it is involved in the processing of norm violations,[66]emotional processing,[67]empathy,[68]and orgasms.[69]

The insula is active during social decision making. Tiziana Quarto et al. measuredemotional intelligence(EI) (the ability to identify, regulate, and process emotions of themselves and of others) of sixty-three healthy subjects. UsingfMRIEI was measured in correlation with left insular activity. The subjects were shown various pictures offacial expressionsand 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.[70]

Emotions

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The insular cortex, in particular its most anterior portion, is considered alimbic-related cortex. The insula has increasingly become the focus of attention for its role in body representation and subjective emotional experience. In particular,Antonio Damasiohas 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 ofWilliam 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 ofembodied cognition.

In terms of function, the insula is believed to process convergent information to produce anemotionally relevant context for sensory experience.To be specific, the anterior insula is related more toolfactory, gustatory, viscero-autonomic, and limbic function,whereas the posterior insula is related more toauditory-somesthetic-skeletomotorfunction.Functional imaging experimentshave revealed that the insula has an important role inpainexperience and the experience of a number of basicemotions,includinganger,fear,disgust,happiness,andsadness.[71]

The anterior insular cortex (AIC) is believed to be responsible for emotional feelings, including maternal and romantic love, anger, fear, sadness, happiness, sexual arousal, disgust, aversion, unfairness, inequity, indignation, uncertainty,[72]disbelief, social exclusion, trust, empathy, sculptural beauty, a ‘state of union with God’, and hallucinogenic states.[73]

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 theamygdala,theventral striatum,and theorbitofrontal cortex,as well as tomotor cortices.[74]

A study usingmagnetic resonance imagingfound that the right anterior insula is significantly thicker in people thatmeditate.[75]Other research intobrain activity and meditationhas shown an increase in grey matter in areas of the brain including the insular cortex.[76]

Another study using voxel-based morphometry and MRI on experiencedVipassana meditatorswas 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.[77]

The strongest evidence against a causative role for the insula cortex in emotion comes from Damasio et al. (2012)[78]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

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Functional neuroimagingresearch suggests the insula is involved in two types ofsalience.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 thepregenual anterior cingulate cortex(Brodmann area 33) and the anterior andposterior mid-cingulate cortices,and, second, a generalsalience networkconcerned with environmental monitoring, response selection, and skeletomotor body orientation that involves all of the insular cortex and the mid-cingulate cortex.[79]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.[80]

An alternative or perhaps complementary proposal is that the right anterior insular regulates the interaction between the salience of theselective attentioncreated to achieve a task (the dorsal attention system) and the salience ofarousalcreated to keep focused upon the relevant part of the environment (ventral attention system).[81]This regulation of salience might be particularly important during challenging tasks where attention mightfatigueand so cause careless mistakes but if there is too much arousal it risks creating poor performance by turning intoanxiety.[81]

Decision making

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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.[82]It plays a significant role in integrating internal and external cues to facilitate adaptive choices.

Auditory perception

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Research indicates that the insular cortex is involved inauditory perception.Responses to sound stimuli were obtained usingintracranial EEGrecordings acquired from patients with epilepsy. The posterior part of the insula showed auditory responses that resemble those observed inHeschl's gyrus,whereas the anterior part responded to the emotional contents of the auditory stimuli.[83]Clinical data additionally shows that bilateral damage to the insula after ischemic injury or trauma can lead to auditory agnosia.[84]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.[85]

Direct recordings from the posterior part of the insula showed responses to unexpected sounds within regular auditory streams, a process known asauditory deviance detection.Researchers observed amismatch negativity(MMN) potential, a well knownevent related potential,as well as the high frequency activity signals originating from local neurons.[86]

Simple auditory illusions and hallucinations were elicited by electrical functional mapping.[87][83]

Clinical significance

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Progressive expressive aphasia

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Progressiveexpressive aphasiais the deterioration of normallanguage functionthat causes individuals to lose the ability to communicate fluently while still being able to comprehend single words and intact other non-linguistic cognition. It is found in a variety of degenerative neurological conditions includingPick's disease,motor neuron disease,corticobasal degeneration,frontotemporal dementia,andAlzheimer's disease.It is associated with hypometabolism[88]and atrophy of the left anterior insular cortex.[89]

Addiction

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A number of functional brain imaging studies have shown that the insular cortex is activated when drug users are exposed to environmental cues that trigger cravings. This has been shown for a variety of drugs, includingcocaine,alcohol,opiates,andnicotine.Despite these findings, the insula has been ignored within the drug addiction literature, perhaps because it is not known to be a direct target of the mesocorticaldopaminesystem, which is central to current dopamine reward theories of addiction. Research published in 2007[90]has shown thatcigarettesmokers suffering damage to the insular cortex, from astrokefor instance, have their addiction to cigarettes practically eliminated. These individuals were found to be up to 136 times more likely to undergo a disruption of smoking addiction than smokers with damage in other areas. Disruption of addiction was evidenced by self-reported behavior changes such as quitting smoking less than one day after the brain injury, quitting smoking with great ease, not smoking again after quitting, and having no urge to resume smoking since quitting. The study was conducted on average eight years after the strokes, which opens up the possibility thatrecall biascould have affected the results.[91]More recent prospective studies, which overcome this limitation, have corroborated these findings[92][93]This suggests a significant role for the insular cortex in the neurological mechanisms underlying addiction tonicotineand other drugs, and would make this area of the brain a possible target for novel anti-addiction medication. In addition, this finding suggests that functions mediated by the insula, especially conscious feelings, may be particularly important for maintaining drug addiction, although this view is not represented in any modern research or reviews of the subject.[94]

A recent study in rats by Contreras et al.[95]corroborates these findings by showing that reversible inactivation of the insula disrupts amphetamineconditioned place preference,an animal model of cue-induced drug craving. In this study, insula inactivation also disrupted "malaise" responses tolithium chlorideinjection, suggesting that the representation of negative interoceptive states by the insula plays a role in addiction. However, in this same study, the conditioned place preference took place immediately after the injection of amphetamine, suggesting that it is the immediate, pleasurable interoceptive effects of amphetamine administration, rather than the delayed, aversive effects of amphetamine withdrawal that are represented within the insula.

A model proposed by Naqvi et al. (see above) is that the insula stores a representation of the pleasurable interoceptive effects of drug use (e.g., the airway sensory effects of nicotine, the cardiovascular effects of amphetamine), and that this representation is activated by exposure to cues that have previously been associated with drug use. A number of functional imaging studies have shown the insula to be activated during the administration of addictive psychoactive drugs. Several functional imaging studies have also shown that the insula is activated when drug users are exposed to drug cues, and that this activity is correlated with subjective urges. In the cue-exposure studies, insula activity is elicited when there is no actual change in the level of drug in the body. Therefore, rather than merely representing the interoceptive effects of drug use as it occurs, the insula may play a role in memory for the pleasurable interoceptive effects of past drug use, anticipation of these effects in the future, or both. Such a representation may give rise to conscious urges that feel as if they arise from within the body. This may make addicts feel as if their bodies need to use a drug, and may result in persons with lesions in the insula reporting that their bodies have forgotten the urge to use, according to this study.

Subjective certainty in ecstatic seizures

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Main article:Ecstatic seizures

A common quality in mystical experiences is a strong feeling of certainty whichcannot be expressed in words.Fabienne Picard proposes a neurological explanation for this subjective certainty, based on clinical research of epilepsy.[96][97] According to Picard, this feeling of certainty may be caused by a dysfunction of the anterior insula, a part of the brain which is involved ininteroception,self-reflection, and in avoiding uncertainty about the internal representations of the world by "anticipation of resolution of uncertainty or risk". This avoidance of uncertainty functions through the comparison between predicted states and actual states, that is, "signaling that we do not understand, i.e., that there is ambiguity."[98]Picard notes that "the concept of insight is very close to that of certainty," and refers to Archimedes' "Eureka!"[99][100]Picard hypothesizes that during ecstatic seizures the comparison between predicted states and actual states no longer functions, and that mismatches between predicted state and actual state are no longer processed, blocking "negative emotionsand negative arousal arising from predictive uncertainty, "which will be experienced as emotional confidence.[101]Picard concludes that "[t]his could lead to a spiritual interpretation in some individuals."[101]

Other clinical conditions

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The insular cortex has been suggested to have a role in anxiety disorders,[102]emotion dysregulation,[103]andanorexia nervosa.[104]

History

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The insula was first described byJohann Christian Reilwhile describing cranial and spinal nerves and plexuses.[105]Henry GrayinGray's Anatomyis responsible for it being known as theIsland of Reil.[105]John Allmanand colleagues showed that anterior insular cortex containsspindle neurons.

Additional images

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  • Location and structure of the insular cortex
    Location and structure of the insular cortex
  • Coronal section of brain immediately in front of pons (Insula labeled at upper right)
    Coronal section of brain immediately in front of pons (Insula labeled at upper right)
  • Horizontal section of left cerebral hemisphere
    Horizontal section of left cerebral hemisphere
  • Insular cortex highlighted in green on coronal T1 MRI images
    Insular cortex highlighted in green on coronal T1 MRI images
  • Insular cortex highlighted in green on sagittal T1 MRI images
    Insular cortex highlighted in green on sagittal T1 MRI images
  • Insular cortex highlighted in green on transversal T1 MRI images
    Insular cortex highlighted in green on transversal T1 MRI images

See also

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This article usesanatomical terminology.

References

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