Editing Mirror neuron (section)

== Possible functions ==

=== Understanding intentions ===
Many studies link mirror neurons to understanding goals and intentions. Fogassi et al. (2005)<ref name="pmid15860620" /> recorded the activity of 41 mirror neurons in the inferior parietal lobe (IPL) of two rhesus macaques. The IPL has long been recognized as an association cortex that integrates sensory information. The monkeys watched an experimenter either grasp an apple and bring it to his mouth or grasp an object and place it in a cup.
* In total, 15 mirror neurons fired vigorously when the monkey observed the "grasp-to-eat" motion, but registered no activity while exposed to the "grasp-to-place" condition.
* For 4 other mirror neurons, the reverse held true: they activated in response to the experimenter eventually placing the apple in the cup but not to eating it.

Only the type of action, and not the kinematic force with which models manipulated objects, determined neuron activity. It was also significant that neurons fired before the monkey observed the human model starting the second motor act (bringing the object to the mouth or placing it in a cup). Therefore, IPL neurons "code the same act (grasping) in a different way according to the final goal of the action in which the act is embedded."<ref name="pmid15860620"/> They may furnish a neural basis for predicting another individual's subsequent actions and inferring intention.<ref name="pmid15860620"/>

Understanding intention can be broken down into various stages such as body perception and action identification. These stages correlate with various regions of the brain, for example for body parts/shapes match with the extrastriate and fusiform body areas of the brain. The action itself is identified and facilitated by the mirror neuron system.<ref> author = Huang, L, Du, F, Huang, W, Ren, H, Qiu, W, Zhang, J, & Wang, Y | title = Three-stage dynamic brain-cognitive model of understanding action intention displayed by human body movements | journal = Brain Topography | date = 2024 | doi = 10.1007/s10548-024-01061-3</ref> Action understanding falls into two different processing levels, the mirror neuron system and the mentalizing system. Expected actions are primarily processed by the mirror neuron system and unexpected actions are processed by a combination of the mirror neuron system and the mentalizing system.<ref> author = Mou, H Liu, L, Zhou, T< Yan, Z, & Wang< Y | title = Action expectancy modulates activity in the mirror neuron system and mentalizing system | journal = Neuroimage | year = 2024 | doi = 10.1016/j.neuroimage.2024.120876</ref>

=== Learning facilitation ===
Another possible function of mirror neurons would be facilitation of learning. The mirror neurons code the concrete representation of the action, i.e., the representation that would be activated if the observer acted. This would allow us to simulate (to repeat internally) the observed action implicitly (in the brain) to collect our own motor programs of observed actions and to get ready to reproduce the actions later. It is implicit training. Due to this, the observer will produce the action explicitly (in his/her behavior) with agility and finesse. This happens due to associative learning processes. The more frequently a synaptic connection is activated, the stronger it becomes.<ref name="Kosonogov2012"/>

=== Empathy ===
Stephanie Preston and [[Frans de Waal]],<ref>{{cite journal | vauthors = Preston SD, de Waal FB | title = Empathy: Its ultimate and proximate bases | journal = The Behavioral and Brain Sciences | volume = 25 | issue = 1 | pages = 1–72 | date = February 2002 | pmid = 12625087 | doi = 10.1017/s0140525x02000018 | citeseerx = 10.1.1.554.2794 }}</ref> [[Jean Decety]],<ref>{{cite journal |last1=Decety |first1=J |title=Naturaliser l'empathie |trans-title=Empathy naturalized |language=fr |journal=L' Encéphale |date=2002 |volume=28 |issue=1 |pages=9–20 |id={{INIST|13554627}} |oclc=110778688 }}</ref><ref>{{cite journal | vauthors = Decety J, Jackson PL | title = The functional architecture of human empathy | journal = Behavioral and Cognitive Neuroscience Reviews | volume = 3 | issue = 2 | pages = 71–100 | date = June 2004 | pmid = 15537986 | doi = 10.1177/1534582304267187 | s2cid = 145310279 }}</ref> and [[Vittorio Gallese]]<ref name="pmid21227300">{{cite journal | vauthors = Gallese V, Goldman A | title = Mirror neurons and the simulation theory of mind-reading | journal = Trends in Cognitive Sciences | volume = 2 | issue = 12 | pages = 493–501 | date = December 1998 | pmid = 21227300 | doi = 10.1016/s1364-6613(98)01262-5 | s2cid = 10108122 }}</ref><ref>{{cite journal |vauthors = Gallese V |year=2001 |title=The "Shared Manifold" hypothesis: from mirror neurons to empathy |url=http://www.ingentaconnect.com/content/imp/jcs/2001/00000008/F0030005/1208 |journal=Journal of Consciousness Studies |volume=8 |pages=33–50}}</ref> and [[Christian Keysers]]<ref name="EmpathicBrain" /> have independently argued that the mirror neuron system is involved in [[empathy]]. A large number of experiments using fMRI, [[electroencephalography]] (EEG) and [[magnetoencephalography]] (MEG) have shown that certain brain regions (in particular the anterior [[Insular cortex|insula]], [[anterior cingulate cortex]], and inferior frontal cortex) are active when people experience an emotion (disgust, happiness, pain, etc.) and when they see another person experiencing an emotion.<ref>{{cite journal | vauthors = Botvinick M, Jha AP, Bylsma LM, Fabian SA, Solomon PE, Prkachin KM | title = Viewing facial expressions of pain engages cortical areas involved in the direct experience of pain | journal = NeuroImage | volume = 25 | issue = 1 | pages = 312–319 | date = March 2005 | pmid = 15734365 | doi = 10.1016/j.neuroimage.2004.11.043 | s2cid = 24988672 }}</ref><ref name="pmid18353686">{{cite journal | vauthors = Cheng Y, Yang CY, Lin CP, Lee PL, Decety J | title = The perception of pain in others suppresses somatosensory oscillations: a magnetoencephalography study | journal = NeuroImage | volume = 40 | issue = 4 | pages = 1833–1840 | date = May 2008 | pmid = 18353686 | doi = 10.1016/j.neuroimage.2008.01.064 | s2cid = 1827514 }}</ref><ref>{{cite journal | vauthors = Morrison I, Lloyd D, di Pellegrino G, Roberts N | title = Vicarious responses to pain in anterior cingulate cortex: is empathy a multisensory issue? | journal = Cognitive, Affective & Behavioral Neuroscience | volume = 4 | issue = 2 | pages = 270–278 | date = June 2004 | pmid = 15460933 | doi = 10.3758/CABN.4.2.270 | doi-access = free }}</ref><ref name="pmid14642287">{{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><ref>{{cite journal | vauthors = Singer T, Seymour B, O'Doherty J, Kaube H, Dolan RJ, Frith CD | title = Empathy for pain involves the affective but not sensory components of pain | journal = Science | location = New York, N.Y. | volume = 303 | issue = 5661 | pages = 1157–1162 | date = February 2004 | pmid = 14976305 | doi = 10.1126/science.1093535 | bibcode = 2004Sci...303.1157S | hdl = 21.11116/0000-0001-A020-5 | s2cid = 14727944 | hdl-access = free }}</ref><ref name="Jabbi_2007">{{cite journal | vauthors = Jabbi M, Swart M, Keysers C | title = Empathy for positive and negative emotions in the gustatory cortex | journal = NeuroImage | volume = 34 | issue = 4 | pages = 1744–1753 | date = February 2007 | pmid = 17175173 | doi = 10.1016/j.neuroimage.2006.10.032 | s2cid = 13988152 }}</ref><ref name="pmid17214562">{{cite journal | vauthors = Lamm C, Batson CD, Decety J | title = The neural substrate of human empathy: effects of perspective-taking and cognitive appraisal | journal = Journal of Cognitive Neuroscience | volume = 19 | issue = 1 | pages = 42–58 | date = January 2007 | pmid = 17214562 | doi = 10.1162/jocn.2007.19.1.42 | s2cid = 2828843 }}</ref>
[[David Freedberg]] and [[Vittorio Gallese]] have also put forward the idea that this function of the mirror neuron system is crucial for [[Experimental aesthetics|aesthetic]] experiences.<ref>{{cite journal | vauthors = Freedberg D, Gallese V | title = Motion, emotion and empathy in esthetic experience | journal = Trends in Cognitive Sciences | volume = 11 | issue = 5 | pages = 197–203 | date = May 2007 | pmid = 17347026 | doi = 10.1016/j.tics.2007.02.003 | s2cid = 1996468 }}</ref> Nevertheless, an experiment aimed at investigating the activity of mirror neurons in empathy conducted by Soukayna Bekkali and Peter Enticott at the University of Deakin yielded a different result. After analyzing the report's data, they came up with two conclusions about motor empathy and emotional empathy. First, there is no relationship between motor empathy and the activity of mirror neurons. Second, there is only weak evidence of these neurons' activity in the inferior frontal gyrus (IFG), and no evidence of emotional empathy associated with mirror neurons in key brain regions (inferior parietal lobule: IPL). In other words, there has not been an exact conclusion about the role of mirror neurons in empathy and if they are essential for human empathy.<ref>{{Cite web| vauthors = Jarrett C |date=March 25, 2019|title=There is Only Weak Evidence that Mirror Neurons Underlie Human Empathy- New Review and Meta-Analysis|url=https://digest.bps.org.uk/2019/03/25/there-is-only-weak-evidence-that-mirror-neurons-underlie-human-empathy-new-review-and-meta-analysis/|archive-url=https://web.archive.org/web/20190325134613/https://digest.bps.org.uk/2019/03/25/there-is-only-weak-evidence-that-mirror-neurons-underlie-human-empathy-new-review-and-meta-analysis/|url-status=dead|archive-date=March 25, 2019|website=Research Digest}}</ref> However, these brain regions are not quite the same as the ones which mirror hand actions, and mirror neurons for emotional states or empathy have not yet been described in monkeys.

In a recent study, done in 2022, sixteen hand actions were given for each assignment. The assignment pictured both an activity word phase and the intended word phase. The hand actions were selected in "trails" each introduced twice. One of the times was with a matching phase and the other time was with a misleading word phase. The action words were depicted in two to three words with each beginning with the word "to". For instance, "to point"  (action) or "to spin" (intention).

Participants were expected to answer whether the correct word phase matched the corresponding action or intention word. The word phase had to be answered within 3000 ms, with a 1000 ms black screen between each image. The black screens purpose was for an adequate amount of time in between responses. Participants pressed on the keyboard "x" or "m" to indicate their responses in a yes/no format.<ref>{{cite journal | vauthors = Thompson EL, Bird G, Catmur C | title = Mirror neuron brain regions contribute to identifying actions, but not intentions | journal = Human Brain Mapping | volume = 43 | issue = 16 | pages = 4901–4913 | date = November 2022 | pmid = 35906896 | doi = 10.1002/hbm.26036 | pmc = 9582378 }}</ref>

Christian Keysers at the [[Social Brain Lab]] and colleagues have shown that people who are more empathic according to self-report questionnaires have stronger activations both in the mirror system for hand actions<ref>{{cite web |author=Gazzola, Aziz-Zadeh and Keysers |title=Current Biology |year=2006 |url=http://www.bcn-nic.nl/txt/people/publications/gazzola2006sound.pdf |url-status=dead |archive-url=https://web.archive.org/web/20070630021007/http://www.bcn-nic.nl/txt/people/publications/gazzola2006sound.pdf |archive-date=2007-06-30 }}</ref> and the mirror system for emotions,<ref name="Jabbi_2007" /> providing more direct support for the idea that the mirror system is linked to empathy.
Some researchers observed that the human mirror system does not passively respond to the observation of actions but is influenced by the mindset of the observer.<ref>{{cite journal | vauthors = Molenberghs P, Hayward L, Mattingley JB, Cunnington R | title = Activation patterns during action observation are modulated by context in mirror system areas | journal = NeuroImage | volume = 59 | issue = 1 | pages = 608–615 | date = January 2012 | pmid = 21840404 | doi = 10.1016/j.neuroimage.2011.07.080 | s2cid = 13951700 }}</ref> Researchers observed the link of the mirror neurons during empathetic engagement in patient care.<ref name="pmid23288854">{{cite journal | vauthors = Hojat M, Louis DZ, Maio V, Gonnella JS | title = Empathy and health care quality | journal = American Journal of Medical Quality| volume = 28 | issue = 1 | pages = 6–7 | date = 2013 | pmid = 23288854 | doi = 10.1177/1062860612464731 | s2cid = 12645544 }}</ref>

Studies in rats have shown that the [[anterior cingulate cortex]] contains mirror neurons for pain, i.e. neurons responding both during the first-hand experience of pain and while witnessing the pain of others,<ref name="pmid30982647" /> and inhibition of this region leads to reduced [[emotional contagion]] in rats<ref name="pmid30982647" /> and mice,<ref name="pmid31600517" /> and reduced aversion towards harming others.<ref name="pmid32142701">{{cite journal | vauthors = Hernandez-Lallement J, Attah AT, Soyman E, Pinhal CM, Gazzola V, Keysers C | title = Harm to Others Acts as a Negative Reinforcer in Rats | journal = Current Biology | volume = 30 | issue = 6 | pages = 949–961.e7 | date = March 2020 | pmid = 32142701 | doi = 10.1016/j.cub.2020.01.017 | s2cid = 212424287 | doi-access = free | bibcode = 2020CBio...30E.949H | hdl = 20.500.11755/ee7ae8ac-7393-4276-84ce-1bad1b8e5e0d | hdl-access = free }}</ref> This provides causal evidence for a link between pain mirror neurons, and [[emotional contagion]] and [[prosocial behavior]], two phenomena associated with empathy, in rodents. That brain activity in the homologous brain region is associated with individual variability in empathy in humans<ref name="pmid20946964" /> suggests that a similar mechanism may be at play across mammals.

===Human self awareness===
[[Vilayanur S. Ramachandran|V. S. Ramachandran]] has speculated that mirror neurons may provide the neurological basis of human self-awareness.<ref>{{cite book | vauthors = Oberman L, Ramachandran VS |chapter=Reflections on the Mirror Neuron System: Their Evolutionary Functions Beyond Motor Representation | veditors = Pineda JA |title=Mirror Neuron Systems: The Role of Mirroring Processes in Social Cognition |url=https://archive.org/details/mirrorneuronsyst00pine |url-access=limited |publisher=Humana Press |year=2009 |pages=[https://archive.org/details/mirrorneuronsyst00pine/page/n49 39]–62 |isbn=978-1-934115-34-3}}</ref> In an essay written for the Edge Foundation in 2009 Ramachandran gave the following explanation of his theory: "... I also speculated that these neurons can not only help simulate other people's behavior but can be turned 'inward'—as it were—to create second-order representations or meta-representations of your ''own'' earlier brain processes. This could be the neural basis of introspection, and of the reciprocity of self awareness and other awareness. There is obviously a chicken-or-egg question here as to which evolved first, but... The main point is that the two co-evolved, mutually enriching each other to create the mature representation of self that characterizes modern humans."<ref>{{cite web | vauthors = Ramachandran VS |title=Self Awareness: The Last Frontier, Edge Foundation web essay |url=http://www.edge.org/3rd_culture/rama08/rama08_index.html |date=January 1, 2009 |access-date=July 26, 2011}}</ref>

===Language===
In humans, functional MRI studies have reported finding areas homologous to the monkey mirror neuron system in the inferior frontal cortex, close to Broca's area, one of the hypothesized language regions of the brain. This has led to suggestions that human language evolved from a gesture performance/understanding system implemented in mirror neurons. Mirror neurons have been said to have the potential to provide a mechanism for action-understanding, imitation-learning, and the simulation of other people's behaviour.<ref>{{cite journal | vauthors = Skoyles JR | author-link1 = John Skoyles (scientist) | title = Gesture, language origins, and right handedness. | journal = Psycoloquy | date = 2000 | volume = 11 | issue = 24 | url = https://courses.washington.edu/lingclas/200/Lectures/Biol/Psycoloquy_2000_Gesture,_language_and_right_handedness.pdf }}</ref> This hypothesis is supported by some [[Cytoarchitectonics|cytoarchitectonic]] homologies between monkey premotor area F5 and human Broca's area.<ref name="pmid15988526">{{cite journal | vauthors = Petrides M, Cadoret G, Mackey S | title = Orofacial somatomotor responses in the macaque monkey homologue of Broca's area | journal = Nature | volume = 435 | issue = 7046 | pages = 1235–1238 | date = June 2005 | pmid = 15988526 | doi = 10.1038/nature03628 | bibcode = 2005Natur.435.1235P | s2cid = 4397762 }}</ref> Rates of [[vocabulary]] expansion link to the ability of [[children]] to vocally mirror non-words and so to acquire the new word pronunciations. Such [[speech repetition]] occurs automatically, fast<ref name="Porter">{{cite journal | vauthors = Porter RJ, Lubker JF | title = Rapid reproduction of vowel-vowel sequences: evidence for a fast and direct acoustic-motoric linkage in speech | journal = Journal of Speech and Hearing Research | volume = 23 | issue = 3 | pages = 593–602 | date = September 1980 | pmid = 7421161 | doi = 10.1044/jshr.2303.593 }}</ref> and separately in the brain to [[speech perception]].<ref name="McCarthy">{{cite journal | vauthors = McCarthy R, Warrington EK | title = A two-route model of speech production. Evidence from aphasia | journal = Brain | volume = 107 | issue = 2 | pages = 463–485 | date = June 1984 | pmid = 6722512 | doi = 10.1093/brain/107.2.463 | doi-access = free }}</ref><ref name="McCarthy2">{{cite journal | vauthors = McCarthy RA, Warrington EK | title = Repeating without semantics: surface dysphasia? | journal = Neurocase | volume = 7 | issue = 1 | pages = 77–87 | year = 2001 | pmid = 11239078 | doi = 10.1093/neucas/7.1.77 | s2cid = 12988855 }}</ref> Moreover, such vocal imitation can occur without comprehension such as in [[speech shadowing]]<ref>{{cite journal | vauthors = Marslen-Wilson W | title = Linguistic structure and speech shadowing at very short latencies | journal = Nature | volume = 244 | issue = 5417 | pages = 522–523 | date = August 1973 | pmid = 4621131 | doi = 10.1038/244522a0 | s2cid = 4220775 | bibcode = 1973Natur.244..522M }}</ref> and [[echolalia]].<ref>
{{cite journal | vauthors = Fay WH, Coleman RO | title = A human sound transducer/reproducer: temporal capabilities of a profoundly echolalic child | journal = Brain and Language | volume = 4 | issue = 3 | pages = 396–402 | date = July 1977 | pmid = 907878 | doi = 10.1016/0093-934X(77)90034-7 | s2cid = 29492873 }}</ref>

Further evidence for this link comes from a recent study in which the brain activity of two participants was measured using fMRI while they were gesturing words to each other using hand gestures with a game of [[charades]] – a modality that some have suggested might represent the evolutionary precursor of human language. Analysis of the data using [[Granger Causality]] revealed that the mirror-neuron system of the observer indeed reflects the pattern of activity in the motor system of the sender, supporting the idea that the motor concept associated with the words is indeed transmitted from one brain to another using the mirror system<ref name="Schippers">{{cite journal | vauthors = Schippers MB, Roebroeck A, Renken R, Nanetti L, Keysers C | title = Mapping the information flow from one brain to another during gestural communication | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 107 | issue = 20 | pages = 9388–9393 | date = May 2010 | pmid = 20439736 | pmc = 2889063 | doi = 10.1073/pnas.1001791107 | doi-access = free | bibcode = 2010PNAS..107.9388S }}</ref>

The mirror neuron system seems to be inherently inadequate to play any role in [[syntax]], given that this definitory property of human languages which is implemented in hierarchical recursive structure is flattened into linear sequences of phonemes making the recursive structure not accessible to sensory detection<ref>{{cite book |url=http://mitpress.mit.edu/catalog/item/default.asp?ttype=2&tid=11488 |title=The Boundaries of Babel. The Brain and the Enigma of Impossible Languages | vauthors = Moro A |publisher=[[MIT Press]] |year=2008 |isbn=978-0-262-13498-9 |pages=257}}</ref>

===Automatic imitation===
The term is commonly used to refer to cases in which an individual, having observed a body movement, unintentionally performs a similar body movement or alters the way that a body movement is performed. Automatic imitation rarely involves overt execution of matching responses. Instead the effects typically consist of reaction time, rather than accuracy, differences between compatible and incompatible trials.
Research reveals that the existence of automatic imitation, which is a covert form of imitation, is distinct from spatial compatibility. It also indicates that, although automatic imitation is subject to input modulation by attentional processes, and output modulation by inhibitory processes, it is mediated by learned, long-term sensorimotor associations that cannot be altered directly by intentional processes. Many researchers believe that automatic imitation is mediated by the mirror neuron system.<ref>{{cite journal | vauthors = Longo MR, Kosobud A, Bertenthal BI | title = Automatic imitation of biomechanically possible and impossible actions: effects of priming movements versus goals | journal = Journal of Experimental Psychology. Human Perception and Performance | volume = 34 | issue = 2 | pages = 489–501 | date = April 2008 | pmid = 18377184 | doi = 10.1037/0096-1523.34.2.489 | url = https://eprints.bbk.ac.uk/id/eprint/5423/1/5423.pdf }}</ref> Additionally, there are data that demonstrate that our postural control is impaired when people listen to sentences about other actions. For example, if the task is to maintain posture, people do it worse when they listen to sentences like this: "I get up, put on my slippers, go to the bathroom."  This phenomenon may be due to the fact that during action perception there is similar motor cortex activation as if a human being performed the same action (mirror neurons system).<ref name="Kosonogov2011">{{cite journal | vauthors = Kosonogov V | title = Listening to action-related sentences impairs postural control | journal = Journal of Electromyography and Kinesiology | volume = 21 | issue = 5 | pages = 742–745 | date = October 2011 | pmid = 21705230 | doi = 10.1016/j.jelekin.2011.05.007 }}</ref>

===Motor mimicry===
In contrast with automatic imitation, [[motor mimicry]] is observed in (1) naturalistic social situations and (2) via measures of action frequency within a session rather than measures of speed and/or accuracy within trials.<ref>{{cite journal | vauthors = Chartrand TL, Bargh JA | title = The chameleon effect: the perception-behavior link and social interaction | journal = Journal of Personality and Social Psychology | volume = 76 | issue = 6 | pages = 893–910 | date = June 1999 | pmid = 10402679 | doi = 10.1037/0022-3514.76.6.893 | s2cid = 11818459 }}</ref>

The integration of research on motor mimicry and automatic imitation could reveal plausible indications that these phenomena depend on the same psychological and neural processes. Preliminary evidence however comes from studies showing that social priming has similar effects on motor mimicry.<ref>{{cite journal | vauthors = Lakin JL, Chartrand TL | title = Using nonconscious behavioral mimicry to create affiliation and rapport | journal = Psychological Science | volume = 14 | issue = 4 | pages = 334–339 | date = July 2003 | pmid = 12807406 | doi = 10.1111/1467-9280.14481 | s2cid = 8458849 }}</ref><ref>{{cite journal | vauthors = van Baaren RB, Maddux WW, Chartrand TL, de Bouter C, van Knippenberg A | title = It takes two to mimic: behavioral consequences of self-construals | journal = Journal of Personality and Social Psychology | volume = 84 | issue = 5 | pages = 1093–1102 | date = May 2003 | pmid = 12757151 | doi = 10.1037/0022-3514.84.5.1093 | s2cid = 729948 | hdl = 2066/63130 | hdl-access = free }}</ref>

Nevertheless, the similarities between automatic imitation, mirror effects, and motor mimicry have led some researchers to propose that automatic imitation is mediated by the mirror neuron system and that it is a tightly controlled laboratory equivalent of the motor mimicry observed in naturalistic social contexts. If true, then automatic imitation can be used as a tool to investigate how the mirror neuron system contributes to cognitive functioning and how motor mimicry promotes prosocial attitudes and behavior.<ref>{{cite journal | vauthors = Heyes C | title = Automatic imitation | journal = Psychological Bulletin | volume = 137 | issue = 3 | pages = 463–483 | date = May 2011 | pmid = 21280938 | doi = 10.1037/a0022288 | s2cid = 6975248 }}</ref><ref name="pmid19679816">{{cite journal | vauthors = Paukner A, Suomi SJ, Visalberghi E, Ferrari PF | title = Capuchin monkeys display affiliation toward humans who imitate them | journal = Science | volume = 325 | issue = 5942 | pages = 880–883 | date = August 2009 | pmid = 19679816 | pmc = 2764469 | doi = 10.1126/science.1176269 | bibcode = 2009Sci...325..880P }}</ref>

Meta-analysis of imitation studies in humans suggest that there is enough evidence of mirror system activation during imitation that mirror neuron involvement is likely, even though no published studies have recorded the activities of singular neurons. However, it is likely insufficient for motor imitation. Studies show that regions of the frontal and parietal lobes that extend beyond the classical mirror system are equally activated during imitation. This suggests that other areas, along with the mirror system are crucial to imitation behaviors.<ref name="pmid19580913"/>

===Autism===

It has also been proposed that problems with the mirror neuron system may underlie cognitive disorders, particularly [[autism]].<ref name="Oberman_2005">{{cite journal | vauthors = Oberman LM, Hubbard EM, McCleery JP, Altschuler EL, Ramachandran VS, Pineda JA | title = EEG evidence for mirror neuron dysfunction in autism spectrum disorders | journal = Brain Research. Cognitive Brain Research | volume = 24 | issue = 2 | pages = 190–198 | date = July 2005 | pmid = 15993757 | doi = 10.1016/j.cogbrainres.2005.01.014 }}</ref><ref name="Mirella Dapretto pp. 28-30">{{cite journal | vauthors = Dapretto M, Davies MS, Pfeifer JH, Scott AA, Sigman M, Bookheimer SY, Iacoboni M | title = Understanding emotions in others: mirror neuron dysfunction in children with autism spectrum disorders | journal = Nature Neuroscience | volume = 9 | issue = 1 | pages = 28–30 | date = January 2006 | pmid = 16327784 | pmc = 3713227 | doi = 10.1038/nn1611 }}</ref> However the connection between mirror neuron dysfunction and autism is tentative and it remains to be demonstrated how mirror neurons are related to many of the important characteristics of autism.<ref name="Dinstein" />

Some researchers claim there is a link between mirror neuron deficiency and [[autism]]. EEG recordings from motor areas are suppressed when someone watches another person move, a signal that may relate to mirror neuron system. Additionally, the correlation can be measured with eye-movement tracking of biological motions, together with EEG recordings, mu suppression index can be calculated.<ref>{{cite journal | vauthors = Sotoodeh MS, Chien SH, Hadjikhani N | title = Visual attention modulates mu suppression during biological motion perception in autistic individuals | journal = European Journal of Neuroscience | volume = 13 | date = November 2024 | issue = 11 | pages = 6668–6685 | pmid = 39537315 | doi = 10.1111/ejn.16596 }}</ref> This suppression was less in children with autism.<ref name="Oberman_2005" /> Although these findings have been replicated by several groups,<ref>{{cite journal | vauthors = Dapretto M, Davies MS, Pfeifer JH, Scott AA, Sigman M, Bookheimer SY, Iacoboni M | title = Understanding emotions in others: mirror neuron dysfunction in children with autism spectrum disorders | journal = Nature Neuroscience | volume = 9 | issue = 1 | pages = 28–30 | date = January 2006 | pmid = 16327784 | pmc = 3713227 | doi = 10.1038/nn1611 | name-list-style = vanc }}</ref><ref name="pmid20598548">{{cite journal | vauthors = Perkins T, Stokes M, McGillivray J, Bittar R | title = Mirror neuron dysfunction in autism spectrum disorders | journal = Journal of Clinical Neuroscience | volume = 17 | issue = 10 | pages = 1239–1243 | date = October 2010 | pmid = 20598548 | doi = 10.1016/j.jocn.2010.01.026 | s2cid = 15141982 }}</ref> other studies have not found evidence of a dysfunctional mirror neuron system in autism.<ref name="Dinstein" /> In 2008, Oberman et al. published a research paper that presented conflicting EEG evidence. Oberman and Ramachandran found typical mu-suppression for familiar stimuli, but not for unfamiliar stimuli, leading them to conclude that the mirror neuron system of children with ASD (Autism Spectrum Disorder) was functional, but less sensitive than that of typical children.<ref name="pmid18304590">{{cite journal | vauthors = Oberman LM, Ramachandran VS, Pineda JA | title = Modulation of mu suppression in children with autism spectrum disorders in response to familiar or unfamiliar stimuli: the mirror neuron hypothesis | journal = Neuropsychologia | volume = 46 | issue = 5 | pages = 1558–1565 | date = April 2008 | pmid = 18304590 | doi = 10.1016/j.neuropsychologia.2008.01.010 | s2cid = 14280719 }}</ref> Based on the conflicting evidence presented by mu-wave suppression experiments, [[Patricia Churchland]] has cautioned that mu-wave suppression results cannot be used as a valid index for measuring the performance of mirror neuron systems.<ref>{{cite book | vauthors = Churcland PS |year=2011 |title=Braintrust |chapter=6 |page=156 |publisher=Princeton University Press}}</ref> Recent research indicates that mirror neurons do not play a role in autism:

{{Blockquote | ...no clear cut evidence emerges for a fundamental mirror system deficit in autism. Behavioural studies have shown that people with autism have a good understanding of action goals. Furthermore, two independent neuroimaging studies have reported that the parietal component of the mirror system is functioning typically in individuals with autism.<ref>{{cite book | vauthors = Hamilton A, Marsh L | chapter = Two systems for action comprehension in autism: mirroring and mentalizing. | veditors = Baron-Cohen S, Lombardo M, Tager-Flusberg H| title = Understanding Other Minds: Perspectives from developmental social neuroscience. | publisher = OUP | location = Oxford | isbn = 978-0-19-166880-7 | date = August 2013 | pages = 380–396 | chapter-url = http://www.antoniahamilton.com/HamiltonMarsh_UoM_preprint.pdf }}</ref>|sign=|source=}}

Some anatomical differences have been found in the mirror neuron related brain areas in adults with autism spectrum disorders, compared to non-autistic adults. All these cortical areas were thinner and the degree of thinning was correlated with autism symptom severity, a correlation nearly restricted to these brain regions.<ref name="Hadjikhani">{{cite journal | vauthors = Hadjikhani N, Joseph RM, Snyder J, Tager-Flusberg H | title = Anatomical differences in the mirror neuron system and social cognition network in autism | journal = Cerebral Cortex | volume = 16 | issue = 9 | pages = 1276–1282 | date = September 2006 | pmid = 16306324 | doi = 10.1093/cercor/bhj069 | doi-access = free }}</ref> Based on these results, some researchers claim that autism is caused by impairments in the mirror neuron system, leading to disabilities in social skills, imitation, empathy and theory of mind.{{Who|date=February 2014}}

Many researchers have pointed out that the "broken mirrors" theory of autism is overly simplistic, and mirror neurons alone cannot explain the differences found in individuals with autism. First of all, as noted above, none of these studies were direct measures of mirror neuron activity - in other words fMRI activity or EEG rhythm suppression do not unequivocally index mirror neurons. Dinstein and colleagues found normal mirror neuron activity in people with autism using fMRI.<ref name = "Callaway_2010">{{Cite web | date = 12 May 2010 | vauthors = Callaway E | work = New Scientist |url= https://www.newscientist.com/article/dn18837-mirror-neurons-seen-behaving-normally-in-autism/|title=Mirror neurons seen behaving normally in autism}}</ref> In individuals with autism, deficits in intention understanding, action understanding and biological motion perception (the key functions of mirror neurons) are not always found,<ref>{{cite journal | vauthors = Hamilton AF | title = Goals, intentions and mental states: challenges for theories of autism | journal = Journal of Child Psychology and Psychiatry, and Allied Disciplines | volume = 50 | issue = 8 | pages = 881–892 | date = August 2009 | pmid = 19508497 | doi = 10.1111/j.1469-7610.2009.02098.x | citeseerx = 10.1.1.621.6275 }}</ref><ref>{{cite journal | vauthors = Murphy P, Brady N, Fitzgerald M, Troje NF | title = No evidence for impaired perception of biological motion in adults with autistic spectrum disorders | journal = Neuropsychologia | volume = 47 | issue = 14 | pages = 3225–3235 | date = December 2009 | pmid = 19666038 | doi = 10.1016/j.neuropsychologia.2009.07.026 | s2cid = 12495492 }}</ref> or are task dependent.<ref>{{cite journal | vauthors = Saygin AP, Cook J, Blakemore SJ | title = Unaffected perceptual thresholds for biological and non-biological form-from-motion perception in autism spectrum conditions | journal = PLOS ONE | volume = 5 | issue = 10 | pages = e13491 | date = October 2010 | pmid = 20976151 | pmc = 2956672 | doi = 10.1371/journal.pone.0013491 | veditors = Baker CI | doi-access = free | bibcode = 2010PLoSO...513491S  }}</ref><ref>{{cite journal | vauthors = Cook J, Saygin AP, Swain R, Blakemore SJ | title = Reduced sensitivity to minimum-jerk biological motion in autism spectrum conditions | journal = Neuropsychologia | volume = 47 | issue = 14 | pages = 3275–3278 | date = December 2009 | pmid = 19632248 | pmc = 2779370 | doi = 10.1016/j.neuropsychologia.2009.07.010 }}</ref> Today, very few people believe an all-or-nothing problem with the mirror system can underlie autism. Instead, "additional research needs to be done, and more caution should be used when reaching out to the media."<ref>{{cite journal | vauthors = Fan YT, Decety J, Yang CY, Liu JL, Cheng Y | title = Unbroken mirror neurons in autism spectrum disorders | journal = Journal of Child Psychology and Psychiatry, and Allied Disciplines | volume = 51 | issue = 9 | pages = 981–988 | date = September 2010 | pmid = 20524939 | doi = 10.1111/j.1469-7610.2010.02269.x }}</ref>

Research from 2010<ref name = "Callaway_2010" /> concluded that autistic individuals do not exhibit mirror neuron dysfunction, although the small sample size limits the extent to which these results can be generalized. A more recent review argued there was not enough neurological evidence to support this  “broken-mirror theory” of autism.<ref>{{cite journal | vauthors = Heyes C, Catmur C | title = What Happened to Mirror Neurons? | journal = Perspectives on Psychological Science | volume = 17 | issue = 1 | pages = 153–168 | date = January 2022 | pmid = 34241539 | pmc = 8785302 | doi = 10.1177/1745691621990638 }}</ref>

===Theory of mind===
In [[Philosophy of mind]], mirror neurons have become the primary rallying call of [[Simulation theory of empathy|simulation theorists]] concerning our "[[theory of mind]]."  "Theory of mind" refers to our ability to infer another person's mental state (i.e., beliefs and desires) from experiences or their behaviour.

There are several competing models which attempt to account for our theory of mind; the most notable in relation to mirror neurons is simulation theory. According to simulation theory, theory of mind is available because we [[subconscious]]ly empathize with the person we're observing and, accounting for relevant differences, imagine what we would desire and believe in that scenario.<ref>{{cite journal | vauthors = Gordon R |year=1986 |title=Folk psychology as simulation |journal=Mind and Language |volume=1 |issue=2 |pages=158–171 |doi=10.1111/j.1468-0017.1986.tb00324.x}}</ref><ref>{{cite journal | vauthors = Goldman A |year=1989 |title=Interpretation psychologized |journal=Mind and Language |volume=4 |issue=3 |pages=161–185 |doi=10.1111/j.1468-0017.1989.tb00249.x}}</ref> Mirror neurons have been interpreted as the mechanism by which we simulate others in order to better understand them, and therefore their discovery has been taken by some as a validation of simulation theory (which appeared a decade before the discovery of mirror neurons).<ref name="pmid21227300"/> More recently, Theory of Mind and Simulation have been seen as complementary systems, with different developmental time courses.<ref name="pmid12689375">{{cite journal | vauthors = Meltzoff AN, Decety J | title = What imitation tells us about social cognition: a rapprochement between developmental psychology and cognitive neuroscience | journal = Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences | volume = 358 | issue = 1431 | pages = 491–500 | date = March 2003 | pmid = 12689375 | pmc = 1351349 | doi = 10.1098/rstb.2002.1261 }}</ref><ref name="pmid16892982">{{cite journal | vauthors = Sommerville JA, Decety J | title = Weaving the fabric of social interaction: articulating developmental psychology and cognitive neuroscience in the domain of motor cognition | journal = Psychonomic Bulletin & Review | volume = 13 | issue = 2 | pages = 179–200 | date = April 2006 | pmid = 16892982 | doi = 10.3758/BF03193831 | s2cid = 14689479 }}</ref><ref>{{cite journal | vauthors = Keysers C, Gazzola V | title = Integrating simulation and theory of mind: from self to social cognition | journal = Trends in Cognitive Sciences | volume = 11 | issue = 5 | pages = 194–196 | date = May 2007 | pmid = 17344090 | doi = 10.1016/j.tics.2007.02.002 | s2cid = 18930071 }}</ref>

At the neuronal-level, in a 2015 study by Keren Haroush and Ziv Williams using jointly interacting primates performing an iterated prisoner's dilemma game, the authors identified neurons in the [[anterior cingulate cortex]] that selectively predicted an opponent's yet unknown decisions or covert state of mind. These "other-predictive neurons" differentiated between self and other decisions and were uniquely sensitive to social context, but they did not encode the opponent's observed actions or receipt of reward. These cingulate cells may therefore importantly complement the function of mirror neurons by providing additional information about other social agents that is not immediately observable or known.<ref name="pmid25728667">{{cite journal | vauthors = Haroush K, Williams ZM | title = Neuronal prediction of opponent's behavior during cooperative social interchange in primates | journal = Cell | volume = 160 | issue = 6 | pages = 1233–1245 | date = March 2015 | pmid = 25728667 | pmc = 4364450 | doi = 10.1016/j.cell.2015.01.045 }}</ref>

===Sex differences===
A series of recent studies conducted by Yawei Cheng, using a variety of neurophysiological measures, including [[Magnetoencephalography|MEG]],<ref name="pmid16837838">{{cite journal | vauthors = Cheng YW, Tzeng OJ, Decety J, Imada T, Hsieh JC | title = Gender differences in the human mirror system: a magnetoencephalography study | journal = NeuroReport | volume = 17 | issue = 11 | pages = 1115–1119 | date = July 2006 | pmid = 16837838 | doi = 10.1097/01.wnr.0000223393.59328.21 | s2cid = 18811017 }}</ref> spinal reflex excitability,<ref name="pmid17515795">{{cite journal | vauthors = Cheng Y, Decety J, Lin CP, Hsieh JC, Hung D, Tzeng OJ | title = Sex differences in spinal excitability during observation of bipedal locomotion | journal = NeuroReport | volume = 18 | issue = 9 | pages = 887–890 | date = June 2007 | pmid = 17515795 | doi = 10.1097/WNR.0b013e3280ebb486 | s2cid = 16295878 }}</ref> electroencephalography,<ref name="pmid19083993">{{cite journal | vauthors = Yang CY, Decety J, Lee S, Chen C, Cheng Y | title = Gender differences in the mu rhythm during empathy for pain: an electroencephalographic study | journal = Brain Research | volume = 1251 | pages = 176–184 | date = January 2009 | pmid = 19083993 | doi = 10.1016/j.brainres.2008.11.062 | s2cid = 40145972 }}</ref><ref name="pmid18461176">{{cite journal | vauthors = Cheng Y, Lee PL, Yang CY, Lin CP, Hung D, Decety J | title = Gender differences in the mu rhythm of the human mirror-neuron system | journal = PLOS ONE | volume = 3 | issue = 5 | pages = e2113 | date = May 2008 | pmid = 18461176 | pmc = 2361218 | doi = 10.1371/journal.pone.0002113 | doi-access = free | bibcode = 2008PLoSO...3.2113C }}</ref> have documented the presence of a gender difference in the human mirror neuron system, with female participants exhibiting stronger motor resonance than male participants.

In another study, sex-based differences among mirror neuron mechanisms was reinforced in that the data showed enhanced empathetic ability in females relative to males{{Citation needed|date=January 2023}}. During an emotional social interaction, females showed a greater ability in emotional perspective taking{{Clarification needed|reason="emotional perspective taking" is meaningless without a definition; intentionally vague|date=January 2023}} than did males when interacting with another person face-to-face. However, in the study, data showed that when it came to recognizing the emotions of others, all participants' abilities were very similar and there was no key difference between the male and female subjects.<ref name="Schulte-Rüther 2008">{{cite journal | vauthors = Schulte-Rüther M, Markowitsch HJ, Shah NJ, Fink GR, Piefke M | title = Gender differences in brain networks supporting empathy | journal = NeuroImage | volume = 42 | issue = 1 | pages = 393–403 | date = August 2008 | pmid = 18514546 | doi = 10.1016/j.neuroimage.2008.04.180 | s2cid = 10461927 }}</ref>

=== Sleep paralysis ===
Baland Jalal and [[Vilayanur S. Ramachandran|V. S. Ramachandran]] have hypothesized that the mirror neuron system is important in giving rise to the intruder hallucination and out-of-body experiences during [[sleep paralysis]].<ref name=":8">{{cite journal | vauthors = Jalal B, Ramachandran VS | title = Sleep Paralysis, "The Ghostly Bedroom Intruder" and Out-of-Body Experiences: The Role of Mirror Neurons | journal = Frontiers in Human Neuroscience | volume = 11 | pages = 92 | date = 2017 | pmid = 28293186 | pmc = 5329044 | doi = 10.3389/fnhum.2017.00092 | doi-access = free }}</ref> According to this theory, sleep paralysis leads to disinhibition of the mirror neuron system, paving the way for hallucinations of human-like shadowy beings. The deafferentation of sensory information during sleep paralysis is proposed as the mechanism for such mirror neuron disinhibition.<ref name=":8" /> The authors suggest that their hypothesis on the role of the mirror neuron system could be tested: {{ blockquote | "These ideas could be explored using neuroimaging, to examine the selective activation of brain regions associated with mirror neuron activity, when the individual is hallucinating an intruder or having an out-of-body experience during sleep paralysis ."<ref name=":8">{{cite journal | vauthors = Jalal B, Ramachandran VS | title = Sleep Paralysis, "The Ghostly Bedroom Intruder" and Out-of-Body Experiences: The Role of Mirror Neurons | journal = Frontiers in Human Neuroscience | volume = 11 | pages = 92 | date = 2017 | pmid = 28293186 | pmc = 5329044 | doi = 10.3389/fnhum.2017.00092 | doi-access = free }}</ref> }}

=== Mirror neuron function, psychosis, and empathy in schizophrenia ===

Recent research, which measured mu-wave suppression, suggests that mirror neuron activity is positively correlated with psychotic symptoms (i.e., greater mu suppression/mirror neuron activity was highest among subjects with the greater severity of psychotic symptoms). Researchers concluded that "higher mirror neuron activity may be the underpinning of schizophrenia sensory gating deficits and may contribute to sensory misattributions particularly in response to socially relevant stimuli, and be a putative mechanism for delusions and hallucinations."<ref name="pmid22510432">{{cite journal | vauthors = McCormick LM, Brumm MC, Beadle JN, Paradiso S, Yamada T, Andreasen N | title = Mirror neuron function, psychosis, and empathy in schizophrenia | journal = Psychiatry Research | volume = 201 | issue = 3 | pages = 233–239 | date = March 2012 | pmid = 22510432 | pmc = 3545445 | doi = 10.1016/j.pscychresns.2012.01.004 }}</ref>