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Mirror neuron
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==Discovery== In the 1980s and 1990s, neurophysiologists [[Giacomo Rizzolatti]], Giuseppe Di Pellegrino, [[Luciano Fadiga]], Leonardo Fogassi, and [[Vittorio Gallese]] at the [[University of Parma]] placed [[electrode]]s in the [[Premotor cortex|ventral premotor cortex]] of the [[macaque]] monkey to study [[neurons]] specialized in the control of hand and mouth actions; for example, taking hold of an object and manipulating it. During each experiment, the researchers allowed the monkey to reach for pieces of food, and recorded from single neurons in the monkey's brain, thus measuring the neuron's response to certain movements.<ref>{{cite journal | vauthors = di Pellegrino G, Fadiga L, Fogassi L, Gallese V, Rizzolatti G | title = Understanding motor events: a neurophysiological study | journal = Experimental Brain Research | volume = 91 | issue = 1 | pages = 176β180 | year = 1992 | pmid = 1301372 | doi = 10.1007/bf00230027 | s2cid = 206772150 }}</ref><ref>{{cite journal | vauthors = Rizzolatti G, Fadiga L, Gallese V, Fogassi L | title = Premotor cortex and the recognition of motor actions | journal = Brain Research. Cognitive Brain Research | volume = 3 | issue = 2 | pages = 131β141 | date = March 1996 | pmid = 8713554 | doi = 10.1016/0926-6410(95)00038-0 }}</ref> They found that some neurons responded when the monkey observed a person picking up a piece of food, and also when the monkey itself picked up the food. The discovery was initially submitted to [[Nature (journal)|''Nature'']], but was rejected for its "lack of general interest" before being published in a less competitive journal.<ref name="pmid19760408">{{cite journal | vauthors = Rizzolatti G, Fabbri-Destro M | title = Mirror neurons: from discovery to autism | journal = Experimental Brain Research | volume = 200 | issue = 3β4 | pages = 223β237 | date = January 2010 | pmid = 19760408 | doi = 10.1007/s00221-009-2002-3 | s2cid = 3342808 }}</ref> A few years later, the same group published another empirical paper, discussing the role of the mirror-neuron system in action recognition, and proposing that the human [[Broca's area]] was the [[Homology (biology)|homologue]] region of the monkey ventral premotor cortex.<ref name="GalleseFadiga1996">{{cite journal | vauthors = Gallese V, Fadiga L, Fogassi L, Rizzolatti G | title = Action recognition in the premotor cortex | journal = Brain | volume = 119 | issue = 2 | pages = 593β609 | date = April 1996 | pmid = 8800951 | doi = 10.1093/brain/119.2.593 | doi-access = free }}</ref> While these papers reported the presence of mirror neurons responding to hand actions, a subsequent study by Pier Francesco Ferrari and colleagues<ref>{{cite journal | vauthors = Ferrari PF, Gallese V, Rizzolatti G, Fogassi L | title = Mirror neurons responding to the observation of ingestive and communicative mouth actions in the monkey ventral premotor cortex | journal = The European Journal of Neuroscience | volume = 17 | issue = 8 | pages = 1703β1714 | date = April 2003 | pmid = 12752388 | doi = 10.1046/j.1460-9568.2003.02601.x | s2cid = 1915143 }}</ref> described the presence of mirror neurons responding to mouth actions and facial gestures. Further experiments confirmed that about 10% of neurons in the monkey inferior frontal and inferior parietal cortex have "mirror" properties and give similar responses to performed hand actions and observed actions. In 2002 [[Christian Keysers]] and colleagues reported that, in both humans and monkeys, the mirror system also responds to the sound of actions.<ref name="EmpathicBrain" /><ref name="pmid12161656">{{cite journal | vauthors = Kohler E, Keysers C, UmiltΓ MA, Fogassi L, Gallese V, Rizzolatti G | title = Hearing sounds, understanding actions: action representation in mirror neurons | journal = Science | location = New York, N.Y. | volume = 297 | issue = 5582 | pages = 846β8 | date = August 2002 | pmid = 12161656 | doi = 10.1126/science.1070311 | bibcode = 2002Sci...297..846K | s2cid = 16923101 }}</ref><ref name="pmid16979560">{{cite journal | vauthors = Gazzola V, Aziz-Zadeh L, Keysers C | title = Empathy and the somatotopic auditory mirror system in humans | journal = Current Biology | volume = 16 | issue = 18 | pages = 1824β9 | date = September 2006 | pmid = 16979560 | doi = 10.1016/j.cub.2006.07.072 | s2cid = 5223812 | doi-access = free | bibcode = 2006CBio...16.1824G }}</ref> Reports on mirror neurons have been widely published<ref name="GalleseFadiga1996"/> and confirmed<ref name="pmid15860620">{{cite journal | vauthors = Fogassi L, Ferrari PF, Gesierich B, Rozzi S, Chersi F, Rizzolatti G | title = Parietal lobe: from action organization to intention understanding | journal = Science | location = New York, N.Y. | volume = 308 | issue = 5722 | pages = 662β7 | date = April 2005 | pmid = 15860620 | doi = 10.1126/science.1106138 | bibcode = 2005Sci...308..662F | s2cid = 5720234 }}</ref> with mirror neurons found in both inferior frontal and inferior parietal regions of the brain. Recently, evidence from [[functional neuroimaging]] strongly suggests that humans have similar mirror neurons systems: researchers have identified brain regions which respond during both action and observation of action. Not surprisingly, these brain regions include those found in the macaque monkey.<ref name="RizzolattiCraighero2004" /> However, [[functional magnetic resonance imaging]] (fMRI) can examine the entire brain at once and suggests that a much wider network of brain areas shows mirror properties in humans than previously thought. These additional areas include the [[somatosensory cortex]] and are thought to make the observer feel what it feels like to move in the observed way.<ref name="GazzolaKeysers2009">{{cite journal | vauthors = Gazzola V, Keysers C | title = The observation and execution of actions share motor and somatosensory voxels in all tested subjects: single-subject analyses of unsmoothed fMRI data | journal = Cerebral Cortex | volume = 19 | issue = 6 | pages = 1239β1255 | date = June 2009 | pmid = 19020203 | pmc = 2677653 | doi = 10.1093/cercor/bhn181 }}</ref><ref name="KeysersKaasGazzola2010">{{cite journal | vauthors = Keysers C, Kaas JH, Gazzola V | title = Somatosensation in social perception | journal = Nature Reviews. Neuroscience | volume = 11 | issue = 6 | pages = 417β428 | date = June 2010 | pmid = 20445542 | doi = 10.1038/nrn2833 | s2cid = 12221575 }}</ref>
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