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Mind, Body, World- Foundations of Cognitive Science, 2013a

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the human brain (Gallese et al., 1996; Iacoboni, 2008; Rizzolatti & Craighero, 2004;<br />

Rizzolatti, Fogassi, & Gallese, 2006) are specialized for both the generation and<br />

interpretation <strong>of</strong> gestures and actions.<br />

Mirror neurons were serendipitously discovered in experiments in which motor<br />

neurons in region F5 were recorded when monkeys performed various reaching<br />

actions (Di Pellegrino et al., 1992). By accident, it was discovered that many <strong>of</strong> the<br />

neurons that were active when a monkey performed an action also responded when<br />

similar actions were observed being performed by another:<br />

After the initial recording experiments, we incidentally observed that some experimenter’s<br />

actions, such as picking up the food or placing it inside the testing box,<br />

activated a relatively large proportion <strong>of</strong> F5 neurons in the absence <strong>of</strong> any overt<br />

movement <strong>of</strong> the monkey. (Di Pellegrino et al., 1992, p. 176)<br />

The chance discovery <strong>of</strong> mirror neurons has led to an explosion <strong>of</strong> research into<br />

their behaviour (Iacoboni, 2008). It has been discovered that when the neurons fire,<br />

they do so for the entire duration <strong>of</strong> the observed action, not just at its onset. They<br />

are grasp specific: some respond to actions involving precision grips, while others<br />

respond to actions involving larger objects. Some are broadly tuned, in the sense<br />

that they will be triggered when a variety <strong>of</strong> actions are observed, while others are<br />

narrowly tuned to specific actions. All seem to be tuned to object-oriented action:<br />

a mirror neuron will respond to a particular action on an object, but it will fail to<br />

respond to the identical action if no object is present.<br />

While most <strong>of</strong> the results described above were obtained from studies <strong>of</strong> the<br />

monkey brain, there is a steadily growing literature indicating that the human brain<br />

also has a mirror system (Buccino et al., 2001; Iacoboni, 2008).<br />

Mirror neurons are not solely concerned with hand and arm movements. For<br />

instance, some monkey mirror neurons respond to mouth movements, such as lip<br />

smacking (Ferrari et al., 2003). Similarly, the human brain has a mirror system<br />

for the act <strong>of</strong> touching (Keysers et al., 2004). Likewise, another part <strong>of</strong> the human<br />

brain, the insula, may be a mirror system for emotion (Wicker et al., 2003). For<br />

example, it generates activity when a subject experiences disgust, and also when a<br />

subject observes the facial expressions <strong>of</strong> someone else having a similar experience.<br />

Two decades after its discovery, extensive research on the mirror neuron<br />

system has led some researchers to claim that it provides the neural substrate<br />

for social cognition and imitative learning (Gallese & Goldman, 1998; Gallese,<br />

Keysers, & Rizzolatti, 2004; Iacoboni, 2008), and that disruptions <strong>of</strong> this system may<br />

be responsible for autism (Williams et al., 2001). The growing understanding <strong>of</strong> the<br />

mirror system and advances in knowledge about the neuroscience <strong>of</strong> face perception<br />

have heralded a new interdisciplinary research program, called social cognitive<br />

neuroscience (Blakemore, Winston, & Frith, 2004; Lieberman, 2007; Ochsner &<br />

Lieberman, 2001).<br />

Elements <strong>of</strong> Embodied <strong>Cognitive</strong> <strong>Science</strong> 247

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