What Are Mirror Neurons?

A mirror neuron is a neuron that fires both when an animal acts and when the animal observes the same action performed by another. Thus, the neuron “mirrors” the behavior of the other, as though the observer were itself acting.

Such neurons have been directly observed in primate species. Birds have been shown to have imitative resonance behaviors and neurological evidence suggests the presence of some form of mirroring system. In humans, brain activity consistent with that of mirror neurons has been found in the premotor cortex, the supplementary motor area, the primary somatosensory cortex and the inferior parietal cortex.

The function of the mirror system in humans is a subject of much speculation. Some researchers in cognitive neuroscience and cognitive psychology consider that this system provides the physiological mechanism for the perception/action coupling. They argue that mirror neurons may be important for understanding the actions of other people, and for learning new skills by imitation.

Some researchers speculate that mirror systems may simulate observed actions, and thus contribute to theory of mind skills, while others relate mirror neurons to language abilities. Neuroscientists such as Marco Iacoboni (UCLA) have argued that mirror neuron systems in the human brain help us understand the actions and intentions of other people. In a study published in March 2005 Iacoboni and his colleagues reported that mirror neurons could discern whether another person who was picking up a cup of tea planned to drink from it or clear it from the table. In addition, Iacoboni has argued that mirror neurons are the neural basis of the human capacity for emotions such as empathy.

However, there are scientists who express skepticism about the theories being advanced to explain the function of mirror neurons. To date, no widely accepted neural or computational models have been put forward to describe how mirror neuron activity supports cognitive functions. The subject of mirror neurons continues to generate intense debate. In 2014, Philosophical Transactions of the Royal Society B published a special issue entirely devoted to mirror neuron research.

Short History Of Mirror Neurons

Giacomo Rizzolatti

Neurophysiologist Giacomo Rizzolatti.
Credit: Cirone-Musi – Giacomo Rizzolatti, CC BY-SA 2.0

In the 1980s and 1990s, neurophysiologists Giacomo Rizzolatti, Giuseppe Di Pellegrino, Luciano Fadiga, Leonardo Fogassi, and Vittorio Gallese at the University of Parma placed electrodes in the ventral premotor cortex of the macaque monkey to study neurons specialize 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.

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 sent to Nature, but was rejected for its “lack of general interest” before being published in a less competitive journal.

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 region was the homologue region of the monkey ventral premotor cortex. While these papers reported the presence of mirror neurons responding to hand actions, a subsequent study by Pier Francesco Ferrari and colleagues 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.

Reports on mirror neurons have been widely published and confirmed 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. 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.

Possible Functions

Many studies link mirror neurons to understanding goals and intentions. Fogassi et al. (2005) 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”. They may furnish a neural basis for predicting another individual’s subsequent actions and inferring intention.

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.


Stephanie Preston and Frans de Waal, Jean Decety, and Vittorio Gallese 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 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.

David Freedberg and Vittorio Gallese have also put forward the idea that this function of the mirror neuron system is crucial for aesthetic experiences. 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.

More recently, 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 and the mirror system for emotions, 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.

Human Self-awareness

V. S. Ramachandran has speculated that mirror neurons may provide the neurological basis of human self-awareness. 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”.


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. This hypothesis is supported by some cytoarchitectonic homologies between monkey premotor area F5 and human Broca’s area.

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 and separately in the brain to speech perception. Moreover, such vocal imitation can occur without comprehension such as in speech shadowing and echolalia.

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.

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.

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. 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).

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.

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.

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.

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.

Doubts About Mirror Neurons

Although many in the scientific community have expressed excitement about the discovery of mirror neurons, there are scientists who have expressed doubts about both the existence and role of mirror neurons in humans.

According to scientists such as Hickok, Pascolo, and Dinstein, it is not clear whether mirror neurons really form a distinct class of cells (as opposed to an occasional phenomenon seen in cells that have other functions), and whether mirror activity is a distinct type of response or simply an artifact of an overall facilitation of the motor system.

In 2008, Ilan Dinstein et al. argued that the original analyses were unconvincing because they were based on qualitative descriptions of individual cell properties, and did not take into account the small number of strongly mirror-selective neurons in motor areas.

Other scientists have argued that the measurements of neuron fire delay seem not to be compatible with standard reaction times, and pointed out that nobody has reported that an interruption of the motor areas in F5 would produce a decrease in action recognition. (Critics of this argument have replied that these authors have missed human neuropsychological and TMS studies reporting disruption of these areas do indeed cause action deficits without affecting other kinds of perception.)

In 2009, Lingnau et al. carried out an experiment in which they compared motor acts that were first observed and then executed to motor acts that were first executed and then observed. They concluded that there was a significant asymmetry between the two processes that indicated that mirror neurons do not exist in humans. They stated that:

“Crucially, we found no signs of adaptation for motor acts that were first executed and then observed.

Failure to find cross-modal adaptation for executed and observed motor acts is not compatible with the core assumption of mirror neuron theory, which holds that action recognition and understanding are based on motor simulation.”.

However, in the same year, Kilner et al. showed that if goal directed actions are used as stimuli, both IPL and premotor regions show the repetition suppression between observation and execution that is predicted by mirror neurons.

In 2009, Greg Hickok published an extensive argument against the claim that mirror neurons are involved in action-understanding: “Eight Problems for the Mirror Neuron Theory of Action Understanding in Monkeys and Humans.” He concluded that

“The early hypothesis that these cells underlie action understanding is likewise an interesting and prima facie reasonable idea. However, despite its widespread acceptance, the proposal has never been adequately tested in monkeys, and in humans there is strong empirical evidence, in the form of physiological and neuropsychological (double-) dissociations, against the claim.”

Vladimir Kosonogov sees another contradiction. The proponents of mirror neuron theory of action understanding postulate that the mirror neurons code the goals of others actions because they are activated if the observed action is goal-directed.

However, the mirror neurons are activated only when the observed action is goal-directed (object-directed action or a communicative gesture, which certainly has a goal too).

How do they “know” that the definite action is goal-directed? At what stage of their activation do they detect a goal of the movement or its absence? In his opinion, the mirror neuron system can be activated only after the goal of the observed action is attributed by some other brain structures.

Neurophilosophers such as Patricia Churchland have expressed both scientific and philosophical objections to the theory that mirror neurons are responsible for understanding the intentions of others. In chapter 5 of her 2011 book, Braintrust, Churchland points out that the claim that mirror neurons are involved in understanding intentions (through simulating observed actions) is based on assumptions that are clouded by unresolved philosophical issues.

She makes the argument that intentions are understood (coded) at a more complex level of neural activity than that of individual neurons. Churchland states that

“A neuron, though computationally complex, is just a neuron. It is not an intelligent homunculus. If a neural network represents something complex, such as an intention [to insult], it must have the right input and be in the right place in the neural circuitry to do that”.

Recently, Cecilia Heyes (Professor of Experimental Psychology, Oxford) has advanced the theory that mirror neurons are the byproduct of associative learning as opposed to evolutionary adaptation. She argues that mirror neurons in humans are the product of social interaction and not an evolutionary adaptation for action-understanding. In particular, Heyes rejects the theory advanced by V.S. Ramachandran that mirror neurons have been “the driving force behind the great leap forward in human evolution.”

Keysers, C.
The Empathic Brain
Social Brain Press (2011). ISBN 1-4637-6906-7

Hickok, Greg
The Myth of Mirror Neurons
W. W. Norton & Company (2014). ISBN 978-0-393-08961-5