Jump to content

Mirror neuron

From Wikipedia, the free encyclopedia

This is an old revision of this page, as edited by 170.213.131.190 (talk) at 20:59, 10 January 2012 (edit summary removed). The present address (URL) is a permanent link to this revision, which may differ significantly from the current revision.

A mirror neuron is a neuron that fires both when an animal acts and when the animal observes the same action performed by another.[1][2][3] Thus, the neuron "mirrors" the behaviour of the other, as though the observer were itself acting. Such neurons have been directly observed in primate and other species including birds. 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.

Mirror neurons were first described in 1992. Some scientists consider this to be one of the most important recent discoveries in neuroscience. Among them is V.S. Ramachandran, who believes they might be very important in imitation and language acquisition. Ramachandran has also speculated that mirror neurons are involved in understanding other people's feelings (empathy) and that they have played a role in the development of human culture.[4] However, scientists such as Greg Hickok have expressed skepticism in regard to the claim that mirror neurons are the basis of the cognitive ability to understand others' intentions or feelings. Hickok, the director of the Center for Cognitive Neuroscience at UC Irvine, has argued that there is little or no evidence to support the claim that mirror neurons are involved in understanding the intentions of others.[5] Despite the excitement generated by these findings, to date no widely accepted neural or computational models have been put forward to describe how mirror neuron activity supports cognitive functions such as imitation.[6]

The function of the mirror system is a subject of much speculation. Many researchers in cognitive neuroscience and cognitive psychology consider that this system provides the physiological mechanism for the perception action coupling (see the common coding theory).[3] These mirror neurons may be important for understanding the actions of other people, and for learning new skills by imitation. Some researchers also speculate that mirror systems may simulate observed actions, and thus contribute to theory of mind skills,[7][8] while others relate mirror neurons to language abilities.[9] It has also been proposed that problems with the mirror system may underlie cognitive disorders, particularly autism.[10][11] However the connection between mirror neuron dysfunction and autism is tentative and it remains to be seen how mirror neurons may be related to many of the important characteristics of autism.[6]

Discovery

In the 1980s and 1990s, Giacomo Rizzolatti was working with Giuseppe Di Pellegrino, Luciano Fadiga, Leonardo Fogassi, and Vittorio Gallese at the University of Parma, Italy. These neurophysiologists had placed electrodes in the ventral premotor cortex of the macaque monkey to study neurons specialized for 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 a single neuron in the monkey's brain, thus measuring the neuron's response to certain movements.[12][13] They found that some of the neurons they recorded from would respond when the monkey saw a person pick up a piece of food as well as when the monkey picked up the food. The discovery was initially sent to Nature but was rejected for its "lack of general interest".[14]

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.[15] While these papers reported the presence of mirror neurons responding to hand actions, a subsequent study by Ferrari Pier Francesco and colleagues[16] 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.[3][17][18]

Reports on mirror neurons have been widely published[19] and confirmed[20] 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[1] 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 [21][22]

In monkeys

Neonatal (newborn) macaque imitating facial expressions

The first animal in which mirror neurons have been studied individually is the macaque monkey. In these monkeys, mirror neurons are found in the inferior frontal gyrus (region F5) and the inferior parietal lobule.[1]

Mirror neurons are believed to mediate the understanding of other animals' behaviour. For example, a mirror neuron which fires when the monkey rips a piece of paper would also fire when the monkey sees a person rip paper, or hears paper ripping (without visual cues). These properties have led researchers to believe that mirror neurons encode abstract concepts of actions like 'ripping paper', whether the action is performed by the monkey or another animal.[1]

The function of mirror neurons in macaques is not known. Adult macaques do not seem to learn by imitation. Recent experiments by Ferrari and colleagues suggest that infant macaqes can imitate a human's face movements, though only as neonates and during a limited temporal window.[23] Even if it has not yet been empirically demonstrated, it has been proposed that mirror neurons underlie this behaviour and other imitative phenomena.[24] Indeed, there is limited understanding of the degree to which monkeys show imitative behaviour.[6]

In adult monkeys, mirror neurons may enable the monkey to understand what another monkey is doing, or to recognise the other monkey's action.[25]

In humans

Diagram of the brain, showing the locations of the frontal and parietal lobes of the cerebrum, viewed from the left. The inferior frontal lobe is the lower part of the blue area, and the superior parietal lobe is the upper part of the yellow area.

It is not normally possible to study single neurons in the human brain, so most evidence for mirror neurons in humans is indirect. Brain imaging experiments using functional magnetic resonance imaging (fMRI) have shown that the human inferior frontal cortex and superior parietal lobe is active when the person performs an action and also when the person sees another individual performing an action. It has been suggested that these brain regions contain mirror neurons, and they have been defined as the human mirror neuron system.[26] More recent experiments have shown that even at the level of single participants, scanned using fMRI, large areas containing multiple fMRI voxels increase their activity both during the observation and execution of actions.[21]

Neuropsychological studies looking at lesion areas that cause action knowledge, pantomime interpretation, and biological motion perception deficits have pointed to a causal link between the integrity of the inferior frontal gyrus and these behaviours.[27][28][29] Transcranial magnetic stimulation studies have confirmed this as well.[30][31] These results indicate the activation in mirror neuron related areas are unlikely to be just epiphenomenal.

A study published in April 2010 reports recordings from single neurons with mirror properties in the human brain.[32] Mukamel et al (Current Biology, 2010) recorded from the brains of 21 patients who were being treated at Ronald Reagan UCLA Medical Center for intractable epilepsy. The patients had been implanted with intracranial depth electrodes to identify seizure foci for potential surgical treatment. Electrode location was based solely on clinical criteria; the researchers, with the patients' consent, used the same electrodes to "piggyback" their research. The researchers found a small number of neurons that fired or showed their greatest activity both when the individual performed a task and when they observed a task. Other neurons had anti-mirror properties, that is, they responded when the participant saw an action but were inhibited when the participant performed that action.

The mirror neurons found were located in the supplementary motor area and medial temporal cortex (other brain regions were not sampled). For purely practical reasons, these regions are not the same as those in which mirror neurons had been recorded from in the monkey: researchers in Parma were studying the ventral premotor cortex and the associated inferior parietal lobe, two regions in which epilepsy rarely occurs, and hence, single cell recordings in these regions are not usually done in humans. On the other hand, no one has to date looked for mirror neurons in the supplementary motor area or the medial temporal lobe in the monkey. Together, this therefore does not suggest that humans and monkeys have mirror neurons in different locations, but rather than they may have mirror neurons both in the ventral premotor cortex and inferior parietal lobe, where they have been recorded in the monkey, and in the supplementary motor areas and medial temporal lobe, where they have been recorded from in human – especially because detailed human fMRI analyses suggest activity compatible with the presence of mirror neurons in all these regions.[21]

Doubts concerning mirror neurons

Although many in the scientific community have expressed excitement about the discovery of mirror neurons, there are scientists who have cautioned that the claims being made for the significance of mirror neurons are not supported by sufficient research. 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.[6] Other scientists have argued that the measurements of neuron fire delay seem not to be compatible with standard reaction times,[33] and pointed out that nobody has reported that an interruption of the motor areas in F5 would produce a decrease in action recognition.[34] (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 [28][30] without affecting other kinds of perception.)[29] 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),[33] and whether mirror activity is a distinct type of response or simply an artifact of an overall facilitation of the motor system.[34]

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."[35]

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."[36]

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."(Ramachandran,2000)[37]

Development

Human infant data using eye-tracking measures suggest that the mirror neuron system develops before 12 months of age, and that this system may help human infants understand other people's actions.[38] A critical question concerns how mirror neurons acquire mirror properties. Two closely related models postulate that mirror neurons are trained through Hebbian[39] or Associative learning[40][41][42] (see Associative Sequence Learning). However, if premotor neurons need to be trained by action in order to acquire mirror properties, it is unclear how newborn babies are able to mimic the facial gestures of another person (imitation of unseen actions), as suggested by the work of Meltzoff and Moore. One possibility is that the sight of tongue protrusion recruits an innate releasing mechanism in neonates. Careful analysis suggests that 'imitation' of this single gesture may account for almost all reports of facial mimicry by new-born infants.[43]

Possible functions

Understanding intentions

Many studies link mirror neurons to understanding goals and intentions. Fogassi et al. (2005)[44] 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".[44] They may furnish a neural basis for predicting another individual’s subsequent actions and inferring intention.[44]

Empathy

Stephanie Preston and Frans de Waal,[45] Jean Decety,[46][47] and Vittorio Gallese[48][49] and Christian Keysers[3] have independently argued that the mirror neuron system is involved in empathy. A large number of experiments using functional MRI, 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.[50][51][52][53][54][55][56] 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[57] and the mirror system for emotions,[55] providing more direct support for the idea that the mirror system is linked to empathy.

Human self awareness

V.S Ramachandran has speculated that mirror neurons may provide the neurological basis of human self awareness.[58] 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." [59]

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.[60] This hypothesis is supported by some cytoarchitectonic homologies between monkey premotor area F5 and human Broca's area.[61] 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[62] and separately in the brain to speech perception.[63][64] Moreover such vocal imitation can occur without comprehension such as in speech shadowing[65] and echolalia.[66]

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 of the 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 [67]

It must be noticed that 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[68]

Automatic Imitation

The term is commonly used to refer to cases in which an individual, having observed a body movement, deliberately performs a topographically similar body movement. Automatic Imitation rarely involves overt behavioral execution of matching responses. Automatic imitation effects typically consist of RT, 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 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.[69]

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. [70]

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.[71][72]

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.[73][74]

Autism

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. This suppression was less in children with autism.[10] Although these findings have been replicated by several groups,[75][76] other studies have not found evidence of a dysfunctional mirror neuron system in autism.[6] 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 was functional, but less sensitive than that of typical children.[77] 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.[78] Finally, 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.[79] 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.

Many researchers have pointed out that the "broken mirrors" theory of autism is overly simplistic, and mirror neurons alone cannot explain the deficits 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.[80] In individuals with autism, deficits in intention understanding, action understanding and biological motion perception (the key functions of mirror neurons) are not always found,[81][82] or are task dependent.[83][84] 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".[85]

Theory of mind

In Philosophy of mind, mirror neurons have become the primary rallying call of 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. For example, if you see a girl reaching into a jar labeled 'cookies,' you might assume that she wants a cookie and believes that there are cookies in the jar (even if you know the jar is empty).

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 subconsciously empathize with the person we're observing and, accounting for relevant differences, imagine what we would desire and believe in that scenario.[86][87] 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).[88] More recently, Theory of Mind and Simulation have been seen as complementary systems, with different developmental time courses.[89][90][91]

Gender differences

The issue of gender differences in empathy is quite controversial and subject to social desirability and stereotypes. However, a series of recent studies conducted by Yawei Cheng, using a variety of neurophysiological measures, including MEG,[92] spinal reflex excitability,[93] electroencephalography,[94][95] have documented the presence of a gender difference in the human mirror neuron system, with female participants exhibiting stronger motor resonance than male participants.

See also

References

  1. ^ a b c d Rizzolatti, Giacomo; Craighero, Laila (2004). "The mirror-neuron system" (PDF). Annual Review of Neuroscience. 27: 169–192. doi:10.1146/annurev.neuro.27.070203.144230. PMID 15217330.
  2. ^ Keysers, Christian (2010). "Mirror Neurons" (PDF). Current Biology. 19 (21): R971–973. doi:10.1016/j.cub.2009.08.026. PMID 19922849.
  3. ^ a b c d Keysers, Christian (2011-06-23). The Empathic Brain. Kindle.
  4. ^ V.S. Ramachandran, "Mirror Neurons and imitation learning as the driving force behind "the great leap forward" in human evolution". Edge Foundation. Retrieved 2006-11-16.
  5. ^ Greg Hickok (2009-02-10). "Eight problems for the mirror neuron theory of action understanding". Talking Brains blog. Retrieved 2010-08-29. {{cite web}}: External link in |publisher= (help)
  6. ^ a b c d e Dinstein I, Thomas C, Behrmann M, Heeger DJ (2008). "A mirror up to nature". Curr Biol. 18 (1): R13–8. doi:10.1016/j.cub.2007.11.004. PMC 2517574. PMID 18177704.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  7. ^ Christian Keysers and Valeria Gazzola, Progress in Brain Research, 2006, [1]
  8. ^ Michael Arbib, The Mirror System Hypothesis. Linking Language to Theory of Mind, 2005, retrieved 2006-02-17
  9. ^ Théoret, Hugo; Pascual-Leone, Alvaro (2002). "Language Acquisition: Do as You Hear". Current Biology. 12 (21): R736–7. doi:10.1016/S0960-9822(02)01251-4. PMID 12419204.
  10. ^ a b Oberman LM, Hubbard EM, McCleery JP, Altschuler EL, Ramachandran VS, Pineda JA., EEG evidence for mirror neuron dysfunction in autism spectral disorders, Brain Res Cogn Brain Res.; 24(2):190-8, 2005-06
  11. ^ Dapretto, M; Davies, MS; Pfeifer, JH; Scott, AA; Sigman, M; Bookheimer, SY; Iacoboni, M (2006). "Understanding emotions in others: mirror neuron dysfunction in children with autism spectrum disorders". Nature neuroscience. 9 (1): 28–30. doi:10.1038/nn1611. PMID 16327784.
  12. ^ Di Pellegrino, G., Fadiga, L., Fogassi, L., Gallese, V., & Rizzolatti, G. (1992). Understanding motor events: a neurophysiological study. Experimental Brain Research, 91, 176-180.
  13. ^ Giacomo Rizzolatti et al. (1996) Premotor cortex and the recognition of motor actions, Cognitive Brain Research 3 131-141
  14. ^ Rizzolatti G, Fabbri-Destro M (2010). "Mirror neurons: from discovery to autism". Exp Brain Res. 200 (3–4): 223–37. doi:10.1007/s00221-009-2002-3. PMID 19760408.
  15. ^ Gallese, V.; Fadiga, L.; Fogassi, L.; Rizzolatti, Giacomo (1996). "Action recognition in the premotor cortex". Brain. 119 (2): 593–609. doi:10.1093/brain/119.2.593. PMID 8800951. {{cite journal}}: Unknown parameter |unused_data= ignored (help)
  16. ^ Ferrari et al. (2003) "Mirror neurons responding to the observation of ingestive and communicative mouth actions in the ventral premotor cortex. European Journal of Neuroscience, 17 (8), 1703-1714
  17. ^ Kohler et al., Science, 2002 [2]
  18. ^ Gazzola et al., Current Biology, 2006 [3]
  19. ^ Gallese et al., Action recognition in the premotor cortex, Brain, 1996
  20. ^ Fogassi et al., Parietal Lobe: From Action Organization to Intention Understanding, Science, 2005
  21. ^ a b c Gazzola, V.; Keysers, C. (2009). "The observation and execution of actions share motor and somatosensory voxels in all tested subjects: single-subject analyses of unsmoothed fMRI data". Cereb Cortex. 19 (6): 1239–1255. doi:10.1093/cercor/bhn181. PMC 2677653. PMID 19020203.
  22. ^ Keysers, Christian; Kaas, John; Gazzola, Valeria (2010). "Somatosensation in Social Cognition" (PDF). Nature Reviews Neuroscience. 11 (6): 417–28. doi:10.1038/nrn2833. PMID 20445542.
  23. ^ Ferrari, P. F.; Visalberghi, E.; Paukner, A.; Fogassi, L.; Ruggiero, A.; Suomi, SJ (2006). "Neonatal Imitation in Rhesus Macaques". PLoS Biology. 4 (9). et al.: e302. doi:10.1371/journal.pbio.0040302. PMC 1560174. PMID 16953662.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  24. ^ Ferrari P.F., Bonini L., Fogassi L. (2009) From monkey mirror neurons to mirror-related behaviours: possible direct and indirect pathways. Philosophical Transactions of the Royal Society, B. .364, 2311-2323
  25. ^ Rizzolatti, Giacomo; Arbib, Michael A. (1998). "Language within our grasp". Trends in Neurosciences. 21 (5): 188–194. doi:10.1016/S0166-2236(98)01260-0. PMID 9610880.
  26. ^ Iacoboni, Marco; Woods, Roger P.; Brass, Marcel; Bekkering, Harold; Mazziotta, John C.; Rizzolatti, Giacomo (1999). "Cortical Mechanisms of Human Imitation". Science. 286 (5449): 2526–2528. doi:10.1126/science.286.5449.2526. PMID 10617472.
  27. ^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 15351628, please use {{cite journal}} with |pmid=15351628 instead.
  28. ^ a b Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 20957578, please use {{cite journal}} with |pmid=20957578 instead.
  29. ^ a b Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 17660183, please use {{cite journal}} with |pmid=17660183 instead.
  30. ^ a b Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 16527749, please use {{cite journal}} with |pmid=16527749 instead.
  31. ^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 18979387, please use {{cite journal}} with |pmid=18979387 instead.
  32. ^ Keysers, Christian; Gazzola, Valeria (2010). "Social Neuroscience: Mirror Neurons recorded in Humans" (PDF). Current Biology. 20 (8): R353–354. doi:10.1016/j.cub.2010.03.013. PMID 21749952.
  33. ^ a b Pascolo PB, Ragogna R, Rossi R, (2009). "The Mirror-Neuron System Paradigm and its consistency". Gait Posture. 30 (Suppl. 1): 65. doi:10.1016/j.gaitpost.2009.07.064.{{cite journal}}: CS1 maint: extra punctuation (link) CS1 maint: multiple names: authors list (link)
  34. ^ a b Hickok G (2009). "Eight problems for the mirror neuron theory of action understanding in monkeys and humans". J Cogn Neurosci. 21 (7): 1229–1243. doi:10.1162/jocn.2009.21189. PMC 2773693. PMID 19199415.
  35. ^ Hickok,G.,Eight Problems for the Mirror Neuron Theory of Action Understanding in Monkeys and Humans, Journal of Cognitive Neuroscience,2009,July,21(7),1229-1243[4]
  36. ^ Churchland,Patricia,Braintrust (2011), Chapter 6, page142
  37. ^ Heyes,Cecilia, Where do mirror neurons come from?, Neuroscience and Biobehavioral Reviews (2009)Article In Press[5]
  38. ^ Terje Falck-Ytter, Gustaf Gredebäck & Claes von Hofsten (2006), Infants predict other people's action goals[6], Nature Neuroscience 9 (2006)
  39. ^ Keysers & Perrett, Trends in Cognitive Sciences 8 (2004)
  40. ^ Heyes C. M. (2001). "Causes and consequences of imitation". Trends in Cognitive Sciences. 5 (6): 253–261. doi:10.1016/S1364-6613(00)01661-2. PMID 11390296.
  41. ^ Brass, M., & Heyes, C. Trends in Cognitive Sciences 9 (2005)
  42. ^ Heyes C. M. (2010). "Where do mirror neurons come from?". Neuroscience and Biobehavioural Reviews. 34: 575–583.
  43. ^ Anisfeld M (1996). "Only tongue protruding modeling is matched by neonates". Developmental Review. 16 (2): 149–161.
  44. ^ a b c Fogassi Leonardo, Pier , Ferrari Francesco, Gesierich Benno, Rozzi Stefano, Chersi Fabian, Rizzolatti Giacomo (2005). "Parietal lobe: from action organization to intention understanding". Science. 308 (5722): 662–667. doi:10.1126/science.1106138. PMID 15860620.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  45. ^ Preston S. D., de Waal F.B.M. (2002). "Empathy: Its ultimate and proximate bases". Behavioral and Brain Sciences. 25: 1–72.
  46. ^ Decety, J. (2002). Naturaliser l’empathie [Empathy naturalized]. L’Encéphale, 28, 9-20.
  47. ^ Decety J., Jackson P.L. (2004). "The functional architecture of human empathy". Behavioral and Cognitive Neuroscience Reviews. 3: 71–100. PMID 15537986.
  48. ^ Gallese V., Goldman A.I. (1998). "Mirror neurons and the simulation theory". Trends in Cognitive Sciences. 2 (12): 493–501. doi:10.1016/S1364-6613(98)01262-5. PMID 21227300.
  49. ^ Gallese V (2001). "The "Shared Manifold" hypothesis: from mirror neurons to empathy". Journal of Consciousness Studies. 8: 33–50.
  50. ^ Botvinick M., Jha A.P., Bylsma L.M., Fabian S.A., Solomon P.E., Prkachin K.M. (2005). "Viewing facial expressions of pain engages cortical areas involved in the direct experience of pain". NeuroImage. 25 (1): 312–319. doi:10.1016/j.neuroimage.2004.11.043. PMID 15734365.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  51. ^ Cheng Y., Yang C.Y., Lin C.P., Lee P.R., Decety J. (2008). "The perception of pain in others suppresses somatosensory oscillations: a magnetoencephalography study". NeuroImage. 40 (4): 1833–1840. doi:10.1016/j.neuroimage.2008.01.064. PMID 18353686.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  52. ^ Morrison, I.; Lloyd, D.; Di Pellegrino, G.; Roberts, N. (2004). "Vicarious responses to pain in anterior cingulate cortex: Is empathy a multisensory issue?". Cognitive, Affective, & Behavioral Neuroscience. 4 (2): 270. doi:10.3758/CABN.4.2.270. PMID 15460933.
  53. ^ Wicker et al., Neuron, 2003 [7]
  54. ^ Singer et al., Science, 2004 [8]
  55. ^ a b Jabbi, Mbemba; Swart, Marte; Keysers, Christian (2007). "Empathy for positive and negative emotions in the gustatory cortex". NeuroImage. 34 (4): 1744–53. doi:10.1016/j.neuroimage.2006.10.032. PMID 17175173.
  56. ^ Lamm C., Batson C.D., Decety J. (2007). "The neural substrate of human empathy: effects of perspective-taking and cognitive appraisal". Journal of Cognitive Neuroscience. 19 (1): 42–58. doi:10.1162/jocn.2007.19.1.42. PMID 17214562.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  57. ^ Gazzola, Aziz-Zadeh and Keysers, Current Biology, 2006 [9]
  58. ^ Oberman, L.; Ramachandran, V.S. (2009). "Reflections on the Mirror Neuron System: Their Evolutionary Functions Beyond Motor Representation". In Pineda, J.A. (ed.). Mirror Neuron Systems: The Role of Mirroring Processes in Social Cognition. Humana Press. pp. 39–62. ISBN 978-1-934115-34-3.
  59. ^ Ramachandran, V.S. (January 1, 2009). "Self Awareness: The Last Frontier, Edge Foundation web essay". Retrieved July 26, 2011.
  60. ^ Skoyles, John R., Gesture, Language Origins, and Right Handedness, Psycholoqy: 11,#24, 2000
  61. ^ Petrides, Michael, Cadoret, Genevieve, Mackey, Scott (2005). Orofacial somatomotor responses in the macaque monkey homologue of Broca's area, Nature: 435,#1235
  62. ^ Porter RJ. Lubker JF. (1980). Rapid reproduction of vowel-vowel sequences, Evidence for a fast and direct acoustic-motoric linkage in speech. Journal of Speech and Hearing Research, 23, 593-602. PMID 7421161
  63. ^ McCarthy R. Warrington EK. (1984). A two-route model of speech production, Evidence from aphasia. Brain, 107, 463-485. PMID 6722512
  64. ^ McCarthy RA, Warrington EK. (2001). Repeating without semantics: surface dysphasia? Neurocase.;7:77-87. PMID 11239078
  65. ^ Marslen-Wilson W. (1973). Linguistic structure and speech shadowing at very short latencies. Nature, 244, 522-523. PMID 4621131
  66. ^ Fay WH. Coleman RO. (1977). A human sound transducer/reproducer: temporal capabilities of a profoundly echolalic child. Brain and Language, 4, 396-402. PMID 907878
  67. ^ Schippers, MB; Roebroeck, A; Renken, R; Nanetti, L; Keysers, C (2010). "Mapping the Information flow from one brain to another during gestural communication" (PDF). Proc Natl Acad Sci U S A. 107 (20): 9388–93. doi:10.1073/pnas.1001791107. PMC 2889063. PMID 20439736.
  68. ^ Moro, Andrea (2008). The Boundaries of Babel. The Brain and the Enigma of Impossible Languages. MIT Press. p. 257. ISBN 978-0-262-13498-9.. {{cite book}}: Check |isbn= value: invalid character (help)
  69. ^ Longo, M. R (2008). "Automatic imitation of biomechanically possible and impossible actions: effects of priming movements versus goals". J Exp Psychol Hum Percept Perform. 34 (2): 489–501. doi:10.1037/0096-1523.34.2.489. PMID 18377184. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  70. ^ Chartrand, T. L (1999). "The chameleon effect: the perception-behavior link and social interaction". J Pers Soc Psychol. 76 (6): 893–910. PMID 10402679. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  71. ^ Lakin, J. L (2003). "Using nonconscious behavioral mimicry to create affiliation and rapport". Psychological Science. 14 (4): 334–9. PMID 12807406. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  72. ^ van Baaren, R. B (2003). "It takes two to mimic: behavioral consequences of self-construals". Journal of Personality and Social Psychology. 84 (5): 1093–102. PMID 12757151. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  73. ^ Heyes, Cecilia (2011). "Automatic imitation". Psychological Bulletin. 137 (3): 463–83. doi:10.1037/a0022288. PMID 21280938.
  74. ^ Paukner A, Suomi SJ, Visalberghi E, Ferrari PF (2009). "Capuchin monkeys display affiliation toward humans who imitate them". Science. 325 (5942): 880–3. doi:10.1126/science.1176269. PMC 2764469. PMID 19679816. {{cite journal}}: Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  75. ^ Dapretto M; Davies MS; Pfeifer JH; et al. (2006). "Understanding emotions in others: mirror neuron dysfunction in children with autism spectrum disorders". Nat. Neurosci. 9 (1): 28–30. doi:10.1038/nn1611. PMID 16327784. {{cite journal}}: Unknown parameter |author-separator= ignored (help); Unknown parameter |month= ignored (help)
  76. ^ Perkins T, Stokes M, McGillivray J, Bittar R. (2010). "Mirror neuron dysfunction in autism spectrum disorders". J. Clin. Neurosci. 17 (10): 1239–1243. doi:10.1016/j.jocn.2010.01.026. PMID 20598548.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  77. ^ Lindsay M. Oberman, Vilayanur S. Ramachandran, Jaime A. Pineda (2008). "Modulation of mu suppression in children with autism spectrum disorders in response to familiar or unfamiliar stimuli: The mirror neuron hypothesis" (PDF). Neuropsychologia. 46 (5): 1558–1565. doi:10.1016/j.neuropsychologia.2008.01.010. PMID 18304590.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  78. ^ Churcland, P.S. (2011). "6". Braintrust. Princeton University Press. p. 156.
  79. ^ Hadjikhani; Joseph, RM; Snyder, J; Tager-Flusberg, H; et al. (2006). "Anatomical Differences in the Mirror Neuron System and Social Cognition Network in Autism". Cerebral Cortex. 16 (9): 1276–82. doi:10.1093/cercor/bhj069. PMID 16306324. {{cite journal}}: Explicit use of et al. in: |author= (help)
  80. ^ http://www.newscientist.com/article/dn18837-mirror-neurons-seen-behaving-normally-in-autism.html
  81. ^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 19508497, please use {{cite journal}} with |pmid=19508497 instead.
  82. ^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 19666038, please use {{cite journal}} with |pmid=19666038 instead.
  83. ^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 20976151, please use {{cite journal}} with |pmid=20976151 instead.
  84. ^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 19632248, please use {{cite journal}} with |pmid=19632248 instead.
  85. ^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 20524939, please use {{cite journal}} with |pmid=20524939 instead.
  86. ^ Gordon R (1986). "Folk psychology as simulation". Mind and Language. 1: 158–171.
  87. ^ Goldman A (1989). "Interpretation psychologized". Mind and Language. 4: 161–185.
  88. ^ Gallese V., Goldman A. (1998). "Mirror neurons and the simulation theory of mindreading". Trends in Cognitive Sciences. 2 (12): 493–501. doi:10.1016/S1364-6613(98)01262-5. PMID 21227300.
  89. ^ Meltzoff A.N., Decety J. (2003). "What imitation tells us about social cognition: A rapprochement between developmental psychology and cognitive neuroscience". The Philosophical Transactions of the Royal Society, London. 358: 491–500.
  90. ^ Sommerville J. A., Decety J. (2006). "Weaving the fabric of social interaction: Articulating developmental psychology and cognitive neuroscience in the domain of motor cognition". Psychonomic Bulletin & Review. 13: 179–200.
  91. ^ Keysers C and Gazzola V(2007) Integrating simulation and theory of mind: from self to social cognition. Trends in Cognitive Sciences 11(5):194-6 [10]
  92. ^ Cheng Y., Tzeng O.J., Decety J., Hsieh J.C. (2006). "Gender differences in the human mirror system: a magnetoencephalography study". NeuroReport. 17 (11): 1115–1119. doi:10.1097/01.wnr.0000223393.59328.21. PMID 16837838.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  93. ^ Cheng Y., Decety J., Hsieh J.C., Hung D., Tzeng O.J. (2007). "Gender differences in spinal excitability during observation of bipedal locomotion". NeuroReport. 18 (9): 887–890. doi:10.1097/WNR.0b013e3280ebb486. PMID 17515795.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  94. ^ Cheng, Y., Decety, J., Yang, C.Y., Lee, S., & Chen, G. (2008). Gender differences in the Mu rhythm during empathy for pain: An electroencephalographic study. Brain Research, in press.
  95. ^ Cheng Y., Lee P., Yang C.Y., Lin C.P., Decety J. (2008). "Gender differences in the mu rhythm of the human mirror-neuron system". PLoS ONE. 3 (5): e2113. doi:10.1371/journal.pone.0002113. PMC 2361218. PMID 18461176.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)

Further reading

Retrieved from "https://en.wikipedia.org/w/index.php?title=Mirror_neuron&oldid=470678528"