Grasping the Intentions of Others with One's Own Mirror Neuron System

“questionnaire adapted from the Edinburgh Handedness Inventory [ 36 ].”

“The co-planar highresolution structural T2-weighted EPI volume was subsequently registered to the Montreal Neurological Institute Talairach-compatible MR atlas averaging 152 normal subjects using FMRIB’s Linear Image Registration Tool (FLIRT) [37].”

“Functional images were first registered to the co-planar high-resolution structural T2weighted EPI volume after non-brain structures had been removed with FMRIB’s Brain Extraction Tool (BET) from the co-planar highresolution T2-weighted EPI volume [38].”

“In the monkey brain, ventral premotor area F5 contains, in addition to mirror neurons, a population of cells called canonical neurons [4].”

“Mirror neurons are premotor neurons that fire when the monkey performs object-directed actions such as grasping, tearing, manipulating, holding, but also when the animal observes somebody else, either a conspecific or a human experimenter, performing the same class of actions [3,4,5].”

“Neurons of this type have indeed been previously reported in F5 and referred to as ‘‘logically related’’ neurons [34].”

“Z image statistics were performed with a threshold of Z = 2.3 at voxel level and a cluster level corrected for the whole brain at p , 0.05 [41, 42 ].”

“Time-series statistical analyses were carried out using FMRIB’s Improved Linear model (FILM) with local autocorrelation correction [39].”

“In the first, contextual features were used to clarify the intention behind a hand-object interaction.(28) In the process of inferring intentions, the frontal node of the mirror system in the right hemisphere was recruited.”

“Evidence that the parieto-frontal mirror circuit in humans is also involved in intention encoding was first provided by an fMRI experiment consisting of three condition”

“A recent fMRI study demonstrated that human mirror neuron areas respond differently to the observation of the same grasping actions, if the actions are embedded in different contexts that suggest different intentions associated with the grasping actions, such as drinking or cleaning u”

“Other studies, however, reported a dominance of left hemispheric areas during imitation tasks, which was interpreted in reference to the lateralisation of language functions (e.g., Aziz-Zadeh et al., 2004; Rizzolatti and Arbib, 1998; Iacoboni et al., 2005).”

“The mirror system goes a long way towards explaining action from directly observable sensory input in terms of immediate and task goals (e.g., Iacoboni et al., 2005).”

“In humans, indirect measures of neural activity support the existence of sensory-motor mirroring mechanisms in homolog frontal and parietal areas [10, 11], other motor regions [12–15], and also the existence of multisensory mirroring mechanisms in nonmotor regions [16–19].”

“…been J. M. Kilner (&) Á K. J. Friston Á C. D. Frith The Wellcome Trust Centre for Neuroimaging, Institute of Neurology, 12 Queen Square, WC1N 3BG London, UK e-mail: j.kilner@fil.ion.ucl.ac.uk demonstrated in humans, both directly (Rushworth et al. 2006) and indirectly (Iacoboni et al. 2001, 2005).”

“…in part because of their purported role in a diverse range of cognitive functions, from imitation and action understanding to social cognition (Iacoboni, 2005, 2009; Iacoboni et al., 2005; Fogassi et al., 2005; Keysers et al., 2010; Rizzolatti & Fabbri-Destro, 2008 ; Rizzolatti & Sinigaglia, 2010).”

“This property, together with the fact that mirror neurons have been found in cortical areas involved in motor control (Keysers and Fadiga, 2008), has led to the proposal that others’ actions can be translated into a motor code exploiting one’s own action knowledge, in terms of an individual’s cortical motor representa…”

“Engagement of the human mirror neuron system occurs even when observers are not directed to attend to a target (Iacoboni et al., 2005) or are put under cognitive load through a concurrent secondary task (Fadiga, Fogassi, Pavesi, & Rizzolatti, 1995; Spunt & Lieberman, 2013).”

“These issues were experimentally addressed with a functional magnetic resonance imaging (fMRI) study (Iacoboni et al. 2005).”

“The direct comparison of premotor representations of action concepts derived through observation and language is important because it suggests that language makes use of the same embodied representations that are thought to be involved in social cognition of the actions and intentions of others [26, 27].”

“Finally in cases where the goal of the actions suggests a series of actions to follow, even if these actions are not seen, occlusion mirror neurons and sequence selective neurons could add to activations (Fogassi et al., 2005; Iacoboni et al., 2005; Umilta et al., 2001).”

“Three studies for instance have looked at where the brain might process why individuals perform certain actions, but have identified 3 different brain regions: the IFG (Iacoboni et al. 2005), the temporoparietal junction (Brass et al. 2007) and midline structures (de Lange et al. 2008).”

“The activity of the right frontal cortex was of different strength for observation of these similar hand actions depending on the context (114).”

“…there is neuroscientific evidence that shows that mirror neuron activation does not necessarily involve a precise match between motor system execution and observed action, but may be involved in ‘‘logically related” actions or in anticipating future action (Csibra, 2005; Iacoboni et al., 2005).”

“, 2003), supramodal sequence predictor (Maess et al., 2001; Kilner et al., 2004; Iacoboni et al., 2005), and internal simulator (Platel et al.”

“Mirror neurons provide an “action-recognition” mechanism: the actor's actions are reproduced in the premotor cortex of the observer (Iacoboni, Molnar-Szakacs, Gallese, Buccino, Mazziotta, & Rizzolatti, 2005 ; Rizzolatti, Fadiga, Fogassi, & Gallese, 1996 ).”

“In particular, mirror neurons have been associated with the comprehension of action goals [e.g. Fogassi et al., 2005; Iacoboni et al., 2005] and imitation [e.g. Iacoboni et al., 1999; Rizzolatti et al., 2001], both of which are thought to be an important stage in the subsequent development of ToM.”

“In a recently published study, Iacoboni et al. (2005) showed participants videos of four different types of actions.”

“Rizzolatti, G. & Fadiga, L. (1998) Grasping objects and grasping action meanings:”

“In fact, two recent studies in macaques [7 ] and humans [27 ] have demonstrated that mirror neurons can code the likely future actions of others so that observers are able to anticipate the intentions of others.”

“The mirror system, found in premotor and parietal cortices of human and monkey brains, is thought to provide the foundation for social understanding, enabling the development of theory of mind and language[5-9].”

“These assumptions are in line with behavioral and neuroimaging evidence for the involvement of mirror neuron mechanisms in social interactions, for example, the detection of emotional states (Carr et al., 2003), intentions (Iacoboni et al., 2005), and the emergence of empathy (Gallese, 2003).”

“Recent studies indeed indicate that not only motor plans but also more abstract intentions (Iacoboni et al., 2005) or emotional states (Carr, Iacoboni, Dubeau, Mazziotta, & Lenzi, 2003) recruit the same areas in the inferior frontal cortex which are supposed to contain mirror neurons.”

“This idea is supported by the discovery of mirror neurons in monkeys [76,77] and of a similar system in humans [78].”

“Evidence for human mirror systems (Craighero et al. 2002; Decety & Chaminade 2005; Decety et al. 1997; Fadiga et al. 2002; Hari et al. 1998; Iacoboni et al. 2005; Rizzolatti et al. 1996; Ruby & Decety 2001) includes brain imaging and demonstrations that observing another person move primes the muscles needed to move i…”

“…Iacoboni, Dubeau, Woods, & Mazziotta, 2003; MolnarSzakacs, Iacoboni, Koski, & Mazziotta, 2005), intention understanding (Gallese & Goldman, 1998; Iacoboni et al., 2005), empathy, and theory of mind (Carr, Iacoboni, Dubeau, Mazziotta, & Lenzi, 2003; Leslie, Johnson-Frey, & Grafton, 2004;…”

“The MNS in humans is directly involved in imitation of simple movements (Iacoboni et al., 1999), imitation learning of complex skills (Buccino et al., 2004a), in the perception of communicative actions (Buccino et al., 2004b), and in the detection of action intentions ( Iacoboni et al., 2005 ).”

“This indicates that mirror areas, in addition to action understanding, also mediate the understanding of others’ intention (Iacoboni et al. 2005).”

“In a recently published functional Magnetic Resonance Imaging (fMRI) study (Iacoboni et al. 2005), these issues were experimentally addressed.”

“In a recently published fMRI study (Iacoboni et al., 2005), we tried to experimentally address these issues.”

“The proposed frontal mirror neuron region has also been shown to be involved in understanding the intentions behind the actions of others (Iacoboni et al., 2005).”

“Neurons in this area of the macaque brain respond when an action is inferred to take place behind a screen (Umilta et al., 2001), and some studies have linked this region to intentionality (Iacoboni et al., 2005).”

“2004), and a more inferior and anterior locus in the pSTS that has been associated with perception of biological motion, action understanding, and changeable aspects of faces such as expression and gaze direction (Iacoboni et al., 2005; Saygin, Wilson, Hagler, Bates, & Sereno, 2004; Beauchamp, Lee, Haxby, & Martin, 20…”

“This result overlaps closely with a study by Iacoboni et al. (2005) showing activation of these same sites when participants were evaluating action videos for intentionality.”

“imitation [50], empathy [21], perspective taking [35], mindreading [ 46 ,48,51,52] and language production and comprehension [53,54]), as suggested by ESC proponents, is still being debated at the empirical level.”

“…some areas of the brain are known to be associated with the perception (but not the execution) of biological motion including the posterior superior temporal sulcus or STS (Allison, Puce, & McCarthy, 2000; Grèzes, Armony, Rowe, & Passingham, 2003; Grèzes et al., 2001; Iacoboni et al., 2005).”

“Recent fMRI and TMS research indicates that areas assumed to constitute the human mirror system (inferior frontal gyrus and inferior parietal cortex) are involved in tasks requiring action understanding [6, 14].”

“…properties, it has been proposed that the mirror neuron system subserves a wide variety of behaviors including social communication and motor learning (Jeannerod, 1994; Gallese and Goldman, 1998; Rizzolatti et al., 2001; Rizzolatti and Craighero, 2004; Fogassi et al., 2005; Iacoboni et al., 2005).”

“46 Evidence now suggests that this bodily resonance (see Glossary) can afford us a unique, first-person 47 understanding of the experiences of others and is central to a number of social processes [7] 48 including intention understanding [19], empathy [20], and emotion recognition [21].”

“Contents lists available at ScienceDirect Journal of Experimental Social Psychology journal homepage: www.elsevier .com/locate / jesp understanding (e.g. Iacoboni et al., 2005; Oberman, Pineda, & Ramachandran, 2006).”

“…e : w w w. e l s e v i e r. c o m / l o c a t e / y n i m g Further studies have shown that in addition to the premotor cortex, brain networks focused around the temporo-parietal junction (Jackson et al., 2006; Lyons et al., 2006) and inferior frontal cortex (Iacoboni et al., 2005 Iacoboni et al."”

“, 2004), and intention understanding (Iacoboni et al., 2005).”

“In an fMRI study [67], participants were presented with the same action embedded in two different contexts.”

“These results suggest a distinct neural network for empathy and emotional responsiveness informed by recent increases in our understanding of “mirror” mechanisms in humans that are implicated in the ability to take another’s emotional perspective (Iacoboni et al., 2005; Kohler, 2002; Miller, 2005).”

“Some years ago, an fMRI experiment showed that when individuals had to infer the intention of the agent from the context, there was a selective activation of the right frontal node of the mirror-system [21].”

“Additionally, previous fMRI studies had indicated that the human MNS was sensitive to intentions and goals, with the IFG modulating its degree of activity based on the intention of the observed action (Iacoboni et al., 2005).”

“Previous studies have shown the involvement of the MNS in action understanding [17, 18].”

“…in imitation of simple movements (Iacoboni et al., 1999), in imitative learning of novel and complex motor skills (Buccino et al., 2004b), in perception of facial communicative actions (Buccino et al., 2004a), and in understanding the intentions of the actions of others (Iacoboni et al., 2005).”

“Further, it was sensitive to the intentions behind the same motor action (Iacoboni et al., 2005).”

“05), replicating the effect observed in our previous study (Iacoboni et al., 2005).”

“The context and the diVerent grip shapes suggested the intention of the agent, i.e., grasping the cup for drinking or grasping it for cleaning the table (Iacoboni et al. 2005).”

“The MNS is located in the frontal brain and helps people understand the intentions of others by means of shared sensorimotor representations (Iacoboni et al., 2005).”

“The increased complexity entails at the same time an (emotional) intention (see Figure 2), which has recently been shown to increase the activation within inferior frontal cortex (Iacoboni et al., 2005).”

“…including theory of mind (Frith & Frith, 1999; Hamilton & Grafton, 2006; Iacoboni et al., 2005), empathy (Oberman & Ramachandran, 2007; Dapretto et al., 2005), interpersonal synchrony (Vicaria and Dickens, 2016), language (Gallese, 2008), and intentional understanding (Iacoboni et al., 2005).”

“[58].”

“Human imaging research using the shared representation logic (Figure 3A) later provided for converging evidence for the existence of a similar human “mirror system” (89,90), although individual mirror neurons have yet to be observed directly.”

“2006) and the passive movement observation protocol (Blakemore et al. 2005; Buccino et al. 2001; Buccino et al. 2004a; Haslinger et al. 2005; Iacoboni et al. 2005), neither of which assess the same criteria used in the monkey.”

“…the findings to humans (Fadiga et al., 1995; Grafton et al., 1996; Rizzolatti et al., 1996b; Grezes et al., 1998, 2003; Binkofski et al., 1999; Iacoboni et al., 1999, 2005; Nishitani and Hari, 2000; Buccino et al., 2001; Aziz- Zadeh et al., 2002; Johnson-Frey et al., 2003; Koski et al., 2003;…”

“(Modified from Iacoboni and colleagues.(37))”

“One approach to study this is to add contextual information that alters the meaning of an observed action (Iacoboni et al. 2005).”

“, 2005b) overlap quite well with the inferior parietal mirror neuron area identified in previous experiments in our lab (Iacoboni et al., 2005).”

“Viewers of human activity consistently activate motor representations corresponding to the actions they observe (Iacoboni et al., 2005).”

“, 2004), and the detection of action intentions (Iacoboni et al., 2005).”

“Imitation of an action that belongs to the motor repertoire of the observer includes activity in the neural circuitry of the superior temporal sulcus and the frontal and parietal mirror areas (Iacoboni, 2005).”

“…(e.g. Carr, Iacoboni, Dubeau, Mazziotta, & Lenzi, 2003; Miall, 2003), action understanding (e.g. Umilta et al., 2001), intention attribution (Iacoboni et al., 2005) and—on the finding of a mirror system for grasping in or near human Broca’s area—(evolution of) language (Rizzolatti & Arbib,…”

“In the light of studies regarding the MNS, and its involvement in the understanding of others’ intentions (Fogassi et al., 2005; Iacoboni et al., 2005), it can be speculated that observers tend to ‘‘resonate’’ more with an intentional agent, also when only assumed, than with a touched passive…”

“Four ROIs were generated, covering the left and right IFG and left and right IPL as defined in previous studies of the MNS (Iacoboni et al. 1999; Johnson-Frey et al. 2003, 2005; Iacoboni et al. 2005).”