The simulating social mind: The role of the mirror neuron system and simulation in the social and communicative deficits of autism spectrum disorders

“In a series of studies conducted by Baron-Cohen, Leslie, and Frith (1985, 1986), typically developing children, children with Down's syndrome, and children with autism were given the " Sally-Anne " test.”

“Indeed, a role of mirror system dysfunction has recently been hypothesized for autism spectrum disorder.(32,33) Autism spectrum disorder is most likely a polygenetic disorder that is expressed as impairment of gray matter architecture and of corticocortical intrahemispheric connections.”

“Interestingly, these regions include the mirror neuron system, which has been hypothesized by some to be critical in autism (Dapretto et al., 2006; Hadjikhani et al., 2006; Oberman et al., 2005; Oberman and Ramachandran, 2007; Williams et al., 2001, 2006).”

“…the concept of simulation has played an important role in developmental research concerning theory of mind and related work on “mentalizing,” or the processes involved in reading the mental states of other individuals (for reviews, see Frith & Frith 2006; Goldman 2006; Oberman & Ramachandran 2007).”

“Just as simulations of the past and future provide a functional benefit, so may daydreams (e.g., keeping the mind occupied at times of boredom; Antrobus & Singer, 1964) and simulations of the minds of others (e.g., social communication; Oberman & Ramachandran, 2007).”

“The lack of empathic engagement displayed by autistic children seems to depend on defective embodied simulation, likely underpinned by malfunctioning and/or altered regulation of the MNS (Gallese, 2003b, 2006; see also Oberman & Ramachandran, 2007).”

“In general, the MNS is assumed to form an important mechanism of shared representation (Decety & Sommerville, 2003; Oberman & Ramachandran, 2007).”

“Certain developmental disorders, such as autism spectrum disorders and conduct disorder, involve deficiencies in empathic responding, albeit for different reasons (Blair, 2005; Decety & Meyer, 2008; Oberman & Ramachandran, 2007; Preston & de Waal, 2002).”

“…of the brain basis of ASD have taken a systemslevel approach, and proposed that ASD may be explained by abnormalities in the mirror neuron system (Oberman and Ramachandran, 2007; Williams et al., 2001), the default-mode network (Kennedy and Courchesne, 2008; Kennedy et al., 2006), or both…”

“…of system may play an integral role in our ability to represent other's actions, as well as potentially the intentions and emotions that guide actions by creating a mechanism that allows observers to simulate the perception within their own motor and limbic systems (Oberman & Ramachandran, 2007).”

“The degree to which such areas may be specifically impaired in autism compared to other disorders may ultimately provide important clues to the nature of language deficits in autism (cf., Oberman & Ramachandran, 2007).”

“…bearing on this broader issue is lacking, although numerous findings of MNS dysfunction in autism (for reviews, see Iacoboni and Dapretto, 2006, and Oberman and Ramachandran, 2007) are consistent with the hypothesis of MNS involvement in social cognition, given that social deficits are a hallmark…”

“Since mirror neurons were initially described, researchers have proposed that the mirrorneuron system is an important component of the social-cognitive network, and there is evidence for impaired functioning of this system in autism (Oberman & Ramachandran, 2007).”

“…social-cognitive and emotional skills 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…”

“…of our own bodily states and the “simulation” and apprehension of such states in others (see Bar-On, Tranel, Denburg, & Bechara, 2003; Decety, 2011; Jabbi et al., 2007; Keysers & Gazzola, 2006; Keysers, Kaas, & Gazzola, 2010; Lamm & Singer, 2010; Oberman & Ramachandran, 2007; Singer et al., 2009).”

“…than mirror neurons, contributes to the autistic1 phenotype, Iacoboni stood by the more specific “broken-mirror theory” (Iacoboni & Dapretto, 2006; Oberman & Ramachandran, 2007; Williams et al., 2001; but see Southgate & Hamilton, 2008), citing a range of brain-imaging studies suggesting…”

“…behavior as evidenced, for example, in the attunement of the facial expressions, the body posture, and the gestures between social interactants (Oberman & Ramachandran, 2007), we hypothesized that another person’s direct and averted gaze would initiate in the observer respectively,…”

“Hence, it has been postulated that impaired simulation processes in shared neural circuits are at the root of social deficits in ASD, such as impaired affective and motor empathy [Gallese, 2003, 2006; Minio-Paluello et al., 2009; Oberman and Ramachandran, 2007].”

“…results show that children with ASD display atypical activation in regions assigned to the mirror neuron system, including the IFG and IPL, when observing facial expressions and hand movements (Dapretto et al., 2006; Martineau et al., 2010; Oberman and Ramachandran, 2007; Williams et al., 2006).”

“The social deficits characterizing autism might be due to a dysfunctional mirror neuron system [190,191].”

“…could stem from the incapacity to establish a motor equivalence between demonstrator and imitator, most likely due to a malfunctioning of the mirror neuron system, and/or because of disrupted emotional– affective regulation of the same system (Gallese, 2003b, 2006; Oberman & Ramachandran, 2007).”

“For example, perspective taking abilities seem to be often impaired in individuals suffering from Autism Spectrum Disorder (ASD; Oberman and Ramachandran, 2007; but cf. Hamilton et al., 2007), while there is also evidence for impaired predictive coding mechanisms in ASD (Friston, 2012a).”

“Many theories have suggested a possible role for mirror neurons in mechanisms of action understanding, imitation, empathy, theory of mind, and language [5–7].”

“Abnormal findings in these areas in subjects with autism have been related to several hypothesis such as the reduced functional output of the mirror neurons [Oberman and Ramachandran, 2007] and the aberrant connectivity theory [Courchesne et al.”

“Moreover, while deficiencies in the social domain are specifically implicated in autism (American Psychiatric Association, 2000; Baron-Cohen, 2001; Frith, 2001, 2003; Klin et al., 2002; Lord et al., 2000a; Oberman and Ramachandran, 2007), the degree to which our data fit with existing theories about ASD remains an int…”

“This research tool is currently being used to examine the potential overconnectivity of cortical neurons in autism [110] and to examine neurologic function in other disorders such as ADHD [111].”

“the recognition of faces and their emotion-conveying expressions [8,60,112,232] as well as difficulty imitating others [185,227]”

“…intentions, and experiences of others, thereby allowing individuals to " use the same systems that process knowledge about selfperformed actions, self-conceived thoughts, and self-experienced emotions to understand actions, thoughts, and emotions in others " (Oberman and Ramachandran, 2007, p.310).”

“Deficits in these domains are central to autism spectrum disorders (Kanner, 1943; Oberman & Ramachandran, 2007).”

“Cerebral Cortex May 2012;22:1025--1037 doi:10.1093/cercor/bhr171 Advance Access publication July 22, 2011”

“Deficits in imitation and mimicry have also been reported in participants with autistic spectrum disorder (Williams et al., 2004), and some have linked this to the functioning of the MNS (Williams et al., 2001; Oberman et al., 2005; Oberman and Ramachandran, 2007).”

“The ‘‘embodied simulation’’ theory (Gallese, 2006; Oberman & Ramachandran, 2007) posits that emotion processing is mediated by an ‘‘internal simulation’’ process, based on a ‘‘direct mapping’’ between observed and experienced emotions.”

“…in MN mechanism functioning could be critical in autism (Dapretto et al., 2006; Gallese, 2003, 2006a; Hadjikhani et al., 2006; Oberman et al., 2005; Oberman & Ramachandran, 2007; Oberman, Ramachandran, & Pineda, 2008; Théoret et al., 2005; Williams et al., 2006; Williams, Whiten, Suddendorf, &…”

“Several functional [Dapretto et al., 2006; Oberman et al., 2005] and anatomical [Hadjikhani et al., 2009] abnormalities have been reported in the IFG in autism by previous studies.”

“…have proposed that an atypically functioning mirror neuron system may be the underlying mechanism (Rogers and Pennington, 1991; Charman et al., 1997; Williams et al., 2001; Rogers et al., 2003; Gallese, 2006; Oberman and Ramachandran, 2007; Colombi et al., 2009; Rizzolatti and Fabbri-Destro, 2010).”

“One prominent hypothesis is that these are caused by a dysfunctional mirror neuron system (Oberman and Ramachandran, 2007).”

“populations like patients with autism [60] or disruptive behavior”

“The results suggest that individuals with ASD do not have a fundamental impairment in catching others’ yawns, such as the impairment to empathize with others [41].”

“An emerging view suggests that, although the MNS and MSAS may be functionally dissociable (Van Overwalle and Baetens, 2009), interactions between these systems and domain-general cognitive control mechanisms best characterize everyday social cognition (Zaki and Ochsner, 2009).”

“The mirror neuron hypothesis of autism has also received considerable attention [Iacoboni & Dapretto, 2006; Oberman & Ramachandran, 2007].”

“It is possible that the relationship between these three social-communicative behaviors represent earlier manifestations of a more global impairment (e.g., motor neurons) with impact in language, object play, motor skills, and empathy ( Oberman and Ramachandran 2007; Williams et al. 2001).”

“These mirror neurons, often called empathy neurons, suggest that the observer has the ability to acknowledge the current state and experience of the other individual (Oberman and Ramachandran 2007).”

“…to action execution become active when the person observes someone else making the same action; in other words, the observer’s brain resonates with the model’s motor behavior (Hogeveen & Obhi, 2011; Iacoboni, 2009; Oberman & Ramachandran, 2007; Obhi & Hogeveen, 2010; Rizzolatti & Sinigaglia, 2010).”

“, & Ochsner, K. (2009). Unpacking the informational bases of empathic accuracy.”

“Such impairments may well affect the ability to communicate and empathize with perspectives of other persons [7].”

“It is assumed that spontaneously imitating observed mental states allows for a direct first person access to that state, thus it can follow that impairments in imitation also affect empathy functions (Gallese & Goldman, 1998; Oberman & Ramachandran, 2007).”

“…autism is premised on the idea that the disruption of basic mirroring mechanisms in autism would have a cascading impact on more complex types of simulation, first goal understanding and later language and mentalising (Iacoboni & Dapretto, 2006; Oberman & Ramachandran, 2007; Williams et al., 2001).”

“Our Psycho Physiological Interaction (PPI) analysis has revealed a significant functional coupling of the MNS with the mentalizing system during imitative interaction (Sperduti et al., 2014).”

“…years, several groups have focused on the functionality of the mirror neuron system (MNS) in observers with ASD (e.g., Dapretto et al., 2006; Gallese, 2006; Iacoboni & Dapretto, 2006; Nishitani, Avikainen, & Hari, 2004; Oberman & Ramachandran, 2007; Williams, 2008; Williams, Whiten, & Singh, 2004).”

“Williams et al. (2001), and also Oberman and Ramachandran (2007), have taken this argument a step further, suggesting that both imitation and the ability to simulate or imagine the mental states of others are mediated by mirror neurons (Gallese and Goldman, 1998; Gallese et al., 2004), and that the…”

“The answer is most likely negative, because simulations are central to processes typically considered to be Type 1, such as perception, categorization, and motor movements (Barsalou, 1999, 2003; Oberman & Ramachandran, 2007).”

“It has been argued, for example, that the mirror neuron system(MNS) is not lateralizedbut that asymmetriesmayappearon the basis of task-dependent functions (Miller and Cummings, 2007).”

“Abnormalities with this system have been suggested to underlie the problems with social cognition observed in ASC [9–12], although there is still debate about whether this is the case [13].”

“It has important implications for the development of such complex and diverse social competences as imitation, language and empathy (Oberman & Ramachandran, 2007).”

“Recently, it has been proposed that mirror neuron dysfunction may underlie some of the symptoms characteristic of ASD, including deficits in social cognition [17– 19].”

“Thus, the shared representations of mouth movements and visual and auditory perceptions within the mirror neuron system may help to strengthen the associations between objects and their names [81].”

“The dominant explanation for action comprehension difficulties in autism is the broken mirror hypothesis, which claims that dysfunction of neural systems for mirroring is a primary cause of poor social skills in autism (Gallese et al., 2009; Iacoboni and Dapretto, 2006; Oberman and Ramachandran, 2007; Rizzolatti and F…”

“Oberman and Ramachandran (2007) note that MNs play a key role in the neural circuitry underlying the simulation of social understanding ; they called this the simulation account of empathy.”

“Due to the fact that individuals with ASD demonstrate impairments in all the aforementioned domains, it is suggested that the system of MN is dysfunctional in ASD [31,32].”