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| Some neurons responded to the sight of an object Rizzolatti, G., Gentilucci, M., Camarda, R. M., Gallese, V., Luppino, G., Matelli, M., & Fogassi, L. (1990). Neurons related to reaching-grasping arm movements in the rostral part of area 6 (area 6aβ). Experimental Brain Research, 82 (2), 337–350.
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| changed our fundamental understanding of how we came to be such social creatures Pellegrino, G. D., Fadiga, L., Fogassi, L., Gallese, V., & Rizzolatti, G. (1992). Understanding motor events: A neurophysiological study. Experimental Brain Research, 91 (1), 176–180.
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| Although some psychologists had argued for this kind of perceptual-motor Prinz, W. (1997). Perception and action planning. European Journal of Cognitive Psychology, 9 (2), 129–154.
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| that hitherto remained mysterious and inaccessible to experiments” Ramachandran, V. S. (2000). Mirror neurons and imitation learning as the driving force behind “the great leap forward” in human evolution. Edge website article: http://www.edge.org/3rd_culture/ramachandran/ramachandran_p1.html .
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| our capacity for language, culture, imitation, mindreading, and empathy Arbib, M. A. (2005). From monkey-like action recognition to human language: An evolutionary framework for neurolinguistics. Behavioral and Brain Sciences, 28 (02), 105–124; Molenberghs, P., Cunnington, R., & Mattingley, J. B. (2009). Is the mirror neuron system involved in imitation? A short review and meta-analysis. Neuroscience & Biobehavioral Reviews, 33 (7), 975–980; Blakeslee, S. (2006). Cells that read minds. New York Times , January 10, p. 1; Fabrega Jr., H. (2005). Biological evolution of cognition and culture: Off Arbib’s mirror-neuron system stage? Behavioral and Brain Sciences, 28 (02), 131–132; Gallese, V. (2001). The shared manifold hypothesis. From mirror neurons to empathy. Journal of Consciousness Studies, 8 (5-7), 33–50.
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| allowing us to keep more abstract ideas in mind at the same time Coolidge, F. L., & Wynn, T. (2005). Working memory, its executive functions, and the emergence of modern thinking. Cambridge Archaeological Journal, 15 (1), 5–26.
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| force behind the ‘great leap forward’ in human evolution” Ramachandran, V. S. (2000). Mirror neurons and imitation learning as the driving force behind “the great leap forward” in human evolution. Edge website article: http://www.edge.org/3rd_culture/ramachandran/ramachandran_p1.html .
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| first evidence regarding the presence of a mirror neuron system in humans Iacoboni, M., Woods, R. P., Brass, M., Bekkering, H., Mazziotta, J. C., & Rizzolatti, G. (1999). Cortical mechanisms of human imitation. Science, 286 (5449), 2526–2528.
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| temporarily “frazzles” the neurons in that area such that the region is essentially taken offline Heiser, M., Iacoboni, M., Maeda, F., Marcus, J., & Mazziotta, J. C. (2003). The essential role of Broca’s area in imitation. European Journal of Neuroscience, 17 (5), 1123–1128.
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| imitating the fingering required to make a set of guitar chords that they were shown Buccino, G., Vogt, S., Ritzl, A., Fink, G. R., Zilles, K., Freund, H. J., & Rizzolatti, G. (2004). Neural circuits underlying imitation learning of hand actions: An event-related fMRI study. Neuron, 42 (2), 323–334.
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| Gordon’s account of the second route Ross, L., Greene, D., & House, P. (1977). The “false consensus effect”: An egocentric bias in social perception and attribution processes. Journal of Experimental Social Psychology, 13 (3), 279–301; Ames, D. R. (2004). Inside the mind reader’s tool kit: Projection and stereotyping in mental state inference. Journal of Personality and Social Psychology, 87 (3), 340.
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| mirror neurons are the neural implementation of Simulation theory Gallese, V., & Goldman, A. (1998). Mirror neurons and the simulation theory of mind-reading. Trends in Cognitive Sciences, 2 (12), 493–501.
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| “the fundamental mechanism that allows us a direct experiential grasp Gallese, V., Keysers, C., & Rizzolatti, G. (2004). A unifying view of the basis of social cognition. Trends in Cognitive Sciences, 8 (9), 396–403; Gordon actually foreshadowed this idea two decades earlier writing that an “interesting possibility for … practical simulation is a prepackaged ‘module’ called upon automatically in the perception of other human beings”: Gordon, R. M. (2007). Folk psychology as simulation. Mind & Language, 1 (2), 158–171.
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| My brain is mirroring your brain Rizzolatti, G., & Sinigaglia, C. (2010). The functional role of the parieto-frontal mirror circuit: Interpretations and misinterpretations. Nature Reviews Neuroscience, 11 (4), 264–274.
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| found mirror neurons that fit the bill Kohler, E., Keysers, C., Umilta, M. A., Fogassi, L., Gallese, V., & Rizzolatti, G. (2002). Hearing sounds, understanding actions: Action representation in mirror neurons. Science, 297 (5582), 846–848.
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| highlighted an important limitation of the sight-sound study Hickok, G. (2009). Eight problems for the mirror neuron theory of action understanding in monkeys and humans. Journal of Cognitive Neuroscience, 21 (7), 1229–1243.
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| that the mirror neurons respond to actions that involve objects that cannot be seen Umilta, M. A., Kohler, E., Gallese, V., Fogassi, L., Fadiga, L., Keysers, C., & Rizzolatti, G. (2001). I know what you are doing: A neurophysiological study. Neuron, 31 (1), 155–166.
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| Humans are certainly capable of seeing an object Lee, H., Simpson, G. V., Logothetis, N. K., & Rainer, G. (2005). Phase locking of single neuron activity to theta oscillations during working memory in monkey extrastriate visual cortex. Neuron, 45 (1), 147–156.
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| She suggests that the purpose of mirror neurons Heyes, C. (2010). Mesmerising mirror neurons. NeuroImage, 51 (2), 789–791.
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| Heyes designed a clever countermirroring procedure Catmur, C., Walsh, V., & Heyes, C. (2007). Sensorimotor learning configures the human mirror system. Current Biology, 17 (17), 1527–1531; Catmur, C., Mars, R. B., Rushworth, M. F., & Heyes, C. (2011). Making mirrors: Premotor cortex stimulation enhances mirror and counter-mirror motor facilitation. Journal of Cognitive Neuroscience, 23 (9), 2352–2362.
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| Another study examined how the mirror system responds Newman-Norlund, R. D., van Schie, H. T., van Zuijlen, A. M., & Bekkering, H. (2007). The mirror neuron system is more active during complementary compared with imitative action. Nature Neuroscience, 10 (7), 817–818.
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| If we look at brains at rest Fox, M. D., Snyder, A. Z., Vincent, J. L., Corbetta, M., Van Essen, D. C., & Raichle, M. E. (2005). The human brain is intrinsically organized into dynamic, anticorrelated functional networks. Proceedings of the National Academy of Sciences of the United States of America, 102 (27), 9673–9678.
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| If he answers, “In order to have a drink” Spunt, R. P., & Lieberman, M. D. (in press). Automaticity, control, and the social brain. In J. Sherman, B. Gawronski, & Y. Trope (Eds.). Dual Process Theories of the Social Mind. New York: Guilford.
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| systematically investigated these distinctions Vallacher, R. R., & Wegner, D. M. (1987). What do people think they’re doing? Action identification and human behavior. Psychological Review, 94 (1), 3.
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| are more likely to focus on the thoughts they are trying to convey Carver, C. S. (1979). A cybernetic model of self-attention processes. Journal of Personality and Social Psychology, 37 (8), 1251.
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| explain another person’s high-level reasons for wanting the light on Jacob, P., & Jeannerod, M. (2005). The motor theory of social cognition: A critique. Trends in Cognitive Sciences, 9 (1).
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| figuring out the how, what, and why of other people’s behavior Spunt, R. P., & Lieberman, M. D. (2012). Dissociating modality-specific and supramodal neural systems for action understanding. Journal of Neuroscience, 32 , 3575–3583; Spunt, R. P., & Lieberman, M. D. (2012). An integrative model of the neural systems supporting the comprehension of observed emotional behavior. NeuroImage, 59 , 3050–3059; Spunt, R. P., Falk, E. B., & Lieberman, M. D. (2010). Dissociable neural systems support retrieval of “how” and “why” action knowledge. Psychological Science, 21 , 1593–1598; Spunt, R. P., Satpute, A. B., & Lieberman, M. D. (2011). Identifying the what, why, and how of an observed action: An fMRI study of mentalizing and mechanizing during action observation. Journal of Cognitive Neuroscience, 23 , 63–74; Brass, M., Schmitt, R. M., Spengler, S., & Gergely, G. (2007). Investigating action understanding: Inferential processes versus action simulation. Current Biology, 17 (24), 2117–2121; de Lange, F. P., Spronk, M., Willems, R. M., Toni, I., & Bekkering, H. (2008). Complementary systems for understanding action intentions. Current Biology, 18 (6), 454–457; Noordzij, M. L., Newman-Norlund, S. E., De Ruiter, J. P., Hagoort, P., Levinson, S. C., & Toni, I. (2009). Brain mechanisms underlying human communication. Frontiers in Human Neuroscience, 3 , 14.
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| there was increased mirror system activity Spunt, R. P., & Lieberman, M. D. (2012). Dissociating modality-specific and supramodal neural systems for action understanding. Journal of Neuroscience, 32 , 3575–3583.
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| this was still the case when participants were distracted Spunt, R. P., & Lieberman, M. D. (2013). The busy social brain: Evidence for automaticity and control in the neural systems supporting social cognition and action understanding. Psychological Science, 24 , 80–86.
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| “blooming, buzzing confusion” James, W. (1890/1950). The Principles of Psychology. New York: Dover, p. 462.
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| The word empathy was introduced into the English language Titchener, E. B. (1909). Lectures on the Experimental Psychology of Thought-Processes. New York: Macmillan.
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| something like a first-person experience from the object’s perspective The modern use of the term to refer to appreciating another person’s experience can be traced to Husserl’s Ideen and his student Edith Stein’s dissertation published as Stein, E. (1989/1916). On the Problem of Empathy. Washington D.C.: ICS Publications.
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| There are at least three kinds of psychological processes Zaki, J., & Ochsner, K. (2012). The neuroscience of empathy: Progress, pitfalls and promise. Nature Neuroscience, 15 , 675-680.
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| individuals watched others receive shocks to their hands or feet Avenanti, A., Bueti, D., Galati, G., & Aglioti, S. M. (2005). Transcranial magnetic stimulation highlights the sensorimotor side of empathy for pain. Nature Neuroscience, 8 (7), 955–960.
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| muscles in our own faces immediately mimic the expression in subtle ways Dimberg, U., Thunberg, M., & Elmehed, K. (2000). Unconscious facial reactions to emotional facial expressions. Psychological Science, 11 (1), 86–89.
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| if a person is unable to mimic those facial expressions Neal, D. T., & Chartrand, T. L. (2011). Embodied emotion perception amplifying and dampening facial feedback modulates emotion perception accuracy. Social Psychological and Personality Science, 2 (6), 673–678.
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| Given that the mirror system is involved in understanding Wicker, B., Keysers, C., Plailly, J., Royet, J. P., Gallese, V., & Rizzolatti, G. (2003). Both of us disgusted in my insula: The common neural basis of seeing and feeling disgust. Neuron, 40 (3), 655–664; Carr, L., Iacoboni, M., Dubeau, M. C., Mazziotta, J. C., & Lenzi, G. L. (2003). Neural mechanisms of empathy in humans: A relay from neural systems for imitation to limbic areas. Proceedings of the National Academy of Sciences, 100 (9), 5497–5502.
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| trying to understand why someone is experiencing a particular emotion Spunt, R. P., & Lieberman, M. D. (2012). An integrative model of the neural systems supporting the comprehension of observed emotional behavior. NeuroImage, 59 , 3050–3059; Mar, R. A. (2011). The neural bases of social cognition and story comprehension. Annual Review of Psychology, 62 , 103–134; Singer, T., Seymour, B., O’Doherty, J., Kaube, H., Dolan, R. J., & Frith, C. D. (2004). Empathy for pain involves the affective but not sensory components of pain. Science, 303 (5661), 1157–1162.
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| affect matching can sometimes lead to avoidance behavior Batson, C. D. (1991). The Altruism Question: Toward a Social-Psychological Answer. Hillsdale, NJ: Lawrence Erlbaum Associates.
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| empathy occurs only when there is an appropriate emotional response Ibid.
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| Given that nearly all of the follow-ups to Singer’s seminal work have focused on empathy Fan, Y., Duncan, N. W., de Greck, M., & Northoff, G. (2011). Is there a core neural network in empathy? An fMRI based quantitative meta-analysis. Neuroscience & Biobehavioral Reviews, 35 (3), 903–911.
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| almost none of the studies that have been done have linked neural responses Hein, G., Silani, G., Preuschoff, K., Batson, C. D., & Singer, T. (2010). Neural responses to ingroup and outgroup members’ suffering predict individual differences in costly helping. Neuron, 68 (1), 149–160.
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| Sylvia Morelli, Lian Rameson, and I ran an fMRI study Morelli, S. A., Rameson, L. T., & Lieberman, M. D. (in press). The neural components of empathy: Predicting daily prosocial behavior. Social Cognitive and Affective Neuroscience.
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| the hot area of study in the next ten years Cf. Moll, J., Zahn, R., de Oliveira-Souza, R., Bramati, I. E., Krueger, F., Tura, B., … , & Grafman, J. (2011). Impairment of prosocial sentiments is associated with frontopolar and septal damage in frontotemporal dementia. NeuroImage, 54 (2), 1735–1742; Krueger, F., McCabe, K., Moll, J., Kriegeskorte, N., Zahn, R., Strenziok, M., … , & Grafman, J. (2007). Neural correlates of trust. Proceedings of the National Academy of Sciences, 104 (50), 20084–20089; Inagaki, T. K., & Eisenberger, N. I. (2012). Neural correlates of giving support to a loved one. Psychosomatic Medicine, 74 , 3–7.
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| the dorsomedial prefrontal cortex (DMPFC)—the CEO of the brain’s mentalizing system Andy, O. J., & Stephan, H. (1966). Septal nuclei in primate phylogeny: A quantitative investigation. Journal of Comparative Neurology, 126 (2), 157–170; Sesack, S. R., Deutch, A. Y., Roth, R. H., & Bunney, B. S. (1989). Topographical organization of the efferent projections of the medial prefrontal cortex in the rat: An anterograde tract-tracing study with Phaseolus vulgaris leucoagglutinin. Journal of Comparative Neurology, 290 (2), 213–242.
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| When a rat pressed the lever Olds, J., & Milner, P. (1954). Positive reinforcement produced by electrical stimulation of septal area and other regions of rat brain. Journal of Comparative and Physiological Psychology, 47 (6), 419.
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| Two decades later a similar study was conducted Heath, R. G. (1972). Pleasure and brain activity in man. Journal of Nervous and Mental Disease, 154 (363), 9.
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| At the same time that researchers were linking the septal area to reward Brady, J. V., & Nauta, W. J. (1953). Subcortical mechanisms in emotional behavior: Affective changes following septal forebrain lesions in the albino rat. Journal of Comparative and Physiological Psychology, 46 (5), 339.
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| Lesion studies in rats, mice, and rabbits suggest Carlson, N. R., & Thomas, G. J. (1968). Maternal behavior of mice with limbic lesions. Journal of Comparative and Physiological Psychology, 66 (3p1), 731; Cruz, M. L., & Beyer, C. (1972). Effects of septal lesions on maternal behavior and lactation in the rabbit. Physiology & Behavior, 9 (3), 361–365; Slotnick, B. M., & Nigrosh, B. J. (1975). Maternal behavior of mice with cingulate cortical, amygdala, or septal lesions. Journal of Comparative and Physiological Psychology, 88 (1), 118.
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| one way to reconcile the findings is to characterize the septal area as shifting the balance Inagaki, T. K., & Eisenberger, N. I. (2012). Neural correlates of giving support to a loved one. Psychosomatic Medicine, 74 , 3–7.
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| The septal area is rich in oxytocin receptors Insel, T. R., Gelhard, R., & Shapiro, L. E. (1991). The comparative distribution of forebrain receptors for neurohypophyseal peptides in monogamous and polygamous mice. Neuroscience, 43 (2), 623–630.
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| Among rodents, pups who receive more parental care Lukas, M., Bredewold, R., Neumann, I. D., & Veenema, A. H. (2010). Maternal separation interferes with developmental changes in brain vasopressin and oxytocin receptor binding in male rats. Neuropharmacology, 58 (1), 78–87; Francis, D. D., Champagne, F. C., & Meaney, M. J. (2001). Variations in maternal behaviour are associated with differences in oxytocin receptor levels in the rat. Journal of Neuroendocrinology, 12 (12), 1145–1148.
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| Just two years after the first Sally-Anne test for Theory of Mind Baron-Cohen, S., Leslie, A. M., & Frith, U. (1985). Does the autistic child have a “theory of mind”? Cognition, 21 (1), 37–46.
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| Subsequent studies have demonstrated other mentalizing deficits Baron-Cohen, S., O’Riordan, M., Stone, V., Jones, R., & Plaisted, K. (1999). Recognition of faux pas by normally developing children and children with Asperger syndrome or high-functioning autism. Journal of Autism and Developmental Disorders, 29 , 407–418; White, S. J., Hill, E. L., Happé, F., & Frith, U. (2009). Revisiting the Strange Stories: Revealing mentalizing impairments in autism. Child Development, 80 , 1097–1117.
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| Autistic individuals are also much less likely to describe Heider, F., & Simmel, M. (1944). An experimental study of apparent behavior. American Journal of Psychology, 57 , 243–259; Klin, A. (2003). Attributing social meaning to ambiguous visual stimuli in higher-functioning autism and Asperger syndrome: The social attribution task. Journal of Child Psychology and Psychiatry, 41 (7), 831–846.
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| The Sally-Anne test isn’t the only or the hardest test of Theory of Mind Frith, U., & Happé, F. (1994). Autism: Beyond “theory of mind.” Cognition, 50 (1), 115–132.
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| Uta Frith gave a group of autistic children Shah, A., & Frith, U. (1983). An islet of ability in autistic children: A research note. Journal of Child Psychology and Psychiatry, 24 (4), 613–620.
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| Autism is associated with a deficit in focusing on high-level meaning Frith, U., & Happé, F. (1994). Autism: Beyond “theory of mind.” Cognition, 50 (1), 115–132.
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| the two deficits do not always go hand in hand in autism Ibid.; Spunt, R. P., Meyer, M. L., & Lieberman, M. D. (under review). Social by default: Brain activity at rest facilitates social cognition; cf. Baron-Cohen, S. (2009). Autism: The Empathizing-Systemizing (E-S) Theory. Annals of the New York Academy of Sciences, 1156 (1), 68–80.
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| Multiple studies have shown that training can lead to sizable gains Hadwin, J., Baron-Cohen, S., Howlin, P., & Hill, K. (1997). Does teaching theory of mind have an effect on the ability to develop conversation in children with autism? Journal of Autism and Developmental Disorders, 27 (5), 519–537; Ozonoff, S., & Miller, J. N. (1995). Teaching theory of mind: A new approach to social skills training for individuals with autism. Journal of Autism and Developmental Disorders, 25 (4), 415–433.
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| We know this because children who are born deaf Peterson, C. C., & Siegal, M. (1999). Deafness, conversation and theory of mind. Journal of Child Psychology and Psychiatry, 36 (3), 459–474; Peterson, C. C., & Siegal, M. (1999). Representing inner worlds: Theory of mind in autistic, deaf, and normal hearing children. Psychological Science, 10 (2), 126–129; Peterson, C. C., & Siegal, M. (2002). Insights into theory of mind from deafness and autism. Mind & Language, 15 (1), 123–145.
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| During the second year, these children tend to ignore others Adrien, J. L., Lenoir, P., Martineau, J., Perrot, A., Hameury, L., Larmande, C., & Sauvage, D. (1993). Blind ratings of early symptoms of autism based upon family home movies. Journal of the American Academy of Child & Adolescent Psychiatry, 32 (3), 617–626; Klin, A., Volkmar, F. R., & Sparrow, S. S. (1992). Autistic social dysfunction: Some limitations of the theory of mind hypothesis. Journal of Child Psychology and Psychiatry, 33 (5), 861–876.
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| experimenters have asked children to imitate various behaviors and hand gestures DeMyer, M. K., Alpern, G. D., Barton, S., DeMyer, W. E., Churchill, D. W., Hingtgen, J. N., … , & Kimberlin, C. (1972). Imitation in autistic, early schizophrenic, and non-psychotic subnormal children. Journal of Autism and Developmental Disorders, 2 (3), 264–287.
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| Children diagnosed with autism consistently perform worse Williams, J. H., Whiten, A., & Singh, T. (2004). A systematic review of action imitation in autistic spectrum disorder. Journal of Autism and Developmental Disorders, 34 (3), 285–299.
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| Once the mirror system was clearly linked with imitation Nishitani, N., Avikainen, S., & Hari, R. (2004). Abnormal imitation-related cortical activation sequences in Asperger’s syndrome. Annals of Neurology, 55 (4), 558–562; Oberman, L. M., Hubbard, E. M., McCleery, J. P., Altshuler, E. L., Ramachandran, V. S., & Pineda, J. A. (2005). EEG evidence for mirror neuron dysfunction in autism spectrum disorders. Cognitive Brain Research, 24 , 190–198; Dapretto, M., Davies, M. S., Pfeifer, J. H., Scott, A. A., Sigman, M., Bookheimer, S. Y., & Iacoboni, M. (2005). Understanding emotions in others: Mirror neuron dysfunction in children with autism spectrum disorders. Nature Neuroscience, 9 (1), 28–30; Williams, J. H., Waiter, G. D., Gilchrist, A., Perrett, D. I., Murray, A. D., & Whiten, A. (2006). Neural mechanisms of imitation and mirror neuron functioning in autistic spectrum disorder. Neuropsychologia, 44 (4), 610–621; Ramachandran, V. S., & Oberman, L. M. (2006). Broken mirrors: A theory of autism. Scientific American, 16 , 62–69; Gallese, V. (2006). Intentional attunement: A neurophysiological perspective on social cognition and its disruption in autism. Brain Research, 1079 (1), 15–24.
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