Single-Neuron Responses in Humans during Execution and Observation of Actions

“In monkeys, however, neurons with mirroring properties have been reported in a variety of areas on the lateral wall of the primate brain [3, 7, 21, 24, 25].”

“This interpretation is in line with the hypothesis ofmultiplemirroring mechanisms in the primate brain, a hypothesis that can easily account for the presence of mirroring cells in many cortical areas [1, 3–5, 7, 8, 24, 25].”

“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].”

“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].”

“For example, the mirror mechanism in the insula might underlie the capacity to understand a specific emotion (disgust) in others [16, 19], whereas the mirror mechanism in the parietofrontal circuit may help understanding the goal of observed motor acts and the intentions behind them [21].”

“, a picture of a smiling face and the execution cue word ‘‘smile’’ [33].”

“This interpretation is in line with the hypothesis ofmultiplemirroring mechanisms in the primate brain, a hypothesis that can easily account for the presence of mirroring cells in many cortical areas [1, 3–5, 7, 8, 24, 25].”

“…such as automatic vs. controlled, emotional vs. neutral, positive vs. negative and affective vs. cognitive (Byrum et al., 1999; Maratos et al., 2001; Kuchinke et al., 2005; Saxe and Wexler, 2005; Keysers and Gazzola, 2007; Mukamel et al., 2010; Bohl and van den Bos, 2012; Kumfor and Piguet, 2012).”

“Because mirror neurons were recorded in these regions (Gallese et al., 1996; Umilt a et al., 2001; Kohler et al., 2002; Keysers et al., 2003; Cisek and Kalaska, 2004; Fogassi et al., 2005; Fujii et al., 2008; Rozzi et al., 2008; Mukamel et al., 2010), this group has been called the (putative) mirror neuron system (pMN…”

“…S A I 1 d e t i p ( I t y a B B i a i 2 e i a h 1 Developmental Cognitive Neuroscience 15 (2015) 1–10 Contents lists available at ScienceDirect Developmental Cognitive Neuroscience j o ur nal ho me pa ge: ht tp : / /www.e lsev ier .com/ locate /dcn n EEG study on the somatotopic organisation of…”

“Nonetheless, the role of posterior SMA in sensory feedback, as described in mirror movements (Mukamel et al. 2010), cannot be excluded.”

“Recent work (Mukamel et al. 2010) suggests that some neurons in the human pre-SMA show mirror properties—that is, discharging when executing a given motor act and when observing the same action being performed by someone else.”

“…neurons in human cortex (Chong et al., 2008; Dinstein et al., 2008; Lingnau et al., 2009), single-cell recordings in humans demonstrate the existence of action-selective and mirror neurons in various areas in the human cortex, including the supplementary motor area (SMA) (Mukamel et al., 2010).”

“…System Group, Lab for Molecular Analysis of Higher Brain Functions, RIKEN Brain Science Institute, Saitama, Japan, 3Division of Sensory and Cognitive Information, National Institute for Physiological Sciences, Aichi, Japan, 4Department of Physiological Sciences, The Graduate University for…”

“Such a result is supported by several fMRI studies of the human mirror system [27] [28] [62], the findings of mirror neurons in supplementary motor area in humans [49], and mirror-like neurons in macaque dorsal premotor cortex [13].”

“Similarly, the study of sensorimotor units in human participants has revealed large proportions of neurons with both non-congruent and ambiguous sensorimotor properties (Mukamel et al., 2010).”

“These deactivations may be a function of suppression of specific cells in the primary motor cortex, possibly through the supplementary motor area during action observation, to prevent imitation (Bien et al., 2009; Kraskov et al., 2009; Mukamel et al., 2010; Vigneswaran et al., 2013).”

“regions has yet to be confirmed [8] (but see [9] for data on human mirror neurons in other brain regions).”

“Neurons with mirroring properties have also been found in the human SMA (Mukamel et al. 2010).”

“both observation and execution of actions (Mukamel et al., 2010).”

“This also provides a kinematic explanation for the recently found parahippocampal activation during motor execution (Mukamel, Ekstrom, Kaplan, Iacoboni, & Fried, 2010 ).”

“For example, action observation interventions could be viewed: (i) passively, without any additional instructions; (ii) with Frontiers in Human Neuroscience | www.frontiersin.org 1 February 2016 | Volume 10 | Article 17 the intent to subsequently imitate the observed movements; or (iii) whilst…”

“…another one is inhibited (Hari et al., 2014; Vigneswaran et al., 2013) reducing the overall output to the spinal cord (Stamos et al., 2010) and thus possibly contributing to the attribution of action to the correct agent (Mukamel et al., 2010; Savaki, 2010; Schütz-Bosbach et al., 2009).”

“Therefore, it has been a topic of considerable debate whether individual neurons in the human brain express a precise sensorimotor correspondence and whether such cells may play a role in linking the perception and execution of vocal signals (Brass et al., 2007; Hari et al., 1998; Mukamel et al., 2010).”

“2005), parietal lobule, and others become active during both the observation and execution of action (Decety et al. 2002; Gazzola and Keysers 2009; Iacoboni et al. 1999; Mukamel et al. 2010).”

“The current findings also provide an interesting new take on theories regarding the ‘action observation system’, where motor skills activate the same neural circuitry in left premotor cortex as observing the same action (Malfait et al., 2010; Mukamel et al., 2010).”

“The same group (Mukamel et al., 2010), by recording extracellular activity, described neurons with mirror properties in the hippocampus (that is neurons which responded to both observation and execution of actions).”

“Interestingly, neurons with mirror properties have also been shown in the hippocampal regions, and their functional significance is still only partly understood, although an involvement of AO in mnestic processes has been suggested (Calvo-Merino et al., 2005; Mukamel et al., 2010).”

“medial temporal lobe, superior parietal lobule (SPL) and even primary motor areas have been described as part of the human MNS [12,15,16,20–25].”

“Mirror systems are both active when an individual performs an action and when another individual performs an action from the same class of actions or an action with a similar goal or meaning (Di Pellegrino et al., 1992; Mukamel et al., 2010; Rizzolatti and Craighero, 2004).”

“Intracranial recordings from patients support the existence of neurons with mirror properties in humans and demonstrate that a much broader cortical and subcortical network may be involved in the mirror system (Mukamel et al., 2010) and its related func-”

“, 1999) and single-cell recording (Mukamel et al., 2010) studies have shown that similar mirror neuron processes can also be observed in humans.”

“It would be interesting to see if the definition of empathy has changed since the discovery of the Mirror Neuron System in the 1990s (Gallese et al., 1996; Mukamel et al., 2010).”

“Keywords: auditory mirror neuron system, action comprehension, mu rhythm, sensorimotor cortex, mu suppression”

“Mukamel et al. (2010) have described overlapping activation patterns for single units during action observation and execution in human supplementary motor area, cingulate cortex and medial temporal lobe, lending further support to the concept of overlapping networks.”

“[54], who reported mirror neurons in humans in the”

“…University of Essex, Brain Science, Department of et al., 1995; Enticott et al., 2010; Sartori et al., 2012), depth electrode recording (Mukamel et al., 2010), and EEG/MEG (e.g., Hari et al., 1998; Nishitani and Hari, 2000; Muthukumaraswamy and Johnson, 2004a,b; Kilner et al., 2009)…”

“Numerous studies have investigated the role of mirror neurons in parieto-frontal circuits in monkeys (Di Pellegrino et al. 1992; Gallese et al. 1996; Caggiano et al. 2009; Bonini et al. 2010; Rizzolatti and Sinigaglia 2010) and humans (Iacoboni et al. 2005; Mukamel et al. 2010).”

“The existence of mirror neurons within the premotor and parietal region [Mukamel et al., 2010], which are fine tuned to different levels of action control, underpins this notion [Fernandino and Iacoboni, 2010].”

“In summary, the present study identified that individuals with HFA/AS possessed increased BOLD in PMC, ACC and medial frontal gyrus during action observation.”

“Mukamel et al. (2010) reported on experiments using single-cell recording in humans. The electrodes were implanted as a prelude to surgery rather than primarily for research, and thus, the location of the electrodes was not selected to test various hypotheses. Nonetheless, these data provide some of the most convincin…”

“…evidence that mirror neurons exist in homologous regions of human brains (de la Rosa et al., 2016; Iacoboni and Dapretto, 2006; Kilner and Lemon, 2013), and these systems are involved in movement recognition, selection of movement, and imitation (Iacoboni and Dapretto, 2006; Mukamel et al., 2010).”

“…1995, 2005), fMRI (e.g., Iacoboni et al., 1999), EEG/MEG (e.g., Hari et al., 1998; Järveläinen et al., 2001; Caetano et al., 2007; Kilner and Frith, 2007; Kilner et al., 2009; Neuper et al., 2009), intracranial recordings (Mukamel et al., 2010) and occasionally with EMG (Leighton et al., 2010).”

“However, one recent neurophysiological study in humans has shown that a significant proportion of neurons in the SMA responded to both the observation and execution of actions (Mukamel et al., 2010).”

“During the observation of facial gestures or during the execution/imitation of the same facial gesture, mirror activation was found in motor (area 6), supplemental motor area (SMA), premotor (BA44 and BA45) and parietal (IGF) cortex (Buccino et al., 2001; Carr et al., 2003).”

“…support has emerged for the presence of mirror neurons in areas that control saccadic activity and overt attention such as the lateral intraparietal area (LIP; Shepherd, Klein, Deaner, & Platt, 2009), and the supplementary motor area including the supplementary eye fields (Mukamel et al., 2010).”

“frontal gyrus (Rizzolatti & Craighero, 2004), although recently, neurons with mirror characteristics have been found outside these areas (Mukamel et al., 2010), in medial”

“Since the original discovery, the existence of mirror neurons in other regions of the human and nonhuman motor pathway (including parietal, and primary motor cortex) has been demonstrated (Filimon et al. 2007; Tkach et al. 2007; Chong et al. 2008; Dushanova and Donoghue 2010; Mukamel et al. 2010; Kilner and Lemon 2013…”

“, 2005), mirror neurons (Mukamel et al., 2011), novelty and familiarity cells (Rutishauser et al.”

“(Fadiga et al., 1995; Mukamel et al., 2010, respectively).”

“More specifically, this has been shown in work reported in research by Mukamel and colleagues (Mukamel et al., 2010; Fried, Mukamel, Kreiman, 2011); see also (Keysers and Gazzola, 2010; Iacoboni, 2008a; Iacoboni, 2008b; Iacoboni & Dapretto, 2006); see also (Iacoboni, 2008a, pp. 201-203).”

“Furthermore, single-unit recordings in humans indicate that viewing grasping actions and facial expressions is associated with greater activity in neurons of the supplementary motor area, hippocampus, parahippocampal gyrus and entorhinal cortex [45].”

“…areas of overlap in the fronto-parietal mirror circuit (Caspers et al. 84 2010; Konoike et al. 2012), and the supplementary motor area (SMA, Vogt et al. 2007; 85 Mukamel et al. 2010; Dayan and Cohen 2011; Hardwick et al. 2013), as well as task-specific 86 differences (research objective 1).”

“MNS predominantly comprises the inferior frontal gyrus, inferior parietal lobule (IPL), and posterior superior temporal sulcus (6).”

“…Oemerged from direct contrasts across tasks (Alario et al., 2006; Barch et al., 2000; Botvinick et al., 2004; Chen et al., 2009; Ketteler et al., 2008; Mukamel et al., 2010; Murphy et al., 1997; Nachev et al., 2007; Simmonds et al., 2008; Sörös et al., 2006; Wahl et al., 2008; Wise et al., 1999).”

“In general, the direct and indirect findings in humans suggest that the human mirror neuron system may be larger than monkey mirror neuron system, thus including somatosensory areas being somatotopicaly organized [62, 70, 79] .”

“32 This opens the possibility that MTL may play a role in the sensori-motor translation of perceived and produced number, 33 a speculation yet to be explored.”