Publications

2013
Frenkel-Toledo, S., Bentin, S., Perry, A., Liebermann, D. G., & Soroker, N. . (2013). Dynamics of the EEG power in the frequency and spatial domains during observation and execution of manual movements. Brain Research, 1509, 43-57. Retrieved from Publisher's VersionAbstract
Mu suppression is the attenuation of EEG power in the alpha frequency range (8–12 Hz) while executing or observing a motor action. Whereas typically observed at central scalp sites, there are diverging reports about the extent of the attenuation over the cortical mantle, its exact frequency range and the specificity of this phenomenon. We investigated the modulation of EEG oscillations in frequency-bands between 4 to 12 Hz at frontal, central, parietal and occipital sites during the execution of manual movements and during observation of similar actions from allocentric (i.e., facing the actor) and egocentric (i.e., seeing the actor from behind) viewpoints. Suppression was determined relative to observation of a non-biological movement. Action observation elicited greater suppression in the lower (8–10 Hz) compared to the higher mu range (10–12 Hz), and greater suppression in the entire range (4–12 Hz) at frontal and central sites compared to parietal and occipital sites. In addition, suppression tended to be greater during observation of a motor action from allocentric compared to egocentric viewpoints. During execution of movement, suppression of the EEG occurred primarily in the higher alpha range and was absent at occipital sites. In the theta range (4–8 Hz), the EEG amplitude was suppressed during action observation and execution. The results suggest a functional distinction between modulation of mu and alpha rhythms, and between the higher and lower ranges of the mu rhythms. The activity of the presumed human mirror-neuron system seems primarily evident in the lower mu range and in the theta range.
Perry, A., Aviezer, H., Goldstein, P., Palgi, S., Klein, E., & Shamay-Tsoory, S. G. . (2013). Face or body? Oxytocin improves perception of emotions from facial expressions in incongruent emotional body context. Psychoneuroendocrinology, 2820. Retrieved from Publisher's VersionAbstract
The neuropeptide oxytocin (OT) has been repeatedly reported to play an essential role in the regulation of social cognition in humans in general, and specifically in enhancing the recognition of emotions from facial expressions. The later was assessed in different paradigms that rely primarily on isolated and decontextualized emotional faces. However, recent evidence has indicated that the perception of basic facial expressions is not context invariant and can be categorically altered by context, especially body context, at early perceptual levels. Body context has a strong effect on our perception of emotional expressions, especially when the actual target face and the contextually expected face are perceptually similar. To examine whether and how OT affects emotion recognition, we investigated the role of OT in categorizing facial expressions in incongruent body contexts. Our results show that in the combined process of deciphering emotions from facial expressions and from context, OT gives an advantage to the face. This advantage is most evident when the target face and the contextually expected face are perceptually similar.
Perry, A., & Shamay-Tsoory, S. . (2013). Understanding emotional and cognitive empathy: A neuropsychological perspective. In Understanding Other Minds: Perspectives from developmental social neuroscience. Oxford University Press. Retrieved from Publisher's VersionAbstract
Empathy is a broad concept that refers to the cognitive as well as the emotional reactions of one individual to the observed experiences of another. Questions regarding how we understand others have intrigued psychologists and philosophers for centuries. In order to answer these questions, two major theories have been proposed, known as Theory Theory and Simulation Theory. In the past two decades, these questions have been re-examined by neuropsychologists and neuroscientists. This chapter reviews the different aspects of emotional and cognitive empathy in light of converging evidence from lesion patients, electrophysiology and neuroimaging studies. Neuropsychiatric disorders, such as Autism or Schizophrenia, although heterogeneous and difficult to study, have also been examined in relation to their deficits in cognitive and emotional empathy, and some of these new findings are discussed in this chapter. Lastly, we propose a model which relates brain mechanisms such as simulation or cognitive and affective ToM, to psychological processes, the empathic responses they lead to, and deficits which may occur when these mechanisms are disrupted.
2012
Aviezer, H., Hassin, R. R., Perry, A., Dudarev, V., & Bentin, S. . (2012). The right place at the right time: Priming facial expressions with emotional face components in developmental visual agnosia. Neuropsychologia, 50, 949-957. Retrieved from Publisher's VersionAbstract
The current study examined the nature of deficits in emotion recognition from facial expressions in case LG, an individual with a rare form of developmental visual agnosia (DVA). LG presents with profoundly impaired recognition of facial expressions, yet the underlying nature of his deficit remains unknown. During typical face processing, normal sighted individuals extract information about expressed emotions from face regions with activity diagnostic for specific emotion categories. Given LG’s impairment, we sought to shed light on his emotion perception by examining if priming facial expressions with diagnostic emotional face components would facilitate his recognition of the emotion expressed by the face. LG and control participants matched isolated face components with components appearing in a subsequently presented full-face and then categorized the face’s emotion. Critically, the matched components were from regions which were diagnostic or non-diagnostic of the emotion portrayed by the full face. In experiment 1, when the full faces were briefly presented (150 ms), LG’s performance was strongly influenced by the diagnosticity of the components: his emotion recognition was boosted within normal limits when diagnostic components were used and was obliterated when non-diagnostic components were used. By contrast, in experiment 2, when the face-exposure duration was extended (2000 ms), the beneficial effect of the diagnostic matching was diminished as was the detrimental effect of the non-diagnostic matching. These data highlight the impact of diagnostic facial features in normal expression recognition and suggest that impaired emotion recognition in DVA results from deficient visual integration across diagnostic face components.
2011
Perry, A., Stein, L., & Bentin, S. . (2011). Motor and attentional mechanisms involved in social interaction—Evidence from mu and alpha EEG suppression. NeuroImage, 58, 895-904. Retrieved from Publisher's VersionAbstract

Mu rhythms are EEG oscillations in the 8–13 Hz recorded at sites located roughly over the sensory-motor cortex. There is reliable evidence that the amplitude of mu rhythms is reduced when the participant performs a motor act (mu suppression). Recent studies found mu suppression not only in response to actual movements but also while the participant observes actions executed by someone else. This finding putatively associates the mu suppression to the activity of a mirror neurons system which, in humans, has been suggested to contribute to social skills. In the present study we explored the effects of different levels of social interaction on mu suppression. Participants observed dynamic displays of hand gestures performing actions used in the Rock–Scissors–Paper game. In different blocks, participants passively viewed identical video clips with no game context and in the context of a game, or while being actually engaged in the game either by imagining actions or by actual playing. As a baseline for calculating mu suppression we used a dynamic display of a rolling ball. In addition, to isolate the social aspect of the actual movements, participants performed the same acts outside the game context. Mu suppression was larger while participants were engaged in the social game than when they passively looked at the “opponent” actions or when they performed movements without the game context. This effect was found while viewing the opponent play as well as while actually playing, which supports the view that mu suppression is affected not only by motion, but also by the social context of the motion. However, we did not find differences in mu suppression between perception segments in which the participant did not actually play. Furthermore, in all perception segments occipital alpha suppression was more robust than mu suppression suggesting the involvement of a strong attentional component. While actually playing, however, mu suppression was stronger than alpha suppression.

2010
Anat, P., & Shlomo, B. . (2010). Does focusing on hand grasping intentions modulate Electroencephalogram mu and alpha suppression?. Neuroreport, 21, 1050-1054. Retrieved from Publisher's VersionAbstract
Understanding the intentions of others presumably involves a human analog of the mirror neuron system. A putative marker of such mirror activity is the suppression of electroencephalographic oscillations in the 8–12 Hz range, which, when recorded over somatosensory areas, is associated with motor activity and labeled μ rhythms. We investigated whether μ-suppression can be modulated by attention to another person’s intention as expressed by her hand movement toward an object and whether this suppression is distinguished from the suppression of α waves that oscillate in the same frequency range and are modulated by attention and cognitive load. Both μ and α suppressions were modulated by task difficulty, and not distinctively by intention, reflecting the recruitment of resources needed for task performance.
Perry, A., Troje, N. F., & Bentin, S. . (2010). Exploring motor system contributions to the perception of social information: Evidence from EEG activity in the mu/alpha frequency range. Social Neuroscience, 5, 272-284. Retrieved from Publisher's VersionAbstract
Putative contributions of a human mirror neuron system (hMNS) to the perception of social information have been assessed by measuring the suppression of EEG oscillations in the mu/alpha (8–12 Hz), beta (15–25 Hz) and low-gamma (25–25 Hz) ranges while participants processed social information revealed by point-light displays of human motion. Identical dynamic displays were presented and participants were instructed to distinguish the intention, the emotion, or the gender of a moving image of a person, while they performed an adapted odd-ball task. Relative to a baseline presenting a nonbiological but meaningful motion display, all three biological motion conditions reduced the EEG amplitude in the mu/alpha and beta ranges, but not in the low-gamma range. Suppression was larger in the intention than in the emotion and gender conditions, with no difference between the latter two. Moreover, the suppression in the intention condition was negatively correlated with an accepted measure of empathy (EQ), revealing that participants high in empathy scores manifested less suppression. For intention and emotion the suppression was larger at occipital than at central sites, suggesting that factors other than motor system were in play while processing social information embedded in the motion of point-light displays.
Perry, A., Bentin, S., Shalev, I., Israel, S., Uzefovsky, F., Bar-On, D., & Ebstein, R. P. . (2010). Intranasal oxytocin modulates EEG mu/alpha and beta rhythms during perception of biological motion. Psychoneuroendocrinology, 35, 1446 - 1453. Retrieved from Publisher's VersionAbstract
Oxytocin (OT) plays a determining role in social and pair bonding in many vertebrates and increasing evidence suggests it is a social hormone also in humans. Indeed, intranasal administration of OT modulates several social cognitive processes in humans. Electrophysiological studies in humans associated the suppression of EEG in the mu/alpha and beta bands with perception of biological motion and social stimuli. It has been suggested that mu and beta suppression over sensory-motor regions reflects a resonance system in the human brain analogous to mirror neurons in the monkey. We therefore hypothesized that OT, a social hormone, would enhance this suppression, hence, for the first time, link the action of this neuropeptide with a human correlate of mirror neuron activity. Twenty-four students were administered 24IU of OT or placebo intranasally in a robust, double-blind within-subject design. 45min later participants were shown a point-light display of continuous biological motion of a
Perry, A., Bentin, S., Ben-Ami Bartal, I., Lamm, C., & Decety, J. . (2010). “Feeling” the pain of those who are different from us: Modulation of EEG in the mu/alpha range. Cognitive, Affective, & Behavioral Neuroscience, 10, 493–504. Retrieved from Publisher's VersionAbstract
We explored how apparently painful stimuli and the ability to identify with the person on whom the pain is inflicted modulate EEG suppression in the mu/alpha range (8-12 Hz). In a 2 × 2 design, we presented pictures of hands either experiencing needle pricks or being touched by a Q-tip. In the dissimilar-other condition, the hand was assigned to a patient suffering from a neurological disease in which Q-tips inflicted pain, whereas needle pricks did not. In the similar-other condition, the hand was assigned to a patient who responded to stimulation in the same way as the healthy participant. Participants [...]
2009
Anat, P., & Shlomo, B. . (2009). Mirror activity in the human brain while observing hand movements: Equivalence between EEG desynchronization in the μ-range and fMRI. Brain Research, 1282, 126-132. Retrieved from Publisher's VersionAbstract
Mu (μ) rhythms are EEG oscillations between 8–13 Hz distinguished from alpha by having more anterior distribution and being desynchronized by motor rather than visual activity. Evidence accumulating during the last decade suggests that the desynchronization of μ rhythms (μ suppression) might be also a manifestation of a human Mirror Neuron System (MNS). To further explore this hypothesis we used a paradigm that, in a previous fMRI study, successfully activated this putative MNS in humans. Our direct goal was to provide further support for a link between modulation of μ rhythms and the MNS, by finding parallels between the reported patterns of fMRI activations and patterns of μ suppression. The EEG power in the μ range has been recorded while participants passively observed either a left or a right hand, reaching to and grasping objects, and compared it with that recorded while participants observed the movement of a ball, and while observing static grasping scenes or still objects. Mirroring fMRI results (Shmuelof, L., Zohary, E., 2005. Dissociation between ventral and dorsal fMRI activation during object and action recognition. Neuron 47, 457–470), μ suppression was larger in the hemisphere contra-lateral to the moving hand and larger when the hands grasped different objects in different ways than when the movement was repetitive. No suppression was found while participants observed still objects but μ suppression was also found while seeing static grasping postures. These data are discussed in light of similar parallels between modulations of alpha waves and fMRI while recording EEG in the magnet. The present data support a link between μ suppression and a human MNS.
Perry, A., & Bentin, S. . (2009). Research Report: Mirror activity in the human brain while observing hand movements: A comparison between EEG desynchronization in the ?-range and previous fMRI results. Brain Research, 1282, 126 - 132. Retrieved from Publisher's VersionAbstract
Mu (?) rhythms are EEG oscillations between 8–13 Hz distinguished from alpha by having more anterior distribution and being desynchronized by motor rather than visual activity. Evidence accumulating during the last decade suggests that the desynchronization of ? rhythms (? suppression) might be also a manifestation of a human Mirror Neuron System (MNS). To further explore this hypothesis we used a paradigm that, in a previous fMRI study, successfully activated this putative MNS in humans. Our direct goal was to provide further support for a link between modulation of ? rhythms and the MNS, by finding parallels between the reported patterns of fMRI activations and patterns of ? suppression. The EEG power in the ? range has been recorded while participants passively observed either a left or a right hand, reaching to and grasping objects, and compared it with that recorded while participants observed the movement of a ball, and while observing static grasping scenes or still objects. Mi
Gilaie-Dotan, S., Perry, A., Bonneh, Y., Malach, R., & Bentin, S. . (2009). Seeing with profoundly deactivated mid-level visual areas: Non-hierarchical functioning in the human visual cortex. Cerebral Cortex, 19, 1687-1703. Retrieved from Publisher's VersionAbstract
A fundamental concept in visual processing is that activity in high-order object-category distinctive regions (e.g., lateral occipital complex, fusiform face area, middle temporal+) is dependent on bottom-up flow of activity in earlier retinotopic areas (V2, V3, V4) whose main input originates from primary visual cortex (V1). Thus, activity in down stream areas should reflect lower-level inputs. Here we qualify this notion reporting case LG, a rare case of developmental object agnosia and prosopagnosia. In this person, V1 was robustly activated by visual stimuli, yet intermediate areas (V2–V4) were strongly deactivated. Despite this intermediate deactivation, activity in down stream visual areas remained robust, showing selectivity for houses and places, while selectivity for faces and objects was impaired. The extent of impairment evident in functional magnetic resonance imaging and electroencephalography activations was somewhat larger in the left hemisphere. This pattern of brain activity, coupled with fairly adequate everyday visual performance is compatible with models emphasizing the role of nonlinear local “amplification” of neuronal inputs in eliciting activity in ventral and dorsal visual pathways as well as perceptual experience in the human brain. Thus, while the proper functioning of intermediate areas appears essential for specialization in the cortex, daily visual behavior and reading are maintained even with deactivated intermediate visual areas.