In Press
D., Cohen, Perry A., Mayseless N., and Shamay-Tsoory S.The role of oxytocin in implicit personal space regulation - an fMRI study..” Psychoneuroendocrinology (In Press).
O., Rubinstein, Korem N., Perry A., and Shamay-Tsoory S.Different neural activations for an approaching friend versus stranger: Linking personal space to numerical cognition in developmental dyscalculia.” Brain & Behavior (In Press).
A., Perry, Stiso J., Chang E.F., Lin J.J., Parvizi J., and Knight R.T.Mirroring in the Human Brain: Deciphering the Spatial-Temporal Patterns of the Human Mirror Neuron System, an ECoG study.” Cerebral Cortex 28, no. 3 (2018): 1039-1048. Publisher's VersionAbstract

2017 FEB 27 (NewsRx) – By a News Reporter-Staff News Editor at Pain Central Nervous System Week – Data detailed on Neurology - Cerebral Cortex Research have been presented. According [...]

N., Ensenberg, Perry A., and Aviezer H.Are you looking at me? Mu suppression modulation by facial expression direction.” Cognitive Affective & Behavioral Neuroscience 17, no. 1 (2017): 174 - 184. Publisher's VersionAbstract

Although we encounter numerous expressive faces on a daily basis, those that are not aimed at us will often be disregarded. Facial expressions aimed at our direction appear far more relevant and evoke an engaging affective experience, while the exact same expressions aimed away from us may not. While the importance of expression directionality is intuitive and commonplace, the neural mechanisms underlying this phenomenon are largely unknown. In the current study we measured EEG mu rhythm suppression, an established measure of mirror neuron activity, while participants viewed short video clips of dynamic facial expressions. Critically, the videos portrayed facial emotions which turned towards or away from the viewer, thus manipulating their degree of social relevance. Mirroring activity increased as a function of social relevance such that expressions turning toward the viewer resulted in increased sensorimotor activation (i.e., stronger mu suppression) compared to identical expression

A., Perry, Saunders S.N., Stiso J., Dewar C., Lubell J., Meling T.R., Solbakk A., Endestad T., and Knight R.T.Effects of prefrontal cortex damage on emotion understanding: EEG and behavioural evidence.” Brain: A Journal of Neurology 140, no. 4 (2017): 1086 - 1099. Publisher's VersionAbstract

Humans are highly social beings that interact with each other on a daily basis. In these complex interactions, we get along by being able to identify others' actions and infer their intentions, thoughts and feelings. One of the major theories accounting for this critical ability assumes that the understanding of social signals is based on a primordial tendency to simulate observed actions by activating a mirror neuron system. If mirror neuron regions are important for action and emotion recognition, damage to regions in this network should lead to deficits in these domains. In the current behavioural and EEG study, we focused on the lateral prefrontal cortex including dorsal and ventral prefrontal cortex and utilized a series of task paradigms, each measuring a different aspect of recognizing others' actions or emotions from body cues. We examined 17 patients with lesions including (n = 8) or not including (n = 9) the inferior frontal gyrus, a core mirror neuron system region, and com

D., Cohen, Perry A., Gilam G., Mayseless N., Gonen T., Hendler T., and Shamay-Tsoory S.G.The role of oxytocin in modulating interpersonal space: A pharmacological fMRI study.” Psychoneuroendocrinology 76 (2017): 77-83. Publisher's VersionAbstract

* We show that intranasal oxytocin (OT) increases personal distance from strangers. * OT reverses the activity of the left dorsomedial prefrontal cortex (dmPFC). * Under OT we observed higher activation in the dmPFC when a friend is approaching. * The effect of OT on personal space depends on the protagonist.

A., Perry, and Ben-Ami Bartal I.The Neurobiology of Empathy.” In Countertransference in Perspective, The double-edged sword of the patient-therapist emotional relationship, 2016.
I., Ben-Ami Bartal, and Perry A.The Evolution of Pro-social Behavior.” In Countertransference in Perspective, The double-edged sword of the patient-therapist emotional relationship, 2016.
A., Perry, Nichiporuk N., and Knight R.T.Where does one stand: a biological account of preferred interpersonal distance.” Social Cognitive & Affective Neuroscience 11, no. 2 (2016): 317 - 326. Publisher's VersionAbstract

What determines how close you choose to stand to someone? Why do some people prefer farther distances than others? We hypothesized that an important factor is one's sensory sensitivity level, i.e. how sensitive one is to nearby visual stimulation, noise, touch or smell. This study characterizes the behavioral, hormonal and electrophysiological metrics of interpersonal distance (IPD) preferences in relation to levels of sensory sensitivity. Using both an ecologically realistic task and electroencephalogram (EEG), we found that sensory sensitivity levels predicted IPD preferences, such that the more sensitive one is the farther distance they prefer. Furthermore, electrophysiological evidence revealed that individuals with higher sensory sensitivity show more alpha suppression for approaching stimuli, strengthening the notion that early sensory cortical excitability is involved in one's social decision of how close to stand to another. The results provide evidence that a core human metri

Z., Gao, Ye T., Shen M., and Perry A.Working memory capacity of biological movements predicts empathy traits.” Psychonomic Bulletin & Review 23, no. 2 (2016): 468 - 475. Publisher's VersionAbstract

Working memory (WM) and empathy are core issues in cognitive and social science, respectively. However, no study so far has explored the relationship between these two constructs. Considering that empathy takes place based on the others' observed experiences, which requires extracting the observed dynamic scene into WM and forming a coherent representation, we hypothesized that a sub-type of WM capacity, i.e., WM for biological movements (BM), should predict one's empathy level. Therefore, WM capacity was measured for three distinct types of stimuli in a change detection task: BM of human beings (BM; Experiment 1), movements of rectangles (Experiment 2), and static colors (Experiment 3). The first two stimuli were dynamic and shared one WM buffer which differed from the WM buffer for colors; yet only the BM conveyed social information. We found that BM-WM capacity was positively correlated with both cognitive and emotional empathy, with no such correlations for WM capacity of movement

L., Peled-Avron, Perry A., and Shamay-Tsoory S.G.The effect of oxytocin on the anthropomorphism of touch.” Psychoneuroendocrinology 66 (2016): 159-165. Publisher's VersionAbstract

* Oxytocin can enhance anthropomorphism of non-human objects. * Intranasal delivery of oxytocin increases the anthropomorphism of touch between two inanimate objects. * Touch plays a key role in the anthropomorphic effect of oxytocin.

A., Perry, Lwi S., Verstaen A., Dewar C., Levenson R.W., and Knight R.T.The role of the orbitofrontal cortex in regulation of interpersonal space: evidence from frontal lesion and frontotemporal dementia patients.” Social Cognitive & Affective Neuroscience 11, no. 12 (2016): 1894 - 1901. Publisher's VersionAbstract

Interpersonal distance is central to communication and complex social behaviors but the neural correlates of interpersonal distance preferences are not defined. Previous studies suggest that damage to the orbitofrontal cortex (OFC) is associated with impaired interpersonal behavior. To examine whether the OFC is critical for maintaining appropriate interpersonal distance, we tested two groups of patients with OFC damage: Patients with OFC lesions and patients with behavioral variant frontotemporal dementia. These two groups were compared to healthy controls and to patients with lesions restricted to the dorsolateral prefrontal cortex. Only patients with OFC damage showed abnormal interpersonal distance preferences, which were significantly different from both controls and patients with dorsolateral prefrontal damage. The comfortable distances these patients chose with strangers were significantly closer than the other groups and resembled distances normally used with close others. The

D., Gotthilf-Nezri, Lev M., Yehezkel O., Gilaie-Dotan S., Perry A., Bentin S., Bonneh Y., and Polat U.Training-induced recovery of undeveloped visual functions may be followed by improved perceptual functions: indication from adult with perceptual impairments.” Developmental Science 18, no. 1 (2015): 50-64.Abstract

Long-term deprivation of normal visual inputs can cause perceptual impairments at various levels of visual function, from basic visual acuity deficits, through mid-level deficits such as contour integration and motion coherence, to high-level face and object agnosia. Yet it is unclear whether training during adulthood, at a post-developmental stage of the adult visual system, can overcome such developmental impairments. Here, we visually trained LG, a developmental object and face agnosic individual. Prior to training, at the age of 20, LG's basic and mid-level visual functions such as visual acuity, crowding effects, and contour integration were underdeveloped relative to normal adult vision, corresponding to or poorer than those of 5–6 year olds (Gilaie-Dotan, Perry, Bonneh, Malach & Bentin, 2009). Intensive visual training, based on lateral interactions, was applied for a period of 9 months. LG's directly trained but also untrained visual functions such as visual acuity, crowding, binocular stereopsis and also mid-level contour integration improved significantly and reached near-age-level performance, with long-term (over 4 years) persistence. Moreover, mid-level functions that were tested post-training were found to be normal in LG. Some possible subtle improvement was observed in LG's higher-order visual functions such as object recognition and part integration, while LG's face perception skills have not improved thus far. These results suggest that corrective training at a post-developmental stage, even in the adult visual system, can prove effective, and its enduring effects are the basis for a revival of a developmental cascade that can lead to reduced perceptual impairments.

A., Perry, Levy-Gigi E., Richter-Levin G., and Shamay-Tsoory S.G.Interpersonal distance and social anxiety in autistic spectrum disorders: A behavioral and ERP study.” Social Neuroscience 10, no. 4 (2015): 354-365. Publisher's VersionAbstract

An inherent feature of social interactions is the use of social space or interpersonal distance-the space between one individual and another. Because social deficits are core symptoms of Autistic Spectrum Disorder (ASD), we hypothesized that individuals on this spectrum will exhibit abnormal interpersonal distance preferences. The literature on interpersonal distance in ASD is not conclusive. While some studies show preferences for closer distances among this group, others show preferences for farther distances than controls. A common symptom of ASD that may explain the variance in responses to interpersonal distance in this population is social anxiety (SA), which has been shown to correlate with interpersonal distance preferences. In the current study, we investigated interpersonal distance preferences in a group of individuals with ASD using both behavioral and ERP measures. We found greater variance in interpersonal distance preferences in the ASD group than in the control group. Furthermore, we showed that this variance can be explained by differences in SA level and can be predicted by the N1 amplitude, an early ERP component related to attention and discrimination processes. These results hint at the early sensory and attentional processes that may be affecting higher social behaviors, both in subclinical and in clinical populations.

A., Perry, Mankuta D., and Shamay-Tsoory S.G.Oxytocin promotes closer interpersonal distance among highly empathic individuals.” Social, Cognitive & Affective Neuroscience 10, no. 1 (2015): 3 - 9. Publisher's VersionAbstract

The space between people, or 'interpersonal distance', creates and defines the dynamics of social interactions and is a salient cue signaling responsiveness and feeling comfortable. This distance is implicit yet clearly felt, especially if someone stands closer or farther away than expected. Increasing evidence suggests that Oxytocin (OT) serves as a social hormone in humans, and that one of its roles may be to alter the perceptual salience of social cues. Considering that empathic ability may shape the way individuals process social stimuli, we predicted that OT will differentially affect preferred interpersonal distance depending on individual differences in empathy. Participants took part in two interpersonal distance experiments: In the first, they had to stop a (computer visualized) protagonist when feeling most comfortable; in the second, they were asked to choose the room in which they would later discuss intimate topics with another. Both experiments revealed an interaction be

S., Frenkel-Toledo, Bentin S., Perry A., Liebermann D.G., and Soroker N.Mirror-neuron system recruitment by action observation: Effects of focal brain damage on mu suppression.” NeuroImage 87 (2014): 127 - 137. Publisher's VersionAbstract

Mu suppression is the attenuation of EEG power in the alpha frequency range (8-12Hz), recorded over the sensorimotor cortex during execution and observation of motor actions. Based on this dual characteristic mu suppression is thought to signalize activation of a human analogue of the mirror neuron system (MNS) found in macaque monkeys. However, much uncertainty remains concerning its specificity and full significance. To further explore the hypothesized relationship between mu suppression and MNS activation, we investigated how it is affected by damage to cortical regions, including areas where the MNS is thought to reside. EEG was recorded in 33 first-event stroke patients during observation of video clips showing reaching and grasping hand movements. We examined the modulation of EEG oscillations at central and occipital sites, and analyzed separately the lower (8-10Hz) and higher (10-12Hz) segments of the alpha/mu range. Suppression was determined relative to observation of a non-

S., Frenkel-Toledo, Bentin S., Perry A., Liebermann D.G., and Soroker N.Dynamics of the EEG power in the frequency and spatial domains during observation and execution of manual movements.” Brain Research 1509 (2013): 43 - 57. Publisher's VersionAbstract

Mu suppression is the attenuation of EEG power in the alpha frequency range (8-12Hz) 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 12Hz 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 (8-10Hz) compared to the higher mu range (10-12Hz), and greater suppression in the entire range (4-12Hz) at frontal and central sites compared to parietal and occipital sites. In addition, supp

A., Perry, and Shamay-Tsoory S.Understanding emotional and cognitive empathy: A neuropsychological perspective.” In Understanding Other Minds: Perspectives from developmental social neuroscience. Oxford University Press, 2013. 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 c

A., Perry, Rubinsten Orly., Peled Leehe., and Shamay-Tsoory S.G.Don't stand so close to me: A behavioral and ERP study of preferred interpersonal distance.” NeuroImage 83 (2013): 761 - 769. Publisher's VersionAbstract

The space between people, or interpersonal distance, creates and defines the dynamics of social interactions. Given that invasion of one's interpersonal space may trigger threat and anxiety, a critical question is if high vulnerability to social anxiety (SA) is associated with avoidance and attentional biases when anticipating invasion to one's interpersonal space. Therefore, the current study sought to examine the behavioral mechanisms, time course and neural correlates underlying the threat of interpersonal distance invasion with a focus on different SA levels, using both a behavioral and an ERP experiment. Preferred interpersonal distance was assessed using a paradigm that involves responding to different virtual protagonists (friend or stranger) approaching the participant by indicating where one would like the protagonist to stop. In addition, participants' level of social anxiety was measured. The behavioral experiment indicated that levels of SA predicted one's preferred interp

A., Perry, Aviezer H., Goldstein P., Palgi S., Klein E., and Shamay-Tsoory S.G.Face or body? Oxytocin improves perception of emotions from facial expressions in incongruent emotional body context.” Psychoneuroendocrinology 38, no. 11 (2013): 2820-2825. 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,