Extrastriate Body Area Research Articles

Engagement of the left extrastriate body area during body-part metaphor comprehension

Grounded cognition explanations of metaphor comprehension predict activation of sensorimotor cortices relevant to the metaphor’s source domain. We tested this prediction for body-part metaphors using functional magnetic resonance imaging while participants heard sentences containing metaphorical or literal references to body parts, and comparable control sentences. Localizer scans identified body-part-specific motor, somatosensory and visual cortical regions. Both subject- and item-wise analyses showed that, relative to control sentences, metaphorical but not literal sentences evoked limb metaphor-specific activity in the left extrastriate body area (EBA), paralleling the EBA’s known visual limb-selectivity. The EBA focus exhibited resting-state functional connectivity with ipsilateral semantic processing regions. In some of these regions, the strength of resting-state connectivity correlated with individual preference for verbal processing. Effective connectivity analyses showed that, during metaphor comprehension, activity in some semantic regions drove that in the EBA. These results provide converging evidence for grounding of metaphor processing in domain-specific sensorimotor cortical activity.

Differential neural encoding of sensorimotor and visual body representations.

Sensorimotor processing specifically impacts mental body representations. In particular, deteriorated somatosensory input (as after complete spinal cord injury) increases the relative weight of visual aspects of body parts’ representations, leading to aberrancies in how images of body parts are mentally manipulated (e.g. mental rotation). This suggests that a sensorimotor or visual reference frame, respectively, can be relatively dominant in local (hands) versus global (full-body) bodily representations. On this basis, we hypothesized that the recruitment of a specific reference frame could be reflected in the activation of sensorimotor versus visual brain networks. To this aim, we directly compared the brain activity associated with mental rotation of hands versus full-bodies. Mental rotation of hands recruited more strongly the supplementary motor area, premotor cortex, and secondary somatosensory cortex. Conversely, mental rotation of full-bodies determined stronger activity in temporo-occipital regions, including the functionally-localized extrastriate body area. These results support that (1) sensorimotor and visual frames of reference are used to represent the body, (2) two distinct brain networks encode local or global bodily representations, and (3) the extrastriate body area is a multimodal region involved in body processing both at the perceptual and representational level.

Altered functional connectivity of interoception in illness anxiety disorder

Interoception collects all information coming from the body and is sustained by several brain areas such as insula and cingulate cortex. Here, we used resting-state functional magnetic resonance imaging to investigate functional connectivity (FC) of networks implied in interoception in patients with Illness anxiety disorders (IADs). We observed significantly reduced FC between the left extrastriate body area (EBA) and the paracentral lobule compared to healthy controls. Moreover, the correlation analysis between behavioural questionnaires and ROI to ROI FC showed that higher levels of illness anxiety were related to hyper-connectivity between EBA and amygdala and hippocampus. Scores on a questionnaire for interoceptive awareness were significantly correlated with higher FC between right hippocampus and nucleus accumbens bilaterally, and with higher connectivity between left anterior cingulate cortex (ACC) and left orbitofrontal cortex (OFC). Last, patients showed increased interoceptive awareness, measured by Self-Awareness Questionnaire (SAQ), and reduced capability in recognizing emotions, indicating inverse correlation between interoception and emotional awareness. Taken together our results suggested that, in absence of structural and micro-structural changes, patients with IADs show functional alteration in the neural network involved in the self-body representation; such functional alteration might be the target of possible treatments.

Recognizing approaching walkers: Neural decoding of person familiarity in cortical areas responsive to faces, bodies, and biological motion

In natural viewing environments, we recognize other people as they move through the world. Behavioral studies indicate that the face, body, and gait all contribute to recognition. We examined the neural basis of person recognition using a decoding approach aimed at discriminating the patterns of neural activity elicited in response to seeing visually familiar versus unfamiliar people in motion. Participants learned 30 identities by viewing multiple videos of the people in action. Recognition was tested inside a functional magnetic resonance imaging (fMRI) scanner using 8-s videos of 60 people (30 learned and 30 novel) approaching from a distance (~13m). Full brain images were taken while participants watched the approach. These images captured neural activity at four time points (TRs) corresponding to progressively closer views of the walker. We used pattern classification techniques to examine familiarity decoding in lateralized ROIs and the combination of left and right (bilateral) regions. Results showed accurate decoding of familiarity at the farthest distance in the bilateral posterior superior temporal sulcus (bpSTS). At a closer distance, familiarity was decoded in the bilateral extrastriate body area (bEBA) and left fusiform body area (lFBA). The most robust decoding was found in the time window during which the average behavioral recognition decision was made – and when the face came into clearer view. Multiple regions, including the right occipital face area (rOFA), bOFA, bFBA, bpSTS, and broadly distributed face- and body-selective voxels in the ventral temporal cortex decoded walker familiarity in this time window. At the closest distance, the lFBA decoded familiarity. These results reveal a broad system of ventral and dorsal visual areas that support person recognition from face, body, and gait. Although the face has been the focus of most person recognition studies, these findings remind us of the evolutionary advantage of being able to differentiate the people we know from strangers at a safe distance.

Left extrastriate body area is sensitive to the meaning of symbolic gesture: evidence from fMRI repetition suppression.

Functional magnetic resonance imaging (fMRI) adaptation (a.k.a. repetition suppression) paradigm was used to test if semantic information contained in object-related (transitive) pantomimes and communicative (intransitive) gestures is represented differently in the occipito-temporal cortex. Participants watched 2.75 s back-to-back videos where the meaning of gesture was either repeated or changed. The just observed (typically second) gesture was then imitated. To maintain participants’ attention, some trials contained a single video. fMRI adaptation –signal decreases– for watching both movement categories were observed particularly in the lateral occipital cortex, including the extrastriate body area (EBA). Yet, intransitive (vs. transitive) gesture specific repetition suppression was found mainly in the left rostral EBA and caudal middle temporal gyrus- the rEBA/cMTG complex. Repetition enhancement (signal increase) was revealed in the precuneus. While the whole brain and region-of-interest analyses indicate that the precuneus is involved only in visuospatial action processing for later imitation, the common EBA repetition suppression discloses sensitivity to the meaning of symbolic gesture, namely the “semantic what” of actions. Moreover, the rEBA/cMTG suppression reveals greater selectivity for conventionalized communicative gesture. Thus, fMRI adaptation shows higher-order functions of EBA, its role in the semantic network, and indicates that its functional repertoire is wider than previously thought.

The relationship between repetition suppression and face perception.

Repetition of identical face stimuli leads to fMRI response attenuation (fMRI adaptation, fMRIa) in the core face-selective occipito-temporal visual cortical network, involving the bilateral fusiform face area (FFA) and the occipital face area (OFA). However, the functional relevance of fMRIa observed in these regions is unclear as of today. Therefore, here we aimed at investigating the relationship between fMRIa and face perception ability by measuring in the same human participants both the repetition-induced reduction of fMRI responses and identity discrimination performance outside the scanner for upright and inverted face stimuli. In the correlation analysis, the behavioral and fMRI results for the inverted faces were used as covariates to control for the individual differences in overall object perception ability and basic visual feature adaptation processes, respectively. The results revealed a significant positive correlation between the participants' identity discrimination performance and the strength of fMRIa in the core face processing network, but not in the extrastriate body area (EBA). Furthermore, we found a strong correlation of the fMRIa between OFA and FFA and also between OFA and EBA, but not between FFA and EBA. These findings suggest that there is a face-selective component of the repetition-induced reduction of fMRI responses within the core face processing network, which reflects functionally relevant adaptation processes involved in face identity perception.

Different contributions of visual and motor brain areas during liking judgments of same- and different-gender bodies

Previous neuroimaging studies have shown that body aesthetic appreciation involves the activation of both visual and motor areas, supporting a role of sensorimotor embodiment in aesthetic processing. Causative evidence, however, that neural activity in these areas is crucial for reliable aesthetic body appreciation has so far provided only for extrastriate body area (EBA), while the functional role played by premotor regions remained less clear. Here, we applied short trains of repetitive transcranial magnetic stimulation (rTMS) over bilateral dorsal premotor cortex (dPMC) and EBA during liking judgments of female and male bodies varying in weight and implied motion. We found that both dPMC and EBA are necessary for aesthetic body appreciation, but their relative contribution depends on the model's gender. While dPMC-rTMS decreased the liking judgments of same-, but not of different-gender models, EBA-rTMS increased the liking judgments of different-, but not of same-gender models. Relative contributions of motor and visual areas may reflect processing of diverse aesthetic properties, respectively implied motion vs. body form, and/or greater sensorimotor embodiment of same- vs. different-gender bodies. Results suggest that aesthetic body processing is subserved by a network of motor and visual areas, whose relative contribution may depend on the specific stimulus and task.

The role of body image and self-perception in anorexianervosa: the neuroimaging perspective.

Anorexia nervosa is a severe psychiatric illness characterized by intense fear of gaining weight, relentless pursuit of thinness, deep concerns about food and a pervasive disturbance of body image. Functional magnetic resonance imaging tries to shed light on the neurobiological underpinnings of anorexia nervosa. This review aims to evaluate the empirical neuroimaging literature about self-perception in anorexia nervosa. This narrative review summarizes a number of task-based and resting-state functional magnetic resonance imaging studies in anorexia nervosa about body image and self-perception. The articles listed in references were searched using electronic databases (PubMed and Google Scholar) from 1990 to February 2016 using specific key words. All studies were reviewed with regard to their quality and eligibility for the review. Differences in brain activity were observed using body image perception and body size estimation tasks showing significant modifications in activity of specific brain areas (extrastriate body area, fusiform body area, inferior parietal lobule). Recent studies highlighted the role of emotions and self-perception in anorexia nervosa and their neural substrate involving resting-state networks and particularly frontal and posterior midline cortical structures within default mode network and insula. These findings open new horizons to understand the neural substrate of anorexia nervosa.

Human-Object Interactions Are More than the Sum of Their Parts.

Understanding human-object interactions is critical for extracting meaning from everyday visual scenes and requires integrating complex relationships between human pose and object identity into a new percept. To understand how the brain builds these representations, we conducted 2 fMRI experiments in which subjects viewed humans interacting with objects, noninteracting human-object pairs, and isolated humans and objects. A number of visual regions process features of human-object interactions, including object identity information in the lateral occipital complex (LOC) and parahippocampal place area (PPA), and human pose information in the extrastriate body area (EBA) and posterior superior temporal sulcus (pSTS). Representations of human-object interactions in some regions, such as the posterior PPA (retinotopic maps PHC1 and PHC2) are well predicted by a simple linear combination of the response to object and pose information. Other regions, however, especially pSTS, exhibit representations for human-object interaction categories that are not predicted by their individual components, indicating that they encode human-object interactions as more than the sum of their parts. These results reveal the distributed networks underlying the emergent representation of human-object interactions necessary for social perception.

Lateral occipitotemporal cortex (LOTC) activity is greatest while viewing dance compared to visualization and movement: learning and expertise effects.

The lateral occipitotemporal cortex (LOTC) is comprised of subregions selectively activated by images of human bodies (extrastriate body area, EBA), objects (lateral occipital complex, LO), and motion (MT+). However, their role in motor imagery and movement processing is unclear, as are the influences of learning and expertise on its recruitment. The purpose of our study was to examine putative changes in LOTC activation during action processing following motor learning of novel choreography in professional ballet dancers. Subjects were scanned with functional magnetic resonance imaging up to four times over 34weeks and performed four tasks: viewing and visualizing a newly learned ballet dance, visualizing a dance that was not being learned, and movement of the foot. EBA, LO, and MT+ were activated most while viewing dance compared to visualization and movement. Significant increases in activation were observed over time in left LO only during visualization of the unlearned dance, and all subregions were activated bilaterally during the viewing task after 34weeks of performance, suggesting learning-induced plasticity. Finally, we provide novel evidence for modulation of EBA with dance experience during the motor task, with significant activation elicited in a comparison group of novice dancers only. These results provide a composite of LOTC activation during action processing of newly learned ballet choreography and movement of the foot. The role of these areas is confirmed as primarily subserving observation of complex sequences of whole-body movement, with new evidence for modification by experience and over the course of real world ballet learning.

The Extrastriate Body Area Computes Desired Goal States during Action Planning.

How do object perception and action interact at a neural level? Here we test the hypothesis that perceptual features, processed by the ventral visuoperceptual stream, are used as priors by the dorsal visuomotor stream to specify goal-directed grasping actions. We present three main findings, which were obtained by combining time-resolved transcranial magnetic stimulation and kinematic tracking of grasp-and-rotate object manipulations, in a group of healthy human participants (N = 22). First, the extrastriate body area (EBA), in the ventral stream, provides an initial structure to motor plans, based on current and desired states of a grasped object and of the grasping hand. Second, the contributions of EBA are earlier in time than those of a caudal intraparietal region known to specify the action plan. Third, the contributions of EBA are particularly important when desired and current object configurations differ, and multiple courses of actions are possible. These findings specify the temporal and functional characteristics for a mechanism that integrates perceptual processing with motor planning.

Discussion paper: Is body processing ‘special’? Insights from the extrastriate body area and face processing literature

Discussion paper: Is body processing ‘special’? Insights from the extrastriate body area and face processing literature

Categorical discrimination of human body parts by magnetoencephalography.

Humans recognize body parts in categories. Previous studies have shown that responses in the fusiform body area (FBA) and extrastriate body area (EBA) are evoked by the perception of the human body, when presented either as whole or as isolated parts. These responses occur approximately 190 ms after body images are visualized. The extent to which body-sensitive responses show specificity for different body part categories remains to be largely clarified. We used a decoding method to quantify neural responses associated with the perception of different categories of body parts. Nine subjects underwent measurements of their brain activities by magnetoencephalography (MEG) while viewing 14 images of feet, hands, mouths, and objects. We decoded categories of the presented images from the MEG signals using a support vector machine (SVM) and calculated their accuracy by 10-fold cross-validation. For each subject, a response that appeared to be a body-sensitive response was observed and the MEG signals corresponding to the three types of body categories were classified based on the signals in the occipitotemporal cortex. The accuracy in decoding body-part categories (with a peak at approximately 48%) was above chance (33.3%) and significantly higher than that for random categories. According to the time course and location, the responses are suggested to be body-sensitive and to include information regarding the body-part category. Finally, this non-invasive method can decode category information of a visual object with high temporal and spatial resolution and this result may have a significant impact in the field of brain–machine interface research.

Neural correlates of different self domains.

The neural substrates of states devoted to processing self-related information ("self-related states") remain not fully elucidated. Besides the complexity of the problem, there is evidence suggesting that self-related states vary according to the information domain being considered. Here, we investigated brain correlates for self-related states concerning historical aspects of one's life (autobiographical self), and one's ongoing body status (core self). We focused on memory-related regions, body-related regions, CMSs (cortical midline structures), and ICs (insular cortices). This was a block-design fMRI study contrasting brain activity for core self (interoception and exteroception) and autobiographical self (personality traits and biographic facts) information domains. It involved 19 participants, who answered questions about each domain (four conditions). All conditions appeared to engage the regions of interest. Nonetheless, autobiographical self compared with core self showed greater activity in memory-related regions (e.g., hippocampus), MPFC (medial prefrontal cortex), superior PMC (posteromedial cortex), and anterior ICs. Core self compared with autobiographical self was associated with greater activity in body-related regions (e.g., somatosensory cortices, and EBA [extrastriate body area]), superior PMC, and posterior ICs. In addition, (1) facts compared with traits showed greater activity in body-related regions, memory-related regions, MPFC, and PMC; (2) traits compared with facts were associated with greater activity in the posterior part of the anterior cingulate cortex; (3) interoception compared with exteroception was associated with greater activity in body-related regions (e.g. postcentral gyrus), memory-related regions, MPFC, inferior PMC and ICs; (4) exteroception compared with interoception showed greater activity in some body-related regions (e.g., premotor cortices and EBA) and superior PMC. The results support the notion that the neural correlates of self-related states depend on the information domain. Those states seem distinguishable in terms of activity in memory-related and body-related regions, and activity in regions that have been associated with self processes (CMSs and the ICs).

Dissociable routes for personal and interpersonal visual enhancement of touch

Seeing a hand can enhance tactile acuity on the hand, even when tactile stimulation is not visible. This visual enhancement of touch (VET) occurs both when participants see their own hand (personal VET), and when they see another person's hand (interpersonal VET). Interpersonal VET occurs irrespective of where the viewed hand appears, while personal VET is eliminated when visual and proprioceptive signals about the location of one's own hand are incongruent. This suggests that the neural mechanisms for VET may differ according to ownership of the seen hand. We used continuous theta-burst transcranial magnetic stimulation (TMS) to disrupt either the human ventral intraparietal area (hVIP), which integrates tactile, proprioceptive, and visual information about one's own body, or the extrastriate body area (EBA), which processes visual body information irrespective of ownership. Participants then judged the orientation of tactile gratings applied to their hand while viewing images of their own hand, another person's hand, or a non-body object on a screen placed over their actual hand. Disrupting the hVIP attenuated personal VET but did not affect interpersonal VET, suggesting the hVIP is only involved in VET when one's own hand is seen. Disrupting the EBA reduced both personal and interpersonal VET, suggesting it is common to both routes.

Personality traits predict brain activation and connectivity when witnessing a violent conflict.

As observers we excel in decoding the emotional signals telling us that a social interaction is turning violent. The neural substrate and its modulation by personality traits remain ill understood. We performed an fMRI experiment in which participants watched videos displaying a violent conflict between two people. Observers’ attention was directed to either the aggressor or the victim. Focusing on the aggressor (vs. focusing on the victim) activated the superior temporal sulcus (STS), extra-striate body area (EBA), occipital poles and centro-medial amygdala (CMA). Stronger instantaneous connectivity occurred between these and the EBA, insula, and the red nucleus. When focusing on the victim, basolateral amygdala (BLA) activation was related to trait empathy and showed increased connectivity with the insula and red nucleus. STS activation was associated with trait aggression and increased connectivity with the hypothalamus. The findings reveal that focusing on the aggressor of a violent conflict triggers more activation in categorical (EBA) and emotion (CMA, STS) areas. This is associated with increased instantaneous connectivity among emotion areas (CMA-insula) and between categorical and emotion (EBA-STS) areas. When the focus is on the victim, personality traits (aggression/empathy) modulate activity in emotion areas (respectively STS and postcentral gyrus/ BLA), along with connectivity in the emotional diencephalon (hypothalamus) and early visual areas (occipital pole).

A causal role for inferior parietal lobule in emotion body perception

Recent investigations of emotion body perception have established that perceiving fearful body expressions critically triggers activity in dorsal stream structures related to action preparation. However, the causal contributions of these areas remain unclear. In the current experiment, we addressed this issue using online transcranial magnetic stimulation (TMS) of the inferior parietal lobule (IPL) in the dorsal stream and visual areas (extrastriate body area – EBA in the ventral stream and early visual cortex – EVC). Participants performed a delayed-match-to-sample task requiring detection of a change in posture of body expressions that were either neutral or fearful. Results revealed a significant interaction between the stimulation site and the emotional valence of stimuli, indicating that processing of emotional versus neutral bodies is affected differentially by stimulation of different central areas in body processing. IPL stimulation specifically enhanced fearful body processing. These findings relate emotion processing to separate processing streams, and moreover provide the first evidence that IPL plays a causal role in processing of fearful bodies.

Brain activity correlates with emotional perception induced by dynamic avatars

An accurate judgment of the emotional state of others is a prerequisite for successful social interaction and hence survival. Thus, it is not surprising that we are highly skilled at recognizing the emotions of others. Here we aimed to examine the neuronal correlates of emotion recognition from gait. To this end we created highly controlled dynamic body-movement stimuli based on real human motion-capture data (Roether et al., 2009). These animated avatars displayed gait in four emotional (happy, angry, fearful, and sad) and speed-matched neutral styles. For each emotional gait and its equivalent neutral gait, avatars were displayed at five morphing levels between the two. Subjects underwent fMRI scanning while classifying the emotions and the emotional intensity levels expressed by the avatars.Our results revealed robust brain selectivity to emotional compared to neutral gait stimuli in brain regions which are involved in emotion and biological motion processing, such as the extrastriate body area (EBA), fusiform body area (FBA), superior temporal sulcus (STS), and the amygdala (AMG). Brain activity in the amygdala reflected emotional awareness: for visually identical stimuli it showed amplified stronger response when the stimulus was perceived as emotional. Notably, in avatars gradually morphed along an emotional expression axis there was a parametric correlation between amygdala activity and emotional intensity.This study extends the mapping of emotional decoding in the human brain to the domain of highly controlled dynamic biological motion. Our results highlight an extensive level of brain processing of emotional information related to body language, which relies mostly on body kinematics.

Body selectivity in occipitotemporal cortex: Causal evidence

Perception of others' bodies provides information that is useful for a number of important social-cognitive processes. Evidence from neuroimaging methods has identified focal cortical regions that are highly selective for perceiving bodies and body parts, including the extrastriate body area (EBA) and fusiform body area (FBA). Our understanding of the functional properties of these regions, and their causal contributions to behavior, has benefitted from the study of neuropsychological patients and particularly from investigations using transcranial magnetic stimulation (TMS). We review this evidence, focusing on TMS studies that are revealing of how (and when) activity in EBA contributes to detecting people in natural scenes; to resolving their body shape, movements, actions, individual parts, and identities; and to guiding goal-directed behavior. These findings are considered in reference to a framework for body perception in which the patterns of neural activity in EBA and FBA jointly serve to make explicit the elements of the visual scene that correspond to the body and its parts. These representations are modulated by other sources of information such as prior knowledge, and are shared with wider brain networks involved in many aspects of social cognition.

Selecting category specific visual information: Top-down and bottom-up control of object based attention.

The ability to select, within the complexity of sensory input, the information most relevant for our purposes is influenced by both internal settings (i.e., top-down control) and salient features of external stimuli (i.e., bottom-up control). We here investigated using fMRI the neural underpinning of the interaction of top-down and bottom-up processes, as well as their effects on extrastriate areas processing visual stimuli in a category-selective fashion. We presented photos of bodies or buildings embedded into frequency-matched visual noise to the subjects. Stimulus saliency changed gradually due to an altered degree to which photos stood-out in relation to the surrounding noise (hence generating stronger bottom-up control signals). Top-down settings were manipulated via instruction: participants were asked to attend one stimulus category (i.e., "is there a body?" or "is there a building?"). Highly salient stimuli that were inconsistent with participants' attentional top-down template activated the inferior frontal junction and dorsal parietal regions bilaterally. Stimuli consistent with participants' current attentional set additionally activated insular cortex and the parietal operculum. Furthermore, the extrastriate body area (EBA) exhibited increased neural activity when attention was directed to bodies. However, the latter effect was found only when stimuli were presented at intermediate saliency levels, thus suggesting a top-down modulation of this region only in the presence of weak bottom-up signals. Taken together, our results highlight the role of the inferior frontal junction and posterior parietal regions in integrating bottom-up and top-down attentional control signals.