Occipital Face Area Research Articles

TMS to the “occipital face area” affects recognition but not categorization of faces

The human cortical system for face perception is comprised of a network of connected regions including the middle fusiform gyrus (“fusiform face area” or FFA), the inferior occipital cortex (“occipital face area” or OFA), and the superior temporal sulcus. The traditional hierarchical feedforward model of visual processing suggests information flows from early visual cortex to the OFA for initial face feature analysis to higher order regions including the FFA for identity recognition. However, patient data suggest an alternative model. Patients with acquired prosopagnosia, an inability to visually recognize faces, have been documented with lesions to the OFA but who nevertheless show face-selective activation in the FFA. Moreover, their ability to categorize faces remains intact. This suggests that the FFA is not solely responsible for face recognition and the network is not strictly hierarchical, but may be organized in a reverse hierarchical fashion. We used transcranial magnetic stimulation (TMS) to temporarily disrupt processing in the OFA in neurologically-intact individuals and found participants’ ability to categorize intact versus scrambled faces was unaffected, however face identity discrimination was significantly impaired. This suggests that face categorization but not recognition can occur without the “earlier” OFA being online and indicates that “lower level” face category processing may be assumed by other intact face network regions such as the FFA. These results are consistent with the patient data and support a non-hierarchical, global-to-local model with re-entrant connections between the OFA and other face processing areas.

Faciotopy - a face-feature map with face-like topology in the occipital face area

Faciotopy - a face-feature map with face-like topology in the occipital face area

Explicating the face perception network with white matter connectivity.

A network of multiple brain regions is recruited in face perception. Our understanding of the functional properties of this network can be facilitated by explicating the structural white matter connections that exist between its functional nodes. We accomplished this using functional MRI (fMRI) in combination with fiber tractography on high angular resolution diffusion weighted imaging data. We identified the three nodes of the core face network: the “occipital face area” (OFA), the “fusiform face area” (mid-fusiform gyrus or mFus), and the superior temporal sulcus (STS). Additionally, a region of the anterior temporal lobe (aIT), implicated as being important for face perception was identified. Our data suggest that we can further divide the OFA into multiple anatomically distinct clusters – a partitioning consistent with several recent neuroimaging results. More generally, structural white matter connectivity within this network revealed: 1) Connectivity between aIT and mFus, and between aIT and occipital regions, consistent with studies implicating this posterior to anterior pathway as critical to normal face processing; 2) Strong connectivity between mFus and each of the occipital face-selective regions, suggesting that these three areas may subserve different functional roles; 3) Almost no connectivity between STS and mFus, or between STS and the other face-selective regions. Overall, our findings suggest a re-evaluation of the “core” face network with respect to what functional areas are or are not included in this network.

Decoding face categories in diagnostic subregions of primary visual cortex

Higher visual areas in the occipitotemporal cortex contain discrete regions for face processing, but it remains unclear if V1 is modulated by top-down influences during face discrimination, and if this is widespread throughout V1 or localized to retinotopic regions processing task-relevant facial features. Employing functional magnetic resonance imaging (fMRI), we mapped the cortical representation of two feature locations that modulate higher visual areas during categorical judgements – the eyes and mouth. Subjects were presented with happy and fearful faces, and we measured the fMRI signal of V1 regions processing the eyes and mouth whilst subjects engaged in gender and expression categorization tasks. In a univariate analysis, we used a region-of-interest-based general linear model approach to reveal changes in activation within these regions as a function of task. We then trained a linear pattern classifier to classify facial expression or gender on the basis of V1 data from ‘eye’ and ‘mouth’ regions, and from the remaining non-diagnostic V1 region. Using multivariate techniques, we show that V1 activity discriminates face categories both in local ‘diagnostic’ and widespread ‘non-diagnostic’ cortical subregions. This indicates that V1 might receive the processed outcome of complex facial feature analysis from other cortical (i.e. fusiform face area, occipital face area) or subcortical areas (amygdala).

Development of a method to present wide-view visual stimuli in MRI for peripheral visual studies

We developed a novel wide-view visual presentation system for fMRI studies. Computer-generated images were projected onto a hemispheric, translucent screen inside the MRI bore and were then back-projected onto a 52mm diameter screen. To achieve a wide field view, a spherical screen with a curvature radius of 30mm was placed 30mm away from the subjects’ eyes. The subjects wore contact lenses that enabled them to focus on the screen, and the resulting visual field reached 120°. To evaluate the clarity and quality of the MRI images, a signal-to-noise ratio valuation experiment was performed. In addition, we successfully applied this visual presentation system to studies of visual retinotopic mapping and object perception neural function in the peripheral visual field. Our study demonstrated that the system is compatible with the MRI environment. Based on the wide-field mapping results, this system was more effective at mapping a checkerboard stimuli in V1–V3 from the central to peripheral visual fields. In higher-level visual areas, we successfully located several classical category-selective areas, including the face-selective area (FFA), occipital face area (OFA), house-selective area (PPA), transverse occipital sulcus (TOS), lateral occipital complex (LOC) and posterior fusiform area (pFs). In these areas, we found that the response amplitudes exhibited different decreasing trends with increasing eccentricity. In conclusion, we developed a simple, effective method for presenting wide-view visual stimuli within the MRI environment that can be applied to many kinds of fMRI studies of peripheral vision.

Neural Correlates of Face Detection

Although face detection likely played an essential adaptive role in our evolutionary past and in contemporary social interactions, there have been few rigorous studies investigating its neural correlates. MJH, a prosopagnosic with bilateral lesions to the ventral temporal-occipital cortices encompassing the posterior face areas (fusiform and occipital face areas), expresses no subjective difficulty in face detection, suggesting that these posterior face areas do not mediate face detection exclusively. Despite his normal contrast sensitivity and visual acuity in foveal vision, the present study nevertheless revealed significant face detection deficits in MJH. Compared with controls, MJH showed a lower tolerance to noise in the phase spectrum for faces (vs. cars), reflected in his higher detection threshold for faces. MJH's lesions in bilateral occipito-temporal cortices thus appear to have produced a deficit not only in face individuation, but also in face detection.

The Occipital Place Area Is Causally and Selectively Involved in Scene Perception

Functional magnetic resonance imaging has revealed a set of regions selectively engaged in visual scene processing: the parahippocampal place area (PPA), the retrosplenial complex (RSC), and a region around the transverse occipital sulcus (previously known as "TOS"), here renamed the "occipital place area" (OPA). Are these regions not only preferentially activated by, but also causally involved in scene perception? Although past neuropsychological data imply a causal role in scene processing for PPA and RSC, no such evidence exists for OPA. Thus, to test the causal role of OPA in human adults, we delivered transcranial magnetic stimulation (TMS) to the right OPA (rOPA) or the nearby face-selective right occipital face area (rOFA) while participants performed fine-grained perceptual discrimination tasks on scenes or faces. TMS over rOPA impaired discrimination of scenes but not faces, while TMS over rOFA impaired discrimination of faces but not scenes. In a second experiment, we delivered TMS to rOPA, or the object-selective right lateral occipital complex (rLOC), while participants performed categorization tasks involving scenes and objects. TMS over rOPA impaired categorization accuracy of scenes but not objects, while TMS over rLOC impaired categorization accuracy of objects but not scenes. These findings provide the first evidence that OPA is causally involved in scene processing, and further show that this causal role is selective for scene perception. Our findings illuminate the functional architecture of the scene perception system, and also argue against the "distributed coding" view in which each category-selective region participates in the representation of all objects.

Implicit Race Bias Decreases the Similarity of Neural Representations of Black and White Faces

Implicit race bias has been shown to affect decisions and behaviors. It may also change perceptual experience by increasing perceived differences between social groups. We investigated how this phenomenon may be expressed at the neural level by testing whether the distributed blood-oxygenation-level-dependent (BOLD) patterns representing Black and White faces are more dissimilar in participants with higher implicit race bias. We used multivoxel pattern analysis to predict the race of faces participants were viewing. We successfully predicted the race of the faces on the basis of BOLD activation patterns in early occipital visual cortex, occipital face area, and fusiform face area (FFA). Whereas BOLD activation patterns in early visual regions, likely reflecting different perceptual features, allowed successful prediction for all participants, successful prediction on the basis of BOLD activation patterns in FFA, a high-level face-processing region, was restricted to participants with high pro-White bias. These findings suggest that stronger implicit pro-White bias decreases the similarity of neural representations of Black and White faces.

An MEG neuroimaging study on the developmental changes of face processing in pre-school aged children

Event Abstract Back to Event An MEG neuroimaging study on the developmental changes of face processing in pre-school aged children Wei He1*, Jon Brock1 and Blake W. Johnson1 1 Macquarie University, Department of Cognitive Science, Australia Introduction: An important research question in developmental neuroscience concerns how the underlying neural mechanisms of face perception develop in early childhood. Magnetoencephalography (MEG) holds great promise for examining this question, because it measures a well-known face sensitive brain response, termed the M170, from young children in an entirely passive manner. The M170 and its electrical equivalent N170 discriminate faces from other non-face objects based on larger amplitudes to face stimuli around 170 ms from stimulus onsets in adults (Rossion & Jacques, 2011). The current study measured the M170 from pre-schoolers using a unique child custom-sized magnetoencephalography (MEG) system (Johnson et al., 2010). Furthermore, we evaluated the effective connectivity of cortical networks underlying the M170 using dynamic causal modelling (DCM); and compared the connectivity patterns in children to those of adults. Methods: Brain responses were measured from 15 young children (aged 4.46 ± 0.93 years) and 15 adults (aged 27.6 ± 6.46 years) during flashing pictures of faces, cars, as well as their phase-scrambled counterparts (Kuefner et al, 2010). There were 64 channels in the child MEG system and 160 channels in the adult MEG system. Surface waveforms were analysed by averaging the strongest response across occipital and temporal sensors separately on each hemisphere based on the time window determined by visual checking on individual data. DCM analyses assumed that the M170 was mediated by sources in the occipital face area (OFA), fusiform face area (FFA), and superior temporal sulcus (STS). Using Bayesian model selection, we evaluated three models of the interconnections between these regions based on previous studies using DCM (Fairhall & Ishai, 2007; Chen, et al., 2009; David et al., 2006). Results: In the surface waveform analysis, the child M170 was slower than the adult M170, but it already showed an adult function pattern (i.e., faster and stronger to faces than cars). Analyses of DCM models showed that the first and simplest model with only forward intra-hemispheric connections from OFA to FFA and STS had the largest model evidence in adults, while in children, the second model with extra inter-hemispheric connections between OFA and contralateral FFA showed the largest model evidence. Conclusions: Early face sensitive brain responses indexed by the M170 are present and functioning in the adult pattern in children as young as 4 years old. However, the functional neural mechanism underlying the M170 undergoes further development and fine-tuning before it reaches adult capacities. Acknowledgements This work was supported by Australian Research Council Linkage Infrastructure Equipment and Facilities Grant LEO668421, Australian Research Council Linkage Project Grant LP0669471, and the Australian Research Council Centre of Excellence for Cognition and its Disorders (CE110001021), http://www.ccd.edu.au. The authors thank Romina Polermo and Douglas Cheyne for helpful comments during the design of the experiment and Bruno Russion for providing the picture stimuli.

The implicit processing of categorical and dimensional strategies: an fMRI study of facial emotion perception

Our understanding of facial emotion perception has been dominated by two seemingly opposing theories: the categorical and dimensional theories. However, we have recently demonstrated that hybrid processing involving both categorical and dimensional perception can be induced in an implicit manner (Fujimura etal., 2012). The underlying neural mechanisms of this hybrid processing remain unknown. In this study, we tested the hypothesis that separate neural loci might intrinsically encode categorical and dimensional processing functions that serve as a basis for hybrid processing. We used functional magnetic resonance imaging to measure neural correlates while subjects passively viewed emotional faces and performed tasks that were unrelated to facial emotion processing. Activity in the right fusiform face area (FFA) increased in response to psychologically obvious emotions and decreased in response to ambiguous expressions, demonstrating the role of the FFA in categorical processing. The amygdala, insula and medial prefrontal cortex exhibited evidence of dimensional (linear) processing that correlated with physical changes in the emotional face stimuli. The occipital face area and superior temporal sulcus did not respond to these changes in the presented stimuli. Our results indicated that distinct neural loci process the physical and psychological aspects of facial emotion perception in a region-specific and implicit manner.

Local Discriminability Determines the Strength of Holistic Processing for Faces in the Fusiform Face Area

Recent evidence suggests that the Fusiform Face Area (FFA) is not exclusively dedicated to the interactive processing of face features, but also contains neurons sensitive to local features. This suggests the existence of both interactive and local processing modes, consistent with recent behavioral findings that the strength of interactive feature processing (IFP) engages most strongly when similar features need to be disambiguated. Here we address whether the engagement of the FFA into interactive versus featural representational modes is governed by local feature discriminability. We scanned human participants while they matched target features within face pairs, independently of the context of distracter features. IFP was operationalized as the failure to match the target without being distracted by distracter features. Picture-plane inversion was used to disrupt IFP while preserving input properties. We found that FFA activation was comparably strong, irrespective of whether similar target features were embedded in dissimilar contexts(i.e., inducing robust IFP) or dissimilar target features were embedded in the same context (i.e., engaging local processing). Second, inversion decreased FFA activation to faces most robustly when similar target features were embedded in dissimilar contexts, indicating that FFA engages into IFP mainly when features cannot be disambiguated at a local level. Third, by means of Spearman rank correlation tests, we show that the local processing of feature differences in the FFA is supported to a large extent by the Occipital Face Area, the Lateral Occipital Complex, and early visual cortex, suggesting that these regions encode the local aspects of face information. The present findings confirm the co-existence of holistic and featural representations in the FFA. Furthermore, they establish FFA as the main contributor to the featural/holistic representational mode switches determined by local discriminability.

Residual fMRI sensitivity for identity changes in acquired prosopagnosia

While a network of cortical regions contribute to face processing, the lesions in acquired prosopagnosia are highly variable, and likely result in different combinations of spared and affected regions of this network. To assess the residual functional sensitivities of spared regions in prosopagnosia, we designed a rapid event-related functional magnetic resonance imaging (fMRI) experiment that included pairs of faces with same or different identities and same or different expressions. By measuring the release from adaptation to these facial changes we determined the residual sensitivity of face-selective regions-of-interest. We tested three patients with acquired prosopagnosia, and all three of these patients demonstrated residual sensitivity for facial identity changes in surviving fusiform and occipital face areas of either the right or left hemisphere, but not in the right posterior superior temporal sulcus. The patients also showed some residual capabilities for facial discrimination with normal performance on the Benton Facial Recognition Test, but impaired performance on more complex tasks of facial discrimination. We conclude that fMRI can demonstrate residual processing of facial identity in acquired prosopagnosia, that this adaptation can occur in the same structures that show similar processing in healthy subjects, and further, that this adaptation may be related to behavioral indices of face perception.

Two Critical and Functionally Distinct Stages of Face and Body Perception

Cortical regions that respond preferentially to particular object categories, such as faces and bodies, are essential for visual perception of these object categories. However, precisely when these regions play a causal role in recognition of their preferred categories is unclear. Here we addressed this question using transcranial magnetic stimulation (TMS). Across a series of experiments, TMS was delivered over the functionally localized right occipital face area (rOFA) or right extrastriate body area (rEBA) at different latencies, up to 150 ms, after stimulus onset while adult human participants performed delayed match-to-sample tasks on face and body stimuli. Results showed that TMS disrupted task performance during two temporally distinct time periods after stimulus onset, the first at 40/50 ms and the second at 100/110 ms. These two time periods exhibited functionally distinct patterns of impairment: TMS delivered during the early time period (at 40/50 ms) disrupted task performance for both preferred (faces at rOFA and bodies at rEBA) and nonpreferred (bodies at rOFA and faces at rEBA) categories. In contrast, TMS delivered during the later time period (at 100/110 ms) disrupted task performance for the preferred category only of each area (faces at rOFA and bodies at rEBA). These results indicate that category-selective cortical regions are critical for two functionally distinct stages of visual object recognition: an early, presumably preparatory stage that is not category selective occurring almost immediately after stimulus onset, followed by a later stage of category-specific perceptual processing.

S2-4: On the (in)Dependence of Visual Cues and Cortical Regions for Judging Trustworthiness and Sex from Faces: TMS, Behavioural, and Human Lesion Evidence

Judging sex from faces relies on cues related to facial morphology and spatial relations between features, whereas judging trustworthiness relies on both structural and expressive cues that signal affective valence. I will present evidence of both asymmetric and symmetric dependence between the processing of a face's sex and trustworthiness. In one study, we used event-related, fMRI-guided repetitive transcranial magnetic stimulation (rTMS) to test whether 2 face-selective regions–right occipital face area (rOFA) and posterior superior temporal sulcus (pSTS)–have functionally dissociable, critical roles in sex and trustworthiness judgments. Sex judgments were disrupted when rTMS was delivered over right OFA (relative to sham stimulation) but not when it was delivered over right or left pSTS, whereas trustworthiness judgments were disrupted when rTMS was delivered over right or left pSTS but not rOFA. Nonetheless, analysis of the reaction time distributions revealed a possible critical role also for rOFA in trustworthiness judgments, limited to faces with longer RTs, perhaps reflecting the later, ancillary use of structural cues related to the sex of the face. New evidence from an individual (DF) with lesions encompassing bilateral occipital cortex indicates that significantly above-chance discrimination of the sex and trustworthiness of faces is possible in the absence of OFA–but only for untrustworthy males and trustworthy females, consistent with the RT data of healthy participants in the TMS study. Further evidence of restricted dependence between perceptions of trustworthiness and sex is provided by a RT-interference experiment with neurologically healthy volunteers using the Garner paradigm.

Intra- and interhemispheric connectivity between face-selective regions in the human brain

Neuroimaging studies have revealed a number of regions in the human brain that respond to faces. However, the way these regions interact is a matter of current debate. The aim of this study was to use functional MRI to define face-selective regions in the human brain and then determine how these regions interact in a large population of subjects (n = 72). We found consistent face selectivity in the core face regions of the occipital and temporal lobes: the fusiform face area (FFA), occipital face area (OFA), and superior temporal sulcus (STS). Face selectivity extended into the intraparietal sulcus (IPS), precuneus (PCu), superior colliculus (SC), amygdala (AMG), and inferior frontal gyrus (IFG). We found evidence for significant functional connectivity between the core face-selective regions, particularly between the OFA and FFA. However, we found that the covariation in activity between corresponding face regions in different hemispheres (e.g., right and left FFA) was higher than between different face regions in the same hemisphere (e.g., right OFA and right FFA). Although functional connectivity was evident between regions in the core and extended network, there were significant differences in the magnitude of the connectivity between regions. Activity in the OFA and FFA were most correlated with the IPS, PCu, and SC. In contrast, activity in the STS was most correlated with the AMG and IFG. Correlations between the extended regions suggest strong functional connectivity between the IPS, PCu, and SC. In contrast, the IFG was only correlated with the AMG. This study reveals that interhemispheric as well as intrahemispheric connections play an important role in face perception.

Prevalence of Selectivity for Mirror-Symmetric Views of Faces in the Ventral and Dorsal Visual Pathways

Although the ability to recognize faces and objects from a variety of viewpoints is crucial to our everyday behavior, the underlying cortical mechanisms are not well understood. Recently, neurons in a face-selective region of the monkey temporal cortex were reported to be selective for mirror-symmetric viewing angles of faces as they were rotated in depth (Freiwald and Tsao, 2010). This property has been suggested to constitute a key computational step in achieving full view-invariance. Here, we measured functional magnetic resonance imaging activity in nine observers as they viewed upright or inverted faces presented at five different angles (-60, -30, 0, 30, and 60°). Using multivariate pattern analysis, we show that sensitivity to viewpoint mirror symmetry is widespread in the human visual system. The effect was observed in a large band of higher order visual areas, including the occipital face area, fusiform face area, lateral occipital cortex, mid fusiform, parahippocampal place area, and extending superiorly to encompass dorsal regions V3A/B and the posterior intraparietal sulcus. In contrast, early retinotopic regions V1-hV4 failed to exhibit sensitivity to viewpoint symmetry, as their responses could be largely explained by a computational model of low-level visual similarity. Our findings suggest that selectivity for mirror-symmetric viewing angles may constitute an intermediate-level processing step shared across multiple higher order areas of the ventral and dorsal streams, setting the stage for complete viewpoint-invariant representations at subsequent levels of visual processing.

The changing landscape of functional brain networks for face processing in typical development.

Greater expertise for faces in adults than in children may be achieved by a dynamic interplay of functional segregation and integration of brain regions throughout development. The present study examined developmental changes in face network functional connectivity in children (5-12 years) and adults (18-43 years) during face-viewing using a graph-theory approach. A face-specific developmental change involved connectivity of the right occipital face area. During childhood, this node increased in strength and within-module clustering based on positive connectivity. These changes reflect an important role of the ROFA in segregation of function during childhood. In addition, strength and diversity of connections within a module that included primary visual areas (left and right calcarine) and limbic regions (left hippocampus and right inferior orbitofrontal cortex) increased from childhood to adulthood, reflecting increased visuo-limbic integration. This integration was pronounced for faces but also emerged for natural objects. Taken together, the primary face-specific developmental changes involved segregation of a posterior visual module during childhood, possibly implicated in early stage perceptual face processing, and greater integration of visuo-limbic connections from childhood to adulthood, which may reflect processing related to development of perceptual expertise for individuation of faces and other visually homogenous categories.

Gender-selective neural populations within the occipital and fusiform face-areas: Evidence from rapid event-related fMRI

Gender-selective neural populations within the occipital and fusiform face-areas: Evidence from rapid event-related fMRI

Impaired Processing of 3D Motion-Defined Faces in Mild Cognitive Impairment and Healthy Aging: An fMRI Study

Mild cognitive impairment (MCI), which shows high risk for conversion to Alzheimer's disease (AD), is accompanied by progressive visual deteriorations that so far are poorly understood. Here, we compared dorsal and ventral visual stream functional magnetic resonance imaging (fMRI) activity among amnestic MCI, healthy elderly, and young participants during structure-from-motion (SFM) face categorization performance. Task performance varied with stimulus depth and duration levels and differences among groups were highly correlated with face-related fMRI activation patterns. Young participants showed larger activation to faces than scrambled faces (face sensitivity) in the right fusiform face area (FFA) and right occipital face area (OFA) whereas in elderly, this difference was reduced. Surprisingly, in MCI, scrambled faces elicited larger activation in right FFA/OFA than faces. The latter observation may be related to the additional finding of elevated depth sensitivity in left FFA/OFA of MCI, suggesting that an increased representation of low-level stimulus aspects may impair face perception in MCI. Discriminant function analysis using face and depth sensitivity indices in FFA/OFA classified MCI and healthy elderly with 88.2% accuracy, marking a fundamental distinction between groups. Potentially related findings include altered activation patterns in dorsal-ventral stream integration regions and attention-related networks of MCI patients. Our results highlight aberrant visual and additional potentially compensatory processes that identify dispositions of (preclinical) AD.

Focal electrical intracerebral stimulation of a face-sensitive area causes transient prosopagnosia

Face perception is subtended by a large set of areas in the human ventral occipito-temporal cortex. However, the role of these areas and their importance for face recognition remain largely unclear. Here we report a case of transient selective impairment in face recognition (prosopagnosia) induced by focal electrical intracerebral stimulation of the right inferior occipital gyrus. This area presents with typical face-sensitivity as evidenced by functional neuroimaging right occipital face area (OFA). A face-sensitive intracerebral N170 was also recorded in this area, supporting its contribution as a source of the well-known N170 component typically recorded on the scalp. Altogether, these observations indicate that face recognition can be selectively impaired by local disruption of a single face-sensitive area of the network subtending this function, the right OFA.