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Face-related activation found in the voxelwise analysis. Activation maps indicate regions where the response was higher for: A) unfamiliar faces than houses. B)acquaintance faces than unfamiliar faces. C) newly-learned faces than unfamiliar faces. These activations are shown on an inflated brain, depicting voxels surviving p,.005 (uncorrected). doi:10.1371/journal.pone.0076100.g001

Face-related activation found in the voxelwise analysis. Activation maps indicate regions where the response was higher for: A) unfamiliar faces than houses. B)acquaintance faces than unfamiliar faces. C) newly-learned faces than unfamiliar faces. These activations are shown on an inflated brain, depicting voxels surviving p,.005 (uncorrected). doi:10.1371/journal.pone.0076100.g001

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Different kinds of known faces activate brain areas to dissimilar degrees. However, the tuning to type of knowledge, and the temporal course of activation, of each area have not been well characterized. Here we measured, with functional magnetic resonance imaging, brain activity elicited by unfamiliar, visually familiar, and personally-familiar fac...

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... order to localize the brain areas responding to different face conditions, a second level random effects analysis over three different contrasts was carried out (Figure 1). Face selective areas were defined by the contrast of unfamiliar faces.houses ...
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... these comparisons we used a subset of unfamiliar face not used further for the HRF estimations.) The results of the random effects model for this contrast are shown in Figure 1A and Table 1. Two main clusters of activation were found bilaterally, which correspond to the FFA and OFA. ...
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... main clusters of activation were found bilaterally, which correspond to the FFA and OFA. Some other areas, PC, mOF, and left FrI, also exhibited significant activations ( Figure 1A, Table 1). Brain areas related to face familiarity processing were located by analyzing the contrasts: acquaintances.unfamiliar ...
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... and newly-learned.unfamiliar faces ( Figure 1B, C). The responses to faces of acquaintances were larger than the responses to unfamiliar faces in several regions, including AC, mOF, PC, left FrI, left and right pSTS, right anterior FFA, and the right parahippocampus ( Figure 1B). ...
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... ( Figure 1B, C). The responses to faces of acquaintances were larger than the responses to unfamiliar faces in several regions, including AC, mOF, PC, left FrI, left and right pSTS, right anterior FFA, and the right parahippocampus ( Figure 1B). Smaller clusters also appeared in other regions (Table 2). ...
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... clusters also appeared in other regions (Table 2). However, the cortical response to newly- learned faces was more limited, evoking larger BOLD responses than unfamiliar faces only in PC, left middle temporal, right hippocampus and the left FrI ( Figure 1C, Table 3). ...
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... with a much smaller differ- ence in the core ROIs (F(1,9) = 10.25.p,0.01) (see Figure S1 in File S1). The contrast for the interaction between lumped core vs. lumped extended ROIs on one hand, and faces of acquaintances vs. unfamiliar faces on the other, was significant (F(1,9) = 7.34, p,0.02). ...
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... S1 Figure S1. Bar graphs showing the median of the height parameter of the core system ROIs(left and right OFA, left and right FFA, and right pSTS) and the extended ROIs (left and right PC, left and right AC, left and right mOF, left and right FrI and left Ins. ...

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... Neuroimaging studies have described that posterior cortical regions (mainly posterior cingulate cortex -PCC-and precuneus -PC) are crucial for tracing the self into autobiographical memory (Burgess et al. 2001;Gobbini et al. 2004Gobbini et al. , 2007Ishai et al. 2000;Sugiura et al. 2005). Additionally, Bobes et al. (2013) observed that anterior regions (e.g., medial prefrontal cortex-mPFC) exhibited stronger and prolonged activity to personalfamiliar faces, being weaker to unknown or visually familiar (learned after repetitive exposition) faces. In a self vs. other identity discrimination, Murray et al. (2015) described mPFC activation to be specific to self-identity, motivationally oriented recognition, while PCC/PC would be specific to other-identity socially oriented recognition. ...
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... On the other hand, representations of face familiarity may reflect a convergence between perceptual information and information stored in declarative recognition memory. This interpretation is consistent with studies of face memory, which have reported enhanced BOLD activations for highly familiar, compared with less familiar faces, in the prefrontal, parietal, and medial temporal cortices (Leveroni et al., 2000;Kosaka et al., 2003;Leube et al., 2003;Gobbini and Haxby, 2006;Bobes et al., 2013;Silson et al., 2019) Pinpointing neural representations of familiarity to the interface of face perception and recognition memory is consistent with the timing of the effects in our study: Representations of face familiarity emerged only after 400 ms of processing, much later than perceptual responses to faces (Eimer et al., 2011). This timing is consistent with a previous ERP study that reported differences between familiar and unfamiliar faces in averaged waveforms after 400 ms (Wiese et al., 2019). ...
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... The extended network also includes the precuneus/posterior cingulate cortex crucial for the perception of (perceptual) face familiarity (Cloutier et al., 2011) and the anterior temporal cortices known to process representations of biographical and autobiographical knowledge (Haxby et al., 2000). The latter regions are more strongly activated by (personally) familiar faces compared to unfamiliar ones (Bobes et al., 2013). ...
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... Therefore, it is hardly surprising that several univariate neuroimaging studies compared the magnitude of neural activity to famous faces to the response to unfamiliar faces (for a review, see Natu & O'Toole, 2011). These studies showed larger activity for famous when compared to unfamiliar faces in both the core and extended face processing systems, including inferior occipital (Ishai, Schmidt, & Boesiger, 2005); fusiform (Bobes et al., 2013;Nielson et al., 2010;Ishai et al., 2005;Gobbini, Leibenluft, Santiago, & Haxby, 2004;Grill-Spector, Knouf, & Kanwisher, 2004;Sergent, Ohta, & MacDonald, 1992); anterior middle temporal (Gorno-Tempini & Price, 2001;Leveroni et al., 2000); medial temporal, such as hippocampal and parahippocampal (Bar, Aminoff, & Ishai, 2008;Leveroni et al., 2000); and superior temporal (Ishai et al., 2005;Leveroni et al., 2000) areas as well as the MPFC, the PC, the TPJ (Nielson et al., 2010;Gobbini et al., 2004;Leveroni et al., 2000), and the amygdala (Elfgren et al., 2006;Bernard et al., 2004). ...
... These studies found a large number of areas from both the core and extended parts of the face processing network. This network of personally familiar face processing includes the MPFC, ACC and PCC, and the anterior paracingulate cortex (Góngora, Castro-Laguardia, Pérez, Valdés-Sosa, & Bobes, 2019;Visconti di Oleggio Castello, Halchenko, Guntupalli, Gors, & Gobbini, 2017;Bobes et al., 2013;Gobbini et al., 2004;Leibenluft, Gobbini, Harrison, & Haxby, 2004); the right iFFA ( Visconti di Oleggio Castello et al., 2017); the IPL and the TPJ (Sugiura, Mano, Sasaki, & Sadato, 2011;Platek et al., 2006); the PC ( Visconti di Oleggio Castello et al., 2017;Sugiura et al., 2011;Gobbini et al., 2004); the middle and superior temporal cortices (Visconti di Oleggio Castello et al., 2017;Sugiura et al., 2011;Platek et al., 2006;Gobbini et al., 2004;Leibenluft et al., 2004); inferior temporal regions, such as the FFA and ATL (Visconti di Oleggio Castello et al., 2017;Ramon, Vizioli, Liu-Shuang, & Rossion, 2015;Sugiura et al., 2011;Gobbini et al., 2004;Leibenluft et al., 2004); the insula (Góngora et al., 2019;Platek et al., 2006;Leibenluft et al., 2004); and MTL regions, such as the amygdala, hippocampus, and perirhinal cortex (Ramon et al., 2015;Sugiura et al., 2011;Taylor et al., 2009), showing stronger neural responses to personally familiar faces. ...
... Probably every face, but the faces of our favorite celebrities and personally familiar faces specially, elicits increased emotional attachments and processing. This is typically related to the differential activation of the amygdala, insula, and limbic areas (Bobes et al., 2013;Natu & O'Toole, 2011;. The fact that only studies applying famous or personally familiar faces found familiarity-dependent responses in the amygdala suggests that the short practice of typical experimental familiarizations is not sufficient to activate these emotion processing areas differentially. ...
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... The other explanation would simply be a better ability to process sensory information derived from the eye region, in other words, sensory expertise caused by familiarity. The pSTS is also engaged in face-selective processing and activates stronger to familiar vs unfamiliar faces 55,56 . In a recent study, the pSTS was found to be related to person-selective processing, irrespectively of modality 57 . ...
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... An important question concerns the potential involvement of the vast and complex semantic, experiential and emotional aspects of our responses to a familiar person (Bobes, Lage Castellanos, Quiñones, García, & Valdes-Sosa, 2013;Gobbini & Haxby, 2007;Wiese et al., 2019); in particular, whether these play a central role in creating the profound differences between recognition of familiar and unfamiliar faces. Accessing semantic, episodic and emotional memories forms a critical aspect of our everyday experience with familiar people, but we have shown here that, from the standpoint of visual recognition, these properties are not essential to differences in recognisability between familiar and unfamiliar faces. ...
Article
We can recognise people that we know across their lifespan. We see family members age, and we can recognise celebrities across long careers. How is this possible, despite the very large facial changes that occur as people get older? Here we analyse the statistical properties of faces as they age, sampling photos of the same people from their 20s to their 70s. Across a number of simulations, we observe that individuals’ faces retain some idiosyncratic physical properties across the adult lifespan that can be used to support moderate levels of age-independent recognition. However, we found that models based exclusively on image-similarity only achieved limited success in recognising faces across age. In contrast, more robust recognition was achieved with the introduction of a minimal top-down familiarisation procedure. Such models can incorporate the within-person variability associated with a particular individual to show a surprisingly high level of generalisation, even across the lifespan. The analysis of this variability reveals a powerful statistical tool for understanding recognition, and demonstrates how visual representations may support operations typically thought to require conceptual properties.
... Most of these areas were composed of more voxels in the left hemisphere (except for mOF/AC). It is important to note that this study used personally-familiar faces instead of famous or learned faces, which must enhance the activation of extended system and recruited areas related to processing the emotional meaning (faces from friends and relatives) of the faces, a subsystem related to the emotion-from identity processing [58] which is not activated in presence of visually familiar faces (learnt in laboratory). On the other hand, an interesting study using multivoxel pattern analysis to fMRI was able to decode identity-independent familiarity information and face identity in a set of overlapping areas in core and extended systems providing evidence that activity in the extended system might also carry information about identity and personal familiarity [59]. ...
... The larger activation of extended system areas in the left hemisphere in response to familiar faces can be explained by the lateralization of positive emotions to the left hemisphere that has been proposed based on the 'valence hypothesis' which assumes an opposite dominance of the left hemisphere for positive emotions and of the right hemisphere for negative emotions [60,61]. Personally familiar faces elicited bilateral activation with important recruitment of the left hemisphere [58]. Leibenluft et al. (2004) by contrasting the face of a mother's own child and a familiar child, which found areas in the anterior paracingulate, pre-frontal cortex, and left insula which was considered like a maternal attachment sign [62]. ...
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Familiar face processing involves face specific regions (the core face system) as well as other non-specific areas related to processing of person-related information (the extended face system). The connections between core and extended face system areas must be critical for face recognition. Some studies have explored the connectivity pattern of unfamiliar face responding area, but none have explored those areas related to face familiarity processing in the extended system. To study these connections, diffusion weighted imaging with probabilistic tractography was used to estimate the white-matter pathways between core and extended system regions, which were defined from functional magnetic resonance imaging responses to personally-familiar faces. Strong white matter connections were found between occipitotemporal face areas (OFA/FFA) with superior temporal sulcus and insula suggesting the possible existence of direct anatomical connections from face-specific areas to frontal nodes that could underlay the processing of emotional information associated to familiar faces.
... We estimated the generators of the early (150-210 ms), middle (300-380 ms) and late (460-580 ms) time windows responses using the BMA method, favoring the solution to the fMRI predefined ROIs, but not strictly excluding all other possible anatomical locations. We hypothesized that a subset of the areas detected in the fMRI study (Bobes et al. 2013) contributes to the generation of each ERP component (Bobes et al. 2007). By using the fMRI-constrained BMA approach we were able to estimate the latency of activation in each face area (rather than only localize ERP generators) thus providing information about the temporal dynamics of the familiar face processing. ...
... In this article, we combined data from a previous ERP study (Bobes et al. 2007), and a previous event related fMRI study (Bobes et al. 2013). We carried out a new source analysis of the ERPs components using the results of the fMRI study for modeling the solution space for source estimation. ...
... The same group of participants was recruited for both experiments, except for two participants, who were not able to participate in the fMRI study and had to be replaced. Details about the event related fMRI experiment (participants, paradigm, acquisition parameters and analysis) could be found in the original paper (Bobes et al. 2013). ...
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Event related potentials (ERPs) provide precise temporal information about cognitive processing, but with poor spatial resolution, while functional magnetic resonance imaging (fMRI) reliably identifies brain areas involved, but with poor temporal resolution. Here we use fMRI to guide source localization of the ERPs at different times for studying the temporal dynamics of the neural system for recognizing familiar faces. fMRI activation areas were defined in a previous experiment applying the same paradigm used for ERPs. The Bayesian model averaging (BMA) method was used to estimate the generators of the ERPs to unfamiliar, visually familiar, and personally-familiar faces constraining the model by fMRI activation results. For this, higher prior probabilities in the solution space were assigned to the fMRI-defined regions, which included face-selective areas and other areas related to “person knowledge” retrieval. Source analysis was carried out in three-time windows: early (150–210 ms), middle (300–380 ms) and late (460–580 ms). The early and middle responses were generated in fMRI-defined areas for all face categories, while these areas do not contribute to the late response. Different areas contributed to the generation of the early and middle ERPs elicited by unfamiliar faces: fusiform (Fus), inferior occipital, superior temporal sulcus and the posterior cingulate (PC) cortices. For familiar faces, the contributing areas were Fus, PC and anterior temporal areas for visually familiar faces, with the addition of the medial orbitofrontal areas and other frontal structures for personally-significant faces. For both unfamiliar and familiar faces, more extended and reliable involvement of contributing areas were obtained for the middle compare with early time window. Our fMRI guide ERP source analysis suggested the recruitment of person-knowledge processing areas as early as 150–210 ms after stimulus onset during recognition of personally-familiar faces. We concluded that fMRI-constrained BMA source analysis provide information regarding the temporal-dynamics in the neural system for cognitive processsing.
... Most of these studies focused on the cognitive processing of familiarity using tasks that required explicit recognition (i.e., providing the name of the person or judging whether a face/name was familiar or unfamiliar) or implicit recognition (i.e., gender-classification, one-back repetition, or inverted-target detection). However, a considerable number of those studies included a category of faces/names called personally familiar (faces/names of family members and friends) that involved stronger familiarity not only in terms of personal and biographical knowledge but also in terms of social attachment and emotion [24], [31], [32], [33] [34]. In addition, a subset of these studies, namely, those on face-identity recognition, focused on emotional processing and attachment by directly examining the faces of loved people, such as romantic partners, parents, or own children [34], [35], [36], [37], [38], [39], [40], [41]. ...
... The posterior cingulate is part of the extended system in Gobbini and Haxby's [24] model, being related to episodic memories. Although not included in the model, the medial orbito-frontal cortex and the anterior cingulate (but also the medial prefrontal cortex) are structures that recent studies have implicated in the processing of personally familiar faces and names [32], [33], [45], [64]. Furthermore, there is evidence that the medial orbito-frontal cortex is involved in emotional processing, a claim supported by data showing (a) suppression of the enhanced skin conductance responses by lesions to this area [63], (b) a significant correlation between BOLD signals and skin conductance amplitudes [65], (c) the role of the medial orbito-frontal cortex in the reward value of stimuli [66], and (d) its activation by emotional facial expressions [67] and pleasant imagery [51]. ...
... The anterior cingulate was one of the areas activated in Bartels & Zeki's studies [35] [36] where people viewed faces of their romantic partner or mothers viewed faces of their own children. These two areas showed intense and prolonged activation in Bobes et al.'s [33] study when participants viewed faces of acquaintances. Combined with the parallel increases in skin conductance responses to faces of acquaintances [70], our present finding of the anterior cingulate as one of the major structures in the processing of loved familiar faces and names -together with our finding of simultaneous increases in heart rate-supports the role that the anterior cingulate plays in the processing of positive emotional experiences. ...
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The neuroscientific study of love has been boosted by an extended corpus of research on face-identity recognition. However, few studies have compared the emotional mechanisms activated by loved faces and names and none have simultaneously examined fMRI and autonomic measures. The present study combined fMRI with the heart rate response when 21 participants (10 males) passively viewed the face or the written name of 4 loved people and 4 unknown people. The results showed accelerative patterns in heart rate, together with brain activations, which were significantly higher for loved people than for unknown people. Significant correlations were found between heart rate and brain activation in frontal areas, for faces, and in temporal areas, for names. The results are discussed in the context of previous studies using the same passive viewing procedure, highlighting the relevance of integrating peripheral and central measures in the scientific study of positive emotion and love.
... Some studies have used personally familiar faces as stimuli, i.e., those that have become familiar through extensive exposure under naturalistic conditions in real life settings (e.g., Arsalidou, Barbeau, Bayless, & Taylor, 2010;Bartels & Zeki, 2000;Bartels & Zeki, 2004;Bobes, Lage Castellanos, Quinones, Garcia, & Valdes-Sosa, 2013;Chauhan et al., 2017;Gobbini et al., 2004;Gobbini et al., 2013;Leibenluft et al., 2004;Nakamura et al., 2000;Ramon, 2015aRamon, , 2015bRamon et al., 2015;Ramon & Van Belle, 2016;Ramon, Busigny, Gosselin, & Rossion, 2017;Ramon, Caharel, & Rossion, 2011;Taylor et al., 2009 Watier & Collin, 2009). This is the type of face familiarity we cover in this review: the one that develops over years of repeated personal interactions and includes a range of familiar individuals, such as relatives, colleagues, and friends. ...
... Collectively, these univariate findings are also in line with previous reports of personally meaningful stimuli eliciting increased temporal pole activation (Nakamura et al., 2000) and selective responses across larger proportions of neurons in the amygdala, entorhinal and perirhinal cortex, and hippocampus of pre-surgical patients (Viskontas, Quiroga, & Fried, 2009). MVPA performed within face-preferential regions revealed that that familiar faces elicited highly similar neural representations in bilateral amygdala and left FFA (Ramon et al., 2015), suggesting categorical signalling of familiarity within these regions. 2 Familiarity-dependent modulation in areas beyond the core system has been reported in numerous earlier studies, which have highlighted regions thought to be involved in spontaneous activation of person knowledge (Arsalidou et al., 2010;Bartels & Zeki, 2000Bobes et al., 2013;Gobbini et al., 2004;Krienen, Tu, & Buckner, 2010;Leibenluft et al., 2004;Taylor et al., 2009), as well as more recent studies (Ramon et al., 2015;. Gobbini et al. (2004) and Leibenluft et al. (2004) reported increased responses for personally familiar faces as compared to famous faces, and for close as compared to more distant personally familiar faces, across a distributed set of areas that are part of the extended system. ...
... The role of ToM areas in the representation of personally familiar faces has been replicated in subsequent fMRI experiments using personally familiar faces characterized by different types of social and emotional attachment (Arsalidou et al., 2010;Bartels & Zeki, 2004;Bobes et al., 2013;Taylor et al., 2009). The role of the ToM areas in retrieval of person knowledge has been corroborated in studies that used experimentally learned person knowledge associated with faces (Cloutier et al., 2011;Todorov et al., 2007) and verbal statements about the personal attributes of friends (Krienen et al., 2010) (see Figure 7). ...
Article
In this review, we synthesize the existing literature investigating personally familiar face processing and highlight the remarkable, enhanced processing efficiency resulting from real-life experience. Highly learned identity-specific visual and semantic information associated with personally familiar face representations facilitates detection, recognition of identity and social cues, and activation of person knowledge. These optimizations afford qualitatively different processing of personally familiar as compared to unfamiliar faces, which manifests on both the behavioural and neural level. (Access the pdf of this paper here: https://www.tandfonline.com/eprint/tV4Q77TzdikFavPThjHr/full)