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The human visual system is very sensitive to the presence of faces in the environment, so much so that it can produce the perception of illusory faces in everyday objects. Growing research suggests that illusory faces and real faces are processed by similar perceptual and neural mechanisms, but whether this similarity extends to visual attention is less clear. A visual search study showed that illusory faces have a search advantage over objects when the types of objects vary to match the objects in the illusory faces (e.g., chair, pepper, clock) (Keys et al., 2021). Here, we examine whether the search advantage for illusory faces over objects remains when compared against objects that belong to a single category (flowers). In three experiments, we compared visual search of illusory faces, real faces, variable objects, and uniform objects (flowers). Search for real faces was best compared with all other types of targets. In contrast, search for illusory faces was only better than search for variable objects, not uniform objects. This result shows a limited visual search advantage for illusory faces and suggests that illusory faces may not be processed like real faces in visual attention.

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Here we examine whether our impressive ability to perceive upright faces arises from evolved orientation-specific mechanisms, our extensive experience with upright faces, or both factors. To do so, we tested Claudio, a man with a congenital joint disorder causing his head to be rotated back so that it is positioned between his shoulder blades. As a result, Claudio has seen more faces reversed in orientation to his own face than matched to it. Controls exhibited large inversion effects on all tasks, but Claudio performed similarly with upright and inverted faces in both detection and identity-matching tasks, indicating these abilities are the product of evolved mechanisms and experience. In contrast, he showed clear upright superiority when detecting "Thatcherized" faces (faces with vertically flipped features), suggesting experience plays a greater role in this judgment. Together, these findings indicate that both evolved orientation-specific mechanisms and experience contribute to our proficiency with upright faces.

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Difficulties in various face processing tasks have been well documented in autism spectrum disorder (ASD). Several meta-analyses and numerous case-control studies have indicated that this population experiences a moderate degree of impairment, with a small percentage of studies failing to detect any impairment. One possible account of this mixed pattern of findings is heterogeneity in face processing abilities stemming from the presence of a subpopulation of prosopagnosic individuals with ASD alongside those with normal face processing skills. Samples randomly drawn from such a population, especially relatively smaller ones, would vary in the proportion of participants with prosopagnosia, resulting in a wide range of group-level deficits from mild (or none) to severe across studies. We test this prosopagnosic subpopulation hypothesis by examining three groups of participants: adults with ASD, adults with developmental prosopagnosia (DP), and a comparison group. Our results show that the prosopagnosic subpopulation hypothesis does not account for the face impairments in the broader autism spectrum. ASD observers show a continuous and graded, rather than categorical, heterogeneity that span a range of face processing skills including many with mild to moderate deficits, inconsistent with a prosopagnosic subtype account. We suggest that pathogenic origins of face deficits for at least some with ASD differ from those of DP.

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The pattern of shadows and shading across a face is determined partly by face shape and may therefore provide a cue for facial recognition. In this study, we measured the ability of human observers to discriminate facial identity based simply on the coarse pattern of contrast produced by the interaction between facial geometry and lighting direction. We used highly realistic 3D models of human heads to create images of faces illuminated from different horizontal and vertical directions, which were then converted to two-tone images ('Mooney faces') to isolate the coarse pattern of contrast. Participants were presented with pairs of two-tone faces and judged whether it was the same person in both images. Participants could discriminate facial identity based on the minimal cues within the two-tone images, though sensitivity depended on the horizontal and vertical lighting direction. Performance on the Mooney recognition task correlated with general facial recognition ability, though the role of face-specific processing in this relationship was not significant. Our results demonstrate that shading information in the form of simple contrast cues is sufficient for discriminating facial identity, and support the idea that visual processing is somewhat optimised for overhead lighting - here, in the relatively high-level context of face identity recognition.

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Individuals with developmental prosopagnosia (DPs) experience severe and lifelong deficits recognising faces, but whether their deficits are selective to the processing of face identity or extend to the processing of face expression remains unclear. Clarifying this issue is important for understanding DP impairments and advancing theories of face processing. We compared identity and expression processing in a large sample of DPs (N = 124) using three different matching tasks that each assessed identity and expression processing with identical experimental formats. We ran each task in upright and inverted orientations and we measured inversion effects to assess the integrity of upright-specific face processes. We report three main results. First, DPs showed large deficits at discriminating identity but only subtle deficits at discriminating expression. Second, DPs showed a reduced inversion effect for identity but a normal inversion effect for expression. Third, DPs' performance on the expression tasks were linked to autism traits, but their performance on the identity tasks were not. These results constitute several dissociations between identity and expression processing in DP, and they are consistent with the view that the core impairment in DP is highly selective to identity.

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Face recognition is strongly influenced by the processing of orientation structure in the face image. Faces are much easier to recognize when they are filtered to include only horizontally oriented information compared with vertically oriented information. Here, we investigate whether preferences for horizontal information in faces are related to face recognition abilities in a typical sample (Experiment 1), and whether such preferences are lacking in people with developmental prosopagnosia (DP; Experiment 2). Experiment 1 shows that preferences for horizontal face information are linked to face recognition abilities in a typical sample, with weak evidence of face-selective contributions. Experiment 2 shows that preferences for horizontal face information are comparable in control and DP groups. Our study suggests that preferences for horizontal face information are related to variations in face recognition abilities in the typical range, and that these preferences are not aberrant in DP.

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Individuals with developmental prosopagnosia (DP) all exhibit impairments in face memory, but the specificity of these face memory impairments is debated. One problem is that standard behavioural tasks are not able to provide independent measurement of face perception, face memory, and face matching (the decision process required to judge whether two instances of a face are of the same individual or different individuals). The present study utilised a new test of face matching, the Oxford Face Matching Test (OFMT), and a novel analysis strategy to derive these independent indices. Twenty-nine individuals with DP and the same number of matched neurotypical controls completed the OFMT, the Glasgow Face Matching Test, and the Cambridge Face Memory Test. Results revealed individuals with DP exhibit impairments in face perception, face memory and face matching. Collectively, these results suggest that face processing impairments in DP are more comprehensive than has previously been suggested.

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Two key functions in human face perception are gaze discrimination and identity recognition. Here we examine whether gaze discrimination can be intact when identity recognition is impaired in developmental prosopagnosia (DP). We ran a large sample of developmental prosopagnosics (DPs) with a series of gaze discrimination tasks that assess various mechanisms in gaze processing. Experiment 1 (N = 101 DP participants) investigates spatial processing using an abnormal eye gaze detection task and a Wollaston illusion task that measures perceptual integration of eye and head direction. Experiment 2 (N = 45 DP participants) investigates temporal processing using an adaptation task and a serial dependence task. Despite their deficits with identity recognition, DPs performed in the normal range across both experiments. These results demonstrate that gaze discrimination can be normal in DP, and that various mechanisms of gaze processing can be spared when identity recognition is impaired. Our findings clarify the highly selective nature of impairments in DP and provide support for neurocognitive models of face perception with distinct mechanisms for gaze and identity processing.

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Tests of face processing are typically designed to identify individuals performing outside of the typical range; either prosopagnosic individuals who exhibit poor face processing ability, or super recognisers, who have superior face processing abilities. Here we describe the development of the Oxford Face Matching Test (OFMT), designed to identify individual differences in face processing across the full range of performance, from prosopagnosia, through the range of typical performance, to super recognisers. Such a test requires items of varying difficulty, but establishing difficulty is problematic when particular populations (e.g., prosopagnosics, individuals with autism spectrum disorder) may use atypical strategies to process faces. If item difficulty is calibrated on neurotypical individuals, then the test may be poorly calibrated for atypical groups, and vice versa. To obtain items of varying difficulty, we used facial recognition algorithms to obtain face pair similarity ratings that are not biased towards specific populations. These face pairs were used as stimuli in the OFMT, and participants were required to judge whether the face images depicted the same individual or different individuals. Across five studies the OFMT was shown to be sensitive to individual differences in the typical population, and in groups of both prosopagnosic individuals and super recognisers. The test-retest reliability of the task was at least equivalent to the Cambridge Face Memory Test and the Glasgow Face Matching Test. Furthermore, results reveal, at least at the group level, that both face perception and face memory are poor in those with prosopagnosia, and are good in super recognisers.

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Developmental prosopagnosia (DP) is a selective neurodevelopmental condition defined by lifelong impairments in face recognition. Despite much research, the extent to which DP is associated with broader visual deficits beyond face processing is unclear. Here we investigate whether DP is accompanied by deficits in colour perception. We tested a large sample of 92 DP individuals and 92 sex/age-matched controls using the well-validated Ishihara and Farnsworth-Munsell 100-Hue tests to assess red-green colour deficiencies and hue discrimination abilities. Group-level analyses show comparable performance between DP and control individuals across both tests, and single-case analyses indicate that the prevalence of colour deficits is low and comparable to that in the general population. Our study clarifies that DP is not linked to colour perception deficits and constrains theories of DP that seek to account for a larger range of visual deficits beyond face recognition.

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Eye movement studies show that humans can make very fast saccades towards faces in natural scenes, but the visual mechanisms behind this process remain unclear. Here we investigate whether fast saccades towards faces rely on mechanisms that are sensitive to the orientation or contrast of the face image. We present participants pairs of images each containing a face and a car in the left and right visual field or the reverse, and we ask them to saccade to faces or cars as targets in different blocks. We assign participants to one of three image conditions: normal images, orientation-inverted images, or contrast-negated images. We report three main results that hold regardless of image conditions. First, reliable saccades towards faces are fast - they can occur at 120-130 ms. Second, fast saccades towards faces are selective - they are more accurate and faster by about 60-70 ms than saccades towards cars. Third, saccades towards faces are reflexive - early saccades in the interval of 120-160 ms tend to go to faces, even when cars are the target. These findings suggest that the speed, selectivity, and reflexivity of saccades towards faces do not depend on the orientation or contrast of the face image. Our results accord with studies suggesting that fast saccades towards faces are mainly driven by low-level image properties, such as amplitude spectrum and spatial frequency.

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Whether face and object recognition are dissociated in prosopagnosia continues to be debated: a recent review highlighted deficiencies in prior studies regarding the evidence for such a dissociation. Our goal was to study cohorts with acquired and developmental prosopagnosia with a complementary battery of tests of object recognition that address prior limitations, as well as evaluating for residual effects of object expertise. We studied 15 subjects with acquired and 12 subjects with developmental prosopagnosia on three tests: the Old/New Tests, the Cambridge Bicycle Memory Test, and the Expertise-adjusted Test of Car Recognition. Most subjects with developmental prosopagnosia were normal on the Old/New Tests: for acquired prosopagnosia, subjects with occipitotemporal lesions often showed impairments while those with anterior temporal lesions did not. Ten subjects showed a putative classical dissociation between the Cambridge Face and Bicycle Memory Tests, seven of whom had normal reaction times. Both developmental and acquired groups showed reduced car recognition on the expertise-adjusted test, though residual effects of expertise were still evident. Two subjects with developmental prosopagnosia met criteria for normal object recognition across all tests. We conclude that strong evidence for intact object recognition can be found in a few subjects but the majority show deficits, particularly those with the acquired form. Both acquired and developmental forms show residual but reduced object expertise effects.

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Recent theories suggest that familiar faces have a robust representation in memory because they have been encountered over a wide variety of contexts and image changes (e.g. lighting, viewpoint and expression). By contrast, unfamiliar faces are encountered only once, and so they do not benefit from such richness of experience and are represented based on image-specific details. In this registered report, we used a repeat detection task to test whether familiar faces are recognized better than unfamiliar faces across image changes. Participants viewed a stream of more than 1000 celebrity face images for 0.5 s each, any of which might be repeated at a later point and has to be detected. Some participants saw the same image at repeats, while others saw a different image of the same face. A post-experimental familiarity check allowed us to determine which celebrities were and were not familiar to each participant. We had three predictions: (i) detection would be better for familiar than unfamiliar faces, (ii) detection would be better across same rather than different images, and (iii) detection of familiar faces would be comparable across same and different images, but detection of unfamiliar faces would be poorer across different images. We obtained support for the first two predictions but not the last. Instead, we found that repeat detection of faces, regardless of familiarity, was poorer across different images. Our study suggests that the robustness of familiar face recognition may have limits, and that under some conditions, familiar face recognition can be just as influenced by image changes as unfamiliar face recognition.

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Face recognition is thought to rely on specific mechanisms underlying a perceptual bias toward processing faces as undecomposable wholes. This face-specific "holistic processing" is commonly quantified using 3 measures: the inversion, part-whole, and composite effects. Consequently, many researchers assume that these 3 effects measure the same cognitive mechanism(s) and these mechanisms contribute to the wide range of individual differences seen in face recognition ability. We test these assumptions in a large sample (N = 282), with individual face recognition abilities measured by the well-validated Cambridge Face Perception Test. Our results provide little support for either assumption. The small to nonexistent correlations among inversion, part-whole, and composite effects (correlations between -.03 and .28) fail to support the first assumption. As for the second assumption, only the inversion effect moderately predicts face recognition (r = .42); face recognition was weakly correlated with the part-whole effect (r = .25) and not correlated with the composite effect (r = .04). We rule out multiple artifactual explanations for our results by using valid tasks that produce standard effects at the group level, demonstrating that our tasks exhibit psychometric properties suitable for individual differences studies, and demonstrating that other predicted correlations (e.g., between face perception measures) are robust. Our results show that inversion, part-whole, and composite effects reflect distinct perceptual mechanisms, and we argue against the use of the single, generic term holistic processing when referring to these effects. Our results also question the contribution of these mechanisms to individual differences in face recognition. (PsycINFO Database Record

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The right posterior superior temporal sulcus (pSTS) shows a strong response to voices, but the cognitive processes generating this response are unclear. One possibility is that this activity reflects basic voice processing. However, several fMRI and magnetoencephalography findings suggest instead that pSTS serves as an integrative hub that combines voice and face information. Here we investigate whether right pSTS contributes to basic voice processing by testing Faith, a patient whose right pSTS was resected, with eight behavioral tasks assessing voice identity perception and recognition, voice sex perception, and voice expression perception. Faith performed normally on all the tasks. Her normal performance indicates right pSTS is not necessary for intact voice recognition and suggests that pSTS activations to voices reflect higher-level processes.

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Synaesthesia is a rare phenomenon in which stimulation in one modality (e.g., audition) evokes a secondary percept not associated with the first (e.g., colour). Prior work has suggested links between synaesthesia and other neurodevelopmental conditions that are linked to altered social perception abilities. With this in mind, here we sought to examine social perception abilities in grapheme-colour synaesthesia (where achromatic graphemes evoke colour experiences) by examining facial identity and facial emotion perception in synaesthetes and controls. Our results indicate that individuals who experience grapheme-colour synaesthesia outperformed controls on tasks involving fine visual discrimination of facial identity and emotion, but not on tasks involving holistic face processing. These findings are discussed in the context of broader perceptual and cognitive traits previously associated with synaesthesia for colour, with the suggestion that performance benefits shown by grapheme-colour synaesthetes may be related to domain-general visual discrimination biases observed in this group.

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Macaque neurophysiology found image-invariant representations of face identity in a face-selective patch in anterior temporal cortex. A face-selective area in human anterior temporal lobe (fATL) has been reported, but has not been reliably identified, and its function and relationship with posterior face areas is poorly understood. Here, we used fMRI adaptation and neuropsychology to ask whether fATL contains image-invariant representations of face identity, and if so, whether these representations require normal functioning of fusiform face area (FFA) and occipital face area (OFA). We first used a dynamic localizer to demonstrate that 14 of 16 normal subjects exhibit a highly selective right fATL. Next, we found evidence that this area subserves image-invariant representation of identity: Right fATL showed repetition suppression to the same identity across different images, while other areas did not. Finally, to examine fATL's relationship with posterior areas, we used the same procedures with Galen, an acquired prosopagnosic who lost right FFA and OFA. Despite the absence of posterior face areas, Galen's right fATL preserved its face selectivity and showed repetition suppression comparable to that in controls. Our findings suggest that right fATL contains image-invariant face representations that can persist despite the absence of right FFA and OFA, but these representations are not sufficient for normal face recognition.

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It has long been suggested that face recognition relies on specialized mechanisms that are not involved in visual recognition of other object categories, including those that require expert, fine-grained discrimination at the exemplar level such as written words. But according to the recently proposed many-to-many theory of object recognition (MTMT), visual recognition of faces and words are carried out by common mechanisms [Behrmann, M., & Plaut, D. C. ( 2013 ). Distributed circuits, not circumscribed centers, mediate visual recognition. Trends in Cognitive Sciences, 17, 210-219]. MTMT acknowledges that face and word recognition are lateralized, but posits that the mechanisms that predominantly carry out face recognition still contribute to word recognition and vice versa. MTMT makes a key prediction, namely that acquired prosopagnosics should exhibit some measure of word recognition deficits. We tested this prediction by assessing written word recognition in five acquired prosopagnosic patients. Four patients had lesions limited to the right hemisphere while one had bilateral lesions with more pronounced lesions in the right hemisphere. The patients completed a total of seven word recognition tasks: two lexical decision tasks and five reading aloud tasks totalling more than 1200 trials. The performances of the four older patients (3 female, age range 50-64 years) were compared to those of 12 older controls (8 female, age range 56-66 years), while the performances of the younger prosopagnosic (male, 31 years) were compared to those of 14 younger controls (9 female, age range 20-33 years). We analysed all results at the single-patient level using Crawford's t-test. Across seven tasks, four prosopagnosics performed as quickly and accurately as controls. Our results demonstrate that acquired prosopagnosia can exist without word recognition deficits. These findings are inconsistent with a key prediction of MTMT. They instead support the hypothesis that face recognition is carried out by specialized mechanisms that do not contribute to recognition of written words.

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Recently, a number of studies have demonstrated the utility of transcranial current stimulation as a tool to facilitate a variety of cognitive and perceptual abilities. Few studies, though, have examined the utility of this approach for the processing of social information. Here, we conducted 2 experiments to explore whether a single session of high-frequency transcranial random noise stimulation (tRNS) targeted at lateral occipitotemporal cortices would enhance facial identity perception. In Experiment 1, participants received 20 min of active high-frequency tRNS or sham stimulation prior to completing the tasks examining facial identity perception or trustworthiness perception. Active high-frequency tRNS facilitated facial identity perception, but not trustworthiness perception. Experiment 2 assessed the spatial specificity of this effect by delivering 20 min of active high-frequency tRNS to lateral occipitotemporal cortices or sensorimotor cortices prior to participants completing the same facial identity perception task used in Experiment 1. High-frequency tRNS targeted at lateral occipitotemporal cortices enhanced performance relative to motor cortex stimulation. These findings show that high-frequency tRNS to lateral occipitotemporal cortices produces task-specific and site-specific enhancements in face perception.

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