Human Face Processing Research Articles

The hows and whys of face processing: Level of construal influences the holistic processing of human faces.

Face recognition and identification are optimized by holistic processing. Various visual-spatial manipulations appear to have transfer effects on holistic face processing. The present experiment tests the effects of a semantic manipulation--of construal level--on holistic processing as measured by composite congruency effects. Participants completed two blocks of trials. The first served as a baseline, whereas the second included a manipulation of construal level. High-level construal resulted in stronger congruency effects, indicative of greater holistic processing (relative to baseline and to low-level construal). These results have implications for conceptualizations of both construal level and holistic processing.

Lateralization for dynamic facial expressions in human superior temporal sulcus

Most face processing studies in humans show stronger activation in the right compared to the left hemisphere. Evidence is largely based on studies with static stimuli focusing on the fusiform face area (FFA). Hence, the pattern of lateralization for dynamic faces is less clear. Furthermore, it is unclear whether this property is common to human and non-human primates due to predisposing processing strategies in the right hemisphere or that alternatively left sided specialization for language in humans could be the driving force behind this phenomenon.We aimed to address both issues by studying lateralization for dynamic facial expressions in monkeys and humans. Therefore, we conducted an event-related fMRI experiment in three macaques and twenty right handed humans. We presented human and monkey dynamic facial expressions (chewing and fear) as well as scrambled versions to both species. We studied lateralization in independently defined face-responsive and face-selective regions by calculating a weighted lateralization index (LIwm) using a bootstrapping method. In order to examine if lateralization in humans is related to language, we performed a separate fMRI experiment in ten human volunteers including a ‘speech’ expression (one syllable non-word) and its scrambled version.Both within face-responsive and selective regions, we found consistent lateralization for dynamic faces (chewing and fear) versus scrambled versions in the right human posterior superior temporal sulcus (pSTS), but not in FFA nor in ventral temporal cortex. Conversely, in monkeys no consistent pattern of lateralization for dynamic facial expressions was observed. Finally, LIwms based on the contrast between different types of dynamic facial expressions (relative to scrambled versions) revealed left-sided lateralization in human pSTS for speech-related expressions compared to chewing and emotional expressions.To conclude, we found consistent laterality effects in human posterior STS but not in visual cortex of monkeys. Based on our results, it is tempting to speculate that lateralization for dynamic face processing in humans may be driven by left-hemispheric language specialization which may not have been present yet in the common ancestor of human and macaque monkeys.

Eye coding mechanisms in early human face event-related potentials.

In humans, the N170 event-related potential (ERP) is an integrated measure of cortical activity that varies in amplitude and latency across trials. Researchers often conjecture that N170 variations reflect cortical mechanisms of stimulus coding for recognition. Here, to settle the conjecture and understand cortical information processing mechanisms, we unraveled the coding function of N170 latency and amplitude variations in possibly the simplest socially important natural visual task: face detection. On each experimental trial, 16 observers saw face and noise pictures sparsely sampled with small Gaussian apertures. Reverse-correlation methods coupled with information theory revealed that the presence of the eye specifically covaries with behavioral and neural measurements: the left eye strongly modulates reaction times and lateral electrodes represent mainly the presence of the contralateral eye during the rising part of the N170, with maximum sensitivity before the N170 peak. Furthermore, single-trial N170 latencies code more about the presence of the contralateral eye than N170 amplitudes and early latencies are associated with faster reaction times. The absence of these effects in control images that did not contain a face refutes alternative accounts based on retinal biases or allocation of attention to the eye location on the face. We conclude that the rising part of the N170, roughly 120-170 ms post-stimulus, is a critical time-window in human face processing mechanisms, reflecting predominantly, in a face detection task, the encoding of a single feature: the contralateral eye.

Disturbed spatial cognitive processing of body-related stimuli in a case of a lesion to the right fusiform gyrus

The fusiform gyrus (FG) is well known as one of the main neural sites of human face and body processing. We report the case of a young male patient with epilepsy and a circumscribed lesion in the right FG who presented with isolated impairments in spatial cognitive processing of body-related stimuli. However, he did not show any clinical signs of prosopagnosia. In particular, handling/processing of body and face stimuli was impaired, when stimuli were presented in unconventional views and orientations, thus requiring additional spatial cognitive operations. In this case study, we discuss the patient’s selective impairment from the view of current empirical and theoretical work on the segregation of functions in the FG.

Part-based and configural processing of owner's face in dogs.

Dogs exhibit characteristic looking patterns when looking at human faces but little is known about what the underlying cognitive mechanisms are and how much these are influenced by individual experience. In Experiment 1, seven dogs were trained in a simultaneous discrimination procedure to assess whether they could discriminate a) the owner's face parts (eyes, nose or mouth) presented in isolation and b) whole faces where the same parts were covered. Dogs discriminated all the three parts of the owner's face presented in isolation, but needed fewer sessions to reach the learning criterion for the eyes than for both nose and mouth. Moreover, covering the eyes region significantly disrupted face discriminability compared to the whole face condition while such difference was not found when the nose or mouth was hidden. In Experiment 2, dogs were presented with manipulated images of the owner's face (inverted, blurred, scrambled, grey-scale) to test the relative contribution of part-based and configural processing in the discrimination of human faces. Furthermore, by comparing the dogs enrolled in the previous experiment and seven ‘naïve’ dogs we examined if the relative contribution of part-based and configural processing was affected by dogs' experience with the face stimuli. Naïve dogs discriminated the owner only when configural information was provided, whereas expert dogs could discriminate the owner also when part-based processing was necessary. The present study provides the first evidence that dogs can discriminate isolated internal features of a human face and corroborate previous reports of salience of the eyes region for human face processing. Although the reliance on part-perception may be increased by specific experience, our findings suggest that human face discrimination by dogs relies mainly on configural rather than on part-based elaboration.

Face, the final frontier: An ERP study probing processing of human and alien faces in Trekkies and non-Trekkies

Face, the final frontier: An ERP study probing processing of human and alien faces in Trekkies and non-Trekkies

Finding faces among faces: human faces are located more quickly and accurately than other primate and mammal faces

We tested the specificity of human face search efficiency by examining whether there is a broad window of detection for various face-like stimuli-human and animal faces-or whether own-species faces receive greater attentional allocation. We assessed the strength of the own-species face detection bias by testing whether human faces are located more efficiently than other animal faces, when presented among various other species' faces, in heterogeneous 16-, 36-, and 64-item arrays. Across all array sizes, we found that, controlling for distractor type, human faces were located faster and more accurately than primate and mammal faces, and that, controlling for target type, searches were faster when distractors were human faces compared to animal faces, revealing more efficient processing of human faces regardless of their role as targets or distractors (Experiment 1). Critically, these effects remained when searches were for specific species' faces (human, chimpanzee, otter), ruling out a category-level explanation (Experiment 2). Together, these results suggest that human faces may be processed more efficiently than animal faces, both when task-relevant (targets) and task-irrelevant (distractors), even in direct competition with other faces. These results suggest that there is not a broad window of detection for all face-like patterns but that human adults process own-species' faces more efficiently than other species' faces. Such own-species search efficiencies may arise through experience with own-species faces throughout development or may be privileged early in development, due to the evolutionary importance of conspecifics' faces.

On the facilitative effects of face motion on face recognition and its development.

For the past century, researchers have extensively studied human face processing and its development. These studies have advanced our understanding of not only face processing, but also visual processing in general. However, most of what we know about face processing was investigated using static face images as stimuli. Therefore, an important question arises: to what extent does our understanding of static face processing generalize to face processing in real-life contexts in which faces are mostly moving? The present article addresses this question by examining recent studies on moving face processing to uncover the influence of facial movements on face processing and its development. First, we describe evidence on the facilitative effects of facial movements on face recognition and two related theoretical hypotheses: the supplementary information hypothesis and the representation enhancement hypothesis. We then highlight several recent studies suggesting that facial movements optimize face processing by activating specific face processing strategies that accommodate to task requirements. Lastly, we review the influence of facial movements on the development of face processing in the first year of life. We focus on infants' sensitivity to facial movements and explore the facilitative effects of facial movements on infants' face recognition performance. We conclude by outlining several future directions to investigate moving face processing and emphasize the importance of including dynamic aspects of facial information to further understand face processing in real-life contexts.

Neural networks related to dysfunctional face processing in autism spectrum disorder.

One of the most consistent neuropsychological findings in autism spectrum disorders (ASD) is a reduced interest in and impaired processing of human faces. We conducted an activation likelihood estimation meta-analysis on 14 functional imaging studies on neural correlates of face processing enrolling a total of 164 ASD patients. Subsequently, normative whole-brain functional connectivity maps for the identified regions of significant convergence were computed for the task-independent (resting-state) and task-dependent (co-activations) state in healthy subjects. Quantitative functional decoding was performed by reference to the BrainMap database. Finally, we examined the overlap of the delineated network with the results of a previous meta-analysis on structural abnormalities in ASD as well as with brain regions involved in human action observation/imitation. We found a single cluster in the left fusiform gyrus showing significantly reduced activation during face processing in ASD across all studies. Both task-dependent and task-independent analyses indicated significant functional connectivity of this region with the temporo-occipital and lateral occipital cortex, the inferior frontal and parietal cortices, the thalamus and the amygdala. Quantitative reverse inference then indicated an association of these regions mainly with face processing, affective processing, and language-related tasks. Moreover, we found that the cortex in the region of right area V5 displaying structural changes in ASD patients showed consistent connectivity with the region showing aberrant responses in the context of face processing. Finally, this network was also implicated in the human action observation/imitation network. In summary, our findings thus suggest a functionally and structurally disturbed network of occipital regions related primarily to face (but potentially also language) processing, which interact with inferior frontal as well as limbic regions and may be the core of aberrant face processing and reduced interest in faces in ASD.

Human face processing is tuned to sexual age preferences.

Human faces can motivate nurturing behaviour or sexual behaviour when adults see a child or an adult face, respectively. This suggests that face processing is tuned to detecting age cues of sexual maturity to stimulate the appropriate reproductive behaviour: either caretaking or mating. In paedophilia, sexual attraction is directed to sexually immature children. Therefore, we hypothesized that brain networks that normally are tuned to mature faces of the preferred gender show an abnormal tuning to sexual immature faces in paedophilia. Here, we use functional magnetic resonance imaging (fMRI) to test directly for the existence of a network which is tuned to face cues of sexual maturity. During fMRI, participants sexually attracted to either adults or children were exposed to various face images. In individuals attracted to adults, adult faces activated several brain regions significantly more than child faces. These brain regions comprised areas known to be implicated in face processing, and sexual processing, including occipital areas, the ventrolateral prefrontal cortex and, subcortically, the putamen and nucleus caudatus. The same regions were activated in paedophiles, but with a reversed preferential response pattern.

Neural responses to emotional expression information in high- and low-spatial frequency in autism: evidence for a cortical dysfunction

Despite an overall consensus that Autism Spectrum Disorder (ASD) entails atypical processing of human faces and emotional expressions, the role of neural structures involved in early facial processing remains unresolved. An influential model for the neurotypical brain suggests that face processing in the fusiform gyrus and the amygdala is based on both high-spatial frequency (HSF) information carried by a parvocellular pathway, and low-spatial frequency (LSF) information separately conveyed by a magnocellular pathway. Here, we tested the fusiform gyrus and amygdala sensitivity to emotional face information conveyed by these distinct pathways in ASD individuals (and matched Controls). During functional Magnetical Resonance Imaging (fMRI), participants reported the apparent gender of hybrid face stimuli, made by merging two different faces (one in LSF and the other in HSF), out of which one displayed an emotional expression (fearful or happy) and the other was neutral. Controls exhibited increased fusiform activity to hybrid faces with an emotional expression (relative to hybrids composed only with neutral faces), regardless of whether this was conveyed by LSFs or HSFs in hybrid stimuli. ASD individuals showed intact fusiform response to LSF, but not HSF, expressions. Furthermore, the amygdala (and the ventral occipital cortex) was more sensitive to HSF than LSF expressions in Controls, but exhibited an opposite preference in ASD. Our data suggest spared LSF face processing in ASD, while cortical analysis of HSF expression cues appears affected. These findings converge with recent accounts suggesting that ASD might be characterized by a difficulty in integrating multiple local information and cause global processing troubles unexplained by losses in low spatial frequency inputs.

Seeing Jesus in toast: Neural and behavioral correlates of face pareidolia

Face pareidolia is the illusory perception of non-existent faces. The present study, for the first time, contrasted behavioral and neural responses of face pareidolia with those of letter pareidolia to explore face-specific behavioral and neural responses during illusory face processing. Participants were shown pure-noise images but were led to believe that 50% of them contained either faces or letters; they reported seeing faces or letters illusorily 34% and 38% of the time, respectively. The right fusiform face area (rFFA) showed a specific response when participants “saw” faces as opposed to letters in the pure-noise images. Behavioral responses during face pareidolia produced a classification image (CI) that resembled a face, whereas those during letter pareidolia produced a CI that was letter-like. Further, the extent to which such behavioral CIs resembled faces was directly related to the level of face-specific activations in the rFFA. This finding suggests that the rFFA plays a specific role not only in processing of real faces but also in illusory face perception, perhaps serving to facilitate the interaction between bottom-up information from the primary visual cortex and top-down signals from the prefrontal cortex (PFC). Whole brain analyses revealed a network specialized in face pareidolia, including both the frontal and occipitotemporal regions. Our findings suggest that human face processing has a strong top-down component whereby sensory input with even the slightest suggestion of a face can result in the interpretation of a face.

Steady-state visual-evoked response to upright and inverted geometrical faces: A magnetoencephalography study

The face is one of the most important visual stimuli in human life, and inverted faces are known to elicit different brain responses than upright faces. This study analyzed steady-state visual-evoked magnetic fields (SSVEFs) in eleven healthy participants when they viewed upright and inverted geometrical faces presented at 6Hz. Steady-state visual-evoked responses are useful measurements and have the advantages of robustness and a high signal-to-noise ratio. Spectrum analysis revealed clear responses to both upright and inverted faces at the fundamental stimulation frequency (6Hz) and harmonics, i.e. SSVEFs. No significant difference was observed in the SSVEF amplitude at 6Hz between upright and inverted faces, which was different from the transient visual-evoked response, N170. On the other hand, SSVEFs were delayed with the inverted face in the right temporal area, which was similar to N170 and the results of previous steady-state visual-evoked potentials studies. These results suggest that different mechanisms underlie the larger amplitude and delayed latency observed with face inversion, though further studies are needed to fully elucidate these mechanisms. Our study revealed that SSVEFs, which have practical advantages for measurements, could provide novel findings in human face processing.

Neuroanatomic correlates of the feature-salience hierarchy in face processing: An fMRI -adaptation study

Previous fMRI studies suggest that faces are represented holistically in human face processing regions. On the other hand, behavioral studies have also shown that some facial features are more salient than others for face recognition: the neural basis of this feature-salience hierarchy is not known. We used fMRI-adaptation together with a behavioral discrimination task and an ideal observer analysis to ask (1) whether different face parts contribute different amounts to the neural signal in face responsive regions, and (2) whether this response correlates more with the behavioral performance of human subjects or with the physical properties of the face stimuli. Twenty-three subjects performed a same/different discrimination experiment to characterize their ability to detect changes to different face parts. The same subjects underwent an fMRI-adaptation study, in which limited portions of the faces were repeated or changed between alternating stimuli. The behavioral study showed high efficiency in identity discrimination when the whole face, top half, or eyes changed, and low efficiency when the bottom half, nose, or mouth changed. During fMRI, there was a release of adaptation in the right and left fusiform face area (FFA) with changes to the whole face, top face-half, or the eyes. Changes to the bottom half, nose or mouth did not result in a significant release of adaptation in the right FFA, although bottom-half changes resulted in a release of adaptation in the left FFA. Adaptation in the right and left FFA and the right pSTS was correlated with human perceptual efficiency but not with ideal observer measures of the physical image differences between face parts. The feature-salience hierarchy of human face perception is therefore reflected in the activity in the right and left FFA and right pSTS, further supporting the key role of these structures in our perceptual experience of faces.

Neural tuning of human face processing

Within the first 6 mo of life, our vision improves many times over and continues to improve during childhood (1). During these early years, we encounter multiple and varied sensory experiences, whose overall patterns can differ considerably between individuals. For example, some of us grow up among a large social group with many races and languages, whereas others grow within a smaller, more uniform cohort. Nevertheless, despite the multitude of environments in which we are raised, there is a surprising convergence in our behavioral capacities. One such capacity is recognizing individual faces. Like other visual skills, face recognition requires experience. Although we have a reasonable idea of when our perception of faces gets refined behaviorally and even some of its neural correlates (2, 3), we don’t really know the mechanisms by which experience tunes our face-processing circuits. Part of the problem, of course, is that one cannot do perturbation experiments in humans as we would with an animal model system (4). A new electrophysiology study by Roder et al. (5) capitalizes on a naturally occurring experiment: a unique clinical population of individuals born blind because of cataracts, who then later have their sight restored following cataract removal (5). The study revealed that the neural signature for face processing is present in these individuals but not sufficiently tuned by experience.

Sensitive periods for the functional specialization of the neural system for human face processing

The aim of the study was to identify possible sensitive phases in the development of the processing system for human faces. We tested the neural processing of faces in 11 humans who had been blind from birth and had undergone cataract surgery between 2 mo and 14 y of age. Pictures of faces and houses, scrambled versions of these pictures, and pictures of butterflies were presented while event-related potentials were recorded. Participants had to respond to the pictures of butterflies (targets) only. All participants, even those who had been blind from birth for several years, were able to categorize the pictures and to detect the targets. In healthy controls and in a group of visually impaired individuals with a history of developmental or incomplete congenital cataracts, the well-known enhancement of the N170 (negative peak around 170 ms) event-related potential to faces emerged, but a face-sensitive response was not observed in humans with a history of congenital dense cataracts. By contrast, this group showed a similar N170 response to all visual stimuli, which was indistinguishable from the N170 response to faces in the controls. The face-sensitive N170 response has been associated with the structural encoding of faces. Therefore, these data provide evidence for the hypothesis that the functional differentiation of category-specific neural representations in humans, presumably involving the elaboration of inhibitory circuits, is dependent on experience and linked to a sensitive period. Such functional specialization of neural systems seems necessary to archive high processing proficiency.

Humans Process Dog and Human Facial Affect in Similar Ways

Humans share aspects of their facial affect with other species such as dogs. Here we asked whether untrained human observers with and without dog experience are sensitive to these aspects and recognize dog affect with better-than-chance accuracy. Additionally, we explored similarities in the way observers process dog and human expressions. The stimulus material comprised naturalistic facial expressions of pet dogs and human infants obtained through positive (i.e., play) and negative (i.e., social isolation) provocation. Affect recognition was assessed explicitly in a rating task using full face images and images cropped to reveal the eye region only. Additionally, affect recognition was assessed implicitly in a lexical decision task using full faces as primes and emotional words and pseudowords as targets. We found that untrained human observers rated full face dog expressions from the positive and negative condition more accurately than would be expected by chance. Although dog experience was unnecessary for this effect, it significantly facilitated performance. Additionally, we observed a range of similarities between human and dog face processing. First, the facial expressions of both species facilitated lexical decisions to affectively congruous target words suggesting that their processing was equally automatic. Second, both dog and human negative expressions were recognized from both full and cropped faces. Third, female observers were more sensitive to affective information than were male observers and this difference was comparable for dog and human expressions. Together, these results extend existing work on cross-species similarities in facial emotions and provide evidence that these similarities are naturally exploited when humans interact with dogs.

Human Face Processing: From Recognition to Emotion

Human Face Processing: From Recognition to Emotion

Faces in Motion: Selectivity of Macaque and Human Face Processing Areas for Dynamic Stimuli

Face recognition mechanisms need to extract information from static and dynamic faces. It has been hypothesized that the analysis of dynamic face attributes is performed by different face areas than the analysis of static facial attributes. To date, there is no evidence for such a division of labor in macaque monkeys. We used fMRI to determine specializations of macaque face areas for motion. Face areas in the fundus of the superior temporal sulcus responded to general object motion; face areas outside of the superior temporal sulcus fundus responded more to facial motion than general object motion. Thus, the macaque face-processing system exhibits regional specialization for facial motion. Human face areas, processing the same stimuli, exhibited specializations for facial motion as well. Yet the spatial patterns of facial motion selectivity differed across species, suggesting that facial dynamics are analyzed differently in humans and macaques.

Effects of aging on face identification and holistic face processing

Several studies have shown that face identification accuracy is lower in older than younger adults. This effect of aging might be due to age differences in holistic processing, which is thought to be an important component of human face processing. Currently, however, there is conflicting evidence as to whether holistic face processing is impaired in older adults. The current study therefore re-examined this issue by measuring response accuracy in a 1-of-4 face identification task and the composite face effect (CFE), a common index of holistic processing, in older adults. Consistent with previous reports, we found that face identification accuracy was lower in older adults than in younger adults tested in the same task. We also found a significant CFE in older adults that was similar in magnitude to the CFE measured in younger subjects with the same task. Finally, we found that there was a significant positive correlation between the CFE and face identification accuracy. This last result differs from the results obtained in a previous study that used the same tasks and which found no evidence of an association between the CFE and face identification accuracy in younger adults. Furthermore, the age difference was found with subtraction-, regression-, and ratio-based estimates of the CFE. The current findings are consistent with previous claims that older adults rely more heavily on holistic processing to identify objects in conditions of limited processing resources.