Face perception

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Several other studies indicate that early perceptual experience is crucial to the development of capacities characteristic of adult visual perception, including the ability to identify familiar people and to recognize and comprehend facial expressions.[11] The capacity to discern between faces, much like language, appears to have a broad potential in early life that is whittled down to kinds of faces that are experienced in early life.[11] Infants can discern between macaque faces at six months of age, but, without continued exposure, cannot at nine months of age. Being shown photographs of macaques during this three-month period gave nine-month-olds the ability to reliably distinguish between unfamiliar macaque faces.[12]

The neural substrates of face perception in infants are likely similar to those of adults, but the limits of imaging technology that are feasible for use with infants currently prevent very specific localization of function as well as specific information from subcortical areas[13] like the amygdala, which is active in the perception of facial expression in adults.[11] In a study on healthy adults, it was shown that faces are likely to be processed, in part, via a retinotectal (subcortical) pathway.[14]

However, there is activity near the fusiform gyrus,[13] as well as in occipital areas.[7] when infants are exposed to faces, and it varies depending on factors including facial expression and eye gaze direction.[3][7]



One of the most widely accepted theories of face perception argues that understanding faces involves several stages:[15] from basic perceptual manipulations on the sensory information to derive details about the person (such as age, gender or attractiveness), to being able to recall meaningful details such as their name and any relevant past experiences of the individual.





Neuroanatomy of facial processing[edit]

There are several parts of the brain that play a role in face perception. Rossion, Hanseeuw, and Dricot[17] used BOLD fMRI mapping to identify activation in the brain when subjects viewed both cars and faces. The majority of BOLD fMRI studies use blood oxygen level dependent (BOLD) contrast to determine which areas of the brain are activated by various cognitive functions.[18] They found that the occipital face area, located in the occipital lobe, the fusiform face area, the superior temporal sulcus, the amygdala, and the anterior/inferior cortex of the temporal lobe, all played roles in contrasting the faces from the cars, with the initial face perception beginning in the fusiform face area and occipital face areas. This entire region links to form a network that acts to distinguish faces. The processing of faces in the brain is known as a "sum of parts" perception.[19] However, the individual parts of the face must be processed first in order to put all of the pieces together. In early processing, the occipital face area contributes to face perception by recognizing the eyes, nose, and mouth as individual pieces.[20] Furthermore, Arcurio, Gold, and James[21] used BOLD fMRI mapping to determine the patterns of activation in the brain when parts of the face were presented in combination and when they were presented singly. The occipital face area is activated by the visual perception of single features of the face, for example, the nose and mouth, and preferred combination of two-eyes over other combinations. This research supports that the occipital face area recognizes the parts of the face at the early stages of recognition. On the contrary, the fusiform face area shows no preference for single features, because the fusiform face area is responsible for "holistic/configural" information,[22] meaning that it puts all of the processed pieces of the face together in later processing. This theory is supported by the work of Gold et al.[19] who found that regardless of the orientation of a face, subjects were impacted by the configuration of the individual facial features. Subjects were also impacted by the coding of the relationships between those features. This shows that processing is done by a summation of the parts in the later stages of recognition.



The OFA is located in the inferior occipital gyrus.[24] Similar to the FFA, this area is also active during successful face detection and identification, a finding that is supported by fMRI activation.[23] The OFA is involved and necessary in the analysis of facial parts but not in the spacing or configuration of facial parts. This suggests that the OFA may be involved in a facial processing step that occurs prior to the FFA processing.[23]

The fSTS is involved in recognition of facial parts and is not sensitive to the configuration of these parts. It is also thought that this area is involved in gaze perception.[26] The fSTS has demonstrated increased activation when attending to gaze direction.[23]

Bilateral activation is generally shown in all of these specialized facial areas.[27][28][29][30][31][32] However, there are some studies that include increased activation in one side over the other. For instance McCarthy (1997) has shown that the right fusiform gyrus is more important for facial processing in complex situations.[25]


It is important to note that while certain areas respond selectively to faces, facial processing involves many neural networks. These networks include visual and emotional processing systems as well. Emotional face processing research has demonstrated that there are some of the other functions at work. While looking at faces displaying emotions (especially those with fear facial expressions) compared to neutral faces there is increased activity in the right fusiform gyrus. This increased activity also correlates with increased amygdala activity in the same situations.[34] The emotional processing effects observed in the fusiform gyrus are decreased in patients with amygdala lesions.[34] This demonstrates possible connections between the amygdala and facial processing areas.[34]


Ishai and colleagues have proposed the object form topology hypothesis, which posits that there is a topological organization of neural substrates for object and facial processing.[35] However, Gauthier disagrees and suggests that the category-specific and process-map models could accommodate most other proposed models for the neural underpinnings of facial processing.[36]

Most neuroanatomical substrates for facial processing are perfused by the middle cerebral artery (MCA). Therefore, facial processing has been studied using measurements of mean cerebral blood flow velocity in the middle cerebral arteries bilaterally. During facial recognition tasks, greater changes in the right middle cerebral artery (RMCA) than the left (LMCA) have been observed.[37][38] It has been demonstrated that men were right lateralized and women left lateralized during facial processing tasks.[39]

Just as memory and cognitive function separate the abilities of children and adults to recognize faces, the familiarity of a face may also play a role in the perception of faces.[19] Zheng, Mondloch, and Segalowitz recorded event-related potentials in the brain to determine the timing of recognition of faces in the brain.[40] The results of the study showed that familiar faces are indicated and recognized by a stronger N250,[40] a specific wavelength response that plays a role in the visual memory of faces.[41] Similarly, Moulson et al.[42] found that all faces elicit the N170 response in the brain.

Hemispheric asymmetries in facial processing capability[edit]


Studies using electrophysiological techniques have demonstrated gender-related differences during a face recognition memory (FRM) task and a facial affect identification task (FAIT). The male subjects used a right, while the female subjects used a left, hemisphere neural activation system in the processing of faces and facial affect.[43] Moreover, in facial perception there was no association to estimated intelligence, suggesting that face recognition performance in women is unrelated to several basic cognitive processes.[44] Gender-related differences[45] may suggest a role for sex hormones. In females there may be variability for psychological functions[46] related to differences in hormonal levels during different phases of the menstrual cycle.[47]

Data obtained in norm and in pathology support asymmetric face processing.[48][49][50] Gorno-Tempini and others in 2001, suggested that the left inferior frontal cortex and the bilateral occipitotemporal junction respond equally to all face conditions. Some neuroscientists contend that both the left inferior frontal cortex (Brodmann area 47) and the occipitotemporal junction are implicated in facial memory.[51][52][53] The right inferior temporal/fusiform gyrus responds selectively to faces but not to non-faces. The right temporal pole is activated during the discrimination of familiar faces and scenes from unfamiliar ones.[54] Right asymmetry in the mid temporal lobe for faces has also been shown using 133-Xenon measured cerebral blood flow (CBF).[55] Other investigators have observed right lateralization for facial recognition in previous electrophysiological and imaging studies.[56]

The implication of the observation of asymmetry for facial perception would be that different hemispheric strategies would be implemented. The right hemisphere would be expected to employ a holistic strategy, and the left an analytic strategy.[57][58][59][60] In 2007, Philip Njemanze, using a novel functional transcranial Doppler (fTCD) technique called functional transcranial Doppler spectroscopy (fTCDS), demonstrated that men were right lateralized for object and facial perception, while women were left lateralized for facial tasks but showed a right tendency or no lateralization for object perception.[61] Njemanze demonstrated using fTCDS, summation of responses related to facial stimulus complexity, which could be presumed as evidence for topological organization of these cortical areas in men. It may suggest that the latter extends from the area implicated in object perception to a much greater area involved in facial perception.



Cognitive neuroscience[edit]


Studies by Gauthier have shown that an area of the brain known as the fusiform gyrus (sometimes called the "fusiform face area, (FFA)" because it is active during face recognition) is also active when study participants are asked to discriminate between different types of birds and cars,[62] and even when participants become expert at distinguishing computer generated nonsense shapes known as greebles.[63] This suggests that the fusiform gyrus may have a general role in the recognition of similar visual objects. Yaoda Xu, then a post doctoral fellow with Nancy Kanwisher, replicated the car and bird expertise study using an improved fMRI design that was less susceptible to attentional accounts.





Face advantage in memory recall[edit]

During face perception, neural networks make connections with the brain to recall memories.[69] According to the Seminal Model of face perception, there are three stages of face processing including recognition of the face, the recall of memories and information that are linked with that face, and finally name recall.[69][70] There are, however, exceptions to this order. For example, names are recalled faster than semantic information in cases of highly familiar stimuli.[71] While the face is powerful identifier of individuals, the voice also helps in the recognition of people and is an identifier for important information.[72][73]

Research has been conducted to see if faces or voices make it easier to identify individuals and recall semantic memory and episodic memory.[74] These experiments look at all three stages of face processing. The experiment method was to show two groups celebrity and familiar faces or voices with a between-group design and ask the participants to recall information about them.[74] The participants are first asked if the stimulus is familiar. If they answer yes then they are asked to information (semantic memory) and memories they have of the person (episodic memory) that fits the face or voice presented. These experiments all demonstrate the strong phenomenon of the face advantage and how it persists through different follow-up studies with different experimental controls and variables.[74]

Recognition-performance issue[edit]

After the first experiments on the advantage of faces over voices in memory recall, errors and gaps were found in the methods used.[74] For one, there was not a clear face advantage for the recognition stage of face processing. Participants showed a familiarity-only response to voices more often than faces.[75] In other words, when voices were recognized (about 60-70% of the time) they were much harder to recall biographical information but very good at being recognized.[74] The results were looked at as remember versus know judgements. A lot more remember results (or familiarity) occurred with voices, and more know (or memory recall) responses happened with faces.[73] This phenomenon persists through experiments dealing with criminal line-ups in prisons. Witnesses are more likely to say that a suspect's voice sounded familiar than his/her face even though they cannot remember anything about the suspect.[76] This discrepancy is due to a larger amount of guesswork and false alarms that occur with voices.[73]

To give faces a similar ambiguity to that of voices, the face stimuli were blurred in the follow-up experiment[75] This experiment followed the same procedures as the first, presenting two groups with sets of stimuli made up of half celebrity faces and half unfamiliar faces.[74] The only difference was that the face stimuli were blurred so that detailed features could not be seen. Participants were than asked to say if they recognized the person, if they could recall specific biographical information about the them, and finally if they knew the person's name. The results were completely different from those of the original experiment, supporting the view that there were problems in the first experiment's methods.[74] According to the results of the followup, the same amount of information and memory could be recalled through voices and faces, dismantling the face advantage. However, these results are flawed and premature because other methodological issues in the experiment still needed to be fixed.[74]

Content of speech[edit]

The process of controlling the content of speech extract has proven to be more difficult than the elimination of non facial cues in photographs.[74] Thus the findings of experiments that did not control this factor lead to misleading conclusions regarding the voice recognition over the face recognition.[74] For example, in an experiment it was found that 40% of the time participants could easily pair the celebrity-voice with their occupation just by guessing.[75] In order to eliminate these errors, experimenters removed parts of the voice samples that could possibly give clues to the identity of the target, such as catchphrases.[77] Even after controlling the voice samples as well as the face samples (using blurred faces), studies have shown that semantic information can be more accessible to retrieve when individuals are recognizing faces than voices.[78]

Another technique to control the content of the speech extracts is to present the faces and voices of personally familiar individuals, like the participant's teachers or neighbors, instead of the faces and voices of celebrities.[74] In this way alike words are used for the speech extracts.[74] For example, the familiar targets are asked to read exactly the same scripted speech for their voice extracts. The results showed again that semantic information is easier to retrieve when individuals are recognizing faces than voices.[74]

Frequency-of-exposure issue[edit]

Another factor that has to be controlled in order for the results to be reliable is the frequency of exposure.[74] If we take the example of celebrities, people are exposed to celebrities' faces more often than their voices because of the mass media.[74] Through magazines, newspapers and the Internet, individuals are exposed to celebrities' faces without their voices on an everyday basis rather than their voices without their faces.[74] Thus, someone could argue that for all of the experiments that were done until now the findings were a result of the frequency of exposure to the faces of celebrities rather than their voices.[79]

To overcome this problem researchers decided to use personally familiar individuals as stimuli instead of celebrities.[74] Personally familiar individuals, such as participant's teachers, are for the most part heard as well as seen.[80] Studies that used this type of control also demonstrated the face advantage.[80] Students were able to retrieve semantic information more readily when recognizing their teachers faces (both normal and blurred)rather their voices.[78]

However, researchers over the years have found an even more effective way to control not only the frequency of exposure but also the content of the speech extracts, the associative learning paradigm.[74] Participants are asked to link semantic information as well as names with pre-experimentally unknown voices and faces.[81][82] In a current experiment that used this paradigm, a name and a profession were given together with, accordingly, a voice, a face or both to three participant groups.[81] The associations described above were repeated four times.[81] The next step was a cued recall task in which every stimulus that was learned in the previous phase was introduced and participants were asked to tell the profession and the name for every stimulus.[81][83] Again, the results showed that semantic information can be more accessible to retrieve when individuals are recognizing faces than voices even when the frequency of exposure was controlled.[74][81]

Extension to episodic memory and explanation for existence[edit]

Episodic memory is our ability to remember specific, previously experienced events.[84] In recognition of faces as it pertains to episodic memory, there has been shown to be activation in the left lateral prefrontal cortex, parietal lobe, and the left medial frontal/anterior cingulate cortex.[85][86] It was also found that a left lateralization during episodic memory retrieval in the parietal cortex correlated strongly with success in retrieval.[85] This may possibly be due to the hypothesis that the link between face recognition and episodic memory were stronger than those of voice and episodic memory.[75] This hypothesis can also be supported by the existence of specialized face recognition devices thought to be located in the temporal lobes.[85][87] There is also evidence of the existence of two separate neural systems for face recognition: one for familiar faces and another for newly learned faces.[85] One explanation for this link between face recognition and episodic memory is that since face recognition is a major part of human existence, the brain creates a link between the two in order to be better able to communicate with others.[88]


Main article: cross-race effect

Differences in own- versus other-race face recognition and perceptual discrimination was first researched in 1914[89] Humans tend to perceive people of other races than their own to all look alike:


This phenomenon is known as the cross-race effect, own-race effect, other-race effect, own race bias or interracial-face-recognition-deficit.[90] The effect occurs as early as 170ms in the brain with the N170 brain response to faces.[91][clarification needed]



In addition to other-race contact, there is suggestion that the own-race effect is linked to increased ability to extract information about the spatial relationships between different features.[96] Richard Ferraro writes that facial recognition is an example of a neuropsychological measure that can be used to assess cognitive abilities that are salient within African-American culture.[97] Daniel T. Levin writes that the deficit occurs because people emphasize visual information specifying race at the expense of individuating information when recognizing faces of other races.[98] Further research using perceptual tasks could shed light on the specific cognitive processes involved in the other-race effect.[95] The question if the own-race effect can be overcome was already indirectly answered by Ekman & Friesen in 1976 and Ducci, Arcuri, Georgis & Sineshaw in 1982. They had observed that people from New Guinea and Ethiopia who had had contact with white people before had a significantly better emotional recognition rate.


In individuals with autism spectrum disorder[edit]

Autism spectrum disorder (ASD) is a comprehensive neural developmental disorder that produces many deficits including social, communicative,[100] and perceptual deficits.[101] Of specific interest, individuals with autism exhibit difficulties in various aspects of facial perception, including facial identity recognition and recognition of emotional expressions.[102][103] These deficits are suspected to be a product of abnormalities occurring in both the early and late stages of facial processing.[104]

Speed and methods[edit]

People with ASD process face and non-face stimuli with the same speed.[104][105] In typically developing individuals, there is a preference for face processing, thus resulting in a faster processing speed in comparison to non-face stimuli.[104][105] These individuals primarily utilize holistic processing when perceiving faces.[101] Contrastingly, individuals with ASD employ part-based processing or bottom-up processing, focusing on individual features rather than the face as a whole.[106][107] When focusing on the individual parts of the face, persons with ASD direct their gaze primarily to the lower half of the face, specifically the mouth, varying from the eye trained gaze of typically developing people.[106][107][108][109][110] This deviation from holistic face processing does not employ the use of facial prototypes, which are templates stored in memory that make for easy retrieval.[103][111]

Additionally, individuals with ASD display difficulty with recognition memory, specifically memory that aids in identifying faces. The memory deficit is selective for faces and does not extend to other objects or visual inputs.[103] Some evidence lends support to the theory that these face-memory deficits are products of interference between connections of face processing regions.[103]

Associated difficulties[edit]

The atypical facial processing style of people with ASD often manifests in constrained social ability, due to decreased eye contact, joint attention, interpretation of emotional expression, and communicative skills.[112] These deficiencies can be seen in infants as young as 9 months; specifically in terms of poor eye contact and difficulties engaging in joint attention.[104] Some experts have even used the term 'face avoidance' to describe the phenomena where infants who are later diagnosed with ASD preferentially attend to non-face objects over faces.[100] Furthermore, some have proposed that the demonstrated impairment in children with ASD's ability to grasp emotional content of faces is not a reflection of the incapacity to process emotional information, but rather, the result of a general inattentiveness to facial expression.[100] The constraints of these processes that are essential to the development of communicative and social-cognitive abilities are viewed to be the cause of impaired social engagement and responsivity.[113] Furthermore, research suggests that there exists a link between decreased face processing abilities in individuals with ASD and later deficits in Theory of Mind; for example, while typically developing individuals are able to relate others' emotional expressions to their actions, individuals with ASD do not demonstrate this skill to the same extent.[114]

There is some contention about this causation however, resembling the chicken or the egg dispute. Others theorize that social impairment leads to perceptual problems rather than vice versa.[106] In this perspective, a biological lack of social interest inherent to ASD inhibits developments of facial recognition and perception processes due to underutilization.[106] Continued research is necessary to determine which theory is best supported.



Studies are not conclusive as to which brain areas people with ASD use instead. One study found that, when looking at faces, people with ASD exhibit activity in brain regions normally active when typically developing individuals perceive objects.[106] Another study found that during facial perception, people with ASD use different neural systems, with each one of them using their own unique neural circuitry.[115]

Compensation mechanisms[edit]

As ASD individuals age, scores on behavioral tests assessing ability to perform face-emotion recognition increase to levels similar to controls.[104][116] Yet, it is apparent that the recognition mechanisms of these individuals are still atypical, though often effective.[116] In terms of face identity-recognition, compensation can take many forms including a more pattern-based strategy which was first seen in face inversion tasks.[109] Alternatively, evidence suggests that older individuals compensate by using mimicry of other’s facial expressions and rely on their motor feedback of facial muscles for face emotion-recognition.[117] These strategies help overcome the obstacles individuals with ASD face in interacting within social contexts.




See also[edit]

Further reading[edit]



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