Running head: age face perception

Age-differences in face perception:

A review of N170 event-related potential studies

Simon van Rysewyk*

University of Tasmania

van Rysewyk, S. (2012). Age-differences in face perception: A review of N170 event-related

potential studies. In: A. Freitas-Magalhães (Ed.), ‘Emotional Expression: The Brain and the

Face’ (V. III, Second Series), University of Fernando Pessoa Press, Oporto.

*Correspondence should be sent to: Simon van Rysewyk, School of Psychology, University
of Tasmania, Private Bag 30, Hobart, Tasmania 7001, Australia.
Email: simon.vanrysewyk@utas.edu.au Phone: +61 3 6226 2661

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Running head: age face perception

Abstract

Age-differences in face perception have been widely reported in experimental studies for

several decades. One explanatory theory proposes that age-differences in higher-level

perceptual processes partly explain declined face perception in healthy older adults.

Plausibly, changes in face perception may indicate general age-differences in visual

perception, and may account for impairments in face recognition. Several event-related

potential studies have used the N170-effect to investigate age-differences in the perception of

specific information in the face. This chapter describes and assesses the impact of these

studies on the explanatory theory described above.

Keywords: healthy aging, human face, face perception, event-related potential, N170-effect

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Running head: age face perception

Introduction

Age-differences in face perception have been widely reported in experimental studies

for several decades. Behavioural age-differences include a higher proportion of false alarms

to unfamiliar faces in healthy older adults (HOAs) (e.g., Bartlett & Fulton, 1989), and

declined memory for faces compared to the memory of objects (Boutet & Faubert, 2006).

Neuroscientific studies using electroencephalography (EEG) (e.g., Pfutze et al. 2002) and

functional magnetic resonance imaging (fMRI) (e.g., Grady et al., 2007) also reliably

demonstrate declined neural processing of faces in HOAs (e.g., Lott et al., 2005; Searcy et al.,

1996). Why is this?

One explanation identifies the origin of age-differences in face perception in reduced

efficiency of visual perception (e.g., Bian & Andersen, 2008; Fiorentini et al., 1996). There is

evidence that declined low-level visual functions such as acuity (Tejeria et al., 2002) and

contrast sensitivity is correlated with age-differences in face perception. For example,

Owsley et al. (1981) showed that increasing the contrast of faces improved face perception

(matching unfamiliar faces) in HOAs. However, other studies using healthy younger adults

(HYAs) have revealed that face perception relies on middle-range spatial frequencies for

which sensitivity is very high and the perception of which should be influenced by low-level

vision (Costen et al. 1996, for review). Although decreased contrast sensitivity may be a

factor in age-differences of face perception, other factors must also be involved.

Another explanation proposes that age-differences in perceptual processes partly

explain declined face perception in HOAs. These perceptual processes include declined

neural specialization for faces (e.g., Payer et al., 2006), changes in face-scanning strategies

(Firestone et al., 2007), reduced categorical perception (Kiffel et al., 2005), declined

spatiotemporal integration (Norton et al., 2009), impaired sensory processing speed

(Salthouse, 1996, 2000), declined attention and working memory capacity (e.g., Lange &

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Running head: age face perception

Verhaeghen, 2009), increased distractibility (e.g., Healey et al., 2008), declined prefrontal

inhibitory control (e.g., Chao and Knight, 1997), reduced cognitive flexibility as revealed by

a general decline in task switching performance (Reimers & Maylor, 2005), and declined

ability to rapidly convert perceptual representations into familiar templates (Habak et al.,

2008). Plausibly, changes in face perception may indicate general age-differences in visual

perception, and may explain impairments in face perception (e.g., Bentin et al., 2007).

Several studies have therefore investigated whether age-differences in face perception

relate to age-differences in the perception of specific information in faces. Based on studies

using HYAs, four types of visual information are perceived in the (upright) human face. The

face as a basic sensory category relies on perception of its general profile, or ‘global face

perception’ (Bentin et al. 2006). The general face profile and intrinsic face-parts (i.e., eyes,

nose and mouth) perceived as a unified whole is termed ‘holistic face perception’ (e.g.,

Tanaka & Farah, 1993). Spatial relations between intrinsic face-parts and of these parts to the

face profile are classed as ‘configural face perception’ (e.g., Rhodes et al. 1993). The

perception of intrinsic face-part shape and surface qualities rely on ‘featural face perception’

(Bentin et al. 2006) (see Table 1).

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Insert Table 1 here

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Since all faces share the same profile (two eyes above a nose above a mouth), the

perception of individual faces is based on the processing of individual features (the shape of

the eyes, nose and mouth) and on the processing of the configuration of these features (the

distance between the eyes, or between the mouth and nose, and so on). The information

perceived in a face may constitute a processing ‘style’ unique to human faces not available

for non-face objects (Yin, 1969).

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Running head: age face perception

The N170 ERP and age-differences in face perception

An experiment designed to study age-differences in face perception must be able to

measure processes specific to face perception (hypothesized to be reduced and slower in

HOAs than HYAs), and processes not age-related. The method chosen must also be capable

of measuring the continuous sequence of processes involved in face perception. Reaction

time and accuracy methods by their nature are unsuitable to study continuous processes in

face perception since they can only provide intermittent (i.e., non-continuous) measures. The

method typically used to address this problem in cognitive research is the event-related

potential (ERP).

ERPs are continuous changes of averaged EEG cortical brain activity. They are time-

locked to the presentation of a sensory event, such as the central presentation of a visual

stimulus to an experimental subject. Research has identified a series of ERPs distinguished by

scalp distribution and peak latency and correlated with different cognitive neural mechanisms.

This review considers the N170 ERP.

The N170 is a negative ERP peaking between 150 and 180 ms from stimulus onset.

This response is maximal over occipito-temporal electrode sites. This response is consistent

with a neural source at the fusiform and inferior-temporal gyri. The N170 shows greater

amplitudes to faces than to any other stimulus category (e.g., non-face objects), and is

correlated with early face perceptual processes (Rossion & Jacques, 2008, for review). The

N170-effect is greater and slightly delayed when intrinsic face-parts (especially eyes) are

presented outside the face profile (e.g., Bentin et al. 2006) as well as by face inversion

(Bentin et al., 1996; Rossion et al., 2000). Since both face inversion and the isolation of

components from the face profile both impair configural processing, the N170-effect

observed in these conditions suggests that it is evoked even when the face-specific structural

encoding is interrupted. Accordingly, Bentin and colleagues have proposed that the N170

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Running head: age face perception

represents a neural mechanism sensitive to faces (or face-related information) and initiates

the perceptual processes to individuate faces (Bentin et al., 1996; Bentin et al., 2006; Sagiv &

Bentin, 2001; Zion-Golumbic & Bentin, 2007). This explanatory theory proposes that the

latency delay in some conditions is due to an increased difficulty to relate a visual stimulus to

the faces, when relevant information is spatially distorted or missing.

Can the N170-effect reveal age-differences in face perception? The earliest studies of

this question are Pfütze et al. (2002) and Chaby et al. (2003). Pfütze et al. (2002) did not find

any N170 age-differences stimulated by either famous or unfamiliar faces. In addition, the

authors found a shift from a right-hemisphere (RH) N170 distribution in the HYAs, towards

greater hemispheric symmetry in the HOAs. Chaby et al. (2003) found that N170 latency was

not affected by healthy aging, but its amplitude was higher for HOAs than HYAs and did not

display the typical larger RH than left hemisphere (LH) asymmetry. Unfortunately, these

studies are both limited since they did not use a non-face control condition. The very

sensitivity of the N170 to faces is characterized by the difference between its amplitude in

response to faces compared to other objects (e.g., Bentin et al. 1996). Investigating age-

differences in face perception as shown by the N170 should therefore assess the N170-effect,

rather than the N170 amplitude alone. Several studies have taken the N170-effect to

investigate whether there are age-differences in the perception of information in faces. This

remainder of this chapter now reviews and evaluates these studies.

Age-differences in basic perceptual sensitivity to faces

The phrase ‘basic perceptual sensitivity’ refers to the detection of a face by the

perceptual system as a face and not as a non-face object. Only two studies have assessed age-

differences in basic perceptual sensitivity to faces using the N170-effect (Gao et al. 2009;

Daniel & Bentin, 2010). Both studies used an oddball paradigm in which faces and non-face

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Running head: age face perception

objects (tables in Gao et al. 2009, wristwatches in Daniel and Bentin, 2010) were presented

sequentially. Participants were instructed to press a button each time the target appeared on

the screen (a butterfly in Gao et al. 2009, a flower in Daniel and Bentin, 2010).

Gao et al. (2009) did not make an experimental hypothesis per se, but reasoned that a

basic face-specific effect in HOAs should stimulate a N170-effect evoked by faces and

objects. Gao et al. (2009) found that the N170 was significantly larger in the HOAs than in

the HYAs, but the peak latency of the N170 did not differ between groups. The N170

amplitude was larger for faces than tables, but the effect did not interact with age. Finally, the

normalized N170-effect was larger over the RH than the LH, and was RH dominant in both

groups. Hence, basic perceptual sensitivity of faces is spared in HOAs.

Daniel and Bentin (2010) hypothesized that while both groups would show higher

N170 amplitudes and shorter latencies to faces than to watches, the N170 amplitudes would

be larger in the HOAs than the HYAs. Consistent with available behavioral data, Daniel and

Bentin (2010) found general age-related slowing of perceptual speed in performance (e.g.,

Boutet and Faubert, 2006; Salthouse, 1996) as well as in N170 quantities (e.g., Rousselet et

al., 2009). Consistent with Gao et al. (2009), the authors found that the basic ability to

perceive faces as measured by the N170 was comparable in HOAs and HYAs, although

HOAs were slower than HYAs. Both groups exhibited higher N170 amplitudes and shorter

latencies to faces than to watches. Consistent with Gao et al. (2009), the N170 amplitudes

were generally larger in the HOAs compared to the HYAs.

Unlike Gao et al. (2009), no brain hemispheric differences were found for the HOAs,

whereas the N170 amplitudes were higher at RH than LH sites for HYAs. The finding of

Daniel and Bentin (2010) challenges studies proposing a general age-related loss of neural

specialization in the ventral extrastriate system (e.g., Park et al., 2004). Instead, neural

representations in the ventral visual cortex may become less domain-specific with healthy

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Running head: age face perception

aging. Although this experiment of basic sensitivity to faces demonstrated age-related

slowing of visual processing, it also showed that the ability of the perceptual system to

distinguish faces from non-face objects stimuli is preserved in old age.

Age-differences in global face perception

The perception of the human face as a basic sensory category is sufficiently informed

by its general profile (‘global’ face perception). Global face perception implies an integrated

percept without detailed analysis of its components. ERP studies have tested this sufficiency

claim in HYAs by measuring N170 amplitude to faces with intrinsic parts in normal (i.e.,

‘unscrambled’) or abnormal (i.e., ‘scrambled’) spatial face locations. Significant N170-

effects to normal schematic faces (Sagiv & Bentin, 2001), but not to scrambled schematic

faces (Bentin & Golland, 2002) were found in HYAs. However, these studies do not support

the sufficiency hypothesis since the intrinsic parts of schematic faces were not perceived as

such at scrambled spatial locations (i.e., they were not perceived as intrinsic face-parts when

scrambled). This question was resolved by Zion-Golumbic and Bentin (2007) who found

significant N170-effects in normal and scrambled faces when the intrinsic face-parts were

perceived by HYAs. This result implies that the global face structure is not a necessary

condition for activating perceptual neural mechanisms.

Daniel and Bentin (2010) compared the sensitivity of perceptual mechanisms

controlling the N170 to normal and scrambled faces in HOAs and HYAs. The authors

predicted that the N170 evoked by scrambled faces in HOAs would be reduced compared to

normal faces based on the hypothesis that HOAs rely more on global face perception. This

claim partly derives from findings that HYAs do not differentially perceive normal and

scrambled faces. Overall, the results of Daniel and Bentin (2010) showed that behaviour and

electrophysiological measurements were slower in the HOAs. Concerning the ERP measures,

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Running head: age face perception

the authors found that scrambling global face structure reduced the N170 amplitude in HOAs

compared to HYAs, as expected. This implies that HOAs are more sensitive to global face

perception than HYAs. However, the latency of the N170 was delayed for scrambled faces in

both groups. Why is this? Since HOAs are more sensitive to global faces, processing intrinsic

face-parts would be slowed in this population. Moreover, the authors found that presenting

normal and scrambled face-parts outside of the face profile increased the N170 in the HYAs

compared to the HOAs. The increase in the N170 amplitude in response to isolated face-parts

in HYAs has been shown in many studies (e.g., Itier et al., 2006, 2007). These findings show

that first-order relations (i.e., global face perception) are more essential in HOAs for face

perception and for activating face-correlated neural mechanisms than HYAs. HOAs are less

effective and less efficient than HYAs in integrating intrinsic face-parts into the global face

profile.

Daniel and Bentin (2010) also found a symmetrical distribution of the N170 across

both brain hemispheres in the HOAs compared to a dominant RH N170-effect in the HYAs.

The symmetrical distribution of the N170 in the HOAs was not caused by a reduction of RH

amplitudes but by greater N170 amplitudes at LH electrode sites (compared to the HYAs).

This finding is compatible with reduced hemispheric asymmetry found in HOAs (e.g.,

Cabeza, 2002; Gao et al., 2009; Pfutze et al., 2002), but incompatible with the theory that

proposes LH control of perceptual details, and RH control of global percepts (e.g., Weissman

& Woldorff, 2005). The activation of the LH in face perception in HOAs suggests that the

relative larger decline of visuospatial compared with verbal abilities (e.g., Goldstein & Shelly,

1981) indicates factors other than faster aging of the RH are involved (see also Bentin et al.,

1981).

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Running head: age face perception

Age-differences in holistic face perception

The general face profile and intrinsic face-parts (i.e., eyes, nose and mouth) perceived

as a unified whole is termed ‘holistic face perception’. Holistic face perception proposes that

intrinsic face-parts are not represented in isolation but only as a unitary representation of the

face as a whole (Tanaka & Farah, 1993). In the landmark study establishing this effect,

Tanaka and Farah (1993) found that perception of general face profiles was superior to

intrinsic face-parts in upright compared to inverted positions. This indicates that inverted

faces are perceived according to intrinsic face-parts, not the face as a whole. Scrambled faces

and non-face objects also do not show this effect. This may mean that such visual stimuli are

perceived according to isolated features. Holistic perception may be important for face

perception because (upright) faces are homogeneous and are perceived individually.

No ERP studies have used the N170-effect to investigate holistic face perception.

However, Boutet and Faubert (2006) conducted a behavioural study. In Experiment 4, Boutet

and Faubert (2006) presented HYAs and HOAs with a studied face, followed by a test

window showing either a target and a distracter face, or a target and a distracter intrinsic face-

part. The visual stimuli were either upright or inverted. The authors found superior perception

of general face profiles compared to intrinsic face-parts in upright and inverted conditions in

both HOAs and HYAs. Moreover, the difference between the two conditions was greater for

the upright than inverted condition. Boutet and Faubert (2006) concluded that the ability to

apply holistic processing is intact in HOAs.

However, this inference may be premature since the HOAs in Boutet and Faubert

(2006) did not show the composite effect (Experiment 2), which is also a measure of holistic

processing (Young et al., 1987). The composite effect is observed when perception of

composite faces is inferior to that of non-composites in the upright compared to the inverted

position (Young et al., 1987). The absence of the composite effect in HOAs in Boutet and

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Running head: age face perception

Faubert (2006) may signify a more general age-related reduction in face profile integration

than holistic perception (Roudaia et al., 2008). Further behavioral and neuroscientific

research is needed to investigate this possibility.

Age-differences in configural face perception

Spatial relations between intrinsic face-parts and of these parts to the face profile are

grouped as ‘configural face perception’ (e.g., Rhodes et al. 1993). Configural analysis is

based on distinction among the details of the face and a relational perception of their spatial

location. The face inversion effect (FIE) is a signature of configural face perception. Face

perception in HYAs is more accurate when faces are presented (and tested) upright than

inverted (Leder et al. 2001). The inversion effect is larger for faces than for any other non-

face object (Yin, 1969), and thus has been termed a ‘disproportionate inversion effect’.

The FIE is defined by an interaction between stimulus category (i.e., face or non-face

object) and orientation (i.e., upright or inverted) such that the difference between the

perception of upright or inverted faces is greater than the difference between the perception

of upright than inverted non-face objects. The FIE is largely confined to second-order

properties of the human face (Maurer et al. 2002). Visual perception of inverted faces may be

especially challenging because face landmarks used to perceive second-order relations

become difficult following inversion. In contrast, non-face objects may be perceived

(Moscovitch & Moscovitch, 2000). Since salient features can be easily perceived whether a

non-face object is upright or inverted, inversion has minimal effect on the perception of such

stimuli. Thus, differences in the FIE indicate the degree to which faces are perceived on the

basis of configural perception.

Concerning age-differences in configural face perception, it may be hypothesized that

if the FIE effect is comparable for HOAs and HYAs, it follows that both groups rely equally

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Running head: age face perception

on configural face perception. In this case, both groups are equally likely to apply configural

perception to upright faces. However, if the FIE effect is smaller in HOAs, then HYAs rely

more heavily on configural face perception compared to HOAs. Thus, HOAs are less likely to

apply configural perception to upright faces.

Boutet and Faubert (2006) assessed age-differences in long-term (Experiments 1) and

working-memory (Experiments 2) of upright and inverted faces, houses and chairs (i.e., non-

face objects). The paradigm consisted of a study phase followed by a perception (test) phase.

The authors found that perception of inverted faces was significantly impaired in both age

groups, but object perception was not. Perception of upright faces was superior in HYAs than

HOAs, but there was no significant age-difference in the perception of upright objects. These

findings were not attributable to task difficulty because post hoc analysis did not reveal an

age-difference between perception of upright faces and upright objects. The authors inferred

that aging does not affect configural face perception. However, normal configural perception

of faces does not ensure that face processing is intact. Plausibly, age may have a detrimental

effect on configural processing that does not rely on memory.

To address this question, Gao et al. (2009) investigated N170 FIE effects in which

faces and tables were randomly interspersed with butterfly pictures (target stimuli). The

participants retained a mental count of the occurrence of the target. The authors hypothesized

that a face-specific effect in visual perception of HOAs would be shown in the difference

between the N170 ERP evoked by faces and objects. If healthy aging does not impair face

configural processing, there would be no age-related N170 inversion effect. Gao et al. (2009)

found that face inversion increased and delayed the N170 peak in the HYAs while in the

HOAs inversion delayed the N170 peak, but had no effect on amplitude. The N170 amplitude

was lateralized to the RH in the HYAs, but not in the HOAs. The absence of face-inversion

effects on N170 amplitudes in HOAs shows that face individuation, which is plausibly the

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Running head: age face perception

default strategy in HOAs, might not be applied by default in HOAs, at least when they

perceive young faces. This implies that HOAs are less likely to apply configural processing

when faces are presented in the normal, upright orientation relying primarily on global or

featural information.

Daniel and Bentin (2010) found that face inversion increased the amplitude and

delayed the latency of the N170 in both HOAs and HYAs. However, this effect differed

between the HYAs and the HOAs. Whereas face inversion significantly delayed the N170

peak similarly for both age groups, the amplitude of the N170 was increased by face

inversion in the HYAs but not in the HOAs. Moreover, the normalized N170-effect on

amplitudes was significant only for the HYAs and higher than the normalized face inversion

effect in the HOAs. The absence of the N170 FIE effect in HOAs indicates that the

hypothetical neural mechanism of configural processing is not as efficient in this group

compared to HYAs. As with Gao et al. (2009), this leads to the prediction that HOAs are less

likely to apply configural perception to faces, even when perceived upright.

One logical consideration challenges the interpretation of these studies. Logically, the

ability to derive the relative spatial locations of intrinsic face-parts relies on the ability to

perceive individual face-parts. Featural face perception is necessary for configural face

perception. Conceivably, the reduced tendency of HOAs to use configural perception while

processing faces may arise from a more fundamental age-difference in processing face details.

This argument is put to the test and answered in the next section.

Age-differences in featural face perception

The perception of intrinsic face-part shape and surface qualities depends on ‘featural

face perception’ (Bentin et al. 2006). Daniel and Bentin (2010) assessed age-differences in

featural face perception by comparing the N170 evoked by isolated intrinsic face-parts with

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Running head: age face perception

those evoked by full upright faces. Specifically, the N170 was compared to (1) upright faces

with intrinsic face-parts presented normally; (2) upright faces with scrambled intrinsic parts

outside (i.e., isolated) the face profile. While the parts in (2) are perceived as intrinsic face-

parts, they are not perceived as a face in any way. The authors reasoned that if there is an

age-difference in featural face perception, it is conceivable that when the intrinsic face-parts

are viewed outside of a full face, the visual stimulus is not immediately perceived as a face.

Thus, the N170 evoked by face-parts outside the face contour should be smaller than that

evoked by full faces (in contrast to HYAs; e.g., Itier et al., 2007; Zion-Golumbic & Bentin,

2007). Furthermore, this reduction should be greater when the face-parts are scrambled.

Daniel and Bentin (2010) found that the effects of isolating and scrambling intrinsic

face-parts had different effects in HOAs and HYAs. For the HOAs, the N170 amplitude

evoked by isolated face-parts was similar to that evoked by full faces when their spatial

organization was normal and reduced when the face-parts were scrambled. In contrast, for the

HYAs, the isolation of face-parts produced a larger N170 peak than full faces without further

sensitivity to their spatial organization. N170 latency was delayed by isolating the

components in both age groups, but only in the HYAs was this delay reduced when the

spatial organization of face-parts was normal. What do these results mean?

The results show that HOAs are relatively impaired while deriving a global face from

individual face-parts. They have difficulty in featural face perception because, as outlined

above, they rely more on global face perception (Daniel & Bentin 2010; Gao et al. 2009).

This interpretation is supported in Daniel and Bentin (2010) by the considerable reduction of

the N170 amplitude in HOAs when the contour of the face was eliminated, and when the

normal spatial location of the isolated face-parts was altered. Finally, this interpretation

answers the logical challenge made at the end of the previous section: since featural face

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Running head: age face perception

perception is logically necessary for configural face perception, an age-difference found in

the latter should also be found in the former, which is precisely what the N170 studies found.

Conclusions

This review has outlined studies using the N170-effect to investigate age-differences

in face perception, and has assessed their impact on the hypothesis that age-differences in

higher-level perceptual processes partly explain declined face perception in HOAs. Although

there is debate concerning the face perception mechanism the N170-effect represents, there is

no question that the N170 is stimulated by the content (mostly human faces) rather than the

low-level properties of visual stimuli.

The N170 studies discussed in this review support the hypothesis that age-differences

in higher-level perceptual processes are a factor in declined face perception in HOAs.

Primarily, it appears the HOAs have problems integrating intrinsic face-parts into global face

profiles. In addition, HOAs seem not to use configural perception by default while processing

faces, perhaps because they perceive faces only at a basic or global level. These changes in

perception may cause slower perception of face details. HOAs show no differential amplitude

in N170 to inverted versus upright faces, despite previous behavioral findings that HOAs are

affected by face inversion as much as HYAs.

At the neural level, face perception in HOAs is not lateralized to the RH, and reflects

greater involvement of the LH leading to a general reduction of interhemispheric asymmetry.

It appears that regions in the HOA LH assist the corresponding RH face-processing regions

during face perception. This finding is compatible with reduced hemispheric asymmetry

found in HOAs, and challenges the theory of a general age-related loss of neural

specialization in the ventral extrastriate system. Instead, neural representations in the ventral

visual cortex may become less domain-specific with age.

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Running head: age face perception

The studies reviewed above reveal the exquisite sensitivity of the N170 to the human

face, and confirm the validity of the N170-effect to investigate face perception and

neurodevelopment in a mutually informative way. Studies of face perception during infancy

and childhood have contributed critical evidence to mechanistic theories of human learning

and declines in performance during puberty. Similarly, future studies of age-differences in

face perception may use the N170 in combination with ERPs such as the P300 or N400, and

other neuroimaging techniques, to assess age-differences between visual perception and

perceptual factors such as executive control, decision-making or processing speed.

Developmental studies of face perception have also provided rich data for assessing theories

of functional specialization in cortex. Face perception data from infants and children have

added to the continuing debate in cognitive neuroscience about whether the fusiform face

area is specialised for face perception due to experience and training or innate biological

endowment. Studies using HOAs may contribute valuable data to this enduring debate, and

prompt fertile new theories.

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Running head: age face perception

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Appendix I

Table 1: Types of information perceived in the human face

Type Description

 The basic arrangement of the face consisting in the placement of the eyes on

both sides of a vertical axis above the nose and the mouth, within an elliptical

Global profile (Bentin et al. 2006)

 Used to group the human face as a discrete class

 Indicates first-order relations (Maurer et al. 2002)

 The integration of the facial features and the face contour as a unitary form

(e.g., Farah et al. 1998)

Holistic  Used to individuate a face (Bentin et al. 2006)

 The ‘composite effect’ shows a prevalence of the whole face over its

constituent parts (Young et al. 1987)

 The spatial relations of the facial features to each other and to the elliptical face

form (Bentin et al. 2006)

Configural
 Used to individuate and identify a face (e.g., Rhodes et al. 1993)

 Indicates second-order relations (Maurer et al. 2002)

 The shape of the facial features (Bentin et al. 2006)

Featural
 Used to individuate and identify a face (Bentin et al. 2006)

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