Social Cognition, Vol. 24, No. 5, 2006, pp. 657-701
FACES FIND FAVOR
ZEBROWITZ
FINALLY, FACES FIND FAVOR
Leslie A. Zebrowitz
Brandeis University
Historical trends in face perception research during the past half century are
summarized. The dual process model offered by cognitive neuroscientists to
account for the perception of identity and emotion expression in the face is
briefly reviewed together with supporting evidence. An ecological approach
that incorporates missing pieces of face perception is offered as an additional,
broader conceptual framework. It is argued that a comprehensive theory of
face perception must account for the extraction from a facial image of all the
attributes that are perceived (social category, identity, emotion, psychological
and physical traits) as well as the interdependence of attributes as shown in
moderating influences of one attribute on the perception of others. It also must
specify the stimulus information that conveys each attribute, perceiver
attunements that influence detection of each attribute, the development of
these attunements, the neural mechanisms engaged in processing various attributes, and the behavioral consequences of face perception. Social, personality, and developmental psychologists are urged to join cognitive
neuroscientists in filling in the missing pieces.
Poets, philosophers, and pseudo–scientists have shown a fascination
with faces that dates to ancient times and spans diverse cultures (cf.
Zebrowitz, 1997). However, social psychologists have been slow to
follow this lead with a scientific scrutiny of the role faces play in human life. Two notable exceptions are research on facial attractiveness
(Eagly, Ashmore, Makhijani, & Longo, 1991; Hatfield & Sprecher,
1986; Langlois et al., 2000) and research on emotion communication
(Ekman & Freiesen, 1975; Rosenberg & Ekman, 2005). Although the
path–breaking and programmatic research on these topics dates
Preparation of this article was supported in part by NIH grants MH066836 and
K02MH72603 and NSF grant 0315307. The author expresses her appreciation to P. Matthew Bronstad for his very helpful comments on earlier drafts of this article.
Address correspondence to Leslie A. Zebrowitz, Brandeis University, Department of
Psychology, MS 062, Waltham, MA 02454; E-mail: zebrowitz@brandeis.edu
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back to the 1960s, psychology research examining other aspects of
faces has only recently begun to blossom. The 590 citations to “face
perception” found in PsycINFO in the quarter century from
1956–1980 swelled to 6,782 citations in the next 25 years, an increase
of over 1000% in comparison with only a 210% increase in citations
for “impression formation.”1 This burgeoning scientific interest in
faces has addressed a range of questions, including the contents of
face perception, influencing stimuli, individual differences, disorders, developmental trajectories, behavioral consequences, and neural mechanisms. This article begins with an overview of the early and
current face perception literature, which is necessarily selective
rather than exhaustive. Two theoretical approaches to face perception are then discussed together with research relevant to each,
including the interesting work that appears in this special issue.
HISTORICAL RESEARCH TRENDS
THEN
Fifty years ago, there were only 11 citations for face perception in
PsycINFO. The topics covered concerned the contents of face perception, indexed by perceived psychological traits (Stritch & Secord,
1956), influencing stimuli, including social context (Cline, 1956) and
face orientation (Engel, 1956) and perceiver individual differences, including perceiver goals (Atkinson & Walker, 1956; Bevan Secord, &
Richards, 1956) and personality traits (Greenbaum, 1956; Weiner &
Ross, 1956). Medline citations become available in 1966, revealing an
article on facial recognition in brain-damaged patients (De Renzi &
Spinnler, 1966), a topic that received increased attention throughout
the next 50 years, yielding a large literature on various disorders in
face perception as well as the underlying neural mechanisms.
AND NOW
Current research shows a continuation and expansion of research
concerning the contents of face perception, influencing stimuli,
1. “Impression formation” citations grew from 583 in the years 1956–1980 to 1,811 in the
years 1980–2005; in the same time periods, “social perception” grew from 13,587 to 49,380,
an increase of 263%.
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perceiver individual differences, disorders, and neural mechanisms. In addition, research examining the development of face perception and its
behavioral consequences has emerged. The following paragraphs provide a brief overview of the burgeoning research in all of these areas
that was published in the year 2005. Although the extant research is
somewhat more ordered than it would appear from this “laundry
list,” the fact is that there is no theoretical framework that integrates
all facets of face perception.
In the year 2005, publications concerning the contents of face perception continued the early interest in the communication of psychological traits with investigations of perceived affiliativeness,
competence, dominance, health, and trustworthiness (DeBruine,
2005; Marsh, Adams, & Kleck., 2005; Todorov, Mandisodza, Goren,
& Hall, 2005; Zaidel, Aarde, & Baig, 2005; Zebrowitz & Montepare,
2005) as well as actual mental health (Buckley et al., 2005). Other
contents also were examined in 2005, including the facial communication of social categories, such as sex and race (Cloutier, Mason, &
Macrae, 2005; Abastado, Guiramand, and Bosquiet, 2005;
Eberhardt, 2005; Humphreys, Hodsoll, & Campell, 2005; Macrae,
Quinn, & Mason, 2005; Little, DeBruine, & Jones, 2005; Ramsey,
Langlois, & Marti, 2005; Vuilleumier, George, Lister, Armony, &
Driver, 2005), emotion (Calder & Young, 2005; Shibui & Shigemasu,
2005), identity (Calder & Young, 2005; DeJong, Wagenaar, Wolters,
& Verstijnen, 2005); likability (Kramer & Parkinson, 2005), and attractiveness (Rhodes, Peters, Lee, Morrone, & Burr, 2005; Zaidel,
Aarde, & Baig, 2005). 2
Research on the influencing stimuli for face perception has also expanded in the last 50 years. Face orientation remains a variable of interest (Kramer & Parkinson, 2005; Rhodes et al., 2005; Yamamoto,
Kowatari, Ueno, Yamane, & Kitazawa, 2005), as does the social context, evidenced in studies of the effects of information about targets’
personality on their perceived facial structure (Veenvliet &
Paunonen, 2005) and targets’ likebility on race categorization
(Richeson & Trawalter, 2005). Other stimulus information to draw
research attention is found in studies examining signs of aging in
self–portraits (Abastado et al., 2005), effects of hair loss on perceived
age (Rexbye et al. 2005); influence of mouth structure on smile attrac-
2. In some cases, facial communication of an attribute was accurate; in others, accuracy
was absent or unexamined.
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tiveness (Moore, Southard, Casko, Qian, & Southard, 2005); dynamic
vs. static facial cues on perceived attractiveness (Rubeinstein, 2005);
subliminal presentation on attractiveness detection (Olson &
Marshuetz, 2005); eye distortions on judgments of identity and normality (McKone, Aitkin, & Edwards, 2005); pigmentation on face recognition (Vuong Peissig, Harrison, & Tarr, 2005); luminance reversal
on face detection (Lewis & Edmonds, 2005); gaze direction, facial dynamics, and face race on emotion perception (Adams & Kleck, 2005;
Ambadar, Schooler, & Cohn, 2005; Ganel, Goshen–Gottstein, &
Goodale, 2005; Hugenberg, 2005); eye region inversion on gaze processing (Schwaninger & Mast, 2005); availability of feature vs. holistic
information when perceiving inverted and upright faces (Carbon,
Schweinberger, Kaufman, & Leder, 2005); left–side perceptual bias on
sex identification (Butler et al., 2005); and multimodal stimulus inform a t i o n o n f a c e r e c o g n i t i o n a n d p r e f e r e n c e s (B a h r i c k ,
Hernandez–Reif, & Flom, 2005; Sai, 2005; von Kriegstein,
Kleinschmidt, Sterzer, & Giraud, 2005).
Research on perceiver individual differences has been sustained
over the past 50 years. There is continued attention to effects of
perceiver goals (Maner et al., 2005; Wheeler & Fiske, 2005) and personality traits (Bayliss & Tipper, 2005; Coles & Heimberg 2005; Eastwood et al., 2005; Gardner, Pickett, Jefferis, & Knowles, 2005; Maier
et al., 2005; Montagne, von Honk, et al., 2005; Pine et al., 2005; Saito
Nakamura, & Endo, 2005). Other individual differences garnering
attention include perceiver sex (Bourne, 2005; Platek, Keenan, &
Mohamed, 2005), hormonal levels (DeBruine, Jones, & Perrett, 2005;
Pearson & Lewis, 2005), and perceiver age (Kiffel, Campanella, &
Bruyer, 2005).
Related to perceiver age is research on the development of face perception. Although absent 50 years ago, the topic received substantial attention in 2005, including studies of infant face perception
(Bhatt, Bertin, Hayden, & Reed, 2005; Pascalis et al., 2005; Ramsey et
al., 2005; Turati, Valenza, Leo, & Simion, 2005), plasticity in early
childhood (Sangriogoli, Pallier, Argenti, Ventureyra, & de
Schonen, 2005), developmental changes from childhood through
maturity (Deruelle & Fagot, 2005), and effects of normal aging from
young to older adulthood (Kiffel et al., 2005). Developmental effects have also been elucidated by investigations of short–term perceptual learning effects (Behrmann & Avidan, 2005; Fahle, 2005;
Morikawa, 2005) and by comparative animal research (Evans,
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Howell, & Westergaard, 2005; Myowa–Yamakoshi, Yamaguchi,
Tomonaga, Tanaka, & Matsuzawa, 2005; Pinsk, DeSimone, Moore,
Gross, & Kastner, 2005).
Research on disorders in face perception shown by people with various pathologies has grown significantly from the single early study
by De Renzi and Spinnler (1966). Disorders in face perception have
been studied in Alzheimer’s disease (Lavenu & Pasquier, 2005), autism
(Castelli, 2005; Dawson, Webb, & McPartland, 2005; Schultz, 2005;
Williams, Goldstein, & Minshew, 2005); brain lesions (de Schonen,
Mancini, Camps, Maes, & Laurent, 2005; Pegna, Khateb, Lazeyras, &
Seghier, 2005), depression (Langenecker et al., 2005); Parkinson’s disease (Biseul et al., 2005; Pell & Leonard, 2005), prosopagnosia
(Behrmann & Avidan, 2005; Caldara et al., 2005; Duchaine &
Nakayama, 2005); schizophrenia (Bediou et al., 2005; Martin,
Baudouin, Tiberghien, & Franck, 2005; Scholten, Aleman, Montagne,
& Kahn, 2005; Suslow, Droste, Roestel, & Arolt, 2005), and social phobias (Straube, Mentzel, & Miltner, 2005).
Research examining the behavioral consequences of face perception,
absent in 1956, is represented in 2005, although the study of overt
motor behaviors is rare. Rather, most investigations are limited to an
examination of orienting and cognitive responses, such as attention
to faces (Georgiou et al., 2005; Hershler & Hochstein, 2005;
Lundqvist & Ohman, 2005; Williams, Moss, Bradshaw, &
Mattingley, 2005), orienting to eye gaze in upright and inverted faces
(Tipples, 2005), eye gaze localization to faces varying in familiarity
(Stacey, Walker, & Underwood, 2005); reaction time to faces varying
in similarity to self (Yoon, & Kircher, 2005); and effects of emotion expressions on implicit learning and memory (Schultheiss, Pang, Torges,
Wirth, Treynor, & Derryberry, 2005; Sergerie Lepage, & Armony,
2005). Other research has examined affective responses, including
heart rate increases to emotion expressions (Critchley et al., 2005); infants’ smiling, gaze aversion, and crying in response to their
mother’s face (Field et al., 2005); and satisfaction with one’s own face
after viewing highly attractive faces (Newton & Minhas, 2005). Demonstrated effects on overt motor behaviors include effects of candidate
facial appearance on voter behavior (Todorov et al., 2005), effects of
primed emotion expressions on beverage consumption by thirsty
people (Winkielman, Berridge, & Wilbarger, 2005), and effects of fear
and anger facial expressions on approach– and avoidance–related
behaviors (Marsh, Ambady, & Kleck, 2005).
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A large growth of face perception research is found in the domain of
neural mechanisms (cf. Gross, 2005). This topic has been informed not
only by the above-cited research documenting disorders in face perception among individuals with various pathologies, but also by research investigating the neural mechanisms that contribute to these
disorders. Event–related potentials (ERPs) and functional magnetic
resonance imaging (fMRI) have been used to study the neural mechanisms in emotion face processing by individuals with Asperger’s syndrome (Welchew et al., 2005), autism (Grelotti et al., 2005; Grice et al.,
2005; Schultz, 2005), PTSD (Shin et al., 2005); schizophrenia (Holt et al.,
2005; Valkonen–Korhonen et al., 2005; Yoo et al., 2005), and
Wernicke–Korsakoff syndrome (Caulo et al., 2005). These methods
have also been applied to the study of emotion expression processing
in deaf individuals who are American Sign Language signers
(McCullough, Emmorey, & Sereno, 2005) and to the study of face processing by normal individuals. The latter research has sought to elucidate not only the neural substrates of emotion perception (Balconi &
Pozzoli, 2005; Noesselt, Driver, Heinze, & Dolan, 2005; Pessoa &
Padmala, 2005; Pourtois, Dan, Grandjean, Sander, & Vuilleumier,
2005; Strauss et al., 2005; Werheid, Alpay, Jentzsch, & Sommer, 2005),
but also identity recognition (Balconi & Lucchiari, 2005; Boehm &
Sommer, 2005; Eimer & Mazza, 2005; Gauthier & Curby, 2005; Goto,
Kinoe, Nakashima, & Tobimatsu, 2005; Guillaume & Tiberghien,
2005; Nessler, Mecklinger, & Penney, 2005; Rotshtein, Henson, Treves,
Driver, & Dolan, 2005; Xu, Liu, & Kanwisher, 2005), race perception (Ito
& Urland, 2005; Lieberman, Hariri, Jarcho, Eisenberger, &
Bookheimer, 2005), and perceptions of eye and mouth motion and face orientation (Watanabe, Miki, & Kakigi, 2005). Neural mechanisms also
have been investigated in research examining developmental changes in
face perception (Aylward et al., 2005; Fischer et al., 2005), sex and age
differences (Guillem & Mograss, 2005; Fischer et al., 2005), personality
differences (Kilgore & Yurgelun–Todd, 2005), context effects (Walla,
Mayer, Deecke, & Lang, 2005), and general face perception (George,
Jemel, Fiori, Chaby, & Renault, 2005; Herrmann, Ehlis, Ellgring, &
Fallgatter, 2005; Ishai, Schmidt, & Boesiger, 2005).
SUMMARY
Fifty years ago, an investigator interested in face perception could
easily read all the extant research literature. Now, it is a challenge to
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read even those studies published in a single year. The rapid growth
in the research literature highlights the importance of developing a
theoretical framework that can integrate what we know as well as direct future research. Although the questions addressed in the studies
that were briefly reviewed in this section provide a foundation for
building a theory of face perception, the integration across topics is
not yet up to the task. Thus, we know that faces communicate information about a person’s social category, identity, emotion, likebility,
and psychological traits with varying degrees of accuracy and error,
thereby demarcating what face perception does. We also know
something about the stimulus information in faces that communicates each of these qualities, thereby providing a window onto the
raw material on which the face perception system operates. And we
know that perception of various qualities in faces is influenced by
perceiver individual differences ranging from transient goals to
more stable personality traits, demographic qualities, and neurological disorders, suggesting interesting plasticity and vulnerabilities in
the face perception system. We are also learning more about the neural substrate for some aspects of face perception, and we know a little
bit about the development of face perception and its behavioral consequences, which begins to elucidate the functional significance of
the face perception system. Of course, there is still much to be learned
within each of the foregoing topics in face perception. An even
greater need is to learn how everything fits together.
TOWARD A COMPREHENSIVE THEORY
OF FACE PERCEPTION
Although the increasingly large body of research regarding face perception is impressive, theoretical developments are still in the early
stages. A comprehensive conceptual framework for understanding
face perception must account for the extraction from a facial image of
all the various attributes that are perceived, including social category, identity, emotion, and psychological and physical traits, as
well as the interdependence of attributes, as shown in moderating influences of one attribute conveyed by the face on the perception of
others. Such an account also must specify the stimulus information
that specifies each attribute, the perceiver attunements that influence
detection of each attribute, the development of these attunements,
and the neural mechanisms engaged in processing various attrib-
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utes. If face perception is to be placed in its proper functional perspective, then a comprehensive conceptual framework must also
specify its behavioral consequences. Obviously, filling in these details is a tall order, but some progress has been made. The dominant
model of face perception in the cognitive neuroscience literature, the
dual process model, is briefly described below followed by an ecological approach to face perception that offers a conceptual framework for incorporating elements missing in the dominant model,
together with illustrative research consistent with that expanded
framework.
DUAL PROCESS MODEL
Cognitive neuroscience research has contributed to a model of face
perception that posits separate mechanisms for the perception of
identity versus the perception of emotional expressions and other
changeable facial qualities, such as eye gaze and head angle (Bruce &
Young, 1986; Calder & Young, 2005; Haxby, Hoffman, & Gobbini,
2002). Focusing primarily on these two aspects of face perception, the
dual process model does not claim to be a comprehensive model. Although it has been argued that the existing evidence provides
weaker confirmation of this model than is typically assumed (Calder
& Young, 2005), support for the model has been provided by both
cognitive and neurological data.
Research has implicated an area of the occipito–temporal cortex,
the fusiform face area (FFA), in identity recognition. Not only have
human brain imaging studies demonstrated stronger responses to
faces than to other stimuli in the FFA (e.g., Kanwisher, McDermott, &
Chun, 1997; Kanwisher, Stanley, & Harris, 1999), but also this area is
typically damaged in prosopagnosia, the loss of ability to recognize
facial identity despite retaining normal intelligence, language skills,
and ability to recognize other objects (cf. Damasio, Tranel, & Rizzo,
2000; De Renzi, Farah, & Feinberg, 2000; Farah, Levinson, & Klein,
1995). Although recent research suggests that FFA activation also
may reflect expertise with stimuli within a particular category rather
than being specialized for faces per se (Tarr & Gauthier, 2000), other
evidence reveals that the highest FFA activation is elicited by faces
(Xu et al., 2005).
In addition to evidence that the FFA participates in identity recognition, other research suggests that identity recognition is function-
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ally and neurologically separable from other aspects of face
perception. Brain injury can selectively impair the recognition of either facial identity or facial expression (Bruyer,1983; Etcoff, 1984;
Tranel, Damasio, & Damasio, 1988; Young, Newcombe, de Haan,
Small, & Hay, 1993). In addition, neurologically intact individuals
show differential brain activation when responding to facial identity
versus facial expression, with the latter yielding stronger activation
of neural systems for processing emotion, such as the amygdala
(Breiter et al., 1996; Morris, Frith, Perrett, & Rowland,1996; Thomas
et al., 2001) and insula (Phillips et al., 1997, 1999), as well as neural
systems for processing reward, including the orbitofrontal cortex
and the anterior cingulate cortex (Hornak, Rolls, & Wade,1996;
Hornak et al., 2003; Rolls, 2004). The somatosensory cortex also may
be involved in the perception of emotional expressions by enabling
motor mimicry (Adolphs, Damasio, Tranel, Cooper, & Damasio,
2000).
The perception of facial attributes other than expression can also be
differentiated from identity perception. The perception of biological
movement, including movements of the eyes and mouth, elicits enhanced brain activation in the superior temporal sulcus (STS) in human fMRI studies (Hoffman & Haxby, 2000). STS activation also
discriminates morphed expression changes from morphed identity
changes (LaBar, Crupain, Voyvodic, & McCarthy, 2003). Although
STS activation is enhanced during the perception of still facial photos, the effect is stronger when attention is directed to eye gaze direction than to identity, with the reverse shown in FFA activation
(Hoffman & Haxby, 2000). Together these findings indicate that the
STS is more involved in the processing of emotion expressions and
biological movement than facial identity.
In sum, considerable research supports the dual process model by
demonstrating functional and neurological divisions in the processing of identity information in faces versus emotion and other
dynamic information.
AN ECOLOGICAL APPROACH TO FACE PERCEPTION
The components of a comprehensive model of face perception can be
profitably placed within the ecological approach to social perception
(McArthur & Baron, 1983; Zebrowitz & Collins, 1997; Zebrowitz &
Montepare, 2006a, 2006b), which draws on Gibsonian theories of
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perception (E. J. Gibson, 1969; J.J. Gibson, 1966, 1979). Ecological theory adds to the dual process model by emphasizing the function of
face perception to guide adaptive behavior, expanding the attributes
that are perceived in faces, focusing attention on the stimulus information that guides face perception, and systematically considering
the perceiver qualities that moderate face perception. The pertinent
tenets of the ecological approach are discussed in more detail in the
following sections together with an overview of some research
evidence bearing on each tenet.
Perceiving Affordances
The ecological approach expands the facial attributes considered by
the dual process model in its emphasis on the perception of social
affordances—the opportunities for acting or being acted upon that are
provided by other people. A vivid illustration of what Gibson meant
by “affordance” is provided by his quotation from Koffka: “Each
thing says what it is . . . a fruit says ‘eat me’; water says ‘drink me’;
thunder says ‘fear me’; and woman says ‘love me’” (Gibson, 1979, p.
138). Within this framework, face perception is not simply about recognizing a particular identity or emotion or psychological trait.
Rather, it is about the function of the perceptual system to guide actions that serve to solve specific adaptive problems or to facilitate
other goal attainments of individuals. As such, the affordance concept focuses attention on the behavioral responses linked to face perception, whereas the dual process model, like many other theories of
perception, does not. Yet, perceiving identity and emotion, as well as
other contents, clearly has implications for adaptive action.
Although one may question whether J.J. Gibson’s emphasis on perceived affordances can be translated to the social domain, Gibson
himself argued for the relevance of an ecological approach to social
perception (Gibson, 1979). Of course, it must be acknowledged that
social affordances differ from physical ones in that they are not limited to opportunities for physical interactions, such as perceiving
who can move a heavy object, but also include opportunities for social interactions, such as perceiving who will agree to help me. Nevertheless, there are commonalities. Like perception of the affordances
of the physical world, perception of social affordances does not require mediating symbolic representations, like verbal labels, which
is consistent with Gibson’s assertions about the nature of direct per-
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ception (Greeno, 1994). For example, nonverbal infants show behavioral responses to faces that reflect a sensitivity to the social
affordances of angry expressions, youthfulness, attractiveness, and
familiarity (Balaban, 1995; Bigelow, MacLean, Wood, & Smith, 1990;
Kramer, Zebrowitz, San Giovanni, & Sherak, 1995; Langlois,
Roggman, & Rieser–Danner, 1990; Walton, Bower, & Bower, 1992).
In addition, research has demonstrated that the human visual system is finely tuned to social attributes revealed in people’s physical
qualities, including deceptive intent, motivational incentives, personality, relationship quality, sexual orientation, mood, and teaching effectiveness (Ambady, Bernieri, & Richeson, 2000; Brownlow,
Dixon, Egbert, & Radcliffe, 1997; Johansson, 1973; Montepare &
Zebrowitz–McArthur, 1988; Runeson & Frykholm, 1983; Schultheiss
et al., 2005). Research also has demonstrated a neural basis for the
perception of social affordances. Not only is activation in particular
brain regions selectively responsive to human movement
(Beauchamp, Lee, Haxby, & Martin, 2002), but also these regions are
selectively activated when human agency is evoked by the movements of geometric shapes (Martin & Weisberg, 2003). Taken
together, this work suggests that just as the optic array can specify
the physical affordances of inanimate objects, it can also reveal social
affordances.
As noted above, most research examining behavioral
concomitants of face perception has focused on orienting and affective responses. However, some work has examined overt behavioral
effects. Consistent with the concept of social affordances is the Marsh
et al. (2005) finding that attractive or fearful faces facilitate approach
responses, whereas disfigured or angry faces facilitate avoidant responses. Facial mimicry of emotion expressions is another behavioral response to faces that has been documented (Dimberg,
Thunberg, & Elmehed, 2000; Hess & Blairy, 2001). Moreover, consistent with ecological theory’s emphasis on behavioral affordances as
emergent qualities that can depend on perceiver attunements (see below), mimicry of positive and negative expressions varies as a function of the perceiver’s emotional state (Vrana & Gross, 2004). More
research is clearly needed to specify the behavioral effects of
perceiving various facial attributes.
The assumption that face perception functions to guide adaptive
behavior highlights the issue of accuracy. Attention to accuracy and
bias in face perception within the dual process model has been most
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prominent in research on disorders. For example, prosopagnosia is
characterized by a dissociation between identity recognition, which
is impaired, and other contents of face perception, such as emotion
recognition, which may be intact. In autism, on the other hand, identity recognition is intact, while some aspects of emotion recognition
are flawed. A more comprehensive model of face perception requires a more nuanced understanding of the stimulus, experiential,
and neural factors that contribute to accurate and inaccurate perception of identity and emotion as well as social category and psychological traits. Factors that contribute to accurate or biased perception
of facial qualities themselves are also of interest (Hassin & Trope,
2000).
In addition to highlighting the issues of adaptive action and accuracy, the concept of affordance also provides a unifying principle for
conceptualizing the contents of face perception. A perceived identity, social category, emotion, or psychological trait may each specify
the same behavioral affordance. This principle has implications for
the neural mechanisms involved in face perception. Perceiving an
emotion or identity or trait may all activate motivational systems
and motor response systems in the brain, with the particular patterns
of activation depending on the similarities in the affordances of the
particular emotion, identity, or trait that is perceived. Thus, although
specific brain regions involved in the recognition of identity or emotion are fairly well established, much remains to be learned regarding similarities and differences in the overall patterns of neural
activation for particular identities, expressions, social categories,
and psychological traits that may be linked to similarities and
differences in their behavioral affordances (Gorno–Tempini et al.,
2001; Winston, O’Doherty, & Dolan, 2003).
Consistent with the argument that different facial attributes may
specify similar affordances, research has demonstrated connections
across multiple facial attributes in human judgments, stimulus information, and neural activation that cannot be readily predicted by the
dual process model, although they are not necessarily inconsistent
with it. For example, the recognizability of faces is related to their sex
prototypicality as well as their attractiveness (O’Toole et al., 1998).
Excellent additional examples of interdependency among the perception of different facial attributes are provided by articles in this
special issue. Mason, Cloutier, and Macrae (2006) show a link between the mechanisms for processing social category and psycho-
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logical traits. Specifically, they find that the sex of distracter faces
influences reaction times when classifying words as stereotypically
male or female. Distractor faces whose sex was incongruent with a
gender-stereotyped word slowed reaction times as compared with
distractor faces whose sex was congruent with the stereotype. These
results indicate that even when perceivers are trying to ignore a face,
sex–linked facial cues automatically activate gender–related traits.
Hugenberg and Sczesny (2006) in the present issue show a link between the mechanisms for processing social category and emotion
expressions in their finding that the tendency for happy faces to be
accurately categorized more quickly than negative ones is stronger
for female than male targets. They further demonstrate that the moderating effect of sex on categorization of happy expressions is likely
due to the stronger positive valence associated with female than
male faces rather than to stereotypic expectancies that women will
show happiness and men will show anger. The stronger happy face
a d v a n t a g e fo r f e m a l e f a c e s h o l d s t r u e e v e n w h e n t h e
sex–stereotypicality of the negative expressions are equated by using
sad rather than angry expressions. The valence explanation is consistent with results reported by Hugenberg (2005), who found that
White perceivers show the happy face advantage for White, but not
Black targets, for whom they categorize negative expressions more
quickly.
Not only is the perception of social category intertwined both with
the perception of emotion expression and with the perception of psychological traits, but also the perception of emotion expression in
faces is linked to the perception of psychological traits (Knutson,
1996; Montepare & Dobish, 2003). Moreover, perceived trait variation in emotion expressions can be predicted from the extent to
which their physical structure resembles the social category of babies
(Marsh et al., 2005; Zebrowitz, Kikuchi, & Fellous, 2006). Similarly,
perceived trait variation in faces from the same social category (e.g.,
age, gender, or race) can be predicted from the extent to which the
faces physically resemble a typical face from a different category
(Blair, Judd, & Chapleau, 2004; Blair, Judd, Sadler, & Jenkins 2002;
Livingston & Brewer, 2002; Montepare & Zebrowitz, 1998;
Zebrowitz, Fellous, Mignault, & Andreoletti, 2003). Consistent with
this evidence for patterns of stimulus information common to multiple face categories, research has revealed some identifiable stimulus
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qualities in faces that code both their identity and their facial
expression (Calder, Burton, Miller, Young, & Akamatsu, 2001).
Commonalities in the stimulus information that differentiates various facial attributes have potential implications for the neural mechanisms involved in the perception of identity, social category,
emotion, and psychological traits. To the extent that neural perceptual mechanisms have evolved to mirror the stimulus information
provided in the perceiver’s visual world, there may be mechanisms
tuned to multiple facial attributes rather than a set of totally
independent processes.
Pertinent evidence for neural mechanisms tuned to multiple attributes is the finding that the FFA was activated more when processing
emotional faces than neutral faces (Critchley et al., 2000; Lewis, 2003)
despite evidence that FFA lesions can leave emotion recognition intact (e.g., Tranel et al., 1988). Also, investigations of face-sensitive
cells in macaques revealed that in addition to cells that responded to
identity and others that responded to expression, some cells were
sensitive to both qualities (Hasselmo, Rolls, & Baylis, 1989; see Calder & Young, 2005 for a fuller discussion). The FFA also has been implicated in the processing of social category in addition to identity.
Specifically, FFA activation is greater for own–race faces, and the
magnitude of the differential own–race versus other–race activation
predicts the magnitude of the own–race advantage in face recognition (Golby, Gabrieli, Chiao, & Eberhardt, 2001). FFA activation also
is greater for atypical schematic faces and anomalous real faces
(Zhang, Aharon, & Zebrowitz, 2006; Loffler, Yourganov, Wilkinson,
& Wilson, 2005).
Research has revealed shared neural mechanisms for the perception of still other facial attributes. For example, not only do highly attractive faces elicit greater brain activation than average faces in
regions associated with reward function in animals (Aharon et al.,
2001), but also this effect is greater for faces that are looking directly
at the observer (Kampe, Frith, Dolan, & Frith, 2001) or smiling
(O’Doherty et al., 2003), thereby demonstrating that the brain regions that process both facial expression and eye gaze are also involved in the processing of facial attractiveness. Similarly, eye gaze
moderates activation of the amgydala by emotion expressions, with
stronger activation when threat is more apparent due to anger being
paired with direct gaze or fear being paired with averted gaze than
vice versa (Adams, Gordon, Baird, Ambady, & Kleck, 2003). The
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amygdala is involved not only in the recognition of facial expressions of emotion, as discussed above, but also in the perception of
psychological traits and social category. Perceivers show more
amygdala activation to unfamiliar than familiar faces (Gobbini,
Leibenluft, Santiago, & Haxby, 2004) as well as to faces judged as untrustworthy (Winston, Strange, O’Doherty, & Dolan, 2002), and bilateral amygdala damage reduces the tendency to perceive strangers
as low in approachability and trustworthiness (Adolphs, Tranel, &
Damasio, 1998). This evidence for heightened amygdala activation
in response to an unfamiliar and possibly threatening face is reinforced by evidence for slower habituation of amygdala activation
when viewing other-race faces (Hart et al., 2000) as well as by the tendency for greater activation by other-race faces to be stronger among
those for whom those faces produced greater potentiation of
eyeblink startle (Phelps et al., 2000).
In summary, there is considerable evidence that many aspects of
face perception are intertwined, including the perception of identity,
emotion, social category, attractiveness, and traits. The connections
between these different attributes are manifest in perceiver judgments, overlapping stimulus information, and patterns of neural activity. The concept of social affordance offers a possible unifying
mechanism for these connections and focuses attention on the behavioral effects of perceiving various facial attributes and the
question of accuracy.
Identifying the Stimulus Information
A pivotal feature of the ecological approach is an emphasis on identifying the features of the external stimulus environment that inform perception. Gibson argued that understanding the nature of the
stimulus information to which organisms respond will elucidate
how their perceptual systems operate. Thus, although he did not focus on neural mechanisms per se, his theory implies that they should
be elucidated by a deep understanding of the stimuli. Indeed, his
specifications of the kinds of higher order stimulus information that
informs adaptive action stimulated research that has identified neurons tuned to such information (cf. Nakayama, 1994). Gibson (1966,
1979) demonstrated that multimodal, dynamic changes over space
and time are features that provide the most useful information to
perceivers in nonsocial perception, and McArthur and Baron (1983)
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suggested that the same would be true for social perception. Thus,
the dynamic and multifaceted facial information that can be gleaned
from facial structure, pigmentation, texture, and movement should
provide the most useful information about all of the qualities that are
gleaned from faces.
A derivation from ecological theory pertinent to identifying the
s t i m u l u s i n f o r m a t i o n fo r f a c e p e r c e p t i o n i s a se t o f
overgeneralization hypotheses that codify effects of a preparedness
to respond to adaptively significant facial qualities. More specifically, these hypotheses hold that the traits and affordances that are
accurately revealed by the facial information that marks babies, emotion, identity, or low fitness are also perceived in people whose faces
resemble that of babies, a particular emotion, a particular identity, or
a particular level of fitness (cf. Zebrowitz, 1996, 1997; Zebrowitz &
Collins, 1997).3 Consistent with the babyface overgeneralization hypothesis, the facial qualities that mark babyfaced adults, such as a
round face, large eyes, high eyebrows, and small chin and nose
bridge, also mark real babies, and the actual traits of babies are mirrored by impressions of babyfaced adults as physically weak, submissive, warm, and naive (Montepare & Zebrowitz 1998; Zebrowitz,
et al., 2003). Similarly, trait impressions of faces vary with their structural resemblance to emotional expressions, consistent with the emotion overgeneralization hypothesis (Montepare & Dobish, 2003), and
impressions of faces vary with their resemblance to known individuals, consistent with the identity overgeneralization hypothesis
( De B r u i n e , 2 0 0 2 ; L e w i c k i , 1 9 8 5 ) . Co n s i s t e n t w i t h th e
overgeneralization of fitness information, impressions of unattractive people as socially, physically, and cognitively deficient—the
converse of the attractiveness halo effect—can be partly explained by
the extent to which their faces resemble anomalous ones (Zebrowitz
et al., 2003, Zebrowitz & Rhodes, 2004). Complementing the judgment and modeling research that has supported the face
overgeneralization hypotheses, one might also predict that faces
linked by one of the overgeneralization effects would evoke
common neural mechanisms.
3. Another overgeneralization hypothesis is “animal analogies” whereby people may be
perceived to have traits that are associated with the animals that their features resemble
(Zebrowitz, 1997, pp. 58–61).
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Although the dual process model is silent regarding the stimulus
information that fuels identity versus emotion recognition or other
contents of face perception, individuals affiliated with this model as
well as other researchers have provided some answers to this question. Below is a brief summary of various methods that have been
employed in an effort to identify the stimulus information for face
perception together with a brief assessment of how an ecological
perspective bears on each
Brunswik’s Lens Model (Brunswik, 1956) is one of the earliest
methods used to identify the particular physical qualities that inform
person perception, including face perception. Within this model, one
would sample physical qualities as they naturally co–occur in a representative sample of faces, with the aim of identifying correlations
with particular attributes that reveal both the ecological validity of various cues (e.g., correlations with actual age, emotion, traits) and also
perceivers’ utilization of these cues (e.g., correlations with perceived
age, emotion, traits). This approach to studying the stimulus information for face perception is consistent with Brunswik’s claim that
organisms are “probability learners” and that the perceptual system
operates like an intuitive statistician. According to Brunswik’s tenet
of probabilistic functionalism, the cues available in the natural environment are of limited validity, and perceivers achieve some degree
of accuracy by combining information from multiple cues, as specified in the lens model. The implicit assumption is that perceivers
achieve accurate perceptions by weighing each cue in proportion to
its actual diagnosticity of the attribute being judged. Although perceptual learning (Bruce & Burton, 2002) is quite congruent with the
ecological approach, the view that the perceptual system operates on
discrete cues like an intuitive statistician stands in contrast to Gibson’s theory. According to Gibson, the information available in the
natural environment is highly valid when one considers higher order structure as opposed to discrete stimulus cues. The need to
search for such structure highlights the necessity of a theoretical
foundation from which to make predictions concerning the particular facial qualities that convey various human attributes. Once such
theoretically meaningful qualities are identified, researchers might
then test their utilization using the lens model.
Although researchers making explicit use of Brunswik’s lens
model have typically employed rather crude measures of physical
qualities, such as perceivers’ subjective ratings, there are more so-
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phisticated methods that share some features in common with this
model. One example is provided in the article by Penton–Voak,
Pound, Little, and Perrett (2006) in the present issue, which makes an
important contribution to our understanding of the stimulus information that contributes to the accurate perception of personality
traits. Ecologically valid facial cues to personality traits were created
by making composite faces of individuals who shared particular personality traits, including agreeableness, extraversion, and emotional
stability. The utilization of these cues was then assessed by measuring
trait impressions of the composite faces. The results showed that the
valid cues were indeed utilized, yielding accurate trait impressions.
Exactly what facial qualities differentiate the composite faces from
one another and what experiential factors and neural mechanisms
underly perceivers’ sensitivity to these facial qualities are fascinating
questions for future research.
Another variant on the lens model is provided by research using
principal components analysis of facial metrics and/or pixels (PCA).
This research has identified components that represent the complex
visual information in faces in a suitable form for specifying different
contents of face perception, including identity (Hancock, Bruce, &
Burton, 1998), emotion (Calder et al., 2001), sex (Valentin, Abdi,
Edelman, & O’Toole, 1997), and race (O’Toole, Abdi, Deffenbacher,
& Bartlett, 1991). Some PCA research has focused exclusively on the
ecological validity of certain components—that is, their actual relationship to particular human attributes. Other research has also examined the utilization of these factors by perceivers. Although PCA
has provided useful analogues for face perception, it has several acknowledged limitations. The components that PCA identifies are essentially dimensions along which a set of faces vary—for example,
age—rather than a meaningful description of the stimulus information that accounts for that variation—for example, variations in
cardiodal strain (Todd, Mark, Shaw, & Pittenger, 1980). In addition,
it is often difficult to provide meaningful labels for some of the dimensions. Moreover, the components identified are necessarily limited by the variation within a particular set of faces, and it is difficult
to know what a representative set should include. Finally, PCA may
or may not correspond to the way in which the perceptual system actually works. Similar shortcomings accrue to connectionist models
that have used facial metric inputs to differentiate faces based on sex
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675
(Fellous, 1997), emotion (Zebrowitz et al., 2006), age, or abnormality
(Zebrowitz et al., 2003).
Another method for examining the stimulus information is the
Bubbles technique (Gosselin & Schyns, 2001), which focuses on the
stimulus information that is utilized for perceiving different facial attributes. This technique automatically adjusts a number of small
Gaussian Windows (“bubbles”), revealing parts of the face so that
75% correct categorization is achieved. All sets of bubbles that yield
criterion performance are summed to determine the effective stimulus information for that categorization task. This research has shown
that the utilized information is not perfectly correlated with the available information, thereby underscoring a shortcoming of PCA or
connectionist modeling research that has not integrated statistically
differentiable stimulus properties of faces with those that guide perception. Bubbles also has provided a more concrete description of the
stimulus information that conveys various attributes to perceivers.
For example, when categorizing faces based on sex, perceivers use
information around the eyes and the central upper part of the mouth.
When categorizing faces varying in emotion, perceivers use information in different regions to recognize different emotions. When recognizing the identity of faces, perceivers use the entire constellation
of facial features together with the jaw line (Gosselin & Schyns, 2001;
Smith, Cottrell, Gosselin, & Schyns, 2005). Although these stimulus
descriptions are more concrete than those generated by PCA and
connectionist modeling research, they are incomplete. For example,
they do not reveal what it is about the eyes that conveys the sex of the
face or what it is about various facial regions that conveys different
emotions. The answer to questions such as these requires some
guiding theory, like the Facial Affect Coding system (FACS) used to
identify muscle action units that differentiate emotion expressions
(Rosenberg & Ekman, 2005).
Some methods for studying the stimulus information that informs
face perception have been guided by the theoretical view that facial
identity is coded relative to an average face, which lies at the center of
a computational face–space (cf. Clifford & Rhodes, 2005). Within this
conceptual framework, both multidimensional scaling (MDS) methods and the study of visual aftereffects have been used to study the
stimulus information that informs face perception. For example,
MDS has revealed that perceiver judgments of the similarity of faces
maps onto a face–space representing spatial relations among differ-
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ent faces that can explain differences in ease of recognizing those that
are caricatured, veridical, and anticaricatured (Lee, Byatt, & Rhodes,
2000). Evidence to support the face–space theoretical framework
also is provided by visual aftereffects. For example, identification of
a face (Jim) is facilitated by adapting perceivers to the opposite identity (a face morphed away from Jim through the prototype to the opposite side of the stimulus space to become an Anti–Jim), but not to
some other face (Anti–Fred), and this effect holds true even when opposite and nonopposite adapt–test pairs are matched on perceived
dissimilarity (Leopold, O’Toole, Vetter, & Blanz, 2001; Rhodes &
Jeffrey, 2006). The process of adapting to the Anti–Jim face shifts the
norm or average toward that face, so that the opposite face, Jim, now
deviates more from the norm than it did before, becoming easier to
identify. Interesting variants on this method have revealed that aftereffects do not generalize across sex categories, suggesting that male
and female faces are coded with respect to different average faces
(Rhodes et al., 2004). The view that facial identity is coded relative to
an average face is consistent with the ecological emphasis on
configural facial qualities. However, the ecological approach would
also underscore the importance of creating face spaces from
three–dimensional faces as well as faces that are animated and
naturalistically colored. The selection of an appropriately
representative set of faces is also an issue from the ecological
perspective.
The study of visual aftereffects approaches the task of identifying
the stimulus information in faces in part by manipulating that information to determine its effects on perception rather than simply measuring it. In addition to morphing faces closer to or farther from some
standard face, other manipulation methods have been used to isolate
effects of facial coloration, form, luminance, or orientation. Together,
these methods have shed light on the stimulus information that differentiates and communicates emotions (Ekman & Friesen, 1975;
Rosenberg & Ekman, 1997), identities (O’Toole, Vetter, & Blanz,
1999), and ages (O’Toole, Vetter, Volz, & Salter, 1997), as well as the
stimulus information that contributes to the perception of facial attractiveness (Rhodes & Zebrowitz, 2002; Russell, 2003) and
babyfaceness (Montepare & Zebrowitz, 1998). Moreover, it has been
demonstrated that variations in these and other facial cues also influence perceived traits, including health and intelligence (Rhodes,
Chan, Zebrowitz, & Simmons, 2003; Zebrowitz, Hall, Murphy, &
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Rhodes, 2002; Zebrowitz & Rhodes, 2004) as well as dominance, and
warmth (Keating, 2002; Montepare & Zebrowitz, 1998; Zebrowitz &
Collins, 1997) and trustworthiness (deBruine, 2005). Much of the research that has manipulated facial qualities is consistent with the
ecological approach in that it has been guided by theoretically derived hypotheses concerning what stimulus information should
matter for the perception of particular attributes. On the other hand,
manipulations of stimulus information have often been
unidimensional and static, thus lacking the ecological validity that
Gibson advocated. In addition, the use of morphing software raises
the question of exactly what codes specify the morph trajectories and
whether these transformations have psychological meaning
(O’Toole, Wenger, & Townsend, 2001).
Among the investigations using perceiver ratings to identify stimulus information in faces are studies demonstrating that the judged
Afrocentrism of facial cues by White raters influences the extent to
which faces from the same race category elicit negative affect
(Livingston & Brewer, 2002), discrimination in judicial sentencing
(Blair et al., 2004), and impressions consistent with African–American stereotypes (Blair et al., 2002). Moreover, the study by Blair
(2006) reported in this issue demonstrates that Afrocentrism is perceived with sufficient ease that it has as great or greater an impact on
impressions when faces are inverted as when they are upright, and
that the degree to which African American faces are perceived as
having Afrocentric features is highly similar across upright and inverted faces. Although these results could merely reflect attention to
skin tone, which should not be disrupted by inversion, a more interesting possibility is that the facial features that contribute to White
perceivers’ judgments of Afrocentrism are processed piecemeal
rather than holistically. Such an effect provides an interesting contrast to research showing that identity recognition from faces is holistic and severely impaired by face inversion (Tanaka & Farah, 1993;
Young, Hellawell, & Hay, 1987). Consistent with this interpretation,
there is some evidence that perceivers respond to other–race faces
with greater activation in brain regions that are specialized for processing objects rather than faces, such as the ventral temporal cortex
(Haxby et al., 2002; Ishai, Ungerleider, Martin, Schouten, & Haxby,
1999). These findings underscore the need to consider perceiver
qualities when investigating the stimulus information for face perception. As emphasized in the ecological approach, the effective
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stimulus information will depend not only on the tangible qualities
of the face, but also the attunements of the perceiver.
In summary, many different methods have been employed to
identify the stimulus information that guides face perception, including perceiver ratings, PCA, connectionist models, the Bubbles
technique, MDS, and visual aftereffects, morphing, and other facial
manipulations. Typically, particular methods have been applied to a
narrow range of the many attributes that are perceived in faces, and it
may prove instructive to apply some methods more broadly. A more
significant shortcoming of efforts to identify the stimulus information that guides face perception is that much of the work has been
atheoretical, although there are exceptions. For example, the face
overgeneralization hypotheses provide a theoretical rationale for examining the contribution of particular facial qualities to impressions
of traits and affordances. Investigations of the stimulus information
for face perception should be theoretically rather than empirically
motivated if they are to successfully capture the facial qualities that
perceivers actually utilize when perceiving various attributes in
faces. According to Gibson (1966, 1979), identifying the effective
stimulus information requires attention to the structured information in ecologically valid multidimensional, dynamic stimulus configurations. The importance of ecological validity is underscored by
evidence for differential encoding of facial qualities from moving vs.
static images (Bruce & Valentine, 1988; Humphreys, Donnelly, &
Riddoch, 1993; Kilts, Egan, Gideon, Ely, & Hoffman, 2003; O’Toole,
Roark, & Abdi, 2002; Pike, Kemp, Towell, & Phillips, 1997), images lit
from above versus those lit from below (Hill & Bruce, 1996; Johnston,
Hill, & Carman, 1992), 3D versus 2D images (Liu, Ward, & Young,
2006; O’Toole, Vetter, Nikolaus, & Bulthoff, 1997), and frontal versus
profile versus three–quarter images (Hill, Schyns, & Akamatsu,
1997; Zebrowitz & Lee, 1999). Once theoretically meaningful facial
configurations are identified, researchers might then test their
utilization using the lens model.
Perceiver Attunements
A third distinguishing tenet of the ecological approach is that the detection of social affordances depends on the perceivers’
attunements—their sensitivity to the stimulus information that reveals particular affordances. Although Gibson emphasized the ob-
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jective reality of affordances, he also emphasized their emergence
from the interaction of the environment and the perceiver. The truth
of Koffka’s observation that “A woman says love me” will depend on
the attunement of the perceiver. Attunements may reflect an innate
preparedness that is “species specific” (e.g., men but not monkeys
may be attuned to a woman’s sexual availability). Attunements also
may vary with perceivers’ behavioral capabilities (men but not boys
may be attuned to a woman’s sexual availability) or social goals (secular men but not priests may be attuned to a woman’s sexual availability). Regardless of their origin, attunements develop through a
process of perceptual learning that spans a variety of mechanisms
(cf. Goldstone, 1998). Although most research pertinent to these
mechanisms concerns the development of attunements to social category information, the perceptual learning mechanisms may also influence the development of attunements to identity, emotions, and
psychological traits (Zebrowitz & Montepare, 2006).
As already discussed, individual differences in face perception
shown by people with varying brain lesions have played a prominent
role in the dual process model of identity and emotion perception.
There also is evidence for variations in perceiver attunements to the
information conveyed by faces among normal perceivers who vary in
sex (Hall, 1984; Montagne, Kessels, Frigerio, de Haan, & Perrett, 2005),
ethnicity (Schimmack, 1996), demographic similarity to the target face
(Elfenbein & Ambady, 2003; Malatesta, Izard, Culver, & Nicholich,
1987; Meissner & Brigham, 2001), attitudes (Richeson & Trawalter,
2005), emotional state (Niedenthal, Halberstadt, Margolin, &
Innes–Ker, 2000), and personality traits (Barrett & Niedenthal, 2004;
Jones et al., 2005; Leppänen, Milders, Bell, Terriere, & Hietanen, 2004;
Surcinelli, Codispoti, Montebarocci, Rossi, & Baldaro, 2006; Tipples,
2006). A theoretically important individual difference variable is
perceiver age. In addition to evidence for variations between older
and younger adult perceivers (Malatesta et al., 1987), considerable research has documented infants’ attention to faces as well as their differential preferences for faces varying in social category, emotion
expression, babyfaceness, and attractiveness (cf. Zebrowitz &
Montepare, 2006a, for a review). A clear understanding of the development of attunements to the various attributes revealed in faces is
needed for a comprehensive theory of face perception.
Perceiver differences not only can influence the ease of recognizing
faces and the likelihood of attending to or perceiving certain facial at-
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tributes, but also they can moderate the effects of one face content on
the perception of another. For example, the effect of facial masculinity
on women’s judgments of men’s attractiveness is moderated by differences in their position in the menstrual cycle, with a greater preference
for more masculine–looking men near the time of ovulation (Johnston,
Hagel, Franklin, Fink, & Grammer, 2001; Penton–Voak et al., 2001).
One explanation for this effect is that facial masculinity signals good
genes, an affordance to which fertile women are putatively more attuned. However, the findings reported by Koehler, Rhodes, Simmons
and Zebrowitz (2006) in the current issue do not support this interpretation, inasmuch as fertile women showed no greater preference for
faces of men known to be healthier or for faces higher in symmetry or
averageness, both presumed indices of long–term health. Additional
research is needed to ascertain why it is that perceiver differences in
fertility moderate the effect of facial masculinity on perceived attractiveness. Other examples of moderating effects of perceiver individual
differences are that more racially prejudiced White perceivers showed
a greater readiness to perceive anger in emotionally ambiguous Black
faces than in comparable White faces (Hugenberg & Bodenhausen,
2003) and a greater tendency to perceive ethnically ambiguous
angry–looking faces as African American than as White American
(Hugenberg & Bodenhausen, 2004).
Individual differences in face perception are also manifested in
neural activation. For example, Golby et al., (2001) found greater activation of the FFA to same-race than to other-race faces, which predicted the own–race bias in face recognition. The article by
Willadsen–Jensen and Ito (2006) in the present issue also bears on
perceiver differences in the neural activation elicited by faces. Although they did not vary perceiver race, the differences in ERPs to
own– and other–race faces shown by White perceivers suggest that
perceivers from different racial backgrounds may show different
early perceptual processing of the same faces. Specifically, after initial attention to less racially familiar faces, later ERP components indicate greater processing of more familiar faces, with racially
ambiguous faces treated like faces of one’s own race. Still later in the
temporal processing of faces, racially ambiguous faces are differentiated from both of the other groups, capturing the explicit
categorizations provided by perceivers.
Willadsen–Jensen and Ito (2006) link their results to evidence that
prior experience determines the dimensions used to encode new
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faces. Whereas long–term perceptual experience may yield face processing mechanisms that are most sensitive to the features that differentiate among ingroup faces, even short–term experiences can have
an effect. For example, after people were adapted to male faces, a face
that was previously judged to be neutral was rated as female (Webster, Kaping, Mizokami, & Duhamel, 2004). Such adaptation effects
also lead to misperceptions of facial identity (Leopold et al, 2001),
and they can affect facial preferences. When adults were briefly exposed to consistent distortions of normal faces, there was an aftereffect marked by a shift toward the distorted faces in which faces
looked most normal and which faces looked most attractive (Rhodes,
Jeffery, Watson, Clifford, & Nakayama, 2003). These aftereffects
show that the average face used as a referent for coding facial identity may vary across time or perceivers (cf. Clifford & Rhodes, 2005),
and they are consistent with greater behavioral sensitivity to the central tendency of the population of faces encountered by the
perceiver. For example, infants show a visual preference for
own–race faces only if they are living in a racially segregated
environment (Bar–Haim, Ziv, Lamyu, & Hodes, 2006).
In summary, research clearly has shown perceiver differences in
face perception that affect the recognition of identity, social category,
emotions, and judgments of attractiveness. Moreover, neural underpinnings for some of these individual differences have been documented. These findings are consistent with the ecological tenet that
perceivers vary in their sensitivity to the stimulus information that
specifies various affordances. Considering the contribution to face
perception of variations in perceiver goals, behavioral capabilities,
and perceptual experiences promises to provide a more systematic
account of individual differences and to predict others.
CONCLUSIONS
Applying the tenets of an ecological approach to the study of face
perception can increase our understanding of this complex process
in several ways. An ecological approach calls attention to the fact
that similar affordances and stimulus configurations are shared by
different categories of faces, which may be reflected in similar neural processing of these different categories. This perspective contrasts with the emphasis on dissociations between the processing of
different attributes emphasized in the dual process model. The eco-
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logical approach also emphasizes the importance of examining the
behavioral concomitants and accuracy of face perception. In addition, high priority is placed on identifying the effective stimulus information for face perception, and the utility of a theoretically
driven effort to do so is stressed along with an emphasis on ecologically valid stimulus information. Finally, an ecological approach
draws attention to perceiver differences in face perception beyond
those accounted for by brain abnormalities. Variations among normally functioning perceivers are conceptualized as reflecting
differences in perceptual experiences, behavioral capabilities, and
social goals.
FUTURE DIRECTIONS
Face perception research has come a long way over the past half century. As reflected in this brief review of the literature, it is the
neuroscientists who have taken the lead in the burgeoning research
despite their slower start, perhaps due to the early discovery of intriguing deficits in face recognition associated with brain lesions. Although cognitive neuroscientists have made seminal contributions
to the understanding of face perception, there is much to be gained
from the input of other psychology subdisciplines. Indeed, face perception would seem to be at least as much the province of personality, developmental, and social psychology as cognitive
neuroscience. The comprehensive model of face perception envisaged in this article requires adding a heavy dose of the expertise of
psychologists who know how to study individual differences, developmental processes, and behavioral consequences. I hope this special issue will inspire the programmatic research on face perception
that is required if the growth in our understanding during the last 25
years is to continue profitably in the next quarter century. Finally, the
favored place of faces for the naïve psychologist will be fulfilled by
psychological science.
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