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Article is adapted from a monograph of mine – Alex.
Abstract
Beauty is said to be in the eye of the beholder, but perceptions of attractive faces are also represented in the brain of the beholder. Results from ERPs and fMRI implicate regions associated with decision-making processes in assessments of facial attractiveness. Attractiveness influences perceptions of trustworthiness and has implications for real-world behavior in social contexts. Further, attractiveness perceptions are moderated by a number of social cues and individual differences, including the proximity of other attractive or unattractive faces, the sex of the observer, and the setting in which faces are placed. Rather than being a domain-general assessment of beauty, facial attractiveness is processed early and fast, as well as distinctly from perceptions of non-facial beauty found in artwork, landscapes, and architecture. In the ongoing debate over modular versus distributed processing, perceptions of facial attractiveness may lean toward the local, similar to the general perception of faces.
Introduction To Facial Attractiveness
Although it is said “beauty is in the eye of the beholder,” perceptions of what an attractive face is are shared both within and across cultures (Coetzee et al., 2014; Jones & Hill, 1993; Langolis et al., 2000; Zebrowitz et al. 2012). Germane et al. (2015) found moderate heritability of perceptions of facial attractiveness in monozygotic twins. Recognition of facial attractiveness begins at a young age as well, with children and adults sharing perceptions of attractive faces (Ma et al., 2016). Even babies as young as four months old show a preference for more attractive faces (Samuels et al., 1994).
Facial attractiveness is often studied in the context of human mating and romantic relationships. Physical and facial attractiveness consistently rank high in both self-reported (Buss & Barnes, 1986; Foster, 2008) and revealed (Asendorpf et al., 2011; Eastwick et al., 2021; Luo & Zhang, 2009; Riggio & Woll, 1984; Roth et al., 2021; Valentine et al., 2014) mate preferences. Facial attractiveness also predicts both short and long term mate desirability (Valentine et al., 2014).
However, the effects of facial attractiveness do not end with romantic desire. Rather, attractiveness carries over to other areas of life. Dion et al. (1972) first identified the “what is beautiful is good” stereotype, also known as the Halo Effect, where more facially attractive individuals are perceived to have more positive character traits. This effect has since been replicated over the decades (Eagly et al., 1991; Langolis et al. 2000) and across cultures (Batres & Shiramizu, 2022; Kordsmeyer et al., 2022). A meta-analysis of job-related outcomes (Hosoda et al., 2003) found moderate sized biases for physical attractiveness across genders, age groups, and professional status in the workplace. Physically attractive individuals may also receive preferential treatment in the criminal justice system (Mazzella & Feingold, 1994; Sigall & Ostrove, 1975). What is beautiful is indeed perceived as good, but not only in romantic contexts.
Facial symmetry (Langlois & Roggman, 1990; Møller & Thornhill, 1994; Grammer & Thornhill, 1994) and facial averageness (Langlois & Roggman, 1990; Langlois et al., 1994; Rhodes, 2006) are the most studied objective features that predict perceptions of facial attractiveness. Facial markers associated with sex steroids that influence facial dimorphism may also predict perceptions of facial attractiveness, although results have been mixed (Chen et al., 2017; Hester et al., 2021) showing preferences for higher (Johnston & Franklin, 1993) and lower (Perrett et al., 1998; Rhodes, 2006) levels of facial dimorphism.
Objective facial features associated with perceptions of facial attractiveness may further vary by culture. For example, in Perret et al. (1998), a Japanese sample was used. Kočnar et al. (2019) and Marcinkowska et al. (2019) found lower preferences for dimorphic faces in Western nations than in less developed nations.
Given the role of attractiveness in mate selection we should also expect a similar evolved ability to detect or discern attractive faces. This may manifest as a specialized bias for attractive faces, against unattractive faces, or both. If this is the case then it may be reflected in regions highly specific to faces and further specific to attractive (or unattractive) faces. If not, we may expect more generalized activation associated with face perception, with reward, or with the perception of objects in general.
Neuroanatomy Of General Facial Perception
A region of the fusiform gyrus of the inferior temporal cortex, now called the fusiform face area (FFA), has been associated with the perceptions of faces in the classic research of Kanwisher et al. (1997). Previous to this, Damasio et al. (1982) summarized the results of postmortem lesions associated with prosopagnosia, an impairment of facial recognition. This earlier research described bilateral and unilateral lesions of the fusiform gyrus and the occipital lobe. However, right hemisphere lesions were not independently predictive of prosopagnosia. Nor did patients who have undergone right or left hemispherectomy experience prosopagnosia (Damasio et al., 1975; as cited in Damasio et al., 1982). This early research associating lesions with impairments in the ability to perceive faces indicated a link between the FFA and perceptions of faces. However, in some of the cases Damasio et al. (1975) reviewed the lesions were large and nonspecific. It was not until later that more clear associations with the FFA and perceptions of faces were found across neuroimaging and other methodologies (Grill-Spector et al., 2004; Kanwisher & Yovel, 2006).
More recently, trends in neuroscience have begun to shift away from understanding the brain in models of region-specific modularity. Instead, perception and processing is increasingly analyzed as a distributed network. Evidence of this in facial processing, for example, is that prosopagnosia may arise even without lesion disruption to the fusiform face area (Bobes et al., 2021; Cohen et al., 2019; de Gelder et al., 2022).
However, despite recent trends toward looking for distributed rather than modular function, evidence for the region-specific processing within the FAA has remained strong (see review by Kanwisher & Yovel, 2006). The shift away from modular or region-based understanding of brain functions is also relatively new and less research has approached perceptions of faces from this perspective. Research on patients with FFA lesions has found a double dissociation for facial and object recognition (Moscovitch et al., 1997, as cited in Kanwisher & Yovel, 2006), particularly when lesions are highly localized (Wada & Yamamoto, 2001, as cited in Kanwisher & Yovel, 2006). While the brain is undoubtedly not a puzzle-piece collection of interlocking regions with isolated roles, the degree to which perceptions are localized is not the same across tasks. Perceptions of faces remain strongly associated with, and dependent on, localized regions.
Electrophysiological Measures Of Face Perception And Facial Attractiveness
The N170 event-related potential (ERP) is the potential component most closely associated with facial stimuli (Bentin et al., 1996; Hinojosa et al., 2015). The M170 magnetoencephalography (MEG) response has also been associated with perceptions of faces (Proverbio, 2021). While ERPs are not specific anatomical representations of brain activity, the N170 and M170 are spatially correspondent to the location of the fusiform gyrus (Deffke et al., 2007; Monroe et al., 2013), consistent with the localization of the FFA. As these represent regions most implicated in the general perception of faces, they are a starting point for seeing how perceptions of facial attractiveness differ from a general representation of faces.
Given this background for facial processing, we may start with the expectation to see activation for faces in the FFA in neuroimaging and in N170/M170 electrophysiological responses. If facial attractiveness were merely represented as higher salience for a face we might also expect most activation to be localized to the FFA in neuroimaging. Alternatively, if facial attractiveness is processed elsewhere — either in a distributed or localized fashion — we might expect a pattern of activation outside of the FFA. This might be detected in ERPs outside of the N170/M170, or in neuroimaging results that show unique activation outside of the FFA.
Facial Attractiveness ERPs
Jin et al. (2017) found an N200 activation for facial attractiveness. The N200 is an ERP implicated in mismatches in decision-making (Xiao, 2019) and thus may be our first indication that assessments of facial attractiveness interact with, or are in and of themselves, a decision-making process. In addition, there was an interaction effect between lending in an economic experiment and facial attractiveness, with unattractive faces eliciting a larger N200 while lending was reduced. This helps to bring forth the neural underpinnings of the “beauty premium” in decision-making, or a negative bias toward less attractive faces. In an economic ultimatum game with fair and unfair offers, Ma et al. (2015; 2017) found an early frontal LPP, FRN, and early P300 response for facial attractiveness. Offers were accepted more when associated with attractive faces and the FRN and P300 amplitudes were higher for unattractive faces in the unfair offer condition. A classic interpretation of attractiveness interactions in decision-making experiments has been that attractive participants are perceived as more trustworthy (Wilson & Eckel, 2006). However, the results of Jin et al. (2017) and Ma et al. (2015;2017) offer an alternate explanation. Rather than attractive participants being perceived as more trustworthy, cohorts may simply care less when they are uncooperative.
Faces are not static and thus facial expression may change both our perceptions of facial attractiveness and other judgments. Calvo et al. (2018) found an effect of facial expression on judgments both of facial attractiveness and trustworthiness. Happy faces were judged as more attractive and trustworthy than neutral or angry faces. Additionally, neutral faces were judged as more attractive and trustworthy than angry faces. N170 and EPN responses were more pronounced for faces with any emotional expression. The EPN is a component implicated in the processing of emotional stimuli (Schupp et al., 2003), so this may reflect the emotional state the face is presented in. Activation was pronounced in the right parieto-occipital for N170. P3b responses showed enhanced positive activation in both attractive and trustworthy conditions. Importantly, processing of attractiveness preceded trustworthiness. Results from Jin et al. (2017) and Ma et al. (2015;2017) raised the question of how attractiveness and trustworthiness interacted to produce reactions to unfair behavior in decision-making. The results of Calvo et al. (2018) may show that attractiveness acts as a priming effect, biasing subsequent perceptions in favor of attractive individuals.
This is consistent with the observation that perceptions of facial attractiveness are processed quickly and without conscious awareness. Luo et al. (2019) found differential activation associated with the occipito-temporal area with very brief exposure to attractive faces (6 Hz, or six faces per second). Olson & Marshuetz (2005) similarly found rapid processing of attractive faces when presented in quick series and when masked. Although perceptions of facial attractiveness may represent a decision-making process, it is likely that they at least partially precede conscious awareness that a face is subjectively attractive to an observer.
Further related to emotion, Huang et al. (2016) found a large effect of social cognition on perceptions of facial attractiveness. When target faces were paired with faces smiling at them, subjects rated those faces higher in attractiveness. If the paired faces were frowning at the target face, subjects rated the target lower in attractiveness. P3 responses showed a main effect for pair condition, with a larger response for smiling paired faces. The P3 is an ERP closely associated with decision-making tasks (De Rosa et al., 2017), again implicating the role of decision-making in attractiveness assessments. Han et al. (2019) similarly found a large effect of faces paired with a positive sentence perceived as more attractive than those paired with a negative sentence. Thus, the priming effect of a positive stimuli on perceptions of attractiveness may extend past the visual. A positive LPC was elicited, a component associated with memory tasks (Stróżak et al., 2016), which may reflect the involvement of word recall. An interaction effect with facial familiarity was found as well, with a higher positive LPC and EPN for familiar faces paired with a positive sentence. As the LPC has also been implicated in the recall of familiar words (Stróżak et al., 2016), this may also indicate a familiarity effect. This would be consistent with classic behavioral results on perceptions of attractiveness; faces that are familiar to participants are consistently rated as more attractive (Peskin & Newell, 2014).
The interaction effects of unfair offers and unattractive faces across these studies may indicate joint regional processing for negative economic and facial stimuli. This may also be gleaned from interactions for trustworthiness and attractiveness, as well as for more positive emotional faces over negative emotional faces. The effect of social cognition on perceptions of facial attractiveness further supports this.
Differential activation for facial attractiveness in the economic experiments, as well as earlier processing of facial attractiveness than trustworthiness, is consistent with how we think of the Halo Effect or “beauty is good” stereotype. That is, that our perception of attractiveness precedes (and is potentially is casual for) our perceptions of trustworthiness and bias in favor of attractive individuals. The higher ERP activation of faces paired with positive stimuli (smiling faces, positive vignettes) also demonstrate an effect in the opposite direction. Not only do we perceive what is beautiful as good, but we perceive what is good as beautiful as well.
Sex Differences in ERPs
Up to this point, activation associated with facial attractiveness has been discussed in general terms. However, past behavioral research in social psychology has indicated that men and women value attractiveness differently, with men placing more importance on physical attractiveness (Walster et al., 1966; Sprecher, 1989). Are ERPs and behavioral results within neuroscience consistent with these observations?
Zhang and Deng (2012) found longer reaction times for men when rating faces as attractive rather than unattractive. P1 (a component associated with perception of faces) and P2 responses were larger for attractive faces for men and women, although latency was larger for men. Positive N300 and P3b responses were greater for attractive faces, while latency was also longer for men. N170 responses showed an interaction effect for attractiveness and gender for men. Lu et al. (2014) similarly found a higher N170 response for attractive faces in male subjects, but not for female subjects. These results highlight gender differences in the processing of attractive faces, both in response time and in ERP response. While the subjective feeling about a face a participant holds cannot be derived from the magnitude of an ERP, differential activation between men and women is consistent with the difference in subjective weight placed on attractiveness by men and women in behavioral data.
ERP responses to both attractive and unattractive faces are stronger than for average faces. Differential activation of both unattractive and attractive faces when compared with average faces has been consistent in neuroimaging as well (the following section). For example, Schacht et al. (2008) found a higher LPC for both attractive and unattractive faces relative to averagely attractive faces. Trujillo et al. (2014) found a larger N170 response for unattractive faces, as well as faster response times for both attractive and unattractive faces. Zhang et al. (2011) found that attractive female faces generated larger negative N300 responses and higher amplitude LPC responses for attractive faces. Attractive faces were also more easily recognized when mixed with novel facial stimuli.
This effect has been found when viewing the faces of infants as well (Hahn et al., 2014). There was no sex difference in ratings of attractive infant faces. Less attractive infant faces predicted larger amplitude early N170 and P2, as well as larger late LPC responses. This and subsequent neuroimaging results for differential activation when viewing infant faces further indicates that facial attractiveness is not reducible to perceptions of romantic or sexual stimuli. Rewarding faces elicit similar activation even when they are not objects of romantic desire.
Carbon et al. (2018) found that facial gender elicited an earlier LRP and N200 by 300ms than facial attractiveness, indicating that processing the perceived gender of a face precedes our perceptions of an attractive face. The N200 response has been associated with inhibition; this may indicate we may inhibit our judgments of attractiveness based on a gender preference (likely romantic or sexual) subsequent to processing facial attractiveness. Hahn et al. (2014) similarly found higher P2 amplitude for viewing opposite-sex faces.
Brain Imaging of Facial Attractiveness Perceptions
The orbitofrontal cortex (OFC) is frequently implicated in the perception of attractive faces (O’Doherty et al., 2003; Tsukiura & Cabeza, 2011; Ueno et al., 2014; Winston et al., 2007). This region is associated with reward processing and social decision making (Kringelbach & Rolls, 2004; Peters & Büchel, 2010; Rolls & Grabenhorst, 2008). Lesions in the OFC have been shown to impair reward-based learning (Hornak et al., 2004). Adjacent to the OFC, the ventromedial prefrontal cortex (vmPFC) is also activated when viewing both attractive and unattractive faces (Smith et al., 2010; Spreckelmeyer et al., 2013; Yu et al., 2013). Functional interconnectivity between these two regions is seen in tasks involving emotion-mediated and social decision making (Levens et al., 2014; Rudebeck et al., 2008).
The anterior cingulate cortex (ACC) (Rudebeck et al., 2008), amygdala, and ventral striatum (Hiser & Koenigs, 2018; Rudebeck et al., 2008) further constitute part of a network in processing reward and value based decisions. The nucleus accumbens (NAc) specifically within the ventral striatum has been associated with perceptions of facial attractiveness (Kranz & Ishai, 2006; Spreckelmeyer et al., 2013; Zaki et al., 201) and the amygdala with a higher nonlinear response in viewing unattractive and attractive faces (Yu et al., 2013).
Together, these areas constitute the primary regions and a network of processing facial attractiveness. Again it is notable that areas associated with perceptions of facial attractiveness overlap with those implicated in decision-making processes. However, faces are not static and perceptions of facial attractiveness change with context. Additionally, what influences perceived attractiveness of a face varies by individual differences. Neuroimaging results regarding facial attractiveness mirror those seen in ERP studies, in addition to the implication of decision-making processes, with differences emerging for gender, sexuality preferences, infant faces, non-romantic contexts, and social biases.
O’Doherty et al. (2003) found activation in the medial OFC when viewing both attractive and unattractive faces. A smiling facial stimulus resulted in higher activation. Although O’Doherty et al. did not find an association between smiling or non-smiling faces and attractiveness, past research has found that smiling faces are perceived as more attractive (Otta et al., 1996, as cited in O’Doherty et al.; Winston et al., 2007). This is consistent with the ERP results of Huang et al. (2016) in the previous section, where smiling faces were perceived as more attractive and elicited differential ERP responses.
Winston et al. (2007) replicated the findings of O’Doherty et al. (2007) for attractive faces and found a linear response to facial attractiveness in the OFC; more attractive faces were associated with higher OFC activation. However, unattractive faces were not associated with higher OFC activation. Additionally, Winston et al. found a nonlinear response in the amygdala. The least and most attractive faces were associated with a higher amygdala response (see also: Kranz & Ishai, 2006). Interaction with gender was further found for activation in the ACC for men viewing female faces, but not for women viewing male faces. This is consistent with higher N170 results for men when viewing female faces mentioned in the previous section (Lu et al., 2014), in addition to deformations in N170 responses associated with lesions in the ACC (Schäfer et al., 2007). That both the least and most attractive faces elicited the highest response may indicate a role of salience or facial distinctiveness in perceptions.
Individual Differences in Attractiveness Perceptions
Ota & Nakano (2021) similarly found a positive association with facial attractiveness in the NAc and a negative association in the amygdala. Additionally, activation in both areas was higher for viewing the faces of others rather than one’s own face. Shen et al. (2016) found activation in the caudate nucleus and the OFC for more attractive faces. There was nonlinear activation in the amygdala for men, with more attractive faces showing higher activation. Facial attractiveness was manipulated as a factor of facial symmetry; more symmetrical faces predicted higher attractiveness and greater activation in these areas. Individual differences in perceptions of facial attractiveness are associated with differential activation as well. Participants who gave higher ratings to faces on average also had higher activation in the right middle temporal gyrus (Vartanian et al., 2013). The left anterior frontal cortex and right middle occipital gyrus were both activated more when faces were rated as more attractive.
Additional sex differences when observing attractive and unattractive faces have been found as well. When viewing an opposite-sex face, there is higher activation in the ventral tegmental, nucleus accumbens and vmPFC (Spreckelmeyer et al., 2013). The temporo-parietal junction (TPJ) is also recruited when women view opposite-sex faces, but not for men. Winston et al. (2007) found higher activation for men viewing female faces in the ACC. Kocsor et al. (2013) found more activation for attractive faces in the medial gyri, fusiform gyrus and anterior cingulate, but women showed additional activation in the insula.
There are also age related differences associated with processing of opposite-sex faces. Telzer et al. (2015) found higher activation in the amygdala of young and late children when viewing opposite-sex faces. By the mid teens, the trend reversed, with greater activation in the amygdala associated with the viewing of same-sex faces. This mirrors strong same-sex ingroup biases that decrease in magnitude with age (Aydt & Corsaro, 2003, as cited in Telzer et al., 2015). This was labeled the “cooties effect” and reflects observations about how children’s gendered play changes with time. The youngest children form same-gender groups and shun opposite-gender playmates.
What we perceive as attractive is also influenced by our own sexual preferences. Kranz & Ishai (2006) found OFC activation that was consistent with raters sexual orientation. Men with a sexual preference for men had higher activation in the OFC and the mediodorsal thalamic nucleus (mdT) when viewing attractive male faces. The same pattern of activation persisted for women who have a sexual preference for women when viewing attractive female faces. The general pattern of higher activation in the OFC for attractive faces was also found across both genders and sexual preferences. Brain activation reflects the perception of attractive faces across genders and sexuality, but it also shows heightened activation when the faces we view are congruent with our own sexual preferences. This is consistent with perception of facial attractiveness having not only an aesthetic, but a sexual or relational, aspect.
Attractiveness, Attention, and Contextual Cues
Attractiveness has been implicated in attention tasks, with more voluntary attention directed toward more attractive faces (Roth et al., 2023). Contrary to expectations, Winston et al. (2007) did not observe more activation in the OFC when attention was focused on an attractive face. However, Kampe et al. (2001) found higher activation in the ventral striatum for attractive faces in a direct gaze task, but no effect of attractiveness in the ventral striatum without the interaction of gaze. It is possible that activation associated with attention to attractive faces is more distributed and varies by level of attention. This condition in Kampe et al. (2001) roughly mimics eye contact and may have implications for eye contact in assessments of facial attractiveness. Past research has shown higher assessments of facial attractiveness in conditions of sustained gaze (Ewing et al., 2010; Shimojo et al., 2003) and that we prolong our gaze towards attractive facial stimuli (Leder et al., 2016). Even subtle shifts in behavior may change who we view as attractive, or who views us as attractive, which may be in turn reflected in neuroimaging results.
Perceptions of attractiveness may be also assessed as a comparison. In other words, we may perceive attractiveness not as absolute but referential to that which is less attractive. Kedia et al. (2013) found similar activation in the inferior frontal gyrus (IFG) and the dorsomedial prefrontal cortex (dmPFC) for comparisons of facial attractiveness and comparisons of physical size. Additionally, there was a distance effect in behavioral response. Response times for facial attractiveness judgments were longer when faces had a higher disparity in attractiveness, consistent with past research (Winston et al., 2007). That comparisons shift perceptions in attractiveness is consistent with perceptions of facial attractiveness being a decision-making process. Faces that are presented concurrently may represent an anchor or a starting point from which subsequent faces are perceived as more attractive. This is also consistent with behavioral results in naturalistic conditions. For example, Taubert et al. (2016) found that the perceptions of past faces in an online dating paradigm resembling the Tinder application affected perceptions of subsequent faces. Exposure to more or less attractive faces makes those that follow seem more or less attractive.
Similarly, anticipation of viewing an attractive or unattractive face may prime subsequent perceptions of a face. Yu et al. (2013) found activation in the bilateral amygdala and ventral striatum associated with the anticipation of viewing an unattractive or attractive face. When a masked face was presented the amygdala and ventral striatum were activated. Activation was enhanced when the face was revealed to be congruent with the participant’s anticipation. When a participant anticipated the face as being attractive, activation was subsequently higher. If the revealed face was incongruent with the participant’s anticipation, connectivity between the vmPFC showed less activation. Here, anticipation of viewing a face worked similarly to actually perceiving a face.
Beauty and Goodness in the Brain
The classic research of Dion et al. (1972) first identified the “what is beautiful is good” stereotype, also called the Halo Effect. Indeed, our joint perception of facial beauty and moral goodness is also reflected in the neuroanatomy of the brain. Tsukiura & Cabeza (2011) found that assessments of both attractiveness and moral goodness were associated with higher activation in the medial OFC and with lower activation in the insular cortex. Moral judgments represent a decision-making process and this further reinforces the observation that attractiveness assessments themselves may be a form of decision-making. OFC activation for facial attractiveness and goodness was highly correlated (r = .86). Behavioral response times were also faster for faces that were both more and less attractive, a pattern that repeated with faces associated with high or low levels of moral goodness. High activation in the OFC and low activation in the insular cortex may thus implicate a dual process for the beauty is good stereotype: both a bias in favor of attractive faces and goodness, as well as a bias against unattractive faces and what is not good.
Luo et al. (2019) similarly identified a shared network for judgments of moral goodness and facial attractiveness in the middle occipital gyrus (MOG) and the OFC. Activation in the PFC occurred when moral judgments were incongruent with facial judgments, for example when moral badness was paired with an attractive face. This may indicate an additional level of processing recruiting areas more closely associated with decision making when teasing out the conflict between a positive and negative stimuli.
Attractiveness and Social Context
The social environment that we are immersed in may also reflect neurological activation associated with our perceptions of an attractive face. Zaki et al. (2011) found social adjustment to perceptions of facial attractiveness modulate response in the OFC and nucleus accumbens. When participants learned how their peers rated a face, follow-up activation in these regions adjusted up (for more positive peer ratings) or down (for less positive peer ratings). The new value attributed to faces was reflected in these two regions of the brain associated with the processing of value and reward. This was consistent with past research (Klucharev et al., 2009, as cited in Zaki et al., 2011).
We tend to think of perceptions of physical attractiveness as primarily related to sexual attraction or romantic aspirations. Indeed, we have seen that the two are not unrelated (Kranz & Ishai, 2006). However, processing a rewarding face can be dissociated from perceptions of faces or stimuli associated with purely romantic or sexual interest. For example, Nitschke et al. (2004) found higher activation in the OFC when mothers viewed faces of their own infants over the faces of strange infants. This is consistent with the ERP responses when viewing infant faces described in the previous section (Hahn et al., 2014).
Research on the Halo Effect also has meaningful implications for biases in favor of attractiveness in work and nonsexual social interactions. Neurological responses to economic rewards have been shown to be similar to the reward of viewing an attractive face, with similar activation in the vmPFC when receiving economic rewards and when viewing attractive faces (Smith et al., 2010). Perceptions of attractiveness may also bias our empathy. Jankowiak-Siuda (2015) found higher activation in the anterior insula and the anterior cingulate cortex, structures associated with processing empathy, when viewing unattractive men and attractive women in painful situations.
Neuroaesthetics; Nonfacial Stimuli, Facial Stimuli and Beauty
Up to this point we have identified correlates of facial attractiveness in electrophysiological and neuroimaging measures. However, the question of general perceptions of beauty remains. Not all beautiful stimuli are faces. One may also imagine artwork, architecture, or inanimate natural beauty, among other stimuli. If these share correlates with perceptions of facial attractiveness it may indicate that facial attractiveness is recognized as a form of general reward. Conversely, if they are different, this may indicate that there are face-specific perceptions of attractiveness. A recent meta-analysis assessed FMRI activation associated with beautiful faces and beautiful nonhuman stimuli (Chen et al., 2020). The anterior medial prefrontal cortex (aMPFC) was consistently identified in representations of beautiful artwork, while the vmPFC (including the OFC) and ventral striatum were associated with attractive faces. Nonetheless, research often indicates that perceptions of attractiveness in faces and beauty in objects have some overlap.
Research on the aesthetic judgments of abstract shapes has shown activation in the frontal medial cortex (FMC) and left intraparietal sulcus (Jacobson et al., 2006), consistent with areas associated with moral judgments of goodness but less consistent with perception of facial attractiveness. Vartanian & Goel (2004) found activation in the bilateral occipital and fusiform gyri, areas associated with perceptions of faces, with higher ratings of paintings. Aesthetic judgments of paintings similarly elicited activation in the OFC and vmPFC, highly consistent with judgments of faces (Kirk et al., 2009). Additionally, context mediated the neuroanatomical response. When paintings were presented associated with an art gallery, rather than in a computer condition, activation was higher. This is consistent with research up to this point indicating a role of framing and contextual cues on subjective perceptions of attractiveness (Zaki et al., 2011). However, paintings may contain a confound if they include faces within them. To rule this out, one may look to artistic stimuli without the possibility of facial associations. Kirk et al. (2009) found activation in the OFC, NAc, and cingulate gyrus when making aesthetic judgments of both faces and architecture. Activation in this area was different between facial and architecture conditions, as well as between expertise (architects viewing) and non-expertise conditions.
Pegors et al. (2015) found activation in the vmPFC when viewing attractive faces and attractive scenes or locations. However, the OFC responded uniquely to faces both independently of attractiveness and for attractive faces. While aesthetic judgments of faces and non-faces seem to share overlapping brain architecture, it does not seem to be the case that they are exactly the same. The structures involved when processing beauty are shared, while BOLD signals still indicate a face-specific role for regions of the OFC. Further supporting an overlapping but not identical network for aesthetic judgments of faces and non-faces, Troiani et al. (2016) found face-specific activity in the OFC when contrasting responses to attractive faces and attractive foods. While both stimuli recruited the OFC, the medial OFC responded selectively to faces while lateral regions of the OFC responded both to presentations of attractive faces and attractive foods. The OFC may therefore have both a face-specific and general role in processing aesthetic judgments.
In addition to places, art, and food, events may be another pleasing stimuli to contrast with perceptions of rewarding faces. Kawabata & Zeki (2008) found differential but overlapping activation for desirable events, objects and persons in the OFC. This further supports the OFC as a reward center for aesthetic judgments while also reserving a unique pattern of activation when viewing attractive faces.
Discussion
Beauty is said to be in the eye of the beholder, but neuroscience can show us the extent to which beauty is also in the brain of the beholder. We consistently see activation in the vmPFC and OFC across conditions associated with beauty, both facial and non-facial. This may give us reason to suspect that perceptions of beauty are both a process of reward and a process of decision making. However, we must be careful in interpreting the results recursively. To see activation in areas associated with reward and decision making does not necessarily mean that beauty is a function of, or related, to these. Unique processes may share the same brain regions. Areas of the brain may be shared in associated activation, even across unrelated processing or tasks. Future research may look toward behavioral rather than neuroimaging results to distinguish between perceptions of facial attractiveness and reward.
A limitation in attractiveness research is the use of facial photos and what are fundamentally unnatural conditions when assessing attractiveness. When we interact with people we are not interacting with a static, masked, and controlled face. The research above has shown how even subtle changes, such as eye gaze, as well as socially-mediated conditions, may have an effect upon both behavioral ratings of attractiveness and corresponding BOLD or ERP activation. The variables involved in assessments of attractiveness given real life interactions are potentially limitless.
Electrophysiological methodology could be combined with recent speed-dating methodology in the study of attractiveness and mate choice. Electrodes may be placed and worn while engaging in more naturalistic dating interactions. Cooper et al. (2012) used fMRI in conjunction with speed-dating to assess correlations between BOLD signals and who was selected in dating scenarios. However, the nature of fMRI restricts its use in many real-world conditions.
Surprisingly little research has been done within attractiveness using videos rather than photos. This is true both for behavioral assessments of attractiveness in psychology and for facial attractiveness research in neuroscience. Rather than continuing to rely upon still photos of faces, using videos of facial stimuli may help to replicate more natural conditions. This also has its limitations however, in that control for third variables is lower in video-based conditions.
There are two methodologies for assessing attractiveness that should be mentioned: ratings by participants and ratings by observers, or “objective” ratings. Mean agreement on facial ratings may be high, but at the same time individual variation is also high within pairs of raters(Hönekopp, 2006). Responses to faces rated as attractive by participants may be different from responses to faces given an “objective” rating by mean consensus. The studies in this review used both methodologies and differences in results across papers may be due in part to this. Small sample sizes used in neuroimaging may inflate the effect of inter-rater variation as well.
Little research has been done to test the causal relationship of brain regions with perceptions of facial attractiveness. This may be the next frontier in neuroscientific facial attractiveness research. Ferrari et al. (2015) found that transcranial direct current stimulation (tDCS) applied to the dorsolateral PFC increased perceptions of facial attractiveness. Stimulation did not affect other face-specific perceptions, so this may further support the hypothesis of a face-specific region in this area related to perceptions of attractiveness.
In the brain, some functions may have more specific localization and some may be more distributed. A great deal of the research on facial attractiveness, as well as facial perception in general, has been through the lens of regional functionality. Examining networks of connectivity in perceptions of facial attractiveness will be another step forward in this field. This, as well as devising new methodology for studying perceptions of attractiveness in more natural conditions, may be where researchers should focus future attention.
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