David I. Perrett

A specific neural substrate for perceiving facial expressions of disgust

Recognition of facial expressions is critical to our appreciation of the social and physical environment, with separate emotions having distinct facial expressions. Perception of fearful facial expressions has been extensively studied, appearing to depend upon the amygdala. Disgust — literally ‘bad taste’ — is another important emotion, with a distinct evolutionary history, and is conveyed by a characteristic facial expression. We have used functional magnetic resonance imaging (fMRI) to examine the neural substrate for perceiving disgust expressions. Normal volunteers were presented with faces showing mild or strong disgust or fear. Cerebral activation in response to these stimuli was contrasted with that for neutral faces. Results for fear generally confirmed previous positron emission tomography findings of amygdala involvement. Both strong and mild expressions of disgust activated anterior insular cortex but not the amygdala; strong disgust also activated structures linked to a limbic cortico–striatal–thalamic circuit. The anterior insula is known to be involved in responses to offensive tastes. The neural response to facial expressions of disgust in others is thus closely related to appraisal of distasteful stimuli.

Effects of sexual dimorphism on facial attractiveness

Testosterone-dependent secondary sexual characteristics in males may signal immunological competence and are sexually selected for in several species,. In humans, oestrogen-dependent characteristics of the female body correlate with health and reproductive fitness and are found attractive. Enhancing the sexual dimorphism of human faces should raise attractiveness by enhancing sex-hormone-related cues to youth and fertility in females,, and to dominance and immunocompetence in males,,. Here we report the results of asking subjects to choose the most attractive faces from continua that enhanced or diminished differences between the average shape of female and male faces. As predicted, subjects preferred feminized to average shapes of a female face. This preference applied across UK and Japanese populations but was stronger for within-population judgements, which indicates that attractiveness cues are learned. Subjects preferred feminized to average or masculinized shapes of a male face. Enhancing masculine facial characteristics increased both perceived dominance and negative attributions (for example, coldness or dishonesty) relevant to relationships and paternal investment. These results indicate a selection pressure that limits sexual dimorphism and encourages neoteny in humans.

Visual cells in the temporal cortex sensitive to face view and gaze direction

The direction of eye gaze and orientation of the face towards or away from another are important social signals for man and for macaque monkey. We have studied the effects of these signals in a region of the macaque temporal cortex where cells have been found to be responsive to the sight of faces. Of cells selectively responsive to the sight of the face or head but not to other objects (182 cells) 63% were sensitive to the orientation of the head. Different views of the head (full face, profile, back or top of the head, face rotated by 45° up to the ceiling or down to the floor) maximally activated different classes of cell. All classes of cell, however, remained active as the preferred view was rotated isomorphically or was changed in size or distance. Isomorphic rotation by 90–180° increased cell response latencies by 10–60 ms. Sensitivity to gaze direction was found for 64% of the cells tested that were tuned to head orientation. Eighteen cells most responsive to the full face preferred eye contact, while 18 cells tuned to the profile face preferred averted gaze. Sensitivity to gaze was thus compatible with, but could be independent of, sensitivity to head orientation. Results suggest that the recognition of one type of object may proceed via the independent high level analysis of several restricted views of the object (viewer-centred descriptions).

Menstrual cycle alters face preference

Women prefer slightly feminized male facial shapes. Such faces (Fig. 1a) are given positive personality attributions that might correlate with actual behaviour. In contrast, masculine features seem to signal immunological competence. Heritable benefits can be realized only if conception follows copulation, so women might be more attentive to phenotypic markers indicating immunological competence during the follicular phase of the menstrual cycle when conception is most likely,. Consistent with this hypothesis is the observation that womens preference for the odour of men with low fluctuating asymmetry (a correlate of testosterone-facilitated trait size and developmental stability) increases with the probability of conception across the menstrual cycle.Symmetrical men report more extra-pair copulation partners, and extra-pair copulation rates peak in midcycle. Here we show that female preference for secondary sexual traits in male face shapes varies with the probability of conception across the menstrual cycle.

Beauty in a smile: the role of medial orbitofrontal cortex in facial attractiveness

The attractiveness of a face is a highly salient social signal, influencing mate choice and other social judgements. In this study, we used event-related functional magnetic resonance imaging (fMRI) to investigate brain regions that respond to attractive faces which manifested either a neutral or mildly happy face expression. Attractive faces produced activation of medial orbitofrontal cortex (OFC), a region involved in representing stimulus-reward value. Responses in this region were further enhanced by a smiling facial expression, suggesting that the reward value of an attractive face as indexed by medial OFC activity is modulated by a perceiver directed smile.

Prototyping and transforming facial textures for perception research

Transforming facial images along perceived dimensions (such as age, gender, race, or health) has application in areas as diverse as psychology, medicine, and forensics. We can use prototype images to define the salient features of a particular face classification (for example, European female adult or East-Asian male child). We then use the differences between two prototypes to define an axis of transformation, such as younger to older. By applying these changes to a given input face, we can change its apparent age, race, or gender. Psychological investigations reveal a limitation with existing methods thats particularly apparent when changing the age of faces. We relate the problem to the loss of facial textures (such as stubble and wrinkles) in the prototypes due to the blending process. We review the existing face prototyping and transformation methods and present a new, wavelet-based method for prototyping and transforming facial textures.

Demystifying social cognition: a Hebbian perspective

For humans and monkeys, understanding the actions of others is central to survival. Here we review the physiological properties of three cortical areas involved in this capacity: the STS, PF and F5. Based on the anatomical connections of these areas, and the Hebbian learning rule, we propose a simple but powerful account of how the monkey brain can learn to understand the actions of others by associating them with self-produced actions, at the same time discriminating its own actions from those of others. As this system appears also to exist in man, this network model can provide a framework for understanding human social perception.

Symmetry and human facial attractiveness

Abstract Symmetry may act as a marker of phenotypic and genetic quality and is preferred during mate selection in a variety of species. Measures of human body symmetry correlate with attractiveness, but studies manipulating human face images report a preference for asymmetry. These results may reflect unnatural feature shapes and changes in skin textures introduced by image processing. When the shape of facial features is varied (with skin textures held constant), increasing symmetry of face shape increases ratings of attractiveness for both male and female faces. These findings imply facial symmetry may have a positive impact on mate selection in humans.

Responses of anterior superior temporal polysensory (stpa) neurons to “biological motion” stimuli

Cells have been found in the superior temporal polysensory area (STPa) of the macaque temporal cortex that are selectively responsive to the sight of particular whole body movements (e.g., walking) under normal lighting. These cells typically discriminate the direction of walking and the view of the body (e.g., left profile walking left). We investigated the extent to which these cells are responsive under biological motion conditions where the form of the body is defined only by the movement of light patches attached to the points of limb articulation. One-third of the cells (25/72) selective for the form and motion of walking bodies showed sensitivity to the moving light displays. Seven of these cells showed only partial sensitivity to form from motion, in so far as the cells responded more to moving light displays than to moving controls but failed to discriminate body view. These seven cells exhibited directional selectivity. Eighteen cells showed statistical discrimination for both direction of movement and body view under biological motion conditions. Most of these cells showed reduced responses to the impoverished moving light stimuli compared to full light conditions. The 18 cells were thus sensitive to detailed form information (body view) from the pattern of articulating motion. Cellular processing of the global pattern of articulation was indicated by the observations that none of these cells were found sensitive to movement of individual limbs and that jumbling the pattern of moving limbs reduced response magnitude. A further 10 cells were tested for sensitivity to moving light displays of whole body actions other than walking. Of these cells 5/10 showed selectivity for form displayed by biological motion stimuli that paralleled the selectivity under normal lighting conditions. The cell responses thus provide direct evidence for neural mechanisms computing form from nonrigid motion. The selectivity of the cells was for body view, specific direction, and specific type of body motion presented by moving light displays and is not predicted by many current computational approaches to the extraction of form from motion.

A region of right posterior superior temporal sulcus responds to observed intentional actions

Human adults and infants identify the actions of another agent based not only on its intrinsic perceptual features, but critically on the contingent relationship between its motion path and the environmental context [Trends Cogn. Sci. 7 (1995) 287; Cognition 72 (2003) 237]. Functional neuroimaging studies of the perception of agents and intentional actions, on the other hand, have mostly focussed on the perception of intrinsic cues to agency, like a face or articulated body motion (e.g. [J. Neurosci. 17 (1997) 4302; Neuroimage 8 (1998) 221; Trends Cogn. Sci. 4 (2000) 267; Nat. Neurosci. 3 (2000) 80; Neuroimage 13 (2001) 775; Proc. Natl. Acad. Sci. U.S.A. 98 (2001) 11656; Neuron 35 (2002) 1167; Neuron 34 (2002) 149, Neuroscience 15 (2003) 991; J. Neurosci. 23 (2003) 6819; Philos. Trans. R Soc. Lond. B. Biol. Sci. 358 (2003) 435]. Here we describe a region of the right posterior superior temporal sulcus that is sensitive not to articulated body motion per se, but to the relationship between the observed motion and the structure of the surrounding environment. From this and other aspects of the regions response, we hypothesize that this region is involved in the representation of observed intentional actions.