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.

The amygdala is thought to play a crucial role in emotional and social behaviour. Animal studies implicate the amygdala in both fear conditioning and face perception. In humans, lesions of the amygdala can lead to selective deficits in the recognition of fearful facial expressions and impaired fear conditioning, and direct electrical stimulation evokes fearful emotional responses. Here we report direct in vivo evidence of a differential neural response in the human amygdala to facial expressions of fear and happiness. Positron-emission tomography (PET) measures of neural activity were acquired while subjects viewed photographs of fearful or happy faces, varying systematically in emotional intensity. The neuronal response in the left amygdala was significantly greater to fearful as opposed to happy expressions. Furthermore, this response showed a significant interaction with the intensity of emotion (increasing with increasing fearfulness, decreasing with increasing happiness). The findings provide direct evidence that the human amygdala is engaged in processing the emotional salience of faces, with a specificity of response to fearful facial expressions.

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.

We report four experiments investigating the perception of photographic quality continua of interpolated ('morphed') facial expressions derived from prototypes of the 6 emotions in the Ekman and Friesen (1976) series (happiness, surprise, fear, sadness, disgust and anger). In Experiment 1, morphed images made from all possible pairwise combinations of expressions were presented in random order; subjects identified these as belonging to distinct expression categories corresponding to the prototypes at each end of the relevant continuum. This result was replicated in Experiment 2, which also included morphs made from a prototype with a neutral expression, and allowed 'neutral' as a response category. These findings are inconsistent with the view that facial expressions are recognised by locating them along two underlying dimensions, since such a view predicts that at least some transitions between categories should involve neutral regions or identification as a different emotion. Instead, they suggest that facial expressions of basic emotions are recognised by their fit to discrete categories. Experiment 3 used continua involving 6 emotions to demonstrate best discrimination of pairs of stimuli falling across category boundaries; this provides further evidence of categorical perception of facial expressions of emotion. However, in both Experiment 1 and Experiment 2, reaction time data showed that increasing distance from the prototype had a definite cost on ability to identify emotion in the resulting morphed face. Moreover, Experiment 4 showed that subjects had some insight into which emotions were blended to create specific morphed images. Hence, categorical perception effects were found even though subjects were sensitive to physical properties of these morphed facial expressions. We suggest that rapid classification of prototypes and better across boundary discriminability reflect the underlying organisation of human categorisation abilities.

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