Lisa A. Parr

The face inversion effect, or impaired recognition of upside down compared to upright faces, is used as a marker for the configural processing of faces in primates. The inversion effect in humans and chimpanzees is strongest for categories of stimuli for which subjects have considerable expertise, primarily conspecifics’ faces. Moreover, discrimination performance decreases linearly as faces are incrementally rotated from upright to inverted. This suggests that rotated faces must be transformed, or normalized back into their most typical viewpoint before configural processing can ensue, and the greater the required normalization, the greater the likelihood of errors resulting. Previous studies in our lab have demonstrated a general face inversion effect in rhesus monkeys that was not influenced by expertise. Therefore, the present study examined the influence of rotation angle on the visual perception of face and nonface stimuli that varied in their level of expertise to further delineate the processes underlying the inversion effect in rhesus monkeys. Five subjects discriminated images in five orientation angles. Results showed significant linear impairments for all stimulus categories, including houses. However, compared to the upright images, only rhesus faces resulted in worse performance at rotation angles greater than 45°, suggesting stronger configural processing for stimuli for which subjects had the greatest expertise.

Faces are salient stimuli for primates that rely predominantly on visual cues for recognizing conspecifics and maintaining social relationships. While previous studies have shown similar face discrimination processes in chimpanzees and humans, data from monkeys are unclear. Therefore, three studies examined face processing in rhesus monkeys using the face inversion effect, a fractured face task, and an individual recognition task. Unlike chimpanzees and humans, the monkeys showed a general face inversion effect reflected by significantly better performance on upright compared to inverted faces (conspecifics, human and chimpanzees faces) regardless of the subjects' expertise with those categories. Fracturing faces alters first- and second-order configural manipulations whereas previous studies in chimpanzees showed selective deficits for second-order configural manipulations. Finally, when required to individuate conspecific's faces, i.e., matching two different photographs of the same conspecific, monkeys showed poor discrimination and repeated training. These results support evolutionary differences between rhesus monkeys and Hominoids in the importance of configural cues and their ability to individuate conspecifics' faces, suggesting a lack of face expertise in rhesus monkeys.

The ability to recognize and accurately interpret facial expressions are critical social cognition skills in primates, yet very few studies have examined how primates discriminate these social signals and which features are the most salient. Four experiments examined chimpanzee facial expression processing using a set of standardized, prototypical stimuli created using the new ChimpFACS coding system. First, chimpanzees were found to accurately discriminate between these expressions using a computerized matching-to-sample task, and recognition was impaired for all but one expression category when they were inverted. Third, a multidimensional scaling analysis examined the perceived dissimilarity among these facial expressions revealing 2 main dimensions, the degree of mouth closure and extent of lip-puckering and retraction. Finally, subjects were asked to match each facial expression category using only individual component features. For each expression category, at least 1 component movement was more salient or representative of that expression than the others. However, these were not necessarily the only movements implicated in subject's overall pattern of errors. Therefore, similar to humans, both configuration and component movements are important during chimpanzee facial expression processing.

There has been little research over the past few decades focusing on similarities and differences in the form and function of emotional signals in nonhuman primates, or whether these communication systems are homologous with those of humans. This is, in part, due to the fact that detailed and objective measurement tools to answer such questions have not been systematically developed for nonhuman primate research. Despite this, emotion research in humans has benefited for over 30 years from an objective, anatomically based facial-measurement tool: the Facial Action Coding System (FACS). In collaboration with other researchers, we have now developed a similar system for chimpanzees (ChimpFACS) and, in the process, have made exciting new discoveries regarding chimpanzees' perception and categorization of emotional facial expressions and similarities in the facial anatomy of chimpanzees and humans, and we have identified homologous facial movements in the two species. Investigating similarities and differences in primate emotional communication systems is essential if we are to understand unique evolutionary specializations among different species.

The Chimpanzee Facial Action Coding System (ChimpFACS) is an objective, standardized observational tool for measuring facial movement in chimpanzees based on the well-known human Facial Action Coding System (FACS; P. Ekman & W. V. Friesen, 1978). This tool enables direct structural comparisons of facial expressions between humans and chimpanzees in terms of their common underlying musculature. Here the authors provide data on the first application of the ChimpFACS to validate existing categories of chimpanzee facial expressions using discriminant functions analyses. The ChimpFACS validated most existing expression categories (6 of 9) and, where the predicted group memberships were poor, the authors discuss potential problems with ChimpFACS and/or existing categorizations. The authors also report the prototypical movement configurations associated with these 6 expression categories. For all expressions, unique combinations of muscle movements were identified, and these are illustrated as peak intensity prototypical expression configurations. Finally, the authors suggest a potential homology between these prototypical chimpanzee expressions and human expressions based on structural similarities. These results contribute to our understanding of the evolution of emotional communication by suggesting several structural homologies between the facial expressions of chimpanzees and humans and facilitating future research.

To understand the evolution of emotional communication, comparative research on facial expression similarities between humans and related species is essential. Chimpanzees display a complex, flexible facial expression repertoire with many physical and functional similarities to humans. This paper reviews what is known about these facial expression repertoires, discusses the importance of social organization in understanding the meaning of different expressions, and introduces a new coding system, the ChimpFACS, and describes how it can be used to determine homologies between human and chimpanzee facial expressions. Finally, it reviews previous studies on the categorization of facial expressions by chimpanzees using computerized tasks, and discusses the importance of configural processing for this skill in both humans and chimpanzees. Future directions for understanding the evolution of emotional communication will include studies on the social function of facial expressions in ongoing social interactions, the development of facial expression communication and more studies that examine the perception of these important social signals.

The inversion effect, or impaired recognition of upside-down faces, is used as evidence supporting the configural processing of faces. Human studies report a linear relationship between face-discrimination performance and orientation, such that recognition is more difficult as faces are rotated away from their typical viewpoint. Previous studies on chimpanzees also support a configural bias for processing faces, particularly faces for which subjects have developed expertise. In the present study, we examined the influence of expertise and rotation angle on the visual perception of faces in chimpanzees. Six subjects were presented with unaltered and blurred conspecific faces and houses in five orientation angles. A computerized paradigm was used to further delineate the nature of configural face processing in this species. The data were consistent with those reported in humans: chimpanzees showed a significant linear impairment when discriminating conspecific faces as they rotated away from their upright orientation. No inversion effect was observed for discriminations involving houses. Thus, chimpanzees, like humans, show a face-specific inversion effect that is linearly affected by angle of orientation, suggesting that their visual processing of faces is strongly influenced by the extraction of configural cues and closely resembles the perceptual strategies of humans.