Abstract: After Charles Darwin's The Expression of the Emotions in Man and Animals, published in 1872, we had to wait 60 years before the theme of animal expressions was picked up by another astute observer. In 1935, Nadezhda Ladygina-Kohts published a detailed comparison of the expressive behavior of a juvenile chimpanzee and of her own child. After Kohts, we had to wait until the 1960s for modern ethological analyses of primate facial and gestural communication. Again, the focus was on the chimpanzee, but ethograms on other primates appeared as well. Our understanding of the range of expressions in other primates is at present far more advanced than that in Darwin's time. A strong social component has been added: instead of focusing on the expressions per se, they are now often classified according to the social situations in which they typically occur. Initially, quantitative analyses were sequential (i.e., concerned with temporal associations between behavior patterns), and they avoided the language of emotions. I will discuss some of this early work, including my own on the communicative repertoire of the bonobo, a close relative of the chimpanzee (and ourselves). I will provide concrete examples to make the point that there is a much richer matrix of contexts possible than the common behavioral categories of aggression, sex, fear, play, and so on. Primate signaling is a form of negotiation, and previous classifications have ignored the specifics of what animals try to achieve with their exchanges. There is also increasing evidence for signal conventionalization in primates, especially the apes, in both captivity and the field. This process results in group-specific or “cultural” communication patterns.

Abstract: Right hemispheric dominance in unconscious emotional processing has been suggested, but remains controversial. This issue was investigated using the subliminal affective priming paradigm combined with unilateral visual presentation in 40 normal subjects. In either left or right visual fields, angry facial expressions, happy facial expressions, or plain gray images were briefly presented as negative, positive, and control primes, followed by a mosaic mask. Then nonsense target ideographs were presented, and the subjects evaluated their partiality toward the targets. When the stimuli were presented in the left, but not the right, visual fields, the negative primes reduced the subjects' liking for the targets, relative to the case of the positive or control primes. These results provided behavioral evidence supporting the hypothesis that the right hemisphere is dominant for unconscious negative emotional processing.

Abstract: We define temperament as an individual's set of characteristic behavioral responses to novel or challenging stimuli. This study adapted a temperament scale used with rhesus macaques by Schneider and colleagues [American Journal of Primatology 25:137-155, 1991] for use with male pigtailed macaque (Macaca nemestrina, n = 7), longtailed macaque (M. fascicularis, n = 3), and baboon infants (Papio cynocephalus anubis, n = 4). Subjects were evaluated twice weekly for the first 5 months of age during routine removal from their cages for weighing. Behavioral measures were based on the subject's interactions with a familiar human caretaker and included predominant state before capture, response to capture, contact latency, resistance to tester's hold, degree of clinging, attention to environment, defecation/urination, consolability, facial expression, vocalizations, and irritability. Species differences indicated that baboons were more active than macaques in establishing or terminating contact with the tester. Temperament scores decreased over time for the variables Response to Capture and Contact Latency, indicating that as they grew older, subjects became less reactive and more bold in their interactions with the tester. Temperament scores changed slowly with age, with greater change occurring at younger ages. The retention of variability in reactivity between and within species may be advantageous for primates, reflecting the flexibility necessary to survive in a changing environment.

Abstract: Auditory-visual processing of species-specific vocalizations was investigated in a female chimpanzee named Pan. The basic task was auditory-visual matching-to-sample, where Pan was required to choose the vocalizer from two test movies in response to a chimpanzee's vocalization. In experiment 1, movies of vocalizing and silent faces were paired as the test movies. The results revealed that Pan recognized the status of other chimpanzees whether they vocalized or not. In experiment 2, two different types of vocalizing faces of an identical individual were prepared as the test movies. Pan recognized the correspondence between vocalization types and faces. These results suggested that chimpanzees possess crossmodal representations of their vocalizations, as do humans. Together with the ability of vocal individual recognition, this ability might reflect chimpanzees' profound understanding of the status of other individuals.

Abstract: When novel and familiar faces are viewed simultaneously, humans and monkeys show a preference for looking at the novel face. The facial features attended to in familiar and novel faces, were determined by analyzing the visual exploration patterns, or scanpaths, of four monkeys performing a visual paired comparison task. In this task, the viewer was first familiarized with an image and then it was presented simultaneously with a novel and the familiar image. A looking preference for the novel image indicated that the viewer recognized the familiar image and hence differentiates between the familiar and the novel images. Scanpaths and relative looking preference were compared for four types of images: (1) familiar and novel objects, (2) familiar and novel monkey faces with neutral expressions, (3) familiar and novel inverted monkey faces, and (4) faces from the same monkey with different facial expressions. Looking time was significantly longer for the novel face, whether it was neutral, expressing an emotion, or inverted. Monkeys did not show a preference, or an aversion, for looking at aggressive or affiliative facial expressions. The analysis of scanpaths indicated that the eyes were the most explored facial feature in all faces. When faces expressed emotions such as a fear grimace, then monkeys scanned features of the face, which contributed to the uniqueness of the expression. Inverted facial images were scanned similarly to upright images. Precise measurement of eye movements during the visual paired comparison task, allowed a novel and more quantitative assessment of the perceptual processes involved the spontaneous visual exploration of faces and facial expressions. These studies indicate that non-human primates carry out the visual analysis of complex images such as faces in a characteristic and quantifiable manner.