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Plotnik, J., Nelson, P. A., & de Waal, F. B. M. (2003). Visual field information in the face perception of chimpanzees (Pan troglodytes). Ann N Y Acad Sci, 1000, 94–98.
Abstract: Evidence for a visual field advantage (VFA) in the face perception of chimpanzees was investigated using a modification of a free-vision task. Four of six chimpanzee subjects previously trained on a computer joystick match-to-sample paradigm were able to distinguish between images of neutral face chimeras consisting of two left sides (LL) or right sides (RR) of the face. While an individual's ability to make this distinction would be unlikely to determine their suitability for the VFA tests, it was important to establish that distinctive information was available in test images. Data were then recorded on their choice of the LL vs. RR chimera as a match to the true, neutral image; a bias for one of these options would indicate an hemispatial visual field advantage. Results suggest that chimpanzees, unlike humans, do not exhibit a left visual field advantage. These results have important implications for studies on laterality and asymmetry in facial signals and their perception in primates.
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de Waal, F. B. M. (2003). Darwin's legacy and the study of primate visual communication. Ann N Y Acad Sci, 1000, 7–31.
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.
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Parr, L. A., Winslow, J. T., Hopkins, W. D., & de Waal, F. B. (2000). Recognizing facial cues: individual discrimination by chimpanzees (Pan troglodytes) and rhesus monkeys (Macaca mulatta). J Comp Psychol, 114(1), 47–60.
Abstract: Faces are one of the most salient classes of stimuli involved in social communication. Three experiments compared face-recognition abilities in chimpanzees (Pan troglodytes) and rhesus monkeys (Macaca mulatta). In the face-matching task, the chimpanzees matched identical photographs of conspecifics' faces on Trial 1, and the rhesus monkeys did the same after 4 generalization trials. In the individual-recognition task, the chimpanzees matched 2 different photographs of the same individual after 2 trials, and the rhesus monkeys generalized in fewer than 6 trials. The feature-masking task showed that the eyes were the most important cue for individual recognition. Thus, chimpanzees and rhesus monkeys are able to use facial cues to discriminate unfamiliar conspecifics. Although the rhesus monkeys required many trials to learn the tasks, this is not evidence that faces are not as important social stimuli for them as for the chimpanzees.
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Moehlman, P. D. (1998). Behavioral patterns and communication in feral asses (Equus africanus). Appl. Anim. Behav. Sci., 60(2-3), 125–169.
Abstract: The behavior of feral populations of the African wild ass (Equus africanus) were studied in the Northern Panamint Range of Death Valley National Monument for 20 months from 1970 to 1973 [Moehlman, P.D., 1974. Behavior and ecology of feral asses (Equus asinus). PhD dissertation, University of Wisconsin, Madison, 251 pp.; Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Natl. Geogr. Soc. Res. Reports, 1970: 405-411]. Maintenance behavior is described and behavior sequences that were used in social interactions are quantified by sex and age class. Agonistic, sexual, and greeting behavior patterns are described and analyzed in conjunction with the responses they elicited. Mutual grooming mainly occurred between adult males, and between females and their offspring. Five types of vocalizations were distinguished: brays, grunts, growls, snorts, and whuffles. A second population was studied for 1 month on Ossabaw Island, GA (Moehlman, 1979). This population had more permanent social groups and had a higher rate of mutual grooming and foal social play.
Keywords: Equids; Feral asses; Behavior patterns; Facial expressions; Postures; Locomotion
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Kaminski, J., Call, J., & Tomasello, M. (2004). Body orientation and face orientation: two factors controlling apes' behavior from humans. Anim. Cogn., 7(4), 216–223.
Abstract: A number of animal species have evolved the cognitive ability to detect when they are being watched by other individuals. Precisely what kind of information they use to make this determination is unknown. There is particular controversy in the case of the great apes because different studies report conflicting results. In experiment 1, we presented chimpanzees, orangutans, and bonobos with a situation in which they had to request food from a human observer who was in one of various attentional states. She either stared at the ape, faced the ape with her eyes closed, sat with her back towards the ape, or left the room. In experiment 2, we systematically crossed the observer's body and face orientation so that the observer could have her body and/or face oriented either towards or away from the subject. Results indicated that apes produced more behaviors when they were being watched. They did this not only on the basis of whether they could see the experimenter as a whole, but they were sensitive to her body and face orientation separately. These results suggest that body and face orientation encode two different types of information. Whereas face orientation encodes the observer's perceptual access, body orientation encodes the observer's disposition to transfer food. In contrast to the results on body and face orientation, only two of the tested subjects responded to the state of the observer's eyes.
Keywords: Animals; *Attention; *Behavior, Animal; Cognition; *Concept Formation; Face; Facial Expression; Female; Fixation, Ocular; Hominidae/*psychology; Humans; Male; *Nonverbal Communication; *Orientation; Pan paniscus/psychology; Pan troglodytes/psychology; Pongo pygmaeus/psychology; *Posture; Social Perception; Species Specificity
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Izumi, A., & Kojima, S. (2004). Matching vocalizations to vocalizing faces in a chimpanzee (Pan troglodytes). Anim. Cogn., 7(3), 179–184.
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.
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Parr, L. A. (2004). Perceptual biases for multimodal cues in chimpanzee (Pan troglodytes) affect recognition. Anim. Cogn., 7(3), 171–178.
Abstract: The ability of organisms to discriminate social signals, such as affective displays, using different sensory modalities is important for social communication. However, a major problem for understanding the evolution and integration of multimodal signals is determining how humans and animals attend to different sensory modalities, and these different modalities contribute to the perception and categorization of social signals. Using a matching-to-sample procedure, chimpanzees discriminated videos of conspecifics' facial expressions that contained only auditory or only visual cues by selecting one of two facial expression photographs that matched the expression category represented by the sample. Other videos were edited to contain incongruent sensory cues, i.e., visual features of one expression but auditory features of another. In these cases, subjects were free to select the expression that matched either the auditory or visual modality, whichever was more salient for that expression type. Results showed that chimpanzees were able to discriminate facial expressions using only auditory or visual cues, and when these modalities were mixed. However, in these latter trials, depending on the expression category, clear preferences for either the visual or auditory modality emerged. Pant-hoots and play faces were discriminated preferentially using the auditory modality, while screams were discriminated preferentially using the visual modality. Therefore, depending on the type of expressive display, the auditory and visual modalities were differentially salient in ways that appear consistent with the ethological importance of that display's social function.
Keywords: Acoustic Stimulation; *Animal Communication; Animals; Auditory Perception/physiology; Cues; Discrimination Learning/*physiology; Facial Expression; Female; Male; Pan troglodytes/*psychology; Perceptual Masking/*physiology; Photic Stimulation; Recognition (Psychology)/*physiology; Visual Perception/physiology; *Vocalization, Animal
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Gothard, K. M., Erickson, C. A., & Amaral, D. G. (2004). How do rhesus monkeys ( Macaca mulatta) scan faces in a visual paired comparison task? Anim. Cogn., 7(1), 25–36.
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.
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Gácsi, M., Miklósi, Á., Varga, O., Topál, J., & Csányi, V. (2004). Are readers of our face readers of our minds? Dogs (Canis familiaris) show situation-dependent recognition of human's attention. Anim. Cogn., 7(3), 144–153.
Abstract: The ability of animals to use behavioral/facial cues in detection of human attention has been widely investigated. In this test series we studied the ability of dogs to recognize human attention in different experimental situations (ball-fetching game, fetching objects on command, begging from humans). The attentional state of the humans was varied along two variables: (1) facing versus not facing the dog; (2) visible versus non-visible eyes. In the first set of experiments (fetching) the owners were told to take up different body positions (facing or not facing the dog) and to either cover or not cover their eyes with a blindfold. In the second set of experiments (begging) dogs had to choose between two eating humans based on either the visibility of the eyes or direction of the face. Our results show that the efficiency of dogs to discriminate between “attentive” and “inattentive” humans depended on the context of the test, but they could rely on the orientation of the body, the orientation of the head and the visibility of the eyes. With the exception of the fetching-game situation, they brought the object to the front of the human (even if he/she turned his/her back towards the dog), and preferentially begged from the facing (or seeing) human. There were also indications that dogs were sensitive to the visibility of the eyes because they showed increased hesitative behavior when approaching a blindfolded owner, and they also preferred to beg from the person with visible eyes. We conclude that dogs are able to rely on the same set of human facial cues for detection of attention, which form the behavioral basis of understanding attention in humans. Showing the ability of recognizing human attention across different situations dogs proved to be more flexible than chimpanzees investigated in similar circumstances.
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Hopkins, W. D., & Washburn, D. A. (2002). Matching visual stimuli on the basis of global and local features by chimpanzees (Pan troglodytes) and rhesus monkeys (Macaca mulatta). Anim. Cogn., 5(1), 27–31.
Abstract: This study was designed to examine whether chimpanzees and monkeys exhibit a global-to-local precedence in the processing of hierarchically organized compound stimuli, as has been reported for humans. Subjects were tested using a sequential matching-to-sample paradigm using stimuli that differed on the basis of their global configuration or local elements, or on both perceptual attributes. Although both species were able to discriminate stimuli on the basis of their global configuration or local elements, the chimpanzees exhibited a global-to-local processing strategy, whereas the rhesus monkeys exhibited a local-to-global processing strategy. The results suggest that perceptual and attentional mechanisms underlying information-processing strategies may account for differences in learning by primates.
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