Tomasello, M., Hare, B., & Agnetta, B. (1999). Chimpanzees, Pan troglodytes, follow gaze direction geometrically. Anim. Behav., 58(4), 769–777.
Abstract: Two experiments on chimpanzee gaze following are reported. In the first, chimpanzee subjects watched as a human experimenter looked around various types of barriers. The subjects looked around each of the barriers more when the human had done so than in a control condition (in which the human looked in another direction). In the second experiment, chimpanzees watched as a human looked towards the back of their cage. As they turned to follow the human's gaze a distractor object was presented. The chimpanzees looked at the distractor while still following the human's gaze to the back of the cage. These two experiments effectively disconfirm the low-level model of chimpanzee gaze following in which it is claimed that upon seeing another animate being's gaze direction chimpanzees simply turn in that direction and look around for something interesting. Rather, they support the hypothesis that chimpanzees follow the gaze direction of other animate beings geometrically to specific locations, in much the same way as human infants. The degree to which chimpanzees have a mentalistic interpretation of the gaze and/or visual experience of others is still an open question.
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Hare, B., Call, J., & Tomasello, M. (2001). Do chimpanzees know what conspecifics know? Anim. Behav., 61(1), 139–151.
Abstract: We conducted three experiments on social problem solving by chimpanzees, Pan troglodytes. In each experiment a subordinate and a dominant individual competed for food, which was placed in various ways on the subordinate's side of two opaque barriers. In some conditions dominants had not seen the food hidden, or food they had seen hidden was moved elsewhere when they were not watching (whereas in control conditions they saw the food being hidden or moved). At the same time, subordinates always saw the entire baiting procedure and could monitor the visual access of their dominant competitor as well. If subordinates were sensitive to what dominants did or did not see during baiting, they should have preferentially approached and retrieved the food that dominants had not seen hidden or moved. This is what they did in experiment 1 when dominants were either uninformed or misinformed about the food's location. In experiment 2 subordinates recognized, and adjusted their behaviour accordingly, when the dominant individual who witnessed the hiding was replaced with another dominant individual who had not witnessed it, thus demonstrating their ability to keep track of precisely who has witnessed what. In experiment 3 subordinates did not choose consistently between two pieces of hidden food, one of which dominants had seen hidden and one of which they had not seen hidden. However, their failure in this experiment was likely to be due to the changed nature of the competition under these circumstances and not to a failure of social-cognitive skills. These findings suggest that at least in some situations (i.e. competition with conspecifics) chimpanzees know what conspecifics have and have not seen (do and do not know), and that they use this information to devise effective social-cognitive strategies. Copyright 2001 The Association for the Study of Animal Behaviour.
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Byrnl, R. W., & Tomasello, M. (1995). Do rats ape? Anim. Behav., 50(5), 1417–1420.
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Tomasello, M., Call, J., & Hare, B. (1998). Five primate species follow the visual gaze of conspecifics. Anim. Behav., 55(4), 1063–1069.
Abstract: Individuals from five primate species were tested experimentally for their ability to follow the visual gaze of conspecifics to an outside object. Subjects were from captive social groups of chimpanzees,Pan troglodytes, sooty mangabeys,Cercocebus atys torquatus, rhesus macaques,Macaca mulatta, stumptail macaques,M. arctoides, and pigtail macaques,M. nemestrina. Experimental trials consisted of an experimenter inducing one individual to look at food being displayed, and then observing the reaction of another individual (the subject) that was looking at that individual (not the food). Control trials consisted of an experimenter displaying the food in an identical manner when the subject was alone. Individuals from all species reliably followed the gaze of conspecifics, looking to the food about 80% of the time in experimental trials, compared with about 20% of the time in control trials. Results are discussed in terms of both the proximate mechanisms that might be involved and the adaptive functions that might be served by gaze-following.
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Heyes, C. M. (1995). Imitation and flattery: a reply to Byrne & Tomasello. Anim. Behav., 50(5), 1421–1424.
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Tomasello, M., Hare, B., & Fogleman, T. (2001). The ontogeny of gaze following in chimpanzees, Pan troglodytes, and rhesus macaques, Macaca mulatta. Anim. Behav., 61(2), 335–343.
Abstract: Primates follow the gaze direction of conspecifics to outside objects. We followed the ontogeny of this social-cognitive skill for two species: rhesus macaques and chimpanzees. In the first two experiments, using both a cross-sectional and a longitudinal design, we exposed individuals of different ages to a human looking in a specified direction. Rhesus infants first began reliably to follow the direction of this gaze at the end of the early infancy period, at about 5.5 months of age. Chimpanzees did not reliably follow human gaze until 3-4 years; this corresponds to the latter part of the late infancy period for this species. In the third experiment we exposed individuals of the same two species to a human repeatedly looking to the same location (with no special object at that location) to see if subjects would learn to ignore the looks. Only adults of the two species diminished their gaze-following behaviour over trials. This suggests that in the period between infancy and adulthood individuals of both species come to integrate their gaze-following skills with their more general social-cognitive knowledge about other animate beings and their behaviour, and so become able to deploy their gaze-following skills in a more flexible manner.
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Sovrano, V. A., Rainoldi, C., Bisazza, A., & Vallortigara, G. (1999). Roots of brain specializations: preferential left-eye use during mirror-image inspection in six species of teleost fish. Behav. Brain. Res., 106(1-2), 175–180.
Abstract: It has recently been reported that predator inspection is more likely to occur when a companion (i.e. the mirror image of the test animal) is visible on the left rather than on the right side of mosquitofish Gambusia holbrooki. This very unexpected outcome could be consistent with the hypothesis of a preferential use of the right eye during sustained fixation of a predator as well as of a preferential use of the left eye during fixation of conspecifics. We measured the time spent in monocular viewing during inspection of their own mirror images in females of six species of fish, belonging to different families--G. holbrooki, Xenotoca eiseni, Phoxinus phoxinus, Pterophyllum scalare, Xenopoecilus sarasinorum, and Trichogaster trichopterus. Results revealed a consistent left-eye preference during sustained fixation in all of the five species. Males of G. holbrooki, which do not normally show any social behaviour, did not exhibit any eye preferences during mirror-image inspection. We found, however, that they could be induced to manifest a left-eye preference, likewise females, if tested soon after capture, when some affiliative tendencies can be observed. These findings add to current evidence in a variety of vertebrate species for preferential involvement of structures located in the right side of the brain in response to the viewing of conspecifics.
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Weeks, J. W., Crowell-Davis, S. L., & Heusner, G. (2002). Preliminary study of the development of the Flehmen response in Equus caballus. Appl. Anim. Behav. Sci., 78(2-4), 329–335.
Abstract: The flehmen response is commonly seen in most ungulates as well as in several other species (e.g. felids). The behavior is most often thought to be part of the sexual behavioral repertoire of males. One reigning hypothesis suggests that this behavior allows the male to determine the estrous state of a female through the chemosensory functions of the vomeronasal organ. However, females and young of both sexes also exhibit this behavior. Horse foals most frequently show the flehmen response during their first month of life with colts showing the behavior more often than fillies. This study tested the flehmen response on male and female foals throughout their pre-pubertal period. Foals were separately presented estrous and non-estrous urine weekly during the first month of life and then monthly until they were approximately 7 months of age. No significant differences were found between male and female foals for the following variables: latency to flehmen, duration of flehmen, frequency of flehmen and sniffs.
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Hall, C. (2007). The impact of visual perception on equine learning. Behav. Process., 76, 29–33.
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Ninomiya, S. (2007). Social leaning and stereotypy in horses. Behav. Process., 76, 22–23.
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