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Horner, V., Whiten, A., Flynn, E., & de Waal, F. B. M. (2006). Faithful replication of foraging techniques along cultural transmission chains by chimpanzees and children. Proc. Natl. Acad. Sci. U.S.A., 103(37), 13878–13883.
Abstract: Observational studies of wild chimpanzees (Pan troglodytes) have revealed population-specific differences in behavior, thought to represent cultural variation. Field studies have also reported behaviors indicative of cultural learning, such as close observation of adult skills by infants, and the use of similar foraging techniques within a population over many generations. Although experimental studies have shown that chimpanzees are able to learn complex behaviors by observation, it is unclear how closely these studies simulate the learning environment found in the wild. In the present study we have used a diffusion chain paradigm, whereby a behavior is passed from one individual to the next in a linear sequence in an attempt to simulate intergenerational transmission of a foraging skill. Using a powerful three-group, two-action methodology, we found that alternative methods used to obtain food from a foraging device (“lift door” versus “slide door”) were accurately transmitted along two chains of six and five chimpanzees, respectively, such that the last chimpanzee in the chain used the same method as the original trained model. The fidelity of transmission within each chain is remarkable given that several individuals in the no-model control group were able to discover either method by individual exploration. A comparative study with human children revealed similar results. This study is the first to experimentally demonstrate the linear transmission of alternative foraging techniques by non-human primates. Our results show that chimpanzees have a capacity to sustain local traditions across multiple simulated generations.
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Neiworth, J. J., Hassett, J. M., & Sylvester, C. J. (2007). Face processing in humans and new world monkeys: the influence of experiential and ecological factors. Anim. Cogn., 10(2), 125–134.
Abstract: This study tests whether the face-processing system of humans and a nonhuman primate species share characteristics that would allow for early and quick processing of socially salient stimuli: a sensitivity toward conspecific faces, a sensitivity toward highly practiced face stimuli, and an ability to generalize changes in the face that do not suggest a new identity, such as a face differently oriented. The look rates by adult tamarins and humans toward conspecific and other primate faces were examined to determine if these characteristics are shared. A visual paired comparison (VPC) task presented subjects with either a human face, chimpanzee face, tamarin face, or an object as a sample, and then a pair containing the previous stimulus and a novel stimulus was presented. The stimuli were either presented all in an upright orientation, or all in an inverted orientation. The novel stimulus in the pair was either an orientation change of the same face/object or a new example of the same type of face/object, and the stimuli were shown either in an upright orientation or in an inverted orientation. Preference to novelty scores revealed that humans attended most to novel individual human faces, and this effect decreased significantly if the stimuli were inverted. Tamarins showed preferential looking toward novel orientations of previously seen tamarin faces in the upright orientation, but not in an inverted orientation. Similarly, their preference to look longer at novel tamarin and human faces within the pair was reduced significantly with inverted stimuli. The results confirmed prior findings in humans that novel human faces generate more attention in the upright than in the inverted orientation. The monkeys also attended more to faces of conspecifics, but showed an inversion effect to orientation change in tamarin faces and to identity changes in tamarin and human faces. The results indicate configural processing restricted to particular kinds of primate faces by a New World monkey species, with configural processing influenced by life experience (human faces and tamarin faces) and specialized to process orientation changes specific to conspecific faces.
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Shoshani, J., Kupsky, W. J., & Marchant, G. H. (2006). Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution. Brain Res Bull, 70(2), 124–157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.
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Gallup, G. G. J. (1985). Do minds exist in species other than our own? Neurosci Biobehav Rev, 9(4), 631–641.
Abstract: An answer to the question of animal awareness depends on evidence, not intuition, anecdote, or debate. This paper examines some of the problems inherent in an analysis of animal awareness, and whether animals might be aware of being aware is offered as a more meaningful distinction. A framework is presented which can be used to make a determination about the extent to which other species have experiences similar to ours based on their ability to make inferences and attributions about mental states in others. The evidence from both humans and animals is consistent with the idea that the capacity to use experience to infer the experience of others is a byproduct of self-awareness.
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Horowitz, A. C. (2003). Do humans ape? Or do apes human? Imitation and intention in humans (Homo sapiens) and other animals. J Comp Psychol, 117(3), 325–336.
Abstract: A. Whiten, D. M. Custance, J.-C. Gomez, P. Teixidor, and K. A. Bard (1996) tested chimpanzees' (Pan troglodytes) and human children's (Homo sapiens) skills at imitation with a 2-action test on an “artificial fruit.” Chimpanzees imitated to a restricted degree; children were more thoroughly imitative. Such results prompted some to assert that the difference in imitation indicates a difference in the subjects' understanding of the intentions of the demonstrator (M. Tomasello, 1996). In this experiment, 37 adult human subjects were tested with the artificial fruit. Far from being perfect imitators, the adults were less imitative than the children. These results cast doubt on the inference from imitative performance to an ability to understand others' intentions. The results also demonstrate how any test of imitation requires a control group and attention to the level of behavioral analysis.
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Collier-Baker, E., Davis, J. M., Nielsen, M., & Suddendorf, T. (2006). Do chimpanzees (Pan troglodytes) understand single invisible displacement? Anim. Cogn., 9(1), 55–61.
Abstract: Previous research suggests that chimpanzees understand single invisible displacement. However, this Piagetian task may be solvable through the use of simple search strategies rather than through mentally representing the past trajectory of an object. Four control conditions were thus administered to two chimpanzees in order to separate associative search strategies from performance based on mental representation. Strategies involving experimenter cue-use, search at the last or first box visited by the displacement device, and search at boxes adjacent to the displacement device were systematically controlled for. Chimpanzees showed no indications of utilizing these simple strategies, suggesting that their capacity to mentally represent single invisible displacements is comparable to that of 18-24-month-old children.
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Uller, C. (2004). Disposition to recognize goals in infant chimpanzees. Anim. Cogn., 7(3), 154–161.
Abstract: Do nonhuman primates attribute goals to others? Traditional studies with chimpanzees provide equivocal evidence for “mind reading” in nonhuman primates. Here we adopt looking time, a methodology commonly used with human infants to test infant chimpanzees. In this experiment, four infant chimpanzees saw computer-generated stimuli that mimicked a goal-directed behavior. The baby chimps performed as well as human infants, namely, they were sensitive to the trajectories of the objects, thus suggesting that chimpanzees may be endowed with a disposition to understand goal-directed behaviors. The theoretical implications of these results are discussed.
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Hostetter, A. B., Cantero, M., & Hopkins, W. D. (2001). Differential use of vocal and gestural communication by chimpanzees (Pan troglodytes) in response to the attentional status of a human (Homo sapiens). J. Comp. Psychol., 115(4), 337–343.
Abstract: This study examined the communicative behavior of 49 captive chimpanzees (Pan troglodytes), particularly their use of vocalizations, manual gestures, and other auditory- or tactile-based behaviors as a means of gaining an inattentive audience's attention. A human (Homo sapiens) experimenter held a banana while oriented either toward or away from the chimpanzee. The chimpanzees' behavior was recorded for 60 s. Chimpanzees emitted vocalizations faster and were more likely to produce vocalizations as their 1st communicative behavior when a human was oriented away from them. Chimpanzees used manual gestures more frequently and faster when the human was oriented toward them. These results replicate the findings of earlier studies on chimpanzee gestural communication and provide new information about the intentional and functional use of their vocalizations.
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Iversen, I. H., & Matsuzawa, T. (2003). Development of interception of moving targets by chimpanzees (Pan troglodytes) in an automated task. Anim. Cogn., 6(3), 169–183.
Abstract: The experiments investigated how two adult captive chimpanzees learned to navigate in an automated interception task. They had to capture a visual target that moved predictably on a touch monitor. The aim of the study was to determine the learning stages that led to an efficient strategy of intercepting the target. The chimpanzees had prior training in moving a finger on a touch monitor and were exposed to the interception task without any explicit training. With a finger the subject could move a small “ball” at any speed on the screen toward a visual target that moved at a fixed speed either back and forth in a linear path or around the edge of the screen in a rectangular pattern. Initial ball and target locations varied from trial to trial. The subjects received a small fruit reinforcement when they hit the target with the ball. The speed of target movement was increased across training stages up to 38 cm/s. Learning progressed from merely chasing the target to intercepting the target by moving the ball to a point on the screen that coincided with arrival of the target at that point. Performance improvement consisted of reduction in redundancy of the movement path and reduction in the time to target interception. Analysis of the finger's movement path showed that the subjects anticipated the target's movement even before it began to move. Thus, the subjects learned to use the target's initial resting location at trial onset as a predictive signal for where the target would later be when it began moving. During probe trials, where the target unpredictably remained stationary throughout the trial, the subjects first moved the ball in anticipation of expected target movement and then corrected the movement to steer the ball to the resting target. Anticipatory ball movement in probe trials with novel ball and target locations (tested for one subject) showed generalized interception beyond the trained ball and target locations. The experiments illustrate in a laboratory setting the development of a highly complex and adaptive motor performance that resembles navigational skills seen in natural settings where predators intercept the path of moving prey.
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Whiten, A., Goodall, J., McGrew, W. C., Nishida, T., Reynolds, V., Sugiyama, Y., et al. (1999). Cultures in chimpanzees. Nature, 399(6737), 682–685.
Abstract: As an increasing number of field studies of chimpanzees (Pan troglodytes) have achieved long-term status across Africa, differences in the behavioural repertoires described have become apparent that suggest there is significant cultural variation. Here we present a systematic synthesis of this information from the seven most long-term studies, which together have accumulated 151 years of chimpanzee observation. This comprehensive analysis reveals patterns of variation that are far more extensive than have previously been documented for any animal species except humans. We find that 39 different behaviour patterns, including tool usage, grooming and courtship behaviours, are customary or habitual in some communities but are absent in others where ecological explanations have been discounted. Among mammalian and avian species, cultural variation has previously been identified only for single behaviour patterns, such as the local dialects of song-birds. The extensive, multiple variations now documented for chimpanzees are thus without parallel. Moreover, the combined repertoire of these behaviour patterns in each chimpanzee community is itself highly distinctive, a phenomenon characteristic of human cultures but previously unrecognised in non-human species.
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