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Beran, M. J., Pate, J. L., Washburn, D. A., & Rumbaugh, D. M. (2004). Sequential responding and planning in chimpanzees (Pan troglodytes) and rhesus macaques (Macaca mulatta). J Exp Psychol Anim Behav Process, 30(3), 203–212.
Abstract: Chimpanzees (Pan troglodytes) and rhesus macaques (Macaca mulatta) selected either Arabic numerals or colored squares on a computer monitor in a learned sequence. On shift trials, the locations of 2 stimuli were interchanged at some point. More errors were made when this interchange occurred for the next 2 stimuli to be selected than when the interchange was for stimuli later in the sequence. On mask trials, all remaining stimuli were occluded after the 1st selection. Performance exceeded chance levels for only 1 selection after these masks were applied. There was no difference in performance for either stimulus type (numerals or colors). The data indicated that the animals planned only the next selection during these computerized tasks as opposed to planning the entire response sequence.
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Boysen, S. T., Bernston, G. G., Hannan, M. B., & Cacioppo, J. T. (1996). Quantity-based interference and symbolic representations in chimpanzees (Pan troglodytes). J Exp Psychol Anim Behav Process, 22(1), 76–86.
Abstract: Five chimpanzees with training in counting and numerical skills selected between 2 arrays of different amounts of candy or 2 Arabic numerals. A reversed reinforcement contingency was in effect, in which the selected array was removed and the subject received the nonselected candies (or the number of candies represented by the nonselected Arabic numeral). Animals were unable to maximize reward by selecting the smaller array when candies were used as array elements. When Arabic numerals were substituted for the candy arrays, all animals showed an immediate shift to a more optimal response strategy of selecting the smaller numeral, thereby receiving the larger reward. Results suggest that a response disposition to the high-incentive candy stimuli introduced a powerful interference effect on performance, which was effectively overridden by the use of symbolic representations.
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Boysen, S. T., & Berntson, G. G. (1995). Responses to quantity: perceptual versus cognitive mechanisms in chimpanzees (Pan troglodytes). J Exp Psychol Anim Behav Process, 21(1), 82–86.
Abstract: Two chimpanzees were trained to select among 2 different amounts of candy (1-6 items). The task was designed so that selection of either array by the active (selector) chimpanzee resulted in that array being given to the passive (observer) animal, with the remaining (nonselected) array going to the selector. Neither animal was able to select consistently the smaller array, which would reap the larger reward. Rather, both animals preferentially selected the larger array, thereby receiving the smaller number of reinforcers. When Arabic numerals were substituted for the food arrays, however, the selector animal evidenced more optimal performance, immediately selecting the smaller numeral and thus receiving the larger reward. These findings suggest that a basic predisposition to respond to the perceptual-motivational features of incentive stimuli can interfere with task performance and that this interference can be overridden when abstract symbols serve as choice stimuli.
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Rumbaugh, D. M., Savage-Rumbaugh, S., & Hegel, M. T. (1987). Summation in the chimpanzee (Pan troglodytes). J Exp Psychol Anim Behav Process, 13(2), 107–115.
Abstract: In this research, we asked whether 2 chimpanzee (Pan troglodytes) subjects could reliably sum across pairs of quantities to select the greater total. Subjects were allowed to choose between two trays of chocolates. Each tray contained two food wells. To select the tray containing the greater number of chocolates, it was necessary to sum the contents of the food wells on each tray. In experiments where food wells contained from zero to four chocolates, the chimpanzees chose the greater value of the summed wells on more than 90% of the trials. In the final experiment, the maximum number of chocolates assigned to a food well was increased to five. Choice of the tray containing the greater sum still remained above 90%. In all experiments, subjects reliably chose the greater sum, even though on many trials a food well on the “incorrect” tray held more chocolates than either single well on the “correct” tray. It was concluded that without any known ability to count, these chimpanzees used some process of summation to combine spatially separated quantities. Speculation regarding the basis for summation includes consideration of perceptual fusion of pairs of quantities and subitization.
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Matsuzawa, T. (1985). Use of numbers by a chimpanzee. Nature, 315(6014), 57–59.
Abstract: Recent studies have examined linguistic abilities in apes. However, although human mathematical abilities seem to be derived from the same foundation as those in language, we have little evidence for mathematical abilities in apes (but for exceptions see refs 7-10). In the present study, a 5-yr-old female chimpanzee (Pan troglodytes), 'Ai', was trained to use Arabic numerals to name the number of items in a display. Ai mastered numerical naming from one to six and was able to name the number, colour and object of 300 types of samples. Although no particular sequence of describing samples was required, the chimpanzee favoured two sequences (colour/object/number and object/colour/number). The present study demonstrates that the chimpanzee was able to describe the three attributes of the sample items and spontaneously organized the 'word order'.
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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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Poti, P. (2005). Chimpanzees' constructional praxis (Pan paniscus, P. troglodytes). Primates, 46(2), 103–113.
Abstract: This study investigated chimpanzees' spontaneous spatial constructions with objects and especially their ability to repeat inter-object spatial relations, which is basic to understanding spatial relations at a higher level than perception or recognition. Subjects were six chimpanzees-four chimpanzees and two bonobos-aged 6-21 years, all raised in a human environment from an early age. Only minor species differences, but considerable individual differences were found. The effect of different object samples was assessed through a comparison with a previous study. A common overall chimpanzee pattern was also found. Chimpanzees repeated different types of inter-object spatial relations such as insertion (I), or vertical (V), or next-to (H) relations. However chimpanzees repeated I or V relations with more advanced procedures than when repeating H relations. Moreover, chimpanzees never repeated combined HV relations. Compared with children, chimpanzees showed a specific difficulty in repeating H relations. Repeating H relations is crucial for representing and understanding multiple reciprocal spatial relations between detached elements and for coordinating independent positions in space. Therefore, the chimpanzees' difficulty indicates a fundamental difference in constructive space in comparison to humans. The findings are discussed in relation to issues of spatial cognition and tool use.
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Palagi, E., Cordoni, G., & Borgognini Tarli, S. (2006). Possible roles of consolation in captive chimpanzees (Pan troglodytes). Am J Phys Anthropol, 129(1), 105–111.
Abstract: Empathy is a necessary prerequisite for the occurrence of consolation. The term “consolation” contains a hypothesis about function, which is distress alleviation. The present study aims to confirm the occurrence of consolation in captive chimpanzees via the post-conflict/matched-control method (PC-MC) and to suggest its possible roles. We collected 273 PC-MC pairs in the group of Pan troglodytes housed in the ZooParc de Beauval (France). We confirmed the presence of consolatory contacts (mean level of consolation, 49.5% +/- 22.3% SEM) in the colony. Consolation rates were significantly higher than reconciliation levels (mean level of reconciliation, 28.9% +/- 16.8% SEM). The level of consolation was greater in the absence of reconciliation than in the presence of it, suggesting that consolation might be an alternative behavior. As friendship and relatedness did not influence the occurrence of consolation, they did not seem to be the best prerequisites for this behavioral mechanism, at least in this chimpanzee colony. Affinitive contacts with third parties were significantly more frequent when the victim called attention to itself during severe aggressions by screaming. These high-pitched sounds seem to be useful in eliciting aid from conspecifics, as occurs in young humans. The occurrence of consolation reduced the likelihood of further attacks among group-members. From this perspective, both victims and consolers most likely gain potential advantages by interacting with each other when aggression is particularly severe, reconciliation is not immediate, and consequently social stress reaches high levels.
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Koski, S. E., Koops, K., & Sterck, E. H. M. (2007). Reconciliation, relationship quality, and postconflict anxiety: testing the integrated hypothesis in captive chimpanzees. Am. J. Primatol., 69(2), 158–172.
Abstract: Reconciliation is a conflict resolution mechanism that is common to many gregarious species with individualized societies. Reconciliation repairs the damaged relationship between the opponents and decreases postconflict (PC) anxiety. The “integrated hypothesis” links the quality of the opponents' relationship to PC anxiety, since it proposes that conflicts among partners with high relationship quality will yield high levels of PC anxiety, which in turn will lead to an increased likelihood of reconciliation. We tested the integrated hypothesis in captive chimpanzees (Pan troglodytes) in the Arnhem Zoo, The Netherlands. We applied the standard PC/matched control (MC) method. Our results mostly support the integrated hypothesis, in that more valuable and compatible partners (i.e., males and frequent groomers) reconciled more often than less valuable and weakly compatible partners (i.e., females and infrequent groomers). In addition, PC anxiety was higher after conflicts among males than among females. Emotional arousal thus appears to be a mediator facilitating reconciliation. However, in contrast to the predictions derived from the integrated hypothesis, PC anxiety appeared only in aggressees, and not in aggressors, of conflicts. This suggests that while relationship quality determines PC anxiety, it is dependent on the role of the participants in the conflict.
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Kutsukake, N., & Castles, D. L. (2004). Reconciliation and post-conflict third-party affiliation among wild chimpanzees in the Mahale Mountains, Tanzania. Primates, 45(3), 157–165.
Abstract: This study investigated post-conflict (PC) behavior among wild chimpanzees (Pan troglodytes) of the M-group in the Mahale Mountains, Tanzania, and examined what types of behavior characterize the PC situation in this group, and the factors that influence the occurrence of PC affiliation between opponents soon after the end of an aggressive conflict (i.e., reconciliation). We found that the opponents affiliated selectively soon after the end of aggression, suggesting that reconciliation occurred in this group. The mean individual corrected conciliatory tendency (CCT) (Veenema et al. 1994 in Behav Proc 31:29-38) was 14.4%, which is similar to or lower than frequencies observed in studies of captive and wild chimpanzees. The valuable relationship hypothesis predicts that the CCT is higher among individuals who share valuable relationships (e.g., males or affiliative dyads) than among individuals who do not (e.g., females or less-associative dyads). However, the analysis based on data for aggression between unrelated individuals (including one incident between an adult and non-adult) and aggression between unrelated adults, did not uncover this difference. Affiliation by a previously uninvolved individual with the victim (“consolation”) and with the aggressor (“appeasement”) occurred more frequently following aggression than in the control condition. The results are compared with previous studies of captive and wild chimpanzees.
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