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Kitchen, D. M., Cheney, D. L., & Seyfarth, R. M. (2005). Male chacma baboons (Papio hamadryas ursinus) discriminate loud call contests between rivals of different relative ranks. Anim. Cogn., 8(1), 1–6.
Abstract: Males in multi-male groups of chacma baboons (Papio hamadryas ursinus) in Botswana compete for positions in a linear dominance hierarchy. Previous research suggests that males treat different categories of rivals differently; competitive displays between males of similar rank are more frequent and intense than those between disparately ranked males. Here we test whether males also respond differently to male-male interactions in which they are not directly involved, using playbacks of the loud 'wahoo' calls exchanged between competing males in aggressive displays. We played paired sequences of vocal contests between two adjacently ranked and two disparately ranked males to ten subjects, half ranking below the signalers in the call sequences and half above. Subjects who ranked above the two signalers showed stronger responses than lower-ranking subjects. Higher-ranking subjects also responded more strongly to sequences involving disparately ranked, as opposed to adjacently ranked opponents, suggesting that they recognized those individuals' relative ranks. Strong responses to sequences between disparately ranked opponents might have occurred either because such contests typically involve resources of high fitness value (defense of meat, estrous females or infants vulnerable to infanticide) or because they indicate a sudden change in one contestant's condition. In contrast, subjects who ranked lower than the signalers responded equally strongly to both types of sequences. These subjects may have been able to distinguish between the two categories of opponents but did not respond differently to them because they had little to lose or gain by a rank reversal between males that already ranked higher than they did.
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Seyfarth, R. M., & Cheney, D. L. (2002). What are big brains for? Proc. Natl. Acad. Sci. U.S.A., 99(7), 4141–4142.
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Owren, M. J., Dieter, J. A., Seyfarth, R. M., & Cheney, D. L. (1993). Vocalizations of rhesus (Macaca mulatta) and Japanese (M. fuscata) macaques cross-fostered between species show evidence of only limited modification. Dev Psychobiol, 26(7), 389–406.
Abstract: Two rhesus and two Japanese macaque infants were cross-fostered between species in order to study the effects of auditory experience on vocal development. Both the cross-fostered and normally raised control subjects were observed over the first 2 years of life and their vocalizations were tape-recorded. We classified 8053 calls by ear, placed each call in one of six acoustic categories, and calculated the rates at which different call-types were used in different social contexts. Species differences were found in the use of “coo” and “gruff” vocalizations among control subjects. Japanese macaques invariably produced coos almost exclusively. In contrast, rhesus macaques produced a mixture of coos and gruffs and showed considerable interindividual variation in the relative use of one call type or the other. Cross-fostered Japanese macaques adhered to their species-typical behavior, rarely using gruffs. Cross-fostered rhesus subjects also exhibited species-typical behavior in many contexts, but in some situations produced coos and gruffs at rates that were intermediate between those shown by normally raised animals of the two species. This outcome suggests that environmentally mediated modification of vocal behavior may have occurred, but that the resulting changes were quite limited.
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Whiten, A. (2005). The second inheritance system of chimpanzees and humans. Nature, 437(7055), 52–55.
Abstract: Half a century of dedicated field research has brought us from ignorance of our closest relatives to the discovery that chimpanzee communities resemble human cultures in possessing suites of local traditions that uniquely identify them. The collaborative effort required to establish this picture parallels the one set up to sequence the chimpanzee genome, and has revealed a complex social inheritance system that complements the genetic picture we are now developing.
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Horner, V., & Whiten, A. (2005). Causal knowledge and imitation/emulation switching in chimpanzees (Pan troglodytes) and children (Homo sapiens). Anim. Cogn., 8(3), 164–181.
Abstract: This study explored whether the tendency of chimpanzees and children to use emulation or imitation to solve a tool-using task was a response to the availability of causal information. Young wild-born chimpanzees from an African sanctuary and 3- to 4-year-old children observed a human demonstrator use a tool to retrieve a reward from a puzzle-box. The demonstration involved both causally relevant and irrelevant actions, and the box was presented in each of two conditions: opaque and clear. In the opaque condition, causal information about the effect of the tool inside the box was not available, and hence it was impossible to differentiate between the relevant and irrelevant parts of the demonstration. However, in the clear condition causal information was available, and subjects could potentially determine which actions were necessary. When chimpanzees were presented with the opaque box, they reproduced both the relevant and irrelevant actions, thus imitating the overall structure of the task. When the box was presented in the clear condition they instead ignored the irrelevant actions in favour of a more efficient, emulative technique. These results suggest that emulation is the favoured strategy of chimpanzees when sufficient causal information is available. However, if such information is not available, chimpanzees are prone to employ a more comprehensive copy of an observed action. In contrast to the chimpanzees, children employed imitation to solve the task in both conditions, at the expense of efficiency. We suggest that the difference in performance of chimpanzees and children may be due to a greater susceptibility of children to cultural conventions, perhaps combined with a differential focus on the results, actions and goals of the demonstrator.
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Stoinski, T. S., Wrate, J. L., Ure, N., & Whiten, A. (2001). Imitative learning by captive western lowland gorillas (Gorilla gorilla gorilla) in a simulated food-processing task. J Comp Psychol, 115(3), 272–281.
Abstract: Although field studies have suggested the existence of cultural transmission of foraging techniques in primates, identification of transmission mechanisms has remained elusive. To test experimentally for evidence of imitation in the current study, we exposed gorillas (Gorilla gorilla gorilla) to an artificial fruit foraging task designed by A. Whiten and D. M. Custance (1996). Gorillas (n=6) watched a human model remove a series of 3 defenses around a fruit. Each of the defenses was removed using 1 of 2 alternative techniques. Subsequent video analysis of gorillas' behavior showed a significant tendency to copy the observed technique on 1 of the individual defenses and the direction of removal on another defense. This is the first statistically reliable evidence of imitation in gorillas. Sequence of defense removal was not replicated. The gorillas' responses were most similar to those of chimpanzees.
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Whiten, A. (1998). Imitation of the sequential structure of actions by chimpanzees (Pan troglodytes). J Comp Psychol, 112(3), 270–281.
Abstract: Imitation was studied experimentally by allowing chimpanzees (Pan troglodytes) to observe alternative patterns of actions for opening a specially designed “artificial fruit.” Like problematic foods primates deal with naturally, with the test fruit several defenses had to be removed to gain access to an edible core, but the sequential order and method of defense removal could be systematically varied. Each subject repeatedly observed 1 of 2 alternative techniques for removing each defense and 1 of 2 alternative sequential patterns of defense removal. Imitation of sequential organization emerged after repeated cycles of demonstration and attempts at opening the fruit. Imitation in chimpanzees may thus have some power to produce cultural convergence, counter to the supposition that individual learning processes corrupt copied actions. Imitation of sequential organization was accompanied by imitation of some aspects of the techniques that made up the sequence.
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Mills, D. S. (1998). Applying learning theory to the management of the horse: the difference between getting it right and getting it wrong. Equine Vet J Suppl, (27), 44–48.
Abstract: Horses constantly modify their behaviour as a result of experience. This involves the creation of an association between events or stimuli. The influence of people on the modification and generation of certain behaviour patterns extends beyond the intentional training of the horse. The impact of any action depends on how it is perceived by the horse, rather than the motive of the handler. Negative and positive reinforcement increase the probability of specific behaviours recurring i.e. strengthen the association between events, whereas punishment reduces the probable recurrence of a behaviour without providing specific information about the desired alternative. In this paper the term 'punishers' is used to refer to the physical aids, such as a whip or crop, which may be used to bring about the process of punishment. However, if their application ceases when a specific behaviour occurs they may negatively reinforce that action. Intended 'punishers' may also be rewarding (e.g. for attention seeking behaviour). Therefore, contingency factors (which define the relationship between stimuli, such as the level of reinforcement), contiguity factors (which describe the proximity of events in space or time) and choice of reinforcing stimuli are critical in determining the rate of learning. The many problems associated with the application of punishment in practice lead to confusion by both horse and handler and, possibly, abuse of the former. Most behaviour problems relate to handling and management of the horse and can be avoided or treated with a proper analysis of the factors influencing the behaviour.
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Hall, C. A., Cassaday, H. J., Vincent, C. J., & Derrington, A. M. (2006). Cone excitation ratios correlate with color discrimination performance in the horse (Equus caballus). J Comp Psychol, 120(4), 438–448.
Abstract: Six horses (Equus caballus) were trained to discriminate color from grays in a counterbalanced sequence in which lightness cues were irrelevant. Subsequently, the pretrained colors were presented in a different sequence. Two sets of novel colors paired with novel grays were also tested. Performance was just as good in these transfer tests. Once the horse had learned to select the chromatic from the achromatic stimulus, regardless of the specific color, they were immediately able to apply this rule to novel stimuli. In terms of the underlying visual mechanisms, the present study showed for the first time that the spectral sensitivity of horse cone photopigments, measured as cone excitation ratios, was correlated with color discrimination performance, measured as accuracy, repeated errors, and latency of approach.
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Christensen, J. W., Rundgren, M., & Olsson, K. (2006). Training methods for horses: habituation to a frightening stimulus. Equine Vet J, 38(5), 439–443.
Abstract: REASONS FOR PERFORMING STUDY: Responses of horses in frightening situations are important for both equine and human safety. Considerable scientific interest has been shown in development of reactivity tests, but little effort has been dedicated to the development of appropriate training methods for reducing fearfulness. OBJECTIVES: To investigate which of 3 different training methods (habituation, desensitisation and counter-conditioning) was most effective in teaching horses to react calmly in a potentially frightening situation. HYPOTHESES: 1) Horses are able to generalise about the test stimulus such that, once familiar with the test stimulus in one situation, it appears less frightening and elicits a reduced response even when the stimulus intensity is increased or the stimulus is presented differently; and 2) alternative methods such as desensitisation and counter-conditioning would be more efficient than a classic habituation approach. METHODS: Twenty-seven naive 2-year-old Danish Warmblood stallions were trained according to 3 different methods, based on classical learning theory: 1) horses (n = 9) were exposed to the full stimulus (a moving, white nylon bag, 1.2 x 0.75 m) in 5 daily training sessions until they met a predefined habituation criterion (habituation); 2) horses (n = 9) were introduced gradually to the stimulus and habituated to each step before the full stimulus was applied (desensitisation); 3) horses (n = 9) were trained to associate the stimulus with a positive reward before being exposed to the full stimulus (counter-conditioning). Each horse received 5 training sessions of 3 min per day. Heart rate and behavioural responses were recorded. RESULTS: Horses trained with the desensitisation method showed fewer flight responses in total and needed fewer training sessions to learn to react calmly to test stimuli. Variations in heart rate persisted even when behavioural responses had ceased. In addition, all horses on the desensitisation method eventually habituated to the test stimulus whereas some horses on the other methods did not. CONCLUSIONS AND POTENTIAL RELEVANCE: Desensitisation appeared to be the most effective training method for horses in frightening situations. Further research is needed in order to investigate the role of positive reinforcement, such as offering food, in the training of horses.
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