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Lachmann M., & Bergstrom C.T. (1998). Signalling among Relatives II. Beyond the Tower of Babel. Theor. Pop. Biol., 54(2).
Abstract: Models of costly signalling are commonly employed in evolutionary biology in order to explain how honest communication between individuals with conflicting interests can be stable. These models have focused primarily on a single type of honest signalling equilibrium, the separating equilibrium in which any two different signallers send distinct signals, thereby providing signal receivers with complete information. In this paper, we demonstrate that in signalling among relatives (modelled using the Sir Philip Sidney game), there is not one but a large number of possible signalling equilibria, most of which are pooling equilibria in which different types of signallers may share a common signal. We prove that in a general Sir Philip Sidney game, any partition of signallers into equi-signalling classes can have a stable signalling equilibrium if and only if it is a contiguous partition, and provide examples of such partitions. A similar (but slightly stricter) condition is shown to hold when signals are transmitted through a medium with signalling error. These results suggest a solution to a problem faced by previous signalling theory models: when we consider the separating equilibrium, signal cost is independent of the frequency of individuals sending that signal and, consequently, even very rare signaller types can drastically affect signal cost. Here, we show that by allowing these rare signallers to pool with more common signallers, signal cost can be greatly reduced. Copyright 1998 Academic Press.
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Bergstrom, C. T., & Lachmann, M. (1998). Signaling among relatives. III. Talk is cheap. Proc. Natl. Acad. Sci. U.S.A., 95(9), 5100–5105.
Abstract: The Sir Philip Sidney game has been used by numerous authors to show how signal cost can facilitate honest signaling among relatives. Here, we demonstrate that, in this game, honest cost-free signals are possible as well, under very general conditions. Moreover, these cost-free signals are better for all participants than the previously explored alternatives. Recent empirical evidence suggests that begging is energetically inexpensive for nestling birds; this finding led some researchers to question the applicability of the costly signaling framework to nestling begging. Our results show that cost-free or inexpensive signals, as observed empirically, fall within the framework of signaling theory.
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Lindberg, A. C., Kelland, A., & Nicol, C. J. (1999). Effects of observational learning on acquisition of an operant response in horses. Appl. Anim. Behav. Sci., 61(3), 187–199.
Abstract: The effect of observational learning on the acquisition of an operant response was examined in eighteen riding horses and ponies. The test horses were randomly divided into three groups of six and individually exposed to one of three treatments. An additional horse was trained as a demonstrator, to perform the operant response. The observer horses watched either the demonstrator performing the bin-opening response (Group D+B); the demonstrator standing passively (Group D); or the operant bin in the absence of the demonstrator (Group B). Observers had access to and were free to interact with an identical bin during testing. Observers in Groups D+B and D were socially familiar with the demonstrator. Each test horse was tested once a day for 10 days. An ANOVA revealed no significant differences between treatment groups in the number of responses or the time taken to reach the learning criterion. However, there were highly significant differences between breed types, with non-warmbloods performing more bouts of opening the bin and feeding (p=0.02), feeding from the bin sooner (p=0.01) and reaching the criterion for learning sooner than warmbloods (p=0.05). There was also a significant negative linear relationship between horses' ages and time spent investigating the bin, with younger horses performing more investigative behaviour (y=-3.08x+106.86; p=0.02).
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Clarke, J. V., Nicol, C. J., Jones, R., & McGreevy, P. D. (1996). Effects of observational learning on food selection in horses. Appl. Anim. Behav. Sci., 50(2), 177–184.
Abstract: Fourteen riding horses of mixed age and breed were randomly allocated to observer and control treatments. An additional horse was pre-trained as a demonstrator to walk the 13.8 m length of the test arena and select one of two food buckets using colour and pattern cues. Observer horses were exposed to correct performances of the task by the trained demonstrator, for 20 trials held over 2 days. Control horses were subjected to the same handling and placement procedures as the observer horses but without exposure to the behaviour of the demonstrator. The third day for all subjects was designated as a test day. Each subject was released individually in a predetermined place in the arena, and the latency to walk the length of the test arena to the food buckets, the latency to feed, the identity of the bucket approached and the identity of the bucket selected were recorded on ten consecutive trials. During tests both food buckets contained food to minimize the possibility of individual trial and error learning. On the first trial the mean latency to approach the goal area was 18 s for observer horses, compared with 119 s for control horses (t = 2.8, d.f. = 12, P < 0.01) and the mean latency to eat was 35 s for observer horses, compared with 181 s for control horses (t = 4.86, d.f. = 11, P < 0.001). However, observer horses were no more likely to choose the demonstrated bucket than control horses on the first trial. Twelve of the 14 horses decreased their latency to approach the goal area during the series of ten trials, but there were no significant changes in the buckets selected.
Keywords: Horse; Observational learning; Food discrimination
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Nicol, C. J. (2006). How animals learn from each other. Appl. Anim. Behav. Sci., 100(1-2), 58–63.
Abstract: This paper explores ways by which animals may learn from one another, using examples drawn mostly from the chicken, an animal for which social learning is likely to be less dangerous than individual learning. In early life, the behaviour of the hen is important in encouraging chicks to peck at edible items. Maternal display not only attracts chicks to profitable food items, but also redirects their attention away from harmful or non-profitable items. Older chicks can enhance their foraging success by observing the behaviour of conspecifics within their own social group. Hens have been trained to perform a novel behaviour (key-pecking for food) by observation of a trained demonstrator bird. Moreover, observers learnt most from watching dominant demonstrators. Thus the ability to learn from others is not `fixed', but depends on the context and the social identity of both the observer and the demonstrator.
Keywords: Social learning; Chickens; Demonstrators; Dominance
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Nicol, C. J. (1995). The social transmission of information and behaviour. Appl. Anim. Behav. Sci., 44(2-4), 79–98.
Abstract: Social influences on established behaviour and on the acquisition of new information and behaviour are reviewed. Distinctions between social facilitation and contagious behaviour are drawn and suggestions for further research on contagious behaviour are made. Socially derived visual, olfactory and auditory cues are considered as important influences on behaviour and subsequent learning. The evidence supporting two potential mechanisms of social learning, i.e. stimulus enhancement followed by individual learning, and imitation, is reviewed in detail. It is argued that the functions of social learning are similarly heterogeneous and include motor skill acquisition, gathering of environmental information, and social conformity. Factors affecting the spread of socially acquired skills, including the social relationship between demonstrator and observer, are highlighted. Lastly, the few studies of social learning that have been conducted with domestic species are reviewed and potential applied goals that could stimulate further research in this area are suggested.
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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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Hare, B., Brown, M., Williamson, C., & Tomasello, M. (2002). The domestication of social cognition in dogs. Science, 298(5598), 1634–1636.
Abstract: Dogs are more skillful than great apes at a number of tasks in which they must read human communicative signals indicating the location of hidden food. In this study, we found that wolves who were raised by humans do not show these same skills, whereas domestic dog puppies only a few weeks old, even those that have had little human contact, do show these skills. These findings suggest that during the process of domestication, dogs have been selected for a set of social-cognitive abilities that enable them to communicate with humans in unique ways.
Keywords: Animals; *Animals, Domestic; *Behavior, Animal; *Cognition; *Cues; *Dogs; Food; Humans; Memory; Pan troglodytes; *Social Behavior; Species Specificity; Vision; Wolves
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Allen, C. (2006). Transitive inference in animals: Reasoning or conditioned associations? In S. Hurley, & M. Nudds (Eds.), Rational Animals? (pp. 175–186). Oxford: Oxford University Press.
Abstract: It is widely accepted that many species of nonhuman animals appear to engage in transitive inference,
producing appropriate responses to novel pairings of non-adjacent members of an ordered series without previous experience of these pairings. Some researchers have taken this capability as providing direct evidence that these animals reason. Others resist such declarations, favouring instead explanations in terms of associative conditioning. Associative accounts of transitive inference have been refined in application to a simple 5-element learning task that is the main paradigm for laboratory investigations of the phenomenon, but it remains unclear how well those accounts generalise to more information-rich environments such as social hierarchies which may contain scores of individuals, and where rapid learning is important. The case of transitive inference is an example of a more general dispute between proponents of associative accounts and advocates of more cognitive accounts of animal behaviour. Examination of the specific details of transitive inference suggests some lessons for the wider debate. |
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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.
Keywords: Predator fixation; Fish; Left-eye preference
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