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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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Zentall, T. R. (2006). Mental time travel in animals: a challenging question. Behav. Process., 72(2), 173–183.
Abstract: Humans have the ability to mentally recreate past events (using episodic memory) and imagine future events (by planning). The best evidence for such mental time travel is personal and thus subjective. For this reason, it is particularly difficult to study such behavior in animals. There is some indirect evidence, however, that animals have both episodic memory and the ability to plan for the future. When unexpectedly asked to do so, animals can report about their recent past experiences (episodic memory) and they also appear to be able to use the anticipation of a future event as the basis for a present action (planning). Thus, the ability to imagine past and future events may not be uniquely human.
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Sterling, E. J., & Povinelli, D. J. (1999). Tool use, aye-ayes, and sensorimotor intelligence. Folia Primatol (Basel), 70(1), 8–16.
Abstract: Humans, chimpanzees, capuchins and aye-ayes all display an unusually high degree of encephalization and diverse omnivorous extractive foraging. It has been suggested that the high degree of encephalization in aye-ayes may be the result of their diverse, omnivorous extractive foraging behaviors. In combination with certain forms of tool use, omnivorous extractive foraging has been hypothesized to be linked to higher levels of sensorimotor intelligence (stages 5 or 6). Although free-ranging aye-ayes have not been observed to use tools directly in the context of their extractive foraging activities, they have recently been reported to use lianas as tools in a manner that independently suggests that they may possess stage 5 or 6 sensorimotor intelligence. Although other primate species which display diverse, omnivorous extractive foraging have been tested for sensorimotor intelligence, aye-ayes have not. We report a test of captive aye-ayes' comprehension of tool use in a situation designed to simulate natural conditions. The results support the view that aye-ayes do not achieve stage 6 comprehension of tool use, but rather may use trial-and-error learning to develop tool-use behaviors. Other theories for aye-aye encephalization are considered.
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Zentall, T. R. (2006). Imitation: definitions, evidence, and mechanisms. Anim. Cogn., 9(4), 335–353.
Abstract: Imitation can be defined as the copying of behavior. To a biologist, interest in imitation is focused on its adaptive value for the survival of the organism, but to a psychologist, the mechanisms responsible for imitation are the most interesting. For psychologists, the most important cases of imitation are those that involve demonstrated behavior that the imitator cannot see when it performs the behavior (e.g., scratching one's head). Such examples of imitation are sometimes referred to as opaque imitation because they are difficult to account for without positing cognitive mechanisms, such as perspective taking, that most animals have not been acknowledged to have. The present review first identifies various forms of social influence and social learning that do not qualify as opaque imitation, including species-typical mechanisms (e.g., mimicry and contagion), motivational mechanisms (e.g., social facilitation, incentive motivation, transfer of fear), attentional mechanisms (e.g., local enhancement, stimulus enhancement), imprinting, following, observational conditioning, and learning how the environment works (affordance learning). It then presents evidence for different forms of opaque imitation in animals, and identifies characteristics of human imitation that have been proposed to distinguish it from animal imitation. Finally, it examines the role played in opaque imitation by demonstrator reinforcement and observer motivation. Although accounts of imitation have been proposed that vary in their level of analysis from neural to cognitive, at present no theory of imitation appears to be adequate to account for the varied results that have been found.
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Urcuioli, P. J., DeMarse, T. B., & Zentall, T. R. (1998). Transfer across delayed discriminations: II. Differences in the substitutability of initial versus test stimuli. J Exp Psychol Anim Behav Process, 24(1), 47–59.
Abstract: In 2 experiments, pigeons were trained on, and then transferred to, delayed simple discriminations in which the initial stimuli signalled reinforcement versus extinction following a retention interval. Experiment 1 showed that discriminative responding on the retention test transferred to novel test stimuli that had appeared in another delayed simple discrimination but not to stimuli having the same reinforcement history off-baseline. By contrast, Experiment 2 showed that performances transferred to novel initial stimuli whether they had been trained on-baseline or off-baseline. These results suggest that the test stimuli in delayed simple discriminations acquire control over responding only in the memory task itself. On the other hand, control by the initial stimuli, if coded as outcome expectancies, does not require such task-specific training.
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Martin, T. I., & Zentall, T. R. (2005). Post-choice information processing by pigeons. Anim. Cogn., 8(4), 273–278.
Abstract: In a conditional discrimination (matching-to-sample), a sample is followed by two comparison stimuli, one of which is correct, depending on the sample. Evidence from previous research suggests that if the stimulus display is maintained following an incorrect response (the so-called penalty-time procedure), acquisition by pigeons is facilitated. The present research tested the hypothesis that the penalty-time procedure allows the pigeons to review and learn from the maintained stimulus display following an incorrect choice. It did so by including a penalty-time group for which, following an incorrect choice, the sample changed to match the incorrect comparison, thus providing the pigeons with post-choice 'misinformation.' This misinformation group acquired the matching task significantly slower than the standard penalty-time group (that had no change in the sample following an error). Furthermore, acquisition of matching by a control group that received no penalty time fell midway between the other two groups, suggesting that the pigeons did not merely take more care in making choices because of the aversiveness of penalty-time. Thus, it appears that in the acquisition of matching-to-sample, when the stimulus display is maintained following an incorrect choice, the pigeons can review or acquire information from the display. This is the first time that such an effect has been reported for a nonhuman species.
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Soproni, K., Miklósi, Á., Topál, J., & Csányi, V. (2002). Dogs' (Canis familiaris) responsiveness to human pointing gestures. J Comp Psychol, 116(1), 27–34.
Abstract: In a series of 3 experiments, dogs (Canis familiaris) were presented with variations of the human pointing gesture: gestures with reversed direction of movement, cross-pointing, and different arm extensions. Dogs performed at above chance level if they could see the hand (and index finger) protruding from the human body contour. If these minimum requirements were not accessible, dogs still could rely on the body position of the signaler. The direction of movement of the pointing arm did not influence the performance. In summary, these observations suggest that dogs are able to rely on relatively novel gestural forms of the human communicative pointing gesture and that they are able to comprehend to some extent the referential nature of human pointing.
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Broom, M. (2002). A unified model of dominance hierarchy formation and maintenance. J. Theor. Biol., 219(1), 63–72.
Abstract: In many different species it is common for animals to spend large portions of their lives in groups. Such groups need to divide available resources amongst the individuals they contain and this is often achieved by means of a dominance hierarchy. Sometimes hierarchies are stable over a long period of time and new individuals slot into pre-determined positions, but there are many situations where this is not so and a hierarchy is formed out of a group of individuals meeting for the first time. There are several different models both of the formation of such dominance hierarchies and of already existing hierarchies. These models often treat the two phases as entirely separate, whereas in reality, if there is a genuine formation phase to the hierarchy, behaviour in this phase will be governed by the rewards available, which in turn depends upon how the hierarchy operates once it has been formed. This paper describes a method of unifying models of these two distinct phases, assuming that the hierarchy formed is stable. In particular a framework is introduced which allows a variety of different models of each of the two parts to be used in conjunction with each other, thus enabling a wide range of situations to be modelled. Some examples are given to show how this works in practice.
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Christie, J. L., Hewson, C. J., Riley, C. B., McNiven, M. A., Dohoo, I. R., & Bate, L. A. (2006). Management factors affecting stereotypies and body condition score in nonracing horses in Prince Edward Island. Can Vet J, 47(2), 136–143.
Abstract: In North America, there are few representative data about the effects of management practices on equine welfare. In a randomized survey of 312 nonracing horses in Prince Edward Island (response rate 68.4%), owners completed a pretested questionnaire and a veterinarian examined each horse. Regression analyses identified factors affecting 2 welfare markers: body condition score (BCS) and stereotypic behavior. Horses' BCSs were high (mean 5.7, on a 9-point scale) and were associated with sex (males had lower BCSs than females; P < 0.001) and examination date (P = 0.052). Prevalences of crib biting, wind sucking, and weaving were 3.8%, 3.8%, and 4.8%, respectively. Age (OR = 1.07, P = 0.08) and hours worked weekly (OR = 1.12, P = 0.03) were risk factors for weaving. Straw bedding (OR = 0.3, P = 0.03), daily hours at pasture (OR = 0.94, P = 0.02), and horse type (drafts and miniatures had a lower risk than light horses; P = 0.12) reduced the risk of horses showing oral stereotypies. Some of these results contradict those of other studies perhaps because of populations concerned.
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Wolf, M., van Doorn, G. S., Leimar, O., & Weissing, F. J. (2007). Life-history trade-offs favour the evolution of animal personalities. Nature, 447(7144), 581–584.
Abstract: In recent years evidence has been accumulating that personalities are not only found in humans but also in a wide range of other animal species. Individuals differ consistently in their behavioural tendencies and the behaviour in one context is correlated with the behaviour in multiple other contexts. From an adaptive perspective, the evolution of animal personalities is still a mystery, because a more flexible structure of behaviour should provide a selective advantage. Accordingly, many researchers view personalities as resulting from constraints imposed by the architecture of behaviour (but see ref. 12). In contrast, we show here that animal personalities can be given an adaptive explanation. Our argument is based on the insight that the trade-off between current and future reproduction often results in polymorphic populations in which some individuals put more emphasis on future fitness returns than others. Life-history theory predicts that such differences in fitness expectations should result in systematic differences in risk-taking behaviour. Individuals with high future expectations (who have much to lose) should be more risk-averse than individuals with low expectations. This applies to all kinds of risky situations, so individuals should consistently differ in their behaviour. By means of an evolutionary model we demonstrate that this basic principle results in the evolution of animal personalities. It simultaneously explains the coexistence of behavioural types, the consistency of behaviour through time and the structure of behavioural correlations across contexts. Moreover, it explains the common finding that explorative behaviour and risk-related traits like boldness and aggressiveness are common characteristics of animal personalities.
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