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Griffin, A. S., & Guez, D. (2014). Innovation and problem solving: A review of common mechanisms. Behav. Process., 109, 121–134.
Abstract: Behavioural innovations have become central to our thinking about how animals adjust to changing environments. It is now well established that animals vary in their ability to innovate, but understanding why remains a challenge. This is because innovations are rare, so studying innovation requires alternative experimental assays that create opportunities for animals to express their ability to invent new behaviours, or use pre-existing ones in new contexts. Problem solving of extractive foraging tasks has been put forward as a suitable experimental assay. We review the rapidly expanding literature on problem solving of extractive foraging tasks in order to better understand to what extent the processes underpinning problem solving, and the factors influencing problem solving, are in line with those predicted, and found, to underpin and influence innovation in the wild. Our aim is to determine whether problem solving can be used as an experimental proxy of innovation. We find that in most respects, problem solving is determined by the same underpinning mechanisms, and is influenced by the same factors, as those predicted to underpin, and to influence, innovation. We conclude that problem solving is a valid experimental assay for studying innovation, propose a conceptual model of problem solving in which motor diversity plays a more central role than has been considered to date, and provide recommendations for future research using problem solving to investigate innovation. This article is part of a Special Issue entitled: Cognition in the wild.
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Galef, B. G. (2013). Imitation and local enhancement: Detrimental effects of consensus definitions on analyses of social learning in animals. Behavioural Processes, 100, 123–130.
Abstract: Development of a widely accepted vocabulary referring to various types of social learning has made important contributions to decades of progress in analyzing the role of socially acquired information in the development of behavioral repertoires. It is argued here that emergence of a consensus vocabulary, while facilitating both communication and research, has also unnecessarily restricted research on social learning. The article has two parts. In the first, I propose that Thorndike, 1898, Thorndike, 1911 definition of imitation as “learning to do an act from seeing it done” has unduly restricted studies of the behavioral processes involved in the propagation of behavior. In part 2, I consider the possibility that success in labeling social learning processes believed to be less cognitively demanding than imitation (e.g. local and stimulus enhancement, social facilitation, etc.) has been mistaken for understanding of those processes, although essentially nothing is known of their stimulus control, development, phylogeny or substrate either behavioral or physiological.
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Taberlet, P., Waits, L. P., & Luikart, G. (1999). Noninvasive genetic sampling: look before you leap. Trends Ecol. Evol, 14(8), 323–327.
Abstract: Noninvasive sampling allows genetic studies of free-ranging animals without the need to capture or even observe them, and thus allows questions to be addressed that cannot be answered using conventional methods. Initially, this sampling strategy promised to exploit fully the existing DNA-based technology for studies in ethology, conservation biology and population genetics. However, recent work now indicates the need for a more cautious approach, which includes quantifying the genotyping error rate. Despite this, many of the difficulties of noninvasive sampling will probably be overcome with improved methodology.
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Klingel, H. (1998). Observations on social organization and behaviour of African and Asiatic Wild Asses (Equus africanus and Equus hemionus). Appl Anim Behav Sci, 60(2), 103–113.
Abstract: 1This paper appears with kind permission of Verlag Paul Parey, Berlin and Hamburg. It was originally published in Z. Tierpsychol., 44, 323-331 (1977), ISSN 0044-3573/ASTM-Coden: ZETIAG.1
Abstract
African and Asiatic Wild Asses (Equus africanus and Equus hemionus) live in unstable groups or herds of variable composition. Some of the adult stallions are territorial in large territories in which they tolerate other ♂♂. The territorial ♂♂ are dominant over all their conspecifics
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Lesimple, C., Reverchon-Billot, L., Galloux, P., Stomp, M., Boichot, L., Coste, C., et al. (2020). Free movement: A key for welfare improvement in sport horses? Appl. Anim. Behav. Sci., 225, 104972.
Abstract: Horses, and in particular sport horses, remain housed predominantly in single stalls. One of the main reported reasons is the fear that they will become agitated and injure themselves and thereby impair their performance if released in paddocks. The hour spent daily at work is also assumed to be sufficient to satisfy the horses' needs for locomotion. Growing scientific evidence shows that single stall housing has negative consequences on horses' welfare and that time for free movement is necessary. Our aim was to assess the feasibility of allowing sport horses used to staying permanently in their stall (except for 1 h riding/day) daily free time in a paddock and to evaluate its potential impact on their welfare using two behavioural reliable indicators (stereotypic behaviours and ear position), as well as selected blood parameters (blood cell count, oxytocin and serotonin concentrations). Two experiments were conducted on the same site. The first experiment evaluated sport horses' habituation to the novel situation of daily sessions in a paddock, and recorded welfare indicators in their stall before and during the experiment, comparing horses that were taken outdoors (experimental) and those that stayed in their stall (control). The second experiment evaluated the impact of this daily free time in a paddock on horses' welfare and its durability, focusing on positive indicators. Behavioural observations in paddocks showed that even horses that had never experienced free movement outside their stall habituated rapidly to this situation. The presence of hay in the paddock, may have speeded up habituation. Their restricted living conditions were associated with abnormalities in blood cell count that were not overcome during the time of daily paddock sessions but behavioural indicators showed that their welfare improved. In the second study, the experimental horses' welfare improved during the paddock release period, in particular their stereotypic behaviours decreased and oxytocin levels increased. No effects on serotonin concentrations could be evidenced. These effects were directly associated with being in paddock, as the indicators returned rapidly to their previous levels indicating compromised welfare when the paddock release sessions stopped. In conclusion, it can be recommended to release sport horses for free movement in paddocks as welfare is improved and subjective assessment by caretakers indicated minimal risks.
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McVey, A., Wilkinson, A., & Mills, D. S. (2018). Social learning in horses: the effect of using a group leader demonstrator on the performance of familiar conspecifics in a detour task. Applied Animal Behaviour Science, .
Abstract: Learning through the observation of others allows the transfer of information without the costs incurred during individual trial and error learning. Horses (Equus caballus) are a highly social species, which might be expected to be capable of learning from others, but experimental findings are inconsistent, and potentially confounded by social facilitation effects not related directly to the learning of the task. We refined the methods used in previous equine social learning studies, to examine and distinguish specific social influences on learning of a task: we used predefined group leaders rather than agonistically dominant individuals to demonstrate a detour task to familiar conspecific observers; in addition we had two control groups: a non-observer (true control) and a group with the demonstrator simply present at the goal (social facilitation control). 44 socially kept horses were allocated to one of the three test conditions and took part in five trials each. Success rate, latency and detour direction were recorded. There was no significant difference between the three groups in the likelihood of them succeeding in the task nor latency to succeed; however there was a significant difference in the route chosen by the groups, with the true control choosing the side with the entrance gate significantly more than either the observer group or social facilitation group. Both of the latter two groups chose to go in the same direction relative to themselves, regardless of which side the gate was. Seven out of nine horses in the observer group chose the same direction as their demonstrator every time. Our results show a significant role of social facilitation on detour behaviour and highlight the importance of including adequate controls for simpler cognitive influences on behaviour before claims can be made about the specific learning of motor actions or goal directed behaviour. Social cues may be important to horses if the task is sufficiently challenging and motivationally important, so future work should consider more demanding, but ecologically relevant situations, in order to maximise the potential revelation of social learning effects which do not depend on simple local or stimulus enhancement effects.
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McLean, A. N., & Christensen, J. W. (2017). The application of learning theory in horse training. Appl. Anim. Behav. Sci., 190, 18–27.
Abstract: The millennia-old practices of horse training markedly predate and thus were isolated from the mid-twentieth century revelation of animal learning processes. From this standpoint, the progress made in the application and understanding of learning theory in horse training is reviewed including a discussion of how learning processes are employed or otherwise under-utilised in training. This review describes the process of habituation and the most commonly applied desensitisation techniques (systematic desensitisation, counter-conditioning, overshadowing, response prevention) and propose two additional techniques (approach conditioning and stimulus blending). The salience of different types of cues, the interaction of operant and classical conditioning and the impact of stress are also discussed. This paper also exposes the inflexibility and occasional inadequacy of the terminology of learning theory when translated from the research laboratory situation to the practical setting in horse training. While learning theory provides a rich toolbox for riders and trainers, the training process is subject to the simultaneous use of multiple learning processes. In addition, learning/behavioural outcomes and trained responses are not just the result of simple stimulus-response based interactions but are further shaped by arousal, affective and attachment states. More research is needed in these areas. For the field of equitation science to progress and to improve clarity and use of learning processes, changes in nomenclature are required. In particular, the use of the terms 'positive' and 'negative' as descriptive labels in both reinforcement and punishment modalities are unacceptably misleading for everyday use. These labels inhibit the understanding and recognition of the learning processes that these terms supposedly represent, yet the learning processes they describe are vital for horse riders, handlers and trainers to understand. We therefore propose that these labels should be re-labelled more appropriately as 'addition' or 'subtraction' reinforcement/punishment. This would enlighten trainers on the correct application of learning theory, and safety and welfare benefits for people and horses would follow. Finally it is also proposed that the term 'conflict theory' be taken up in equitation science to facilitate diagnosis of training-related behaviour disorders and thus enable the emergence of improved training practices. The optimal use of learning theory should be established as a fundamental principle in equestrian education.
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König v. Borstel, U., Visser, E. K., & Hall, C. (2017). Indicators of stress in equitation. Appl. Anim. Behav. Sci., 190, 43–56.
Abstract: Abstract Stress is a generic concept describing the body's reaction to external stimuli, including both physiological and psychological factors. Therefore, by definition, the assessment of psychological stress in the exercising horse encompasses the problem of teasing apart the psychological and physiological factors both of which result in stress responses. The present study reviews the existing literature on various measures of stress taken specifically in the context of equitation science. Particular attention has been paid to short-term effects, and commonly used measurements of short-term stress include heart rate, a number of heart rate variability parameters, blood or saliva cortisol levels, eye temperature, and various behaviour parameters including in particular behaviour patterns presumably indicative of conflict with the rider's/trainer's aids. Inspection of the individual studies' results revealed that disagreement between these different measures of stress is commonplace. For physiological parameters, the largest proportion of agreement (i.e. both parameters simultaneously indicated either higher, insignificant or lower stress compared to a control treatment) was found for heart rate and heart rate variability parameters, while generally limited agreement was found for cortisol. It appears that cortisol levels may not be particularly useful for assessing/assessment of the valence of a situation in the exercising horse as cortisol levels are predominantly linked to activation and exercise levels. Although heart rate variability parameters reflect in theory more closely sympathovagal balance compared to cortisol levels, great care has to be taken regarding the use of appropriate time-frames, appropriate raw data correction methods as well as the use of appropriate equipment. In spite of its wide-spread and apparently successful use, popular equipment may in fact not be accurate enough under field conditions. Eye temperature is another promising parameter for assessment of psychological stress, but the technique is likewise susceptible to application errors. Given the high susceptibility of physiological parameters to errors at various experimental stages, behavioural rather than physiological parameters may in fact provide more accurate measures of valence when conducting experiments in the exercising horse. Behavioural parameters that appear to be particularly practical in assessing stress in ridden horses' behaviour are associated with frequencies of behaviour indicative of conflict. However, while increased frequencies of are a good indicator of stress, the absence of conflict behaviour does not provide proof of the absence of stress due to the possible occurrence of conditions such as Learned Helplessness. In future studies, the above issues should be taken into consideration when designing experiments to assess psychological stress in ridden horses.
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Mejdell, C. M., Buvik, T., Jørgensen, G. H. M., & Bøe, K. E. (2016). Horses can learn to use symbols to communicate their preferences. Appl. Anim. Behav. Sci., 184, 66–73.
Abstract: This paper describes a method in which horses learn to communicate by touching different neutral visual symbols, in order to tell the handler whether they want to have a blanket on or not. Horses were trained for 10-15min per day, following a training program comprising ten steps in a strategic order. Reward based operant conditioning was used to teach horses to approach and touch a board, and to understand the meaning of three different symbols. Heat and cold challenges were performed to help learning and to check level of understanding. At certain stages, a learning criterion of correct responses for 8-14 successive trials had to be achieved before proceeding. After introducing the free choice situation, on average at training day 11, the horse could choose between a “no change” symbol and the symbol for either “blanket on” or “blanket off” depending on whether the horse already wore a blanket or not. A cut off point for performance or non-performance was set to day 14, and 23/23 horses successfully learned the task within this limit. Horses of warm-blood type needed fewer training days to reach criterion than cold-bloods (P<0.05). Horses were then tested under differing weather conditions. Results show that choices made, i.e. the symbol touched, was not random but dependent on weather. Horses chose to stay without a blanket in nice weather, and they chose to have a blanket on when the weather was wet, windy and cold (χ2=36.67, P<0.005). This indicates that horses both had an understanding of the consequence of their choice on own thermal comfort, and that they successfully had learned to communicate their preference by using the symbols. The method represents a novel tool for studying preferences in horses.
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Koistinen, T., Korhonen, H. T., Hämäläinen, E., & Mononen, J. (2016). Blue foxes' (Vulpes lagopus) motivation to gain access and interact with various resources. Appl. Anim. Behav. Sci., 176, 105–111.
Abstract: We analysed the willingness of blue foxes (Vulpes lagopus) to work for and utilise five resources: a platform, wooden block, sand floor, nest box and empty space. Ten juvenile blue fox males were housed singly in apparatus consisting of three cages connected with one-way doors through the walls in between the cages and subjected to work for each of the five resources, one at a time. The resource was placed in one of the outermost cages of the apparatus. Force needed to open the door leading to the resource cage was increased daily by 0.25 or 0.5kg. The number of daily entries, visit durations and interaction with the resource were recorded on workloads of 0, 0.5, 1.5, 2.5, 3.5, 5, 6.5, and 8kg of extra weight. The latency to start interacting with the resource after entering the resource cage was measured on a workload of 3.5kg. The mean number of daily entries in the resource and the other outermost, i.e. control cage varied from 7 to 28 and from 17 to 44, respectively. The increasing workload decreased the number of entries in the resource cage, increased those in the control cage (Linear Mixed Model: F1,638=79.5, P<0.001) and lengthened the visit durations in both cages (F1,642=7.2, P<0.01). The foxes made most (F4,643=9.0, P<0.001) and shortest (F4,641=2.8, P<0.05) visits to the outermost cages when the available resource was either a platform or empty space. The visit durations were longest when the available resource was a nest box. The foxes interacted regularly with the wooden block, but five foxes were not observed interacting with the platform. The nest box was utilised approximately 50% of the time spent in the resource cage, while the platform was utilised only 1-6% and wooden block 2-17% of the time. The mean latency to start interacting with the resource after entering the resource cage was shortest for the sand floor (8s) and longest for the platform (113s, F3,335=26.3, P<0.001). The results show that the foxes re-scheduled their activities on increasing workloads in the apparatus. Based on the number of entries and visit durations, blue foxes valued the wooden block, nest box and sand floor more than the platform or an empty cage. After entering the resource cage, the foxes started interacting fastest with the sand floor, showing high motivation to interact. After entering the resource cage, the foxes make use of the roof of the nest box more urgently than the interior of the nest box. Long bouts in the cage with nest box indicate resting behaviour.
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