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Frank S. A. (1996). Policing and group cohesion when resources vary. Anim. Behav., 52, 1163–1169.
Abstract: The transition from competing individuals to cooperative groups has occurred several times inevolutionary history. The puzzle is why selfish individuals did not subvert cohesive group behaviour bytaking resources without contributing to the group’s overall success. Kin selection and reciprocal altruism are the two standard explanations for group cohesion. But many groups have evolved into
cooperative units when relatedness was low and opportunities were limited for the strategic alliances required for reciprocity. A new theory was recently proposed in which individuals invest some of their resources into repressing competition between group members. Such policing increases the fair distribution of resources in the group and enhances group cohesion. The surprising aspect of this theory
is that low relatedness is more conducive to the spread of policing traits than is high relatedness. Here a new explanation is developed of the biological processes that favour policing. The model is then extended in two ways. First, more realism is added to the theory by accounting for the full range of costs and benefits associated with competitive and cooperative traits within groups. Second, another
surprising result is introduced about cooperative evolution. Small variations in individual vigour or resources can lead to large variations in individual contributions to policing the group. Stronger individuals often invest all of their excess resources into policing, but weaker individuals do not contribute to group cohesion.
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Lovrovich, P., Sighieri, C., & Baragli, P. (2015). Following human-given cues or not? Horses (Equus caballus) get smarter and change strategy in a delayed three choice task. Appl. Anim. Behav. Sci., 166, 80–88.
Abstract: Highlights
�Horses remember the location of food hidden by the experimenter after a delay.
�They understand the communicative meaning of a human positioned close to the target.
�The same horses are capable of changing their decision-making strategy.
�They are able to shift from accuracy inferred from human given cues to speed.
�Horses can use human cues or not depending on time, cost, experience and reward.
Abstract
To date, horses have seemed capable of using human local enhancement cues only when the experimenter remains close to the reward, since they fail to understand the communicative meaning of the human as momentary local enhancement cue (when the human is not present at the moment of the animal's choice). This study was designed to analyse the ability of horses to understand, remember and use human-given cues in a delayed (10 s) three-choice task. Twelve horses (experimental group) had to find a piece of carrot hidden under one of three overturned buckets after seeing the experimenter hide it. The results were then compared with those of a control group (twelve horses) that had to find the carrot using only the sense of smell or random attempts. At the beginning, the experimental horses made more correct choices at the first attempt, although they took more time to find the carrot. Later the same horses were less accurate but found the carrot in less time. This suggests that the value of the proximal momentary local enhancement cues became less critical. It seemed, in fact, that the experimental and control group had aligned their behaviour as the trials proceeded. Despite this similarity, in the second half of the trials, the experimental group tended to first approach the bucket where they had found the carrot in the immediately preceding trial. Our findings indicate that horses are capable of remembering the location of food hidden by the experimenter after a delay, by using the human positioned close to the target as valuable information. The same horses are also capable of changing their decision-making strategy by shifting from the accuracy inferred from human given cues to speed. Therefore, horses are able to decide whether or not to use human given-cues, depending on a speed-accuracy trade-off.
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Rørvang, M. V., Ahrendt, L. P., & Christensen, J. W. (2015). A trained demonstrator has a calming effect on naïve horses when crossing a novel surface. Appl. Anim. Behav. Sci., 171, 117–120.
Abstract: Abstract Habituated horses have been found to have a calming effect on conspecifics in fear-eliciting situations. In practice, experienced horses are often used as companions when young horses are introduced to potentially frightening situations, like loading onto a trailer. However, studies of social transmission of habituation in horses are scarce. This study investigated if demonstration by a habituated demonstrator horse influenced the willingness of young Icelandic horses (n = 22, 3 years old) to cross a novel surface. Observer horses (n = 11) were allowed to observe the similarly aged demonstrator horse being led five times across a novel surface. Immediately afterwards the observer horses were given the opportunity to cross the novel surface themselves to obtain food on the other side. Controls (n = 11) were allowed to observe the demonstrator eating on the opposite side of the novel surface but not the demonstration of crossing the novel surface. All observers and controls succeeded the task, but observers had significantly lower average and maximum heart rate, compared to controls. This result suggests a calming effect of the demonstration, which could be exploited for habituation training of horses in fear-eliciting situations.
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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: 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 10–15 min 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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Rocha, A. D. de L., Menescal-de-Oliveira, L., & da Silva, L. F. S. (2017). Effects of human contact and intra-specific social learning on tonic immobility in guinea pigs, Cavia porcellus. Appl. Anim. Behav. Sci., .
Abstract: Abstract Social learning is the capacity of animals to acquire adaptive information from others. In the case of fear responses, animals can learn fearful or non-fearful responses by observing the behavior of conspecifics. Tonic immobility (TI) is an anti-predatory behavior elicited during intense fear situations. Studies have revealed that regular contact with humans can reduce TI responses in animals. In our study, we evaluated the effect of human contact on the TI responses in guinea pigs. We also evaluated the effect of cohabitation (non-fearful animals with fearful animals) on their TI responses. To achieve this, we measured the TI responses induced by postural inversion and restraint in guinea pigs as a result of different treatments. In our first experiment, we determined the effect of human contact on TI responses by establishing 3 treatment groups: no contact, handled, and habituated. In our second experiment, we addressed the effect of social learning on TI response by testing TI response in habituated, and unhabituated animals that had cohabitated for 10 days. In the first experiment, 10 days of either handling or habituation did not prevent TI in guinea pigs, but habituation did increase latency [F(2,119) = 14.19; p < 0.0001] and handling or habituation decrease duration [F(2,119) = 15.01; p < 0.0001] of the TI behavior in the guinea pigs. In the second experiment, the cohabitation of unhabituated and habituated animals reduced TI duration [F(2,93) = 5.058; p < 0.008]. These data suggest that both forms of human interaction can reduce experimenter fear in guinea pigs. It therefore seems that unhabituated guinea pigs learn not to fear the experimenter by cohabitating with habituated guinea pigs.
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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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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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Amici, F., Widdig, A., Lehmann, J., & Majolo, B. (2019). A meta-analysis of interindividual differences in innovation. Anim. Behav., 155, 257–268.
Abstract: The ability to innovate and the social transmission of innovations have played a central role in human evolution. However, innovation is also crucial for other animals, by allowing them to cope with novel socioecological challenges. Although innovation plays such a central role in animals' lives, we still do not know the conditions required for innovative behaviour to emerge. Here, we focused on interindividual differences in innovation by (1) extensively reviewing existing literature on innovative behaviour in animals and (2) quantitatively testing the different evolutionary hypotheses that have been proposed to explain interindividual variation in innovation propensity during foraging tasks. We ran a series of phylogenetically controlled mixed-effects meta-regression models to determine which hypotheses (if any) are supported by currently available empirical studies. Our analyses show that innovation is more common in individuals that are older and belong to the larger sex, but also in more neophilic and/or explorative individuals. Moreover, these effects change depending on the study setting (i.e. wild versus captive). Our results provide no clear support to the excess of energy or the bad competitor hypotheses and suggest that study setting and interindividual differences in traits related to personality are also important predictors of innovation.
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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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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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