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Author Heffner, R.S.; Heffner, H.E.
Title Localization of tones by horses: use of binaural cues and the role of the superior olivary complex Type Journal Article
Year 1986 Publication Behavioral Neuroscience Abbreviated Journal Behav Neurosci
Volume 100 Issue 1 Pages 93-103
Keywords Animals; Auditory Pathways/physiology; Auditory Perception/*physiology; Avoidance Learning/physiology; Brain Mapping; Electroshock; Female; Horses/*physiology; Male; Olivary Nucleus/anatomy & histology/*physiology; Orientation/physiology; Pitch Perception/physiology; Sound Localization/*physiology
Abstract The ability of horses to use binaural time and intensity difference cues to localize sound was assessed in free-field localization tests by using pure tones. The animals were required to discriminate the locus of a single tone pip ranging in frequency from 250 Hz to 25 kHz emitted by loudspeakers located 30 degrees to the left and right of the animals' midline (60 degrees total separation). Three animals were tested with a two-choice procedure; 2 additional animals were tested with a conditioned avoidance procedure. All 5 animals were able to localize 250 Hz, 500 Hz, and 1 kHz but were completely unable to localize 2 kHz and above. Because the frequency of ambiguity for the binaural phase cue delta phi for horses in this test was calculated to be 1.5 kHz, these results indicate that horses can use binaural time differences in the form of delta phi but are unable to use binaural intensity differences. This finding was supported by an unconditioned orientation test involving 4 additional horses, which showed that horses correctly orient to a 500-Hz tone pip but not to an 8-kHz tone pip. Analysis of the superior olivary complex, the brain stem nucleus at which binaural interactions first take place, reveals that the lateral superior olive (LSO) is relatively small in the horse and lacks the laminar arrangement of bipolar cells characteristic of the LSO of most mammals that can use binaural delta I.
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Language English Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0735-7044 ISBN Medium
Area Expedition Conference
Notes PMID:3954885 Approved no
Call Number Equine Behaviour @ team @ Serial 5634
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Author Krueger., K.; Farmer, K.
Title Social learning in Horses: Differs from individual learning only in the learning stimulus and not in the learning mechanisms Type Abstract
Year 2018 Publication 14th Meeting of the Internatinoal Society for Equitation Science Abbreviated Journal 14th Meeting ISES
Volume Issue Pages
Keywords horse; individual learning; learning mechanisms; learning stimuli; social learning
Abstract Equine welfare can be enhanced by applying species specific training. This may incorporate social learning, as horses are highly social and social stimuli are of primary importance. Social learning is comparable to individual learning in its learning mechanisms, differing primarily in the way it is stimulated. Our initial study showed that horses of different breeds (N = 38) follow humans after observing other horses doing so, but only if the observed horse was familiar to and higher ranking than the observer (Fisher's exact test: N = 12, P = 0.003). A second study showed that horses and ponies (N = 25) learned to pull a rope to open a feeding apparatus after observing demonstrations by conspecifics, again, only if the demonstrating horse was older and higher ranking than the observer (Fisher's combination test, N = 3, v2 = 27.71, p = 0.006). Our third approach showed that horses and ponies (N = 24) learned to press a switch to open a feeding apparatus after observing a familiar person (GzLM: N = 24, z = 2.33, P = 0.02). Most recently, we confronted horses and ponies (N = 50) with persons demonstrating different techniques for opening a feeding apparatus. In this study we investigated whether the horses would copy the demonstrators' techniques or apply their own. Here only some horses copied the technique, and most of the successful learners used their mouths irrespective of the demonstrators' postures (Chi Square Test: N = 40, df = 2, &#967;2 = 31.4, p < 0.001). In all the approaches social stimuli elicited learning processes in the test horses, while only a few individuals in the control groups mastered the tasks by individual learning. The following behaviour observed in the initial study may have been facilitated by a social stimuli (social facilitation), and the opening of the feed boxes in the subsequent studies appear to be mostly the result of enhancement (social enhancement). Some horses may have used the social stimuli at first and continued their learning process by individual trial and error. However, the horses were also selective in whom and some in how to copy. This may have been conditioned (socially conditioned) or the result of simple forms of reasoning on the reliability of the particular information provided by demonstrators of certain social ranks or social positions, as high ranking and familiar horses and familiar persons were copied and some imitated exactly.

Lay person message: Traditional riding instructions suggest that horses learn by observing other horses. For example, older, more experienced driving horses are used for initial training of young driving horses. We have shown that horses indeed use learning stimuli provided by other horse, as well as by humans. Horses readily accept stimuli observed in high ranking and familiar horses, and familiar persons. Such stimuli elicit learning processes which are comparable to individual learning. We suggest applying social learning whenever possible, as it is much faster and less stressful than individual learning, where learners experience negative outcomes in trial and error learning.
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Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 6405
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Author McGreevy, P.D.; Oddie, C.; Burton, F.L.; McLean, A.N.
Title The horse–human dyad: Can we align horse training and handling activities with the equid social ethogram? Type Journal Article
Year 2009 Publication The Veterinary Journal Abbreviated Journal Special Issue: Equitation Science
Volume 181 Issue 1 Pages 12-18
Keywords Horse training; Social inter- and intra-specific communication; Learning; Counter–predator behaviour
Abstract This article examines the recently completed equid ethogram and shows how analogues of social interactions between horses may occur in various human–horse interactions. It discusses how some specific horse–horse interactions have a corresponding horse–human interaction – some of which may be directly beneficial for the horse while others may be unusual or even abnormal. It also shows how correspondent behaviours sometimes become inappropriate because of their duration, consistency or context. One analogue is unlikely to hold true for all horse–human contexts, so when applying any model from horse–horse interactions to human–horse interactions, the limitations of the model may eclipse the intended outcome of the intervention. These limitations are especially likely when the horse is being ridden. Such analyses may help to determine the validity of extrapolating intra-specific interactions to the inter-specific setting, as is advocated by some popular horse-training methods, and highlight the subsequent limitations where humans play the role of the ‘alpha mare’ or leader in horse handling and training. This examination provides a constructive framework for further informed debate and empirical investigation of the critical features of successful intra-specific interactions.
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Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 1090-0233 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 5729
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Author Christensen, J.W.; Ahrendt, L.P.; Lintrup, R.; Gaillard, C.; Palme, R.; Malmkvist, J.
Title Does learning performance in horses relate to fearfulness, baseline stress hormone, and social rank? Type Abstract
Year 2012 Publication Applied Animal Behaviour Science Abbreviated Journal App Anim Behav Sci
Volume 140 Issue 1 Pages 44-52
Keywords Horse; Learning; Fearfulness; Stress; Reinforcement; Social rank
Abstract The ability of horses to learn and remember new tasks is fundamentally important for their use by humans. Fearfulness may, however, interfere with learning, because stimuli in the environment can overshadow signals from the rider or handler. In addition, prolonged high levels of stress hormones can affect neurons within the hippocampus; a brain region central to learning and memory. In a series of experiments, we aimed to investigate the link between performance in two learning tests, the baseline level of stress hormones, measured as faecal cortisol metabolites (FCM), fearfulness, and social rank. Twenty-five geldings (2 or 3 years old) pastured in one group were included in the study. The learning tests were performed by professional trainers and included a number of predefined stages during which the horses were gradually trained to perform exercises, using either negative (NR) or positive reinforcement (PR). Each of the learning tests lasted 3 days; 7min/horse/day. The NR test was repeated in a novel environment. Performance, measured as final stage in the training programme, and heart rate (HR) were recorded. Faeces were collected on four separate days where the horses had been undisturbed at pasture for 48h. Social rank was determined through observations of social interactions during feeding. The fear test was a novel object test during which behaviour and HR were recorded. Performance in the NR and PR learning tests did not correlate. In the NR test, there was a significant, negative correlation between performance and HR in the novel environment (rS=-0.66, P<0.001, i.e. nervous horses had reduced performance), whereas there was no such correlation in the home environment (both NR and PR). Behavioural reactions in the fear test correlated significantly with performance in the NR test in the novel environment (e.g. object alertness and final stage: rS=-0.43, P=0.04), suggesting that performance under unfamiliar, stressful conditions may be predicted by behavioural responses in a fear test. There was a negative correlation between social rank and baseline stress hormones (rS=-0.43, P=0.04), i.e. high rank corresponded to low FCM concentrations, whereas neither rank nor FCM correlated with fearfulness or learning performance. We conclude that performance under stressful conditions is affected by activation of the sympathetic nervous system during training and related to behavioural responses in a standardised fear test. Learning performance in the home environment, however, appears unrelated to fearfulness, social rank and baseline FCM levels.
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Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0168-1591 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ S0168-1591(12)00168-2 Serial 5769
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Author Ahrendt, L.P.; Christensen, J.W.; Ladewig, J.
Title The ability of horses to learn an instrumental task through social observation Type Abstract
Year 2012 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.
Volume 139 Issue 1 Pages 105-113
Keywords Horse; Social learning; Social interaction; Instrumental task; Investigative behaviour; Aggression
Abstract The ability of horses to learn through social observation may ease the implementation of new management systems, because the use of automatic feeders etc. by naive horses could be facilitated by observation of experienced horses. However, previous studies found no documentation for observational learning abilities in horses. This study aimed to investigate the ability of horses to learn an instrumental task from a familiar conspecific when social interaction was allowed during the demonstration. Two similar experiments were performed. In the first experiment, Observer horses (n=11) participated in ten successive demonstrations, where a trained Demonstrator opened an operant device by pushing a sliding lid aside with the muzzle in order to obtain a food reward. Immediately after the demonstrations the Observer horses were given the opportunity to operate the device alone. Control horses (n=11) were aware that the device contained food but were presented to the operant device without demonstration of the task. The learning criterion was at least two openings. Accomplishment of and latency to accomplish the learning criterion, and investigative behaviour towards the operant device were recorded. Five Observers and one Control, out of the eleven horses in each treatment group, accomplished the learning criterion. Even though this presents a high odds ratio (OR) in favour of the Observer treatment (OR=7.6), there was no significant difference between the treatment groups (P=0.15). Analysis of investigative behaviour showed, however, that the demonstrations increased the motivation of the Observer horses to investigate the device. Subsequently, a similar experiment was performed in a practical setting with 44 test horses (mixed age, gender and breed). We used the same operant device and the same number and type of demonstrations, although the horses were held on a loose rope to minimise aggression. In this second experiment, six of 23 Observer horses and five of 21 Control horses learned the instrumental task, representing no influence of the demonstration. Thus, this study did not demonstrate an ability of horses to learn an instrumental task through observation.
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Corporate Author Thesis
Publisher (down) Place of Publication Editor
Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0168-1591 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ S0168-1591(12)00087-1 Serial 5773
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Author Rørvang, M.V.; Ahrendt, L.P.; Christensen, J.W.
Title A trained demonstrator has a calming effect on naïve horses when crossing a novel surface Type Journal Article
Year 2015 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.
Volume 171 Issue Pages 117-120
Keywords Fear; Habituation; Social learning; Social transmission; Heart rate
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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Publisher (down) Place of Publication Editor
Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0168-1591 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 5922
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Author Leadbeater, E.
Title What evolves in the evolution of social learning? Type Journal Article
Year 2015 Publication Journal of Zoology Abbreviated Journal J Zool
Volume 295 Issue 1 Pages 4-11
Keywords social learning; associative learning; social information use
Abstract Social learning is fundamental to social life across the animal kingdom, but we still know little about how natural selection has shaped social learning abilities on a proximate level. Sometimes, complex social learning phenomena can be entirely explained by Pavlovian processes that have little to do with the evolution of sociality. This implies that the ability to learn socially could be an exaptation, not an adaptation, to social life but not that social learning abilities have been left untouched by natural selection. I discuss new empirical evidence for associative learning in social information use, explain how natural selection might facilitate the associative learning process and discuss why such studies are changing the way that we think about social learning.
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Publisher (down) Place of Publication Editor
Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 1469-7998 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 6015
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Author Brubaker, L.; Udell, M.A.R.
Title Cognition and learning in horses (Equus caballus): What we know and why we should ask more Type Journal Article
Year 2016 Publication Behavioural Processes Abbreviated Journal
Volume 126 Issue Pages 121-131
Keywords Horse behaviour; Horse welfare; Learning; Social cognition
Abstract Abstract Horses (Equus caballus) have a rich history in their relationship with humans. Across different cultures and eras they have been utilized for work, show, cultural rituals, consumption, therapy, and companionship and continue to serve in many of these roles today. As one of the most commonly trained domestic animals, understanding how horses learn and how their relationship with humans and other horses impacts their ability to learn has implications for horse welfare, training, husbandry and management. Given that unlike dogs and cats, domesticated horses have evolved from prey animals, the horse-human relationship poses interesting and unique scientific questions of theoretical value. There is still much to be learned about the cognition and behaviour of horses from a scientific perspective. This review explores current research within three related areas of horse cognition: human-horse interactions, social learning and independent learning in horses. Research on these topics is summarized and suggestions for future research are provided.
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Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0376-6357 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 6021
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Author Boogert, N.J.; Reader, S.M.; Hoppitt, W.; Laland, K.N.
Title The origin and spread of innovations in starlings Type Journal Article
Year 2008 Publication Animal Behaviour Abbreviated Journal Anim. Behav.
Volume 75 Issue 4 Pages 1509-1518
Keywords diffusion dynamics; dominance; foraging; group; innovation; neophobia; social learning; social network; starling; Sturnus vulgaris
Abstract There are numerous reports of novel learned behaviour patterns in animal populations, yet the factors influencing the invention and spread of these innovations remain poorly understood. Here we investigated to what extent the pattern of spread of innovations in captive groups of starlings, Sturnus vulgaris, could be predicted by knowledge of individual and social group variables, including association patterns, social rank orders, measures of neophobia and asocial learning performance. We presented small groups of starlings with a series of novel extractive foraging tasks and recorded the latency for each bird to contact and solve each task, as well as the orders of contacting and solving. We then explored which variables best predicted the observed diffusion patterns. Object neophobia and social rank measures characterized who was the first of the group to contact the novel foraging tasks, and the subsequent spread of contacting tasks was associated with latency to feed in a novel environment. Asocial learning performance, measured in isolation, predicted who was the first solver of the novel foraging tasks in each group. Association patterns did not predict the spread of solving. Contact latency and solving duration were negatively correlated, consistent with social learning underlying the spread of solving. Our findings indicate that we can improve our understanding of the diffusion dynamics of innovations in animal groups by investigating group-dependent and individual variables in combination. We introduce novel methods for exploring predictors of the origin and spread of behavioural innovations that could be widely applied.
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Publisher (down) Place of Publication Editor
Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0003-3472 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 6036
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Author Daniel, J.C.; Mikulka, P.J.
Title Discrimination learning in the white rhinoceros Type Journal Article
Year 1998 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.
Volume 58 Issue 1–2 Pages 197-202
Keywords Rhinoceros; Learning
Abstract This study examined the ability of two adult white rhinoceroses (Ceratotherium simum simum) to develop a visual discrimination between an open circle and a triangle. These stimuli were presented as black symbols on large white cards. The cards were presented 4.6 m apart and a food reward was given if the subject approached the open circle. Ten discrimination choices were given daily until each subject reached the criterion of 80% correct responding over a block of 50 trials. The female reached the criterion over trials 151–200, while the male required considerably longer (trials 501–550). The male's discrimination was dramatically affected by a shift in the food reward. This study demonstrates that these rhinos were able to develop a successful discrimination and this protocol could be used to further examine their visual acuity.
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Corporate Author Thesis
Publisher (down) Place of Publication Editor
Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0168-1591 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 6145
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