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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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	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	Greco, B.J.; Brown, T.K.; Andrews, J.R.M.; Swaisgood, R.R.; Caine, N.G.
	Title	Social learning in captive African elephants (Loxodonta africana africana)			Type	Journal Article
	Year	2013	Publication		Abbreviated Journal	Animal Cognition
	Volume	16	Issue	3	Pages	459-469
	Keywords	Elephants; Loxodonta; Social learning; Imitation; Animal cognition
	Abstract	Social learning is a more efficient method of information acquisition and application than trial and error learning and is prevalent across a variety of animal taxa. Social learning is assumed to be important for elephants, but evidence in support of that claim is mostly anecdotal. Using a herd of six adult female African bush elephants (Loxodonta africana africana) at the San Diego Zoo’s Safari Park, we evaluated whether viewing a conspecific’s interactions facilitated learning of a novel task. The tasks used feeding apparatus that could be solved in one of two distinct ways. Contrary to our hypothesis, the method the demonstrating animal used did not predict the method used by the observer. However, we did find evidence of social learning: After watching the model, subjects spent a greater percentage of their time interacting with the apparatus than they did in unmodeled trials. These results suggest that the demonstrations of a model may increase the motivation of elephants to explore novel foraging tasks.
	Address
	Corporate Author				Thesis
	Publisher	Springer-Verlag	Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5668
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	Author	Russon, A.E.; Galdikas, B.M.F.
	Title	*Constraints on great apes' imitation: Model and action selectivity in rehabilitant orangutan (Pongo pygmaeus) imitation*			Type	Journal Article
	Year	1995	Publication	Journal of Comparative Psychology	Abbreviated Journal	J. Comp. Psychol.
	Volume	109	Issue	1	Pages	5-17
	Keywords	*Imitation (Learning); Primates (Nonhuman)
	Abstract	We discuss selectivity in great ape imitation, on the basis of an observational study of spontaneous imitation in free-ranging rehabilitant orangutans (Pongo pygmaeus). Research on great ape imitation has neglected selectivity, although comparative evidence suggests it may be important. We observed orangutans in central Indonesian Borneo and assessed patterns in the models and actions they spontaneously imitated. The patterns we found resembled those reported in humans. Orangutans preferred models with whom they had positive affective relationships (e.g., important caregiver or older sibling) and actions that reflected their current competence, were receptively familiar, and were relevant to tasks that faced them. Both developmental and individual variability were found. We discuss the probable functions of imitation for great apes and the role of selectivity in directing it. We also make suggestions for more effective elicitation of imitation. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
	Address
	Corporate Author				Thesis
	Publisher	American Psychological Association	Place of Publication	Us	Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1939-2087(Electronic);0735-7036(Print)	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @ 1995-20268-001			Serial	5690
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	Author	Abramson, J.Z.; Hernández-Lloreda, V.; Call, J.; Colmenares, F.
	Title	Experimental evidence for action imitation in killer whales (Orcinus orca)			Type	Journal Article
	Year	2013	Publication		Abbreviated Journal	Animal Cognition
	Volume	16	Issue	1	Pages	11-22
	Keywords	Social learning; Imitation; ‘Do-as-other-does’ test; Animal culture; Killer whales
	Abstract	Comparative experimental studies of imitative learning have focused mainly on primates and birds. However, cetaceans are promising candidates to display imitative learning as they have evolved in socioecological settings that have selected for large brains, complex sociality, and coordinated predatory tactics. Here we tested imitative learning in killer whales, Orcinus orca. We used a ‘do-as-other-does’ paradigm in which 3 subjects witnessed a conspecific demonstrator’s performance that included 15 familiar and 4 novel behaviours. The three subjects (1) learned the copy command signal ‘Do that’ very quickly, that is, 20 trials on average; (2) copied 100 % of the demonstrator’s familiar and novel actions; (3) achieved full matches in the first attempt for 8–13 familiar behaviours (out of 15) and for the 2 novel behaviours (out of 2) in one subject; and (4) took no longer than 8 trials to accurately copy any familiar behaviour, and no longer than 16 trials to copy any novel behaviour. This study provides experimental evidence for body imitation, including production imitation, in killer whales that is comparable to that observed in dolphins tested under similar conditions. These findings suggest that imitative learning may underpin some of the group-specific traditions reported in killer whales in the field.
	Address
	Corporate Author				Thesis
	Publisher	Springer-Verlag	Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5695
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	Author	Morton, F.B.; Lee, P.C.; Buchanan-Smith, H.M.
	Title	Taking personality selection bias seriously in animal cognition research: a case study in capuchin monkeys (Sapajus apella)			Type	Journal Article
	Year	2013	Publication		Abbreviated Journal	Animal Cognition
	Volume	16	Issue	4	Pages	677-684
	Keywords	Platyrrhines; Temperament; Cognitive experiment; Selection bias; Associative learning; Training
	Abstract	In most experimental work on animal cognition, researchers attempt to control for multiple interacting variables by training subjects prior to testing, allowing subjects to participate voluntarily, and providing subjects with food rewards. However, do such methods encourage selection bias from subjects’ personalities? In this study, we trained eighteen zoo-housed capuchin monkeys (Sapajus apella) for two experiments, under conditions of positive reinforcement (i.e. food rewards) and free-choice participation. Using a combination of behavioral and rater-based methods, we identified and validated five personality dimensions in these capuchins (Assertiveness, Openness, Neuroticism, Sociability, and Attentiveness). Scores on Openness were positively related to individual differences in monkey task participation, reflecting previous work showing that such individuals are often more active, curious, and willing to engage in testing. We also found a negative relationship between scores on Assertiveness and performance on tasks, which may reflect the trade-offs between speed and accuracy in these animals’ decision-making. Highly Assertive individuals (the most sociable within monkey groups) may also prioritize social interactions over engaging in research. Lastly, monkeys that consistently participated and performed well on both tasks showed significantly higher Openness and lower Assertiveness compared to others, mirroring relationships found between personality, participation, and performance among all participants. Participation and performance during training was clearly biased toward individuals with particular personalities (i.e. high Openness, low Assertiveness). Results are discussed in light of the need for careful interpretation of comparative data on animal cognition and the need for researchers to take personality selection bias more seriously. Animal Cognition Animal Cognition Look Inside
	Address
	Corporate Author				Thesis
	Publisher	Springer-Verlag	Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5696
Permanent link to this record



	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, χ2 = 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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	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	Valenchon, M.; Lévy, F.; Górecka-Bruzda, A.; Calandreau, L.; Lansade, L.
	Title	Characterization of long-term memory, resistance to extinction, and influence of temperament during two instrumental tasks in horses			Type	Journal Article
	Year	2013	Publication		Abbreviated Journal	Animal Cognition
	Volume	16	Issue	6	Pages	1001-1006
	Keywords	Personality; Learning; Individuality; Fearfulness; Equid; Cognitive flexibility
	Abstract	The present study investigated the influence of temperament on long-term recall and extinction of 2 instrumental tasks in 26 horses. In the first task (backward task), horses learned to walk backward, using commands given by an experimenter, in order to obtain a food reward. In the second task (active avoidance task), horses had to cross an obstacle after a bell rang in order to avoid emission of an air puff. Twenty-two months after acquisition, horses exhibited perfect recall performance in both tasks. Accordingly, no influence of temperament on recall performance could be observed for either task. In contrast, in the absence of positive or negative outcomes, the horses’ ability to extinguish their response to either task was highly variable. Resistance to extinction was related to some indicators of temperament: The most fearful horses tended to be the most resistant to extinction in the backward task, while the least sensitive horses tended to be the most resistant to extinction in the active avoidance task. These findings reveal extensive long-term memory abilities in horses and suggest an influence of temperament on learning processes other than acquisition.
	Address
	Corporate Author				Thesis
	Publisher	Springer Berlin Heidelberg	Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5735
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	Author	Krueger, K.; Farmer, K.; Heinze, J.
	Title	The effects of age, rank and neophobia on social learning in horses			Type	Journal Article
	Year	2014	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	17	Issue	3	Pages	645-655
	Keywords	Horse; Social learning; Sociality; Ecology; Social relationships
	Abstract	Social learning is said to meet the demands of complex environments in which individuals compete over resources and co-operate to share resources. Horses (Equus caballus) were thought to lack social learning skills because they feed on homogenously distributed resources with few reasons for conflict. However, the horse’s social environment is complex, which raises the possibility that its capacity for social transfer of feeding behaviour has been underestimated. We conducted a social learning experiment using 30 socially kept horses of different ages. Five horses, one from each group, were chosen as demonstrators, and the remaining 25 horses were designated observers. Observers from each group were allowed to watch their group demonstrator opening a feeding apparatus. We found that young, low ranking, and more exploratory horses learned by observing older members of their own group, and the older the horse, the more slowly it appeared to learn. Social learning may be an adaptive specialisation to the social environment. Older animals may avoid the potential costs of acquiring complex and potentially disadvantageous feeding behaviours from younger group members. We argue that horses show social learning in the context of their social ecology, and that research procedures must take such contexts into account. Misconceptions about the horse’s sociality may have hampered earlier studies.
	Address
	Corporate Author				Thesis
	Publisher	Springer Berlin Heidelberg	Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5737
Permanent link to this record



	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.
	Address
	Corporate Author				Thesis
	Publisher		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)00168-2			Serial	5769
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