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	Author	Stomp, M.; d'Ingeo, S.; Henry, S.; Cousillas, H.; Hausberger, M.
	Title	Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model			Type	Journal Article
	Year	2021	Publication	Applied Animal Behaviour Science	Abbreviated Journal
	Volume	236	Issue		Pages	105271
	Keywords	Laterality; Electroencephalography; Theta wave; Welfare; Horses
	Abstract	Assessing the animal welfare state is a challenge given the subjective individual cognitive and emotional processing involved. Electroencephalography (EEG) spectrum analysis has proved an ecologically valid recording situation to assess the link between brain processes and affective or cognitive states in humans: a higher slow wave/fast wave ratio has been associated with a positive internal state. In particular, a high production of theta power (3-8 Hz) has been related to positive emotions. On the other hand, it has been hypothesized that a left hemisphere (LH) dominance may be associated with a better welfare state. Here, we test the hypothesis that individual differences in the resting-state quantitative EEG power spectrum of adult horses (N = 18) and its lateralization pattern may reflect individual subjective perception of their conditions of life and welfare state. The results show clear individual differences in the proportions of the different waves and their inter-hemispheric distribution. Three different EEG power spectrum profiles were highlighted, from a bilateral predominance of theta waves in horses in a more positive welfare state to a bilateral predominance of beta waves in horses with clear expressions of compromised welfare. Interestingly, particular correlations were found between wave power activity and welfare parameters. We found a negative correlation between the number of stereotypic behaviours per hour and the median proportion of theta waves in the left hemisphere. and between the overall state (total chronic stress score) of welfare and gamma production in the right hemisphere (RH). These findings go along the hypothesis of a particular involvement of the left hemisphere for positive processing and of the right hemisphere for negative processing. However, the pattern of laterality did not appear as the most important feature here as both extreme clusters in terms of welfare showed bilateral predominance of one wave type. It is possible that hemispheric specialization makes more sense during acute emotion-inducing conditions rather than in this resting-state context (i.e. in absence of any high emotion-inducing stimulation), although the opposition gamma versus theta waves between both hemispheres in the horses with an intermediate welfare state is noticeable and intriguing. It seems that bilateral but also LH theta activity is a promising neurophysiological marker of good welfare in horses, while a bilateral or RH high production of gamma waves should alert about potential welfare alterations. Quantitative resting-state EEG power spectrum appears as a highly promising tool for exploring the brain processes involved in the subjective perception of chronic welfare, as a useful complementary tool for welfare assessment.
	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 @			Serial	6629
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	Author	Wolff, A.; Hausberger, M.; Le Scolan, N.
	Title	Experimental tests to assess emotionality in horses			Type	Journal Article
	Year	1997	Publication	Behavioural Processes	Abbreviated Journal	Behav. Process.
	Volume	40	Issue	3	Pages	209-221
	Keywords	Emotivity; Gregariousness; Horse; Neophobia; Open-field
	Abstract	Different tests were used to assess different aspects of the emotionality of 1-3 year-old horses: arena test; a [`]novel object' test; and a handling test. In reaction to the test situations no important differences were observed according to age or sex in the behaviour patterns, but clear individual differences were observed within these classes. The arena test seemed to reveal the degree of gregariousness of the animals whereas the results in the two other tests were correlated and seemed to reflect an inherent degree of fearfulness in the horse. Indices were developed that enabled to rank the animals, by taking into account all behaviour patterns shown. Such individual characteristics might have some genetic basis: half-siblings tended to behave the same way in most cases.
	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	0376-6357	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5018
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	Author	Rochais, C.; Henry, S.; Fureix, C.; Hausberger, M.
	Title	Investigating attentional processes in depressive-like domestic horses (Equus caballus)			Type	Journal Article
	Year	2016	Publication	Behavioural Processes	Abbreviated Journal
	Volume	124	Issue		Pages	93-96
	Keywords	Horses; Attention; Cognition; Welfare; Depression
	Abstract	Abstract Some captive/domestic animals respond to confinement by becoming inactive and unresponsive to external stimuli. Human inactivity is one of the behavioural markers of clinical depression, a mental disorder diagnosed by the co-occurrence of symptoms including deficit in selective attention. Some riding horses display ‘withdrawn’ states of inactivity and low responsiveness to stimuli that resemble the reduced engagement with their environment of some depressed patients. We hypothesized that ‘withdrawn’ horses experience a depressive-like state and evaluated their level of attention by confronting them with auditory stimuli. Five novel auditory stimuli were broadcasted to 27 horses, including 12 ‘withdrawn’ horses, for 5 days. The horses’ reactions and durations of attention were recorded. Non-withdrawn horses reacted more and their attention lasted longer than that of withdrawn horses on the first day, but their durations of attention decreased over days, but those of withdrawn horses remained stable. These results suggest that the withdrawn horses’ selective attention is altered, adding to already evidenced common features between this horses’ state and human depression.
	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	0376-6357	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6023
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	Author	Hausberger, M.; Bruderer, C.; Le Scolan, N.; Pierre, J.-S.
	Title	*Interplay between environmental and genetic factors in temperament/personality traits in horses (Equus caballus)*			Type	Journal Article
	Year	2004	Publication	Journal of Comparative Psychology	Abbreviated Journal	J Comp Psychol
	Volume	118	Issue	4	Pages	434-446
	Keywords	Affect; Animals; Behavior, Animal/physiology; Environment; Female; Horses/psychology; Learning; Male; Memory/physiology
	Abstract	The aim of the present study was to broach the question of the relative influence of different genetic and environmental factors on different temperament/personality traits of horses (Equus caballus). The researchers submitted 702 horses to standardized experimental tests and investigated 9 factors, either genetic or environmental. Genetic factors, such as sire or breed, seemed to influence more neophobic reactions, whereas environmental factors, such as the type of work, seemed to play a more dominant role in reactions to social separation or learning abilities. Additive effects were evident, showing how environmental factors may modulate behavioral traits. This study constitutes a first step toward understanding the relative weights of genetic factors and how the environment may intervene in determining individual behavioral characteristics.
	Address	Ethologie-Evolution-Ecologie, Universite de Rennes 1, Rennes, France. Martine.Hausberger@univ-rennes1.fr
	Corporate Author				Thesis
	Publisher		Place of Publication	Washington, D.C. : 1983	Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0735-7036	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15584780			Approved	no
	Call Number				Serial	1897
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	Author	Larose, C.; Richard-Yris, M.-A.; Hausberger, M.; Rogers, L.J.
	Title	Laterality of horses associated with emotionality in novel situations			Type	Journal Article
	Year	2006	Publication	Laterality	Abbreviated Journal	Laterality
	Volume	11	Issue	4	Pages	355-367
	Keywords	Affect/physiology; Animals; Brain/physiology; Female; Functional Laterality/physiology; Horses; Male; Social Behavior; *Social Environment
	Abstract	We have established that lateral biases are characteristic of visual behaviour in 65 horses. Two breeds, Trotters and French Saddlebreds aged 2 to 3, were tested on a novel object test. The main finding was a significant correlation between emotionality index and the eye preferred to view the novel stimulus: the higher the emotionality, the more likely that the horse looked with its left eye. The less emotive French Saddlebreds, however, tended to glance at the object using the right eye, a tendency that was not found in the Trotters, although the emotive index was the same for both breeds. The youngest French Saddlebreds did not show this trend. These results are discussed in relation to the different training practices for the breeds and broader findings on lateralisation in different species.
	Address	Universite de Rennes 1, France
	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	1357-650X	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16754236			Approved	no
	Call Number	Equine Behaviour @ team @ room B 3.029			Serial	1826
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	Author	Henry, S.; Fureix, C.; Rowberry, R.; Bateson, M.; Hausberger, M.
	Title	Do horses with poor welfare show 'pessimistic' cognitive biases?			Type	Journal Article
	Year	2017	Publication	The Science of Nature	Abbreviated Journal	Sci. Nat.
	Volume	104	Issue	1	Pages	8
	Keywords
	Abstract	This field study tested the hypothesis that domestic horses living under putatively challenging-to-welfare conditions (for example involving social, spatial, feeding constraints) would present signs of poor welfare and co-occurring pessimistic judgement biases. Our subjects were 34 horses who had been housed for over 3 years in either restricted riding school situations (e.g. kept in single boxes, with limited roughage, ridden by inexperienced riders; N = 25) or under more naturalistic conditions (e.g. access to free-range, kept in stable social groups, leisure riding; N = 9). The horses' welfare was assessed by recording health-related, behavioural and postural indicators. Additionally, after learning a location task to discriminate a bucket containing either edible food ('positive' location) or unpalatable food ('negative' location), the horses were presented with a bucket located near the positive position, near the negative position and halfway between the positive and negative positions to assess their judgement biases. The riding school horses displayed the highest levels of behavioural and health-related problems and a pessimistic judgment bias, whereas the horses living under more naturalistic conditions displayed indications of good welfare and an optimistic bias. Moreover, pessimistic bias data strongly correlated with poor welfare data. This suggests that a lowered mood impacts a non-human species' perception of its environment and highlights cognitive biases as an appropriate tool to assess the impact of chronic living conditions on horse welfare.
	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	1432-1904	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @ Henry2017			Serial	6665
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	Author	Sankey, C.; Richard-Yris, M.-A.; Henry, S.; Fureix, C.; Nassur, F.; Hausberger, M.
	Title	*Reinforcement as a mediator of the perception of humans by horses (Equus caballus)*			Type	Journal Article
	Year	2010	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	13	Issue	5	Pages	753-764-764
	Keywords	Perception of humans – Human/animal relationship – Positive reinforcement – Negative reinforcement – Equus caballus
	Abstract	A central question in the interspecific human/animal relationship is how domestic animals perceive humans as a significant element of their environment. In this study, we tested the hypothesis that the use of positive or negative reinforcement in horse training may have consequences on the animals’ perception of humans, as a positive, negative or neutral element. Two groups of ponies were trained to walk backwards in response to a vocal order using either positive or negative reinforcement. Heart rate monitors and behavioural observations were used to assess the animals’ perception of humans on the short (just after training) and long (5 months later) terms. The results showed that the type of reinforcement had a major effect on the subsequent animals’ perception of familiar and unfamiliar humans. Negative reinforcement was rapidly associated with an increased emotional state, as revealed by heart rate measurements and behavioural observations (head movements and ears laid back position). Its use led the ponies to seek less contact with humans. On the contrary, ponies trained with positive reinforcement showed an increased interest in humans and sought contact after training. This is especially remarkable as it was reached in a maximum of 5 sessions of 1 to 3 min (i.e. 5 to 15 min) and had lasting effects (visible after 5 months). Even learning was positively influenced by positive reinforcement. Overall, horses seem capable of associating humans to particular experiences and display extended long-term memory abilities.
	Address
	Corporate Author				Thesis
	Publisher	Springer Berlin / Heidelberg	Place of Publication		Editor
	Language		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	5175
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	Author	Basile, M.; Boivin, S.; Boutin, A.; Blois-Heulin, C.; Hausberger, M.; Lemasson, A.
	Title	*Socially dependent auditory laterality in domestic horses (Equus caballus)*			Type	Journal Article
	Year	2009	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	12	Issue		Pages	611-619
	Keywords
	Abstract	Laterality is now known to be an ubiquitous phenomenon among the vertebrates. Particularly, laterality of auditory processing has been demonstrated in a variety of species, especially songbirds and primates. Such a hemispheric specialization has been shown to depend on factors such as sound structure, species specificity and types of stimuli. Much less is known on the possible influence of social familiarity although a few studies suggest such an influence. Here we tested the influence of the degree of familiarity on the laterality of the auditory response in the domestic horse. This species is known for its social system and shows visible reactions to sounds, with one or two ears moving towards a sound source. By comparing such responses to the playback of different conspecific whinnies (group member, neighbor and stranger), we could demonstrate a clear left hemisphere (LH) preference for familiar neighbor calls while no preference was found for group member and stranger calls. Yet, we found an opposite pattern of ear side preference for neighbor versus stranger calls. These results are, to our knowledge, the first to demonstrate auditory laterality in an ungulate species. They open further lines of thought on the influence of the social “value” of calls and the listener's arousal on auditory processing and laterality.
	Address	Universite de Rennes I, EthoS, Ethologie animale et humaine-UMR 6552-CNRS Station Biologique de Paimpont, 35380, Paimpont, France, basilemuriel@hotmail.com
	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	1435-9456	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:19283416			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4761
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	Author	Rochais, C.; HENRY, S.; Sankey, C.; Nassur, F.; Gorecka-Bruzda, A.; HAUSBERGER, M.
	Title	Visual attention, an indicator of human-animal relationships? A study of domestic horses (Equus caballus)			Type	Journal Article
	Year	2014	Publication	Frontiers in Psychology	Abbreviated Journal
	Volume	5	Issue		Pages
	Keywords
	Abstract	BACKGROUND: As visual attention is an intrinsic part of social relationships, and because relationships are built on a succession of interactions, their establishment involves learning and attention. The emotional, rewarding or punishing, content can modulate selective attention. In horses, the use of positive/negative reinforcement during training determines short and long-term human-horse relationships. In a recent study in horses, where either food or withers' grooming were used as a reward, it appeared that only the food-rewarded horses learned the task and show better relationship with humans. In the present study, we hypothesized that this differential effect of grooming/food rewards on learning performances could be due to attentional processes. Monitoring, gazes and behaviors directed towards the trainer revealed that the use of a food reward (FR) as positive reinforcement increased horses' selective attention towards their trainer. Conversely, horses trained with grooming reward (GR) expressed more inattentive responses and did not show a decrease of “agitated” behavior. However, individual plotting of attention vs. rate of learning performances revealed a complex pattern. Thus, while all FR horses showed a “window” of attention related to faster learning performances, GR horses' pattern followed an almost normal curve where the extreme animals (i.e., highest and lowest attention) had the slowest learning performances. On the other hand, learning was influenced by attention: at the end of training, the more attentive horses had also better learning performances. This study, based on horses, contributes to the general debate on the place of attentional processes at the interface of emotion and cognition and opens new lines of thought about individual sensitivities (only individuals can tell what an appropriate reward is), attentional processes and 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	1664-1078	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5909
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	Author	Lesimple, C.; Sankey, C.; Richard, M.-A.; HAUSBERGER, M.
	Title	Do Horses Expect Humans to Solve Their Problems?			Type	Journal Article
	Year	2012	Publication	Frontiers in Psychology	Abbreviated Journal	Front. Psychol.
	Volume	3	Issue		Pages	306
	Keywords
	Abstract	Domestic animals are highly capable of detecting human cues, while wild relatives tend to perform less well (e.g. responding to pointing gestures). It is suggested that domestication may have led to the development of such cognitive skills. Here, we hypothesized that because domestic animals are so attentive and dependant to humans' actions for resources, the counter effect may be a decline of self sufficiency, such as individual task solving. Here we show a negative correlation between the performance in a learning task (opening a chest) and the interest shown by horses towards humans, despite high motivation expressed by investigative behaviours directed at the chest. If human-directed attention reflects the development of particular skills in domestic animals, this is to our knowledge the first study highlighting a link between human-directed behaviours and impaired individual solving task skills (ability to solve a task by themselves) in horses.
	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	1664-1078	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6568
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