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Malavasi, R.; Huber, L. |
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Referential communication in the domestic horse (Equus caballus): first exploration in an ungulate species |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
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Proc. 3. Int. Equine. Sci. Mtg |
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domestic horse, referential communication, human-horse communication, intentionality |
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An important question in the study of animal communication is whether non-human animals are able to produce communicative gestures, i.e. nonvocal bodily actions directed to a recipient, physically ineffective but with a meaning shared in the social group [1]. Passive gestures are instrumental, tuned to the mere presence/absence of others, whereas active informers recognize receivers as communicative agents and activate shared-attention mechanisms for identifying their attentional state (SAM [2]; e.g. Schwab and Huber [3]). Six operational criteria must be evaluated to classify a signal as referential and intentional [4]: (1) alternative gazes between the partner and the target; (2) apparent attention-getting behaviours are deployed; (3) an audience is required to exhibit the behaviour; (4) the attentional status of an observer influences the propensity to exhibit behaviours; (5) communication is persistent and (6) there is elaboration of communicative behaviour when apparent attempts to manipulate the partner fail. Dogs [5] and non-human primates (reviewed in Liebal and Call [6]) can tune a human receiver’s attention to the object of interest by combining directional and attention-getting signals, such as turning the head or body, gazing to the receiver, and/or establishing eye contact. Research on other species is scarce.
Horses rely on humans to survive in domestic settings and may have evolved skills for communicating flexibly with them [7]. Horses understand human attentional cues (such as body and head orientation, eyes opened/closed) [8], permanent pointing [9] and, to some extent, gazing [10]. Here we tested the ability of 14 outdoor, herd-living domestic horses to communicate referentially with a human partner about the location of a desired target, a bucket of food out of reach. After the baiting of two buckets placed in opposite, unreachable locations were shown by the experimenter, the subject would walk to one of the two buckets. Because approaching a bucket would reveal that the food is out of reach, we expected the horse to look back to the experimenter, then to the bucket, and alternate this gazing several times to indicate its intention. To test whether our prediction is correct and alternate gazing is indeed the result of the horse's referential communication, we video-recorded the behaviour of the subjects in the test (FORWARD) and three control conditions: (1) FORWARD: experimenter oriented to the center of the arena, (2) BACK: experimenter backward oriented in respect to the arena, (3) ALONE: experimenter absent, (4) MANY: as FORWARD plus a familiar human oriented to the subject behind the bucket (Figure 1). We used a conservative criterion of back gazing by considering only turning the head back more than 90 degrees. The results confirmed our prediction. The horses alternated gazes between the partner and the buck significantly more often in the FORWARD than in all the other conditions (Table 1), thus satisfying operational criteria #1, #3 and #4. They also alternated head nods with gazes to the partner significantly more often during the FORWARD condition. We thus considered head nods not an instrumental signal of arousal, but an attention-getting behaviour with communicative function. Subjects used both head nods and neck stretched toward the buck more often in the FORWARD than in the BACK and the ALONE conditions, thus satisfying criteria #2, #3 and #4. In condition MANY, the frequency of head nods did not differ from condition FORWARD, probably because nods were directed to the additional partner behind the buck. This also satisfies criteria #4. The horses gazed to the partner most often in the FORWARD than in the BACK and the MANY conditions, but not in the ALONE. In this condition, subjects could observe the partner walking further from the test arena. To test for the different functions of gazes in presence and in absence of the partner, we compared their average duration between the two conditions: the significantly longer duration of gazes when the subject was alone suggests the instrumental monitoring function of gazes in this experimental condition.
Altogether, the findings suggest that domestic horses possess the ability to use referential communication in an interspecific context, but additional analyses are needed to test for operational criteria #5 and #6. Flexible and voluntary use of communicative signals reveal sophisticated cognitive processes involved in the strategic emission of these signals, and the finding of referential communication skills in an ungulate species forces us to reconsider the evolutionary path of intelligence. Furthermore, ungulates are used intensively by humans (transportation, meat, agriculture, leisure activities), and their welfare is often compromised. Determining whether ungulates can communicate their needs and preferences is paramount to a proper ethical management. |
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Malavasi, R. |
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Equine Behaviour @ team @ |
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5876 |
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Chapron, G.; Kaczensky, P.; Linnell, J.D.C.; Arx, M.; Huber, D.; Andrén, H. |
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Recovery of large carnivores in Europe's modern human-dominated landscapes |
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2014 |
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Science |
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346 |
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Equine Behaviour @ team @ Chapron2014 |
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6451 |
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Warmuth, V.; Eriksson, A.; Bower, M.A.; Barker, G.; Barrett, E.; Hanks, B.K.; Li, S.; Lomitashvili, D.; Ochir-Goryaeva, M.; Sizonov, G.V.; Soyonov, V.; Manica, A. |
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Reconstructing the origin and spread of horse domestication in the Eurasian steppe |
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Journal Article |
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2012 |
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Proceedings of the National Academy of Sciences |
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Proc Natl Acad Sci U S A |
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Despite decades of research across multiple disciplines, the early history of horse domestication remains poorly understood. On the basis of current evidence from archaeology, mitochondrial DNA, and Y-chromosomal sequencing, a number of different domestication scenarios have been proposed, ranging from the spread of domestic horses out of a restricted primary area of domestication to the domestication of numerous distinct wild horse populations. In this paper, we reconstruct both the population genetic structure of the extinct wild progenitor of domestic horses, Equus ferus, and the origin and spread of horse domestication in the Eurasian steppes by fitting a spatially explicit stepping-stone model to genotype data from >300 horses sampled across northern Eurasia. We find strong evidence for an expansion of E. ferus out of eastern Eurasia about 160 kya, likely reflecting the colonization of Eurasia by this species. Our best-fitting scenario further suggests that horse domestication originated in the western part of the Eurasian steppe and that domestic herds were repeatedly restocked with local wild horses as they spread out of this area. By showing that horse domestication was initiated in the western Eurasian steppe and that the spread of domestic herds across Eurasia involved extensive introgression from the wild, the scenario of horse domestication proposed here unites evidence from archaeology, mitochondrial DNA, and Y-chromosomal DNA. |
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Equine Behaviour @ team @ |
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5612 |
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Rybarczyk, P.; Rushen, J.; de Passille, A.M. |
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Recognition of people by dairy calves using colour of clothing |
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2003 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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81 |
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4 |
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307-319 |
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Cattle handling; Discrimination; Human-animal relationships; Colour cues; Y-maze |
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We examined whether very young dairy calves are able to discriminate between two people, and whether they use the colour of clothing or other indices to do so. During the familiarisation phase, one person (the familiar rewarder), who always wore the same colour clothes, gave milk, spoke gently and patted the calves in their individual pen for 6 days each week. During the test phase, the calf had to make a choice in an Y-maze placed in front of the gate of its stall. When the calves chose the familiar rewarder, they received 200 ml of milk as reinforcement. When they made the incorrect choice, they received nothing and were returned to their stall. On each test day, the calves made eight choices. The criterion of success was that the calf made at least six correct choices in eight trials on each of two consecutive test days (P<0.021 by the binomial law). The first experiment was carried out with fourteen 1-week-old male and female Holstein calves to see if calves could approach a person, who changed position in the maze, in order to obtain a feed reward. The familiar rewarder wore the same clothes as during the period of familiarisation and was in one arm of the Y-maze. The other arm was empty and the position of the familiar rewarder in the maze was randomised. Eleven of the 14 calves reached the criterion for success, after only three tests. The second experiment, carried out with five 2-week-old calves, examined whether the calves can differentiate the familiar rewarder (wearing the same clothing as during the period of familiarisation) from another person (the non-rewarder) wearing clothes of a different colour. The criterion of success was reached by all five calves. The third experiment was carried out with seven 2-week-old calves. It examined whether the calves can differentiate the familiar rewarder and the non-rewarder, when the two people are wearing clothes of the same colour (i.e. the same colour worn by the familiar rewarder during the phase of familiarisation). None of the calves were able to reach the criterion of success within a limited number of four test days. Often, calves would always choose the same arm of the maze. The fourth experiment was carried out on six 1-month-old calves. It was similar to experiment 3 with the difference that the familiar rewarder and the non-rewarder both wore the same colour clothes, but which were not the same colour as worn during the phase of familiarisation. Only one calf achieved the criterion of success within two test days. Results demonstrated that colour cues help very young calves to discriminate between two people, when these people wear different colour clothing. Some calves may be able to use other indicators than the colour of clothing. The Y-maze method is an promising way of examining calves' abilities to recognise people. |
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Aldridge, B.; Lescun, T.; Radcliffe, J.S. |
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Recent Molecular Advances in Equine Nutrition – from Test Tube to Practice |
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Conference Article |
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2012 |
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Proceedings of the 2. International Equine Science Meeting |
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Proc. 2. Int. Equine. Sci. Mtg |
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in press |
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Molecular Biology, Equine, Gut |
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The use of molecular techniques has increased our understanding of biological systems; in particular how genes and proteins operate within a cell or organ in response to various stimuli. These techniques can be used to study modes of action or mechanisms in disease states, or in response to applied management or nutritional changes. While applied and basic research are often conducted independent of one another, the real advantages to the equine industry will come from the use of molecular techniques to address applied challenges and/or opportunities within the industry. Therefore, the overall goal of our research has been to use basic and molecular tools to investigate modes of action resulting from changes in nutrition or management. We investigated active nutrient transport capacities of the small and large intestine, the effects of feed withdrawal on intestinal transporter gene expression and function, as well as other measurements pertaining to gut health (gut barrier function and gastrointestinal morphology). This novel and thorough approach to characterize sectional differences (fore- and hind- gut) of nutrient absorption has resulted in quantitative measures of active transport capacity for glucose, phosphorus, glutamine, lysine, and di-peptides in the horse. This research demonstrates that by-products of fermentation in the cecum such as microbial proteins and hydrolyzed phytate phosphorus, can be absorbed in the colon. Previously unidentified in the hindgut of the horse, we reported the expression of the di-and tri-peptide transporter, PepT1, as well as the neutral amino acid transporter, ASCT2, and transport of their substrates. We observed that nutrient transporters for glucose (SGLT1), glutamine (ASCT2) and for di-peptides (PepT1) can be regulated by diet, implying biological relevance to feeding management strategies. Feed withdrawal in horses pre- or post-surgery, prior to or during horse shows, during transport, or for other management reason is common. It is therefore important to understand how the removals of nutrients impacts gut health and/or disease susceptibility. The use of molecular techniques has provided a basis for the roles of certain core nutrients during times of feed withdrawal. For example, nutrients such as glutamine play a major role in proliferation and repair in the gut, serve as a major energy substrate, and may provide a means of assessing nutritional stress in the gut. Recent data concludes that following a feed withdrawal, the need for glutamine increases up to 116% in the proximal jejunum, and the gene expression of its transporter (ASCT2) increases by 78%. Additional studies utilizing molecular techniques to address nutritional challenges in the horse include how hormone loss due to ovariectomization impacts the homeostatic regulation of calcium and phosphorus in the horse. To improve nutrition and feeding management practices in horses, knowledge regarding how and where nutrients are absorbed, utilized and regulated under varying physiologic stressors (disease, exercise, reproductive status, etc.) is needed. The integration of applied and basic research facilitates a better and more complete understanding of mechanisms underlying common health and nutritional challenges facing the equine industry. |
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Aldridge, B. |
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Xenophon Publishing |
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Wald |
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Krueger, K.; |
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978-3-9808134-26 |
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Equine Behaviour @ team @ |
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5575 |
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Drent, P.J.; van Oers, K.; van Noordwijk, A.J. |
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Realized heritability of personalities in the great tit (Parus major) |
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2003 |
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Proceedings. Biological sciences / The Royal Society |
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Proc Biol Sci |
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270 |
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1510 |
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45-51 |
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Aggression; Animals; Animals, Domestic; Animals, Wild; *Behavior, Animal; Breeding; Exploratory Behavior; Female; *Heredity; Male; Selection (Genetics); Songbirds/*genetics/*physiology; Variation (Genetics) |
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Behaviour under conditions of mild stress shows consistent patterns in all vertebrates: exploratory behaviour, boldness, aggressiveness covary in the same way. The existence of highly consistent individual variation in these behavioural strategies, also referred to as personalities or coping styles, allows us to measure the behaviour under standardized conditions on birds bred in captivity, link the standardized measurements to the behaviour under natural conditions and measure natural selection in the field. We have bred the great tit (Parus major), a classical model species for the study of behaviour under natural conditions, in captivity. Here, we report a realized heritability of 54 +/- 5% for early exploratory behaviour, based on four generations of bi-directional artificial selection. In addition to this, we measured hand-reared juveniles and their wild-caught parents in the laboratory. The heritability found in the mid-offspring-mid-parent regression was significantly different from zero. We have thus established the presence of considerable amounts of genetic variation for personality types in a wild bird. |
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Netherlands Institute of Ecology, PO Box 40, 6666 ZG Heteren, The Netherlands. drent@cto.nioo.knaw.nl |
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0962-8452 |
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PMID:12590770 |
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refbase @ user @ |
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Lansade, L.; Bouissou, M.-F.; Erhard, H.W. |
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Reactivity to isolation and association with conspecifics: A temperament trait stable across time and situations |
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2008 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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106 |
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2-4 |
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355-373 |
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Horse; Equus caballus; Temperament; Trait; Isolation; Conspecific association; Stability |
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A temperament trait is generally defined as individual differences in behaviour that are present early in life and relatively stable across situations and over time. The aim of this study was to test the existence of a trait <<gregariousness>> in horses, by testing the stability across situations and over time of the responses to different social events. Sixty-six Welsh ponies and 44 Anglo-Arab horses were successively tested at 8 months and 1.5 years of age. Among them, 33 ponies and 21 horses were also tested at 2.5 years of age. They were submitted to four test situations: isolation and separation from, attraction towards and passing conspecifics. We carried out the analysis using each of four test groups as a unit (e.g. 33 Welsh ponies born in 2001, tested in isolation). Isolation and separation stood out as tests that showed a high consistency within test, across tests and across time. The most interesting behavioural parameter was the frequency of neighing, which was well correlated with other parameters measured in the same tests, such as defecation, locomotion and vigilance, as well as across the 3 years (e.g. for separation test: 0.41 < R < 0.61). Therefore, the behaviour of neighing observed in separation or isolation tests as early as 8 months of age appears to be a good indicator of similar behaviour in similar situations later in life, but also of other behaviours which can render the horse difficult to use. No parameter recorded during the attraction test presented stability across situations and time: the reactions to this test were not the expression of a stable characteristic of the individual and did not reflect the same characteristic as the three other tests. Of the different parameters recorded during the passing conspecifics test, the time to cross the arrival line near conspecifics showed good stability across years (0.35 < R < 0.68). This parameter was also correlated with many others recorded during the same test, and also, to a certain extent, to the frequency of neighing in the isolation and separation tests. This stability across responses expressed in various social contexts, and this stability over time, from 8 months to 2.5 years of age suggest the existence of a trait of gregariousness in the horse. From a practical point of view, that means it is possible to estimate the level of gregariousness of a horse as early as 8 months of age. Furthermore, additional analysis shows that gregariousness decreases with age. |
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Lansade, L.; Bouissou, M.-F. |
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Reactivity to humans: A temperament trait of horses which is stable across time and situations |
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Journal Article |
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2008 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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114 |
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3-4 |
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492-508 |
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Horse; Equus caballus; Reactivity to humans; Temperament trait; Personality; Behavioural consistency |
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The aim of this study was to test the existence of a “reactivity-to-humans” trait which is stable over time and across situations. For this purpose, four test situations were repeated on the same animals at 8 months, 1.5 years (N = 110 horses) and 2.5 years of age (N = 54 horses). These situations involved a “familiar passive human test” during which a familiar person stayed motionless in the test pen, a “familiar active human test” and an “unfamiliar active human test” during which a familiar or unfamiliar person attempted to touch the horse, and a “halter-fitting and heart-rate measurement test”, during which the horse was haltered and equipped with a surcingle and then had its heart-rate measured. Many behavioural variables were recorded during the tests, and correlations between ages and variables measured during other situations were investigated. When correlations were found, we concluded that the variables corresponded to temperament traits which are stable over time and across situations. The horses tested were divided into four groups according to breed (Anglo-Arab and Welsh) and year of birth (2001 and 2002). Data for each group were analysed separately with Spearman rank correlations. Regarding stability over time, there was a strong correlation between ages in the frequency of licking/nibbling the passive human, the time taken by a familiar or unfamiliar person to touch the horse, and the time taken to fit a halter (e.g. the time taken to touch the horse's muzzle was significantly correlated between ages in 7 out of the 8 possible cases: 0.40 <= R <= 0.67). Regarding stability across situations, results indicate that the more frequently a horse sniffed, licked or nibbled a passive human, the easier it was for either a familiar or unfamiliar active human to touch and halter it. Moreover, a horse which could be touched easily by a familiar human could also be touched and haltered easily by an unfamiliar human. These behaviours, which are linked directly to humans, therefore indicate good stability over time and across situations. In conclusion, this stability suggests the existence of a “reactivity-to-humans” temperament trait, whether the human is passive or active, familiar or unfamiliar. When the animals do not have frequent contact with humans, this reactivity is stable over time and can be measured as early as 8 months of age using the behavioural variables cited above. |
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Equine Behaviour @ team @ |
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Górecka-Bruzda, A.; Jastrzebska, E.; Sosnowska, Z.; Jaworski, Z.; Jezierski, T.; Chruszczewski, M.H. |
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Reactivity to humans and fearfulness tests: Field validation in Polish Cold Blood Horses |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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In Press, Corrected Proof |
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Cold blood horse; Temperament; Reactivity; Behavioural tests; Questionnaire; Validity |
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The aim of the present paper was to examine in field conditions the feasibility of the experimentally developed behavioural tests for assessment of two temperamental traits of cold blood horses: fearfulness and reactivity to humans. The predictive, convergent and discriminant validity of the tests was evaluated. Fifty-three Polish Cold Blood Horses (CBHs) were tested with the battery of behavioural tests involving passive and active contact with humans, handling, startling, novel object and surface tests. The behaviour of horses in real, on-farm conditions was evaluated with the caretaker's questionnaire (CQ). The tests demonstrated that a non-startling object and unforced human manipulations were willingly accepted by horses. In contrast, the horses were more reluctant to interact spontaneously with humans and showed longer persistence of a withdrawal reaction when presented with a startling stimulus or when restrained. CQ confirmed the common opinion of calmness, patience and good manageability of the CBHs. The results of behavioural tests were correlated with the scores derived from factor analysis of the CQ. Moreover, the majority of tests measures were correlated within reactivity to humans (median rs = 0.56) and fear tests (median rs = 0.41). Thus, predictive and convergent validity of the employed tests has been confirmed. However, reactivity to humans and fear measures were also correlated (median rs = 0.36), therefore discriminant validity of the tests used has been not confirmed. Therefore, we suggest that the parameters of tests designed to measure two separate traits: reactivity to humans and fearfulness, measured in fact avoidance of potential harm and withdrawal behaviour. Although selection for the work-appropriate behaviour is almost nonexistent nowadays, we conclude that Polish Cold Blood Horses are still predisposed for disciplines of equitation and driving, which demand an easy to handle horse. Nonetheless, we recommend behavioural testing of horses when approving for breeding. For this purpose, we can confirm usefulness of the experimentally developed tests in field assessment of general behavioural reactivity of horses. |
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0168-1591 |
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Equine Behaviour @ team @ |
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Blokland, A. |
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Reaction time responding in rats |
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1998 |
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Neuroscience and Biobehavioral Reviews |
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Neurosci Biobehav Rev |
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22 |
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6 |
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847-864 |
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Amphetamine/pharmacology; Animals; Behavior, Animal/drug effects/*physiology; Conditioning, Operant/drug effects/*physiology; Dopamine Uptake Inhibitors/pharmacology; Dose-Response Relationship, Drug; Male; Rats; Rats, Inbred Lew; Reaction Time/drug effects/*physiology |
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The use of reaction time has a great tradition in the field of human information processing research. In animal research the use of reaction time test paradigms is mainly limited to two research fields: the role of the striatum in movement initiation; and aging. It was discussed that reaction time responding can be regarded as “single behavior”, this term was used to indicate that only one behavioral category is measured, allowing a better analysis of brain-behavior relationships. Reaction time studies investigating the role of the striatum in motor functions revealed that the initiation of a behavioral response is dependent on the interaction of different neurotransmitters (viz. dopamine, glutamate, GABA). Studies in which lesions were made in different brain structures suggested that motor initiation is dependent on defined brain structures (e.g. medialldorsal striatum, prefrontal cortex). It was concluded that the use of reaction time measures can indeed be a powerful tool in studying brain-behavior relationships. However, there are some methodological constraints with respect to the assessment of reaction time in rats, as was tried to exemplify by the experiments described in the present paper. On the one hand one should try to control for behavioral characteristics of rats that may affect the validity of the parameter reaction time. On the other hand, the mean value of reaction time should be in the range of what has been reported in man. Although these criteria were not always met in several studies, it was concluded that reaction time can be validly assessed in rats. Finally, it was discussed that the use of reaction time may go beyond studies that investigate the role of the basal ganglia in motor output. Since response latency is a direct measure of information processing this parameter may provide insight into basic elements of cognition. Based on the significance of reaction times in human studies the use of this dependent variable in rats may provide a fruitful approach in studying brain-behavior relationships in cognitive functions. |
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Department of Psychology, University of Maastricht, The Netherlands |
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0149-7634 |
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PMID:9809315 |
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Equine Behaviour @ team @ |
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2807 |
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