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Harlow, H. F. (1950). Learning and satiation of response in intrinsically motivated complex puzzle performance by monkeys. J Comp Physiol Psychol, 43(4), 289–294.
Abstract: Two rhesus monkeys, given 60 two-hour sessions with a six-device mechanical puzzle showed clear evidence of learning, the curve showing ratio of incorrect to correct responses appearing quite comparable to similar curves obtained during externally rewarded situations. When, on the thirteenth day of tests, the subjects were presented with the puzzle 100 times at 6-minute intervals, the number of devices manipulated decreased regularly throughout the day, although there was no significant change in the number of times the problem assembly was attacked.
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Birch, H. G. (1945). The relation of previous experience to insightful problem-solving. J Comp Psychol, 38, 367–383.
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Boyce, P. N., & McLoughlin, P. D. (2021). Ecological Interactions Involving Feral Horses and Predators: Review with Implications for Biodiversity Conservation. Jour. Wild. Mgmt., n/a(n/a).
Abstract: ABSTRACT For many ecosystems, feral horses are increasingly becoming an important if not dominant component of ungulate biomass and hence influence on community dynamics. Yet we still know little of how horses contribute to key ecological interactions including predator-prey and indirect competitive relationships at a community level. Notably, feral species like horses can exhibit life-history traits that differ from that of native (mainly artiodactyl) herbivore competitors. Artificial selection for traits like increased, early, or extended reproduction that have yet to be reversed by natural selection, coupled with naturally selected differences in anatomy and behavior, in addition to unique management objectives for horses compared to other species, means that the dynamics of feral horse populations are not likely to align with what might be expected of other large herbivores. Unexpected population dynamics and inherent biological asymmetries between native ungulates and feral horses may therefore influence the former via direct competition for shared resources and through enemy-mediated interactions like apparent competition. In several localities feral horses now co-exist with multiple native prey species, some of which are in decline or are species at risk. Compounding risks to native species from direct or indirect competitive exclusion by horses is the unique nature and socio-political context of feral horse management, which tends towards allowing horse populations to be limited largely by natural, density-dependent factors. We summarize the inherent asymmetries between feral horse biology and that of other ungulate prey species with consequences for conservation, focusing on predator-prey and emerging indirect interactions in multi-prey systems, and highlight future directions to address key knowledge gaps in our understanding of how feral horses may now be contributing to the (re)structuring of food webs. Observations of patterns of rapid growth and decline, and associated skews in sex ratios of feral horse populations, indicate a heightened potential for indirect interactions among large ungulate prey species, where there is a prevalence of feral horses as preferred prey, particularly where native prey are declining. In places like western North America, we expect predator-prey interactions involving feral horses to become an increasingly important factor in the conservation of wildlife. This applies not only to economically or culturally important game species but also at-risk species, both predators (e.g., wolves [Canis lupus], grizzly bears [Ursus arctos]) and prey (e.g., woodland caribou [Rangifer tarandus caribou]), necessitating an ecological understanding of the role of horses in natural environments that goes beyond that of population control. ? 2021 The Wildlife Society.
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Palme, R. (2019). Non-invasive measurement of glucocorticoids: Advances and problems. Physiol. Behav., 199, 229–243.
Abstract: Glucocorticoids (GCs; i.e. cortisol/corticosterone) are a central component of the stress response and thus their measurement is frequently used to evaluate the impact of stressful situations. Their metabolites from faeces of various animal species are more and more taken as a non-invasive aid to assess GC release and thus adrenocortical activity. The current literature review includes an extensive collection (1327 papers) and evaluation (see also Supplementary Tables) of the literature on faecal cortisol/corticosterone metabolite (FCM) analysis published to date. It aims at giving reference for researchers interested in implementing FCM analysis into their study or seeking to improve such methods by providing background knowledge on GC metabolism and excretion, conveying insights into methodological issues and stating caveats of FCM analysis and by highlighting prerequisites for and some examples of a successful application of such methods. Collecting faecal samples and analysing FCMs may appear simple and straightforward, but researchers have to select and apply methods correctly. They also need to be aware of the many pitfalls and potentially confounding factors and, last but not least, have to carefully interpret results. Applied properly, measurement of FCMs is a powerful non-invasive tool in a variety of research areas, such as (stress) biology, ethology, ecology, animal conservation and welfare, but also biomedicine.
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Yarnell, K., Hall, C., & Billett, E. (2013). An assessment of the aversive nature of an animal management procedure (clipping) using behavioral and physiological measures. Physiol. Behav., 118, 32–39.
Abstract: Animal management often involves procedures that, while unlikely to cause physical pain, still cause aversive responses. The domestic horse (Equus caballus) regularly has excessive hair clipped off to facilitate its use as a riding/driving animal and this procedure causes adverse behavioral responses in some animals. The aim of this study was to compare behavioral and physiological measures to assess the aversive effect of this procedure. Ten horses were selected on the basis of being either compliant (C: n=5) or non-compliant (NC: n=5) during this procedure. The horses were subjected to a sham clipping procedure (SC: where the blades had been removed from the clippers) for a period of ten minutes. Measures were taken pre, during and post SC (-10min to +30min) and mean values calculated for ALL horses and for C and NC separately. Behavioral activity was scored (scale 1-5) by twenty students from video footage in (phase/group-blind scoring). Heart rate (HR), salivary cortisol and eye temperature were monitored throughout the procedure. The NC horses were found to be significantly more behaviorally active/less relaxed throughout the trial than C horses (p<0.05) with the greatest difference occurring during the SC procedure (p<0.01). NC horses were more active/less relaxed during, compared with pre or post SC (p<0.05), but showed no behavioral difference pre and post SC. HR of the NC horses was higher than that of the C horses throughout the trial but only significantly so after 10min of SC (p<0.01). ALL horses showed a significant increase in HR between +5 and +10min into the procedure (p<0.05). There was a significant increase in salivary cortisol concentration in ALL horses post procedure (p<0.01) with levels peaking at 20minute post SC. No significant differences in salivary cortisol concentration between C and NC were found at any stage of the trial. Eye temperature increased significantly in ALL horses during SC, peaking at +10min into the procedure (p<0.05) and then decreased substantially when SC had ceased (p<0.01). Although no significant differences were found between C and NC per se, there was a significant interaction between group and phase of trial (p<0.05) with the NC group showing a greater decrease in eye temperature post SC. There was a significant positive correlation between changes in salivary cortisol concentration and eye temperature (p<0.01) but no correlation between any of the other measures. Although the behavioral response of C and NC to this procedure was significantly different the physiological responses indicated that ALL horses found the procedure aversive. Eye temperature could be used as an objective and immediate measure of how an animal is responding to a specific situation in order to evaluate management procedures and adapt them where appropriate to reduce the negative impact on animal health and welfare.
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Lucidi, P., Bacco, G., Sticco, M., Mazzoleni, G., Benvenuti, M., Bernabò, N., et al. (2013). Assessment of motor laterality in foals and young horses (Equus caballus) through an analysis of derailment at trot. Physiol. Behav., 109, 8–13.
Abstract: The conflicting results regarding the study of motor laterality in horses may indicate that there does not exist a proper method to assess the degree and the direction of motor bias in these animals. Unfortunately, even less is known about the development of laterality in horses, and to what extent early manipulations can still exert their effects in adulthood. We propose a new method that can be easily applied at a very early age thus avoiding testing adult horses eventually biased by human handling and/or training. Forty-six horses (29 nine-month-old foals and 17 two-year old horses) were handled since birth bilaterally and housed in groups in wide areas. At the time of the analysis, in order to minimize environmental and sensorial disturbances, each horse was tested in a round pen individually or as dyad mother-foal. The ability/inability to properly execute a circle at trot was then recorded, assuming the direction of derailment, i.e. the cutting of the circle, as an indicator of motor bias. From the results of the study it is arguable that motor laterality in horses is acquired over time: in fact foals tested while their mothers were being subjected to longeing showed a higher percentage of ambidextrous animals, while two-year-old horses appeared biased toward the right (p<0.05). Results are discussed in the light of the scientific knowledge about equine biomechanics, taking into account horses' locomotion that leads to the advancement of the body mass through the activation of a kinetic chain that originates from the hindquarters.
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Marr, I., Stefanski, V., & Krueger, K. (2022). Lateralität – ein Indikator für das Tierwohl?[Laterality – an animal welfare indicator?]. Der Praktische Tierarzt, 103(12/2022), 1246–12757.
Abstract: Ein gutes Tierwohl definiert sich nicht nur durch die Abwesenheit von Stressindikatoren, sondern auch durch das Vorhandensein von Indikatoren, die auf ein gutes Wohlergehen hinweisen. So können stressbedingte Erkrankungen vermieden werden. Zur Bestimmung des Tierwohls bei Pferden wurde daher untersucht, inwieweit sich die sensorische Lateralität (einseitiger Gebrauch von Sinnesorganen) und die motorische Lateralität (einseitiger Gebrauch von Gliedmaßen) als einfach, schnell und kostengünstig zu erhebende Parameter eignen. Hierzu werden neben aktueller Literatur auch die eigenen Untersuchungsergebnisse zusammenfassend dargestellt. Die nach außen sichtbar werdende sensorische und motorische Lateralität sind das Resultat der cerebralen Lateralisierung. Dies beinhaltet nicht nur die Aufgabenteilung beider Gehirnhälften für ein effizienteres Aufnehmen und Speichern von Informationen, sondern sie steht auch in Verbindung mit der Entstehung und Verarbeitung von Emotionen, die maßgeblich am Wohlergehen eines Lebewesens beteiligt sind. Kurzzeitige Stressoren führen zu einer Erregung, die je nach Erfahrungen mit positiven oder negativen Emotionen in Verbindung steht. Emotionen helfen dem Organismus dabei, zu überleben. Andauernde negative Emotionen durch regelmäßige oder anhaltende negative Ereignisse führen zu Stress und reduzieren die Erwartung positiver Ereignisse (negativer cognitive Bias). Das Tier ist im Wohlergehen beeinträchtigt. Jüngst zeigte insbesondere die Messung der motorischen Lateralität Potenzial als Indikator für lang anhaltenden und chronischen Stress, denn gestresste Pferde, deren Stresshormonlevel stark ansteigt, zeigen einen zunehmenden Gebrauch der linken Gliedmaßen über einen längeren Zeitraum. Weiterhin zeigen erste Messungen einen Zusammenhang zwischen einer linksseitigen motorischen Lateralität und einer reduzierten Erwartung positiver Ereignisse (negativer cognitive Bias). Zusammen mit der sensorischen Lateralität, die in einer akuten Stressphase ebenso eine Linksverschiebung zeigt und somit als Indikator für Kurzzeitstress gilt, kann eine generelle, vermehrte Linksseitigkeit auch einen Hinweis auf erhöhte Emotionalität und Stressanfälligkeit sein. Eine sich steigernde Linksseitigkeit bedeutet eine präferierte Informationsverarbeitung durch die rechte Gehirnhälfte, die beispielsweise reaktives Verhalten, starke Emotionen und Stressantworten steuert. Es stellte sich jedoch heraus, dass wie bei allen Stressindikatoren auch in der Lateralitätsmessung ein Vergleichswert aus einer vorangegangenen Messung notwendig ist, denn nur Veränderungen zum häufiger werdenden Gebrauch der linken Seite können auf Stress bei Pferden hindeuten und die parallele Erhebung weiterer Parameter, wie zum Beispiel das Verhalten oder Stresshormone, können die Aussage der Lateralität bekräftigen.
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Boissy, A. (1995). Fear and Fearfulness in Animals. The Quarterly Review of Biology, 70(2), 165–191.
Abstract: Persistence of individual differences in animal behavior in reactions to various environmental challenges could reflect basic divergences in temperament, which might be used to predict details of adaptive response. Although studies have been carried out on fear and anxiety in various species, including laboratory, domestic and wild animals, no consistent definition of fearfulness as a basic trait of temperament has emerged. After a classification of the events that may produce a state of fear, this article describes the great variability in behavior and in physiological patterns generally associated with emotional reactivity. The difficulties of proposing fearfulness-the general capacity to react to a variety of potentially threatening situations-as a valid basic internal variable are then discussed. Although there are many studies showing covariation among the psychobiological responses to different environmental challenges, other studies find no such correlations and raise doubts about the interpretation of fearfulness as a basic personality trait. After a critical assessment of methodologies used in fear and anxiety studies, it is suggested that discrepancies among results are mainly due to the modulation of emotional responses in animals, which depend on numerous genetic and epigenetic factors. It is difficult to compare results obtained by different methods from animals reared under various conditions and with different genetic origins. The concept of fearfulness as an inner trait is best supported by two kinds of investigations. First, an experimental approach combining ethology and experimental psychology produces undeniable indicators of emotional reactivity. Second, genetic lines selected for psychobiological traits prove useful in establishing between behavioral and neuroendocrine aspects of emotional reactivity. It is suggested that fearfulness could be considered a basic feature of the temperament of each individual, one that predisposes it to respond similarly to a variety of potentially alarming challenges, but is nevertheless continually modulated during development by the interaction of genetic traits of reactivity with environmental factors, particularly in the juvenile period. Such interaction may explain much of the interindividual variability observed in adaptive responses.
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Brinkmann, L., Gerken, M., & Riek, A. (2013). Effect of long-term feed restriction on the health status and welfare of a robust horse breed, the Shetland pony (Equus ferus caballus). Res. Vet. Sci., 94(3), 826–831.
Abstract: Outdoor group housing is increasingly recognized as an appropriate housing system for domesticated horses. The objective of this study was therefore to investigate the effect of potential feed shortage in semi-natural horse keeping systems in winter on animal health and welfare. In 10 female Shetland ponies blood concentrations (NEFA, total protein (TP), total bilirubin (TB), beta-hydroxybutyrate (BHB) and thyroxine (T4)), body mass and the body condition score (BCS) were monitored for 7months including a 4months period of feed restriction in five of the 10 ponies. Restrictively fed animals lost 18.4±2.99% of their body mass and the BCS decreased by 2.2±0.8 points (BCS scale: 0=emaciated, 5=obese). Feed restriction led to a continuous increase in TB (P<0.001) and NEFA (P<0.01) concentrations compared to control ponies. The TP and BHB values only differed at the end of the trial with lower concentrations in restricted fed mares (P<0.05). Feed restriction had no effect on thyroxine concentrations. TB concentrations in the feed restricted group were out of the reference range during the entire feeding trial. The increased NEFA concentrations in feed restricted compared to control ponies suggest that fat was mobilized. The BCS, as well as plasma NEFA and TB concentrations were good indicators for a rapid detection of possible health problems caused by undernourishment in horses when kept under semi-natural conditions. In contrast, blood parameters of the control animals were within the reference ranges, suggesting that a year round outdoor housing with additional feed supply is an adequate housing system for a robust horse breed like the Shetland pony.
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Bussières, G., Jacques, C., Lainay, O., Beauchamp, G., Leblond, A., Cadoré, J. - L., et al. (2008). Development of a composite orthopaedic pain scale in horses. Research in Veterinary Science, 85(2), 294–306.
Abstract: This study addresses development and validation of a composite multifactorial pain scale (CPS) in an experimental equine model of acute orthopaedic pain. Eighteen horses were allocated to control (sedation with/without epidural analgesia – mixture of morphine, ropivacaine, detomidine and ketamine) and experimental groups: amphotericin-B injection in the tarsocrural joint induced pain and analgesia was either i.v. phenylbutazone administered post-induction of synovitis, or pre-emptive epidural mixture, or a pre-emptive combination of the 2. Inter- and intra-observer reproducibility was good (0.8<K<1). The key specific and sensitive behavioural indices were response to palpation of the painful area, posture, and, of lesser value, pawing on the floor, kicking at abdomen and head movement. Of particular interest was the statistical correlation observed between the CPS and both non-invasive blood pressure (P<0.0001) and blood cortisol (P<0.002). This study established the value of some behavioural and physiological criteria in determining equine orthopaedic pain intensity and clearly demonstrated that pre-emptive, multimodal analgesia provided better management than the two other protocols tested.
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