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McBride, S. D., Parker, M. O., Roberts, K., & Hemmings, A. (2017). Applied neurophysiology of the horse; implications for training, husbandry and welfare. Appl. Anim. Behav. Sci., 190, 90–101.
Abstract: Understanding the neural circuits underlying equine behaviour has the potential to help optimise strategies of husbandry and training. This review discusses two areas of neurophysiological research in a range of species and relates this information to the horse. The first discussion focuses on mechanisms of learning and motivation and assesses how this information can be applied to improve the training of the horse. The second concerns the identification of the equine neurophysiological phenotype, through behavioural and genetic probes, as a way of improving strategies for optimal equine husbandry and training success. The review finishes by identifying directions for future research with an emphasis on how neurophysiological systems (and thus behaviour) can be modified through strategic husbandry. This review highlights how a neurophysioloigical understanding of horse behaviour can play an important role in attaining the primary objectives of equitation science as well as improving the welfare of the horse.
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Bentley-Condit, V., & Smith, E. O. (2010). Animal tool use: current definitions and an updated comprehensive catalog. Behaviour, 147(2), 185–32.
Abstract: Despite numerous attempts to define animal tool use over the past four decades, the definition remains elusive and the behaviour classification somewhat subjective. Here, we provide a brief review of the definitions of animal tool use and show how those definitions have been modified over time. While some aspects have remained constant (i.e., the distinction between 'true' and 'borderline' tool use), others have been added (i.e., the distinction between 'dynamic' and 'static' behaviours). We present an updated, comprehensive catalog of documented animal tool use that indicates whether the behaviours observed included any 'true' tool use, whether the observations were limited to captive animals, whether tool manufacture has been observed, and whether the observed tool use was limited to only one individual and, thus, 'anecdotal' (i.e., N = 1). Such a catalog has not been attempted since Beck (1980). In addition to being a useful reference for behaviourists, this catalog demonstrates broad tool use and manufacture trends that may be of interest to phylogenists, evolutionary ecologists, and cognitive evolutionists. Tool use and tool manufacture are shown to be widespread across three phyla and seven classes of the animal kingdom. Moreover, there is complete overlap between the Aves and Mammalia orders in terms of the tool use categories (e.g., food extraction, food capture, agonism) arguing against any special abilities of mammals. The majority of tool users, almost 85% of the entries, use tools in only one of the tool use categories. Only members of the Passeriformes and Primates orders have been observed to use tools in four or more of the ten categories. Thus, observed tool use by some members of these two orders (e.g., Corvus, Papio) is qualitatively different from that of all other animal taxa. Finally, although there are similarities between Aves and Mammalia, and Primates and Passeriformes, primate tool use is qualitatively different. Approximately 35% of the entries for this order demonstrate a breadth of tool use (i.e., three or more categories by any one species) compared to other mammals (0%), Aves (2.4%), and the Passeriformes (3.1%). This greater breadth in tool use by some organisms may involve phylogenetic or cognitive differences � or may simply reflect differences in length and intensity of observations. The impact that tool usage may have had on groups' respective ecological niches and, through niche-construction, on their respective evolutionary trajectories remains a subject for future study.
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Beck, B. B. (1980). Animal tool behaviour: The use and manufacture of tools by animals. New York: Garland.
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Bergmüller, R., & Taborsky, M. (2010). Animal personality due to social niche specialisation. Trends in Ecology & Evolution, 25(9), 504–511.
Abstract: The existence of 'animal personality', i.e. consistent individual differences in behaviour across time and contexts, is an evolutionary puzzle that has recently generated considerable research interest. Although social factors are generally considered to be important, it is as yet unclear how they might select for personality. Drawing from ecological niche theory, we explore how social conflict and alternative social options can be key factors in the evolution and development of consistent individual differences in behaviour. We discuss how animal personality research might benefit from insights into the study of alternative tactics and illustrate how selection can favour behavioural diversification and consistency due to fitness benefits resulting from conflict reduction among social partners.
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Reader, S. M., & Laland, K. N. (2003). Animal Innovation. Oxford: Oxford University Press.
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Krueger, K., Esch, L., & Byrne, R. (2019). Animal behaviour in a human world: A crowdsourcing study on horses that open door and gate mechanisms. Plos One, 14(6), e0218954.
Abstract: Anecdotal reports of horses opening fastened doors and gates are an intriguing way of exploring the possible scope of horses' problem-solving capacities. The species' natural environment has no analogues of the mechanisms involved. Scientific studies on the topic are missing, because the rate of occurrence is too low for exploration under controlled conditions. Therefore, we compiled from lay persons case reports of horses opening closed doors and gates. Additionally, we collected video documentations at the internet platform YouTube, taking care to select raw data footage of unedited, clearly described and clearly visible cases of animals with no distinct signs of training or reduced welfare. The data included individuals opening 513 doors or gates on hinges, 49 sliding doors, and 33 barred doors and gateways; mechanisms included 260 cases of horizontal and 155 vertical bars, 43 twist locks, 42 door handles, 34 electric fence handles, 40 carabiners, and 2 locks with keys. Opening was usually for escape, but also for access to food or stable-mates, or out of curiosity or playfulness. While 56 percent of the horses opened a single mechanism at one location, 44 percent opened several types of mechanism (median = 2, min. = 1, max. = 5) at different locations (median = 2, min. = 1, max. = 4). The more complex the mechanism was, the more movements were applied, varying from median 2 for door handles to 10 for carabiners. Mechanisms requiring head- or lip-twisting needed more movements, with significant variation between individuals. 74 horses reported in the questionnaire had options for observing the behaviour in stable mates, 183 did not, which indicates that the latter learned to open doors and gates either individually or from observing humans. Experience favours opening efficiency; subjects which opened several door types applied fewer movements per lock than horses which opened only one door type. We failed to identify a level of complexity of door-fastening mechanism that was beyond the learning capacity of the horse to open. Thus, all devices in frequent use, even carabiners and electric fence handles, are potentially vulnerable to opening by horses, something which needs to be considered in relation to keeping horses safely.
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Gaunitz, C., Fages, A., Hanghøj, K., Albrechtsen, A., Khan, N., Schubert, M., et al. (2018). Ancient genomes revisit the ancestry of domestic and Przewalski's horses. Science, 360(6384), 111–114.
Abstract: The Eneolithic Botai culture of the Central Asian steppes provides the earliest archaeological evidence for horse husbandry, ~5,500 ya, but the exact nature of early horse domestication remains controversial. We generated 42 ancient horse genomes, including 20 from Botai. Compared to 46 published ancient and modern horse genomes, our data indicate that Przewalski's horses are the feral descendants of horses herded at Botai and not truly wild horses. All domestic horses dated from ~4,000 ya to present only show ~2.7% of Botai-related ancestry. This indicates that a massive genomic turnover underpins the expansion of the horse stock that gave rise to modern domesticates, which coincides with large-scale human population expansions during the Early Bronze Age.
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Murphy, M. A., Waits, L. P., Kendall, K. C., Wasser, S. K., Higbee, J. A., & Bogden, R. (2002). An evaluation of long-term preservation methods for brown bear (Ursus arctos) faecal DNA samples. Conservat. Genet., 3(4), 435–440.
Abstract: Relatively few large-scale faecal DNA studieshave been initiated due to difficulties inamplifying low quality and quantity DNAtemplate. To improve brown bear faecal DNA PCRamplification success rates and to determinepost collection sample longevity, fivepreservation methods were evaluated: 90%ethanol, DETs buffer, silica-dried, oven-driedstored at room temperature, and oven-driedstored at -20 °C. Preservationeffectiveness was evaluated for 50 faecalsamples by PCR amplification of a mitochondrialDNA (mtDNA) locus (~146 bp) and a nuclear DNA(nDNA) locus (~200 bp) at time points of oneweek, one month, three months and six months. Preservation method and storage timesignificantly impacted mtDNA and nDNAamplification success rates. For mtDNA, allpreservation methods had >= 75% success atone week, but storage time had a significantimpact on the effectiveness of the silicapreservation method. Ethanol preserved sampleshad the highest success rates for both mtDNA(86.5%) and nDNA (84%). Nuclear DNAamplification success rates ranged from 26-88%, and storage time had a significant impacton all methods but ethanol. Preservationmethod and storage time should be importantconsiderations for researchers planningprojects utilizing faecal DNA. We recommendpreservation of faecal samples in 90% ethanolwhen feasible, although when collecting inremote field conditions or for both DNA andhormone assays a dry collection method may beadvantageous.
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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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Bottom, S. H. Age-related changes in taste and gustatory response and feeding behaviour in the stabled horse.. Nottingham, UK: Nottingham Trent University.
Abstract: There is a paucity of research relating to the anatomy and physiology of gustation and olfaction in the horse. Moreover, whilst an age-related decline in gustation and olfaction has been recorded in humans, no such study has been conducted in the horse. The horse is reliant on gustation and olfaction to make appropriate decisions relating to both short and long term diet selection and thus, any compromise in function, has implications for food intake and potentially welfare. The principal aim of this study was to establish if, and to what extent, taste andgustatory responses are affected by age in the horse. Horses were allocated to the age groups Young (2-5 years), Middle (8-14 years) and Old (16 plus years) for the study of taste (n=18) and to Young (4-6 years), Middle (10-14 years) and Old (16 plus years) for the study of gustation (n=18). Individual taste responses and gustatory responses (taste in the absence of additional olfactory cues) were identified using two-choice preference testing and monadic testing. Statistical analysis was conducted using Minitab 14.0 and behaviour data was analysed using The Observer 5.0 (Noldus, Netherlands). No effect of age on taste response or gustatory response was recorded.
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