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Mizuguchi, M., Arai, M., Ke, Y., Nitta, K., & Kuwajima, K. (1998). Equilibrium and kinetics of the folding of equine lysozyme studied by circular dichroism spectroscopy. Journal of Molecular Biology, 283(1), 265–277.
Abstract: The equilibrium unfolding and the kinetics of unfolding and refolding of equine lysozyme, a Ca2+-binding protein, were studied by means of circular dichroism spectra in the far and near-ultraviolet regions. The transition curves of the guanidine hydrochloride-induced unfolding measured at 230 nm and 292.5 nm, and for the apo and holo forms of the protein have shown that the unfolding is well represented by a three-state mechanism in which the molten globule state is populated as a stable intermediate. The molten globule state of this protein is more stable and more native-like than that of α-lactalbumin, a homologous protein of equine lysozyme. The kinetic unfolding and refolding of the protein were induced by concentration jumps of the denaturant and measured by stopped-flow circular dichroism. The observed unfolding and refolding curves both agreed well with a single-exponential function. However, in the kinetic refolding reactions below 3 M guanidine hydrochloride, a burst-phase change in the circular dichroism was present, and the burst-phase intermediate in the kinetic refolding is shown to be identical with the molten globule state observed in the equilibrium unfolding. Under a strongly native condition, virtually all the molecules of equine lysozyme transform the structure from the unfolded state into the molten globule, and the subsequent refolding takes place from the molten globule state. The transition state of folding, which may exist between the molten globule and the native states, was characterized by investigating the guanidine hydrochloride concentration-dependence of the rate constants of refolding and unfolding. More than 80% of the hydrophobic surface of the protein is buried in the transition state, so that it is much closer to the native state than to the molten globule in which only 36% of the surface is buried in the interior of the molecule. It is concluded that all the present results are best explained by a sequential model of protein folding, in which the molten globule state is an obligatory folding intermediate on the pathway of folding.
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McLean, A. N. (2004). Short-term spatial memory in the domestic horse. Appl. Anim. Behav. Sci., 85(1-2), 93–105.
Abstract: This study investigates the ability of horses to recall a feeding event in a two-point choice apparatus. Twelve horses were individually tested whereby they were maintained immobile in a test arena and visually and aurally experienced the delivery of food into one of two feed goals. The horses were then released to make their choice in two experimental contexts: immediate release after experiencing the delivery of food, and release 10 s after food delivery. Each horse performed 40 immediate-release (IR) trials, followed by forty 10-s release trials over a 3-day period. In addition, the same horses were tested 3 months later in the spring with the same number and sequence of trials. Results were analysed by log-linear analysis of frequencies. Results showed that while horses were able to achieve the correct feed goal choice in the immediate-release trials, they were unsuccessful with the 10-s release trials. This suggests that there are limitations in recall abilities in horses, in that they may not possess a prospective type of memory. There are welfare and training implications in these findings concerning the effects of overestimating the mental abilities of horses during training and the effects of delays in reinforcements.
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Nicol, C. J. (2002). Equine learning: progress and suggestions for future research. Appl. Anim. Behav. Sci., 78(2-4), 193–208.
Abstract: Horses are well able to form classical and instrumental associations and so the focus of much recent research has been on the stimulus control of instrumental learning. Horses appear to discriminate using spatial cues more easily than other stimulus features, as indicated both by the speed of initial task acquisition and by the extent to which acquired discriminations can be reversed. Phenomena associated with discrimination learning in laboratory animals, including generalisation and peak shift, have been demonstrated in horses. However, the ability of horses to classify stimuli into categories is more controversial. Although there is some evidence that horses may be able to form categories based on similarities in the physical appearance of different stimuli, there is currently no evidence that they are able to develop abstract concepts. Their performance on social learning tasks has also been poor. Few correlations are observed between the learning ability of individual horses on different tasks, suggesting that it may not be possible to classify individual horses as `good' or `poor' learners. Better learning performance by horses that are naturally calm is probably due to reduced interference in the learning process. Correct handling procedures can lower reactivity levels in horses, and may facilitate learning in some circumstances. Future research on equine learning needs to take into account the complex nature of equine social interaction. Studies on the effects of stress on learning, and on social and spatial cognition, are also particularly needed.
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Berger, J. (1986). Wild horses of the Great Basin. Chicago: University of Chicago Press.
Abstract: Describes the behavior of wild horses living in the Great Basin Desert of Nevada and discusses the role of the horses in the area's ecology
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McDonnell, S. M., & Poulin, A. (2002). Equid play ethogram. Appl. Anim. Behav. Sci., 78(2-4), 263–290.
Abstract: An ethogram of play behavior among equids was developed. Several key English-language studies on equids were reviewed to derive a preliminary inventory of specific behaviors to be included in the ethogram. Our primary observations were based on a herd of semi-feral Shetland-type ponies kept at New Bolton Center, University of Pennsylvania School of Veterinary Medicine, Kennett Square, PA. Greater than 100 h of direct observation and photo-documentation focused specifically on play in order to identify play behaviors to be added to the preliminary inventory and to obtain detailed descriptions of each behavior. Additionally, these observations were supplemented with photographs obtained during several years of observational study of this herd for other purposes, and with the cumulative equid observational experience and study notes of the principal investigator with other equid species. An initial draft was sent out to 18 equine behavior colleagues for review. A total of 38 individual behaviors classified into four distinct categories were included in the ethogram. These included object play (14 entries), play sexual behavior (3 entries), locomotor play (14 entries) and play fighting (7 entries). All of the behaviors catalogued from direct observation of the herd were also found in the equid literature. The resulting ethogram offers a practical tool as a field guide or reference for quantitative research and other studies of equid play behavior as well as for teaching of equid behavior.
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Spengler M.I., & Rasia M. (2001). Influence of Plasma Proteins on Erythrocyte Aggregation in Three Mammalian Species. Vet.Res.Comm, 25(7), 591–599.
Abstract: The aggregation capacity of human erythrocytes lies between that of the non-aggregating bovine erythrocytes and the remarkably aggregating equine ones. As the ability to aggregate is attributed to cell factors and the composition of the plasma proteins, the role that plasma proteins play in the aggregation process in these three species was studied. Washed erythrocytes were suspended in phosphate-buffered saline (PBS; pH 7.4, 300 mOsm/L) plus polyvinylpyrrolidone (PVP) in a suitable concentration to obtain an average intensity of aggregation (control media). The superimposed effect of replacing 80% of the medium by either autologous plasma, serum or albumin solution was studied. The plasma proteins appeared to enhance aggregation by human and equine erythrocytes, but impaired this process in bovine erythrocytes. Some evidence was obtained supporting the existence of serum factors capable of reducing aggregation of erythrocytes in cattle and it was concluded that the non-aggregating behaviour of bovine erythrocytes may be due to the cells interacting particularly with the macromolecules in the serum.
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Williams, J. L., Friend, T. H., Nevill, C. H., & Archer, G. (2004). The efficacy of a secondary reinforcer (clicker) during acquisition and extinction of an operant task in horses. Appl. Anim. Behav. Sci., 88(3-4), 331–341.
Abstract: “Clicker training” is a popularly promoted training method based on operant conditioning with the use of a secondary reinforcer (the clicker). While this method draws from theories of learning and is used widely, there has been little scientific investigation of its efficacy. We used 60 horses, Equus callabus, and assigned each horse to one of six reinforcement protocols. The reinforcement protocols involved combinations of reinforcers administered (primary versus secondary plus primary), schedule of reinforcement (continuous versus variable ratio), and reinforcers applied during extinction (none or secondary). There were no differences (P>=0.11) between horses which received a secondary reinforcer (click) followed by the primary reinforcer (food) and those which received only the primary reinforcer (food) in the number of trials required to train the horses to touch their noses to a plastic cone (operant response). There also were no differences (P>=0.12) between horses which received the secondary reinforcer plus primary reinforcer and those which received only the primary reinforcer in regards to the number of trials to extinction. We conclude that there is no difference in the amount of training required to learn the operant task or in the task's resistance to extinction between horses that received a secondary reinforcer followed by a primary reinforcer versus horses which received only a primary reinforcer.
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Strand, S. C., Tiefenbacher, S., Haskell, M., Hosmer, T., McDonnell, S. M., & Freeman, D. A. (2002). Behavior and physiologic responses of mares to short-term isolation. Appl. Anim. Behav. Sci., 78(2-4), 145–157.
Abstract: The aim of this study was to evaluate the behavior and physiologic responses of mares to removal from an established pasture herd and to isolation in a pasture setting for 6 h (Group I, n=5). Responses of mares in Group I were compared to mares that were transported and returned to the herd (Group T, n=5) and to mares moved to the isolation pasture with a companion (Group C, n=5). Behavior was recorded continuously for 6 h on the day before the isolation procedures (baseline, Day 0) and again on the day of the procedure (test, Day 1). Plasma cortisol, white blood cell count (WBC), neutrophil:lymphocyte ratio (N:L), and hematocrit (HCT) were measured once on Day 0 (a.m.) and twice on Day 1 (a.m. and p.m.). Heart rate (HR) was monitored continuously during Day 0 and Day 1. Intradermal response to phytohemagglutinin (PHA) injection was measured 18 h following injection, which was administered at the end of Day 1. Average time spent standing alert increased (P<0.05) in Groups I and C and average time spent grazing decreased (P<0.05) in Group C from Day 0 to Day 1. Also, there was a significant difference between groups (based on a calculated χ2-square value) in the proportion of mares that autogroomed, defecated, urinated, rolled, and whinnied on Day 1. Activity shift rate (ASR) and temperament scores increased significantly in Groups I and C from Day 0 to Day 1 (P<0.05). Plasma cortisol increased significantly in all groups from Day 0 to Day 1, a.m. (P<0.05) and decreased significantly from Day 1, a.m. to Day 1, p.m. (P<0.05). HCT significantly increased in all three groups from Day 0 to Day 1, a.m. (P<0.05). WBC significantly increased in Group T from Day 0 to Day 1, a.m. (P<0.05). N:L ratio significantly increased in Groups I and C from Day 0 and Day 1, a.m. to Day 1, p.m. (P<0.05). A variety of measures did indicate a response to removal from the pasture group, however, the overall, short-term response was minimal. Since the responses of Groups I and C were similar, the effects of isolation versus a novel environment or separation from the established herd could not be differentiated.
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Williams, J. L., Friend, T. H., Toscano, M. J., Collins, M. N., Sisto-Burt, A., & Nevill, C. H. (2002). The effects of early training sessions on the reactions of foals at 1, 2, and 3 months of age. Appl. Anim. Behav. Sci., 77(2), 105–114.
Abstract: An early training procedure commonly termed “foal imprint training” is widely promoted in the horse industry. However, there have been no published scientific investigations of its efficacy. This study determined the effects of a training procedure on foals and their reaction to stimuli used in the early training procedure, and to a novel stimulus, at 1, 2 and 3 months of age. Twenty-five foals received a standard training procedure at 2, 12, 24, and 48 h after birth. After the training procedure, the foals received minimal additional handling that included veterinary treatments and occasional relocation. Twenty-two foals born over the same time period served as controls. All 47 (25 trained, 22 control) foals were tested at 1 month of age. Only 20 were available for testing at 2 months of age, and nine were available at 3 months. Percentage change from baseline heart rate, time required to complete exposure to each stimulus (foals that were more reactive took longer) and the behavior of each foal during the introduction of each stimulus were recorded. Overall, the control foals tended to receive lower (better) behavioral scores at 1 and 2 months of age. Foals that underwent the training procedure tended to require less time to complete exposure to the stimulus and had lower heart rates during exposure to the stimuli at 1 and 2 months of age. By 3 months of age, there were no significant differences between trained and control foals for any measures. Early training was not efficacious in this study.
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Pick, D. F., Lovell, G., Brown, S., & Dail, D. (1994). Equine color perception revisited. Appl. Anim. Behav. Sci., 42(1), 61–65.
Abstract: An attempt to replicate Grzimek (1952; Z. Tierpsychol., 27: 330-338) is reported where a Quarter-Horse mare chose between colored and gray stimuli for food reinforcement. Stimuli varied across a broad range of reflectance values. A double-blind procedure with additional controls for auditory, olfactory, tactile, and position cues was used. The subject could reliably discriminate blue (462 nm) vs. gray, and red (700 nm) vs. gray without regard to reflectance (P<0.001), but could not discriminate green (496 nm) vs. gray. It is suggested that horses are dichromats in a manner similar to swine and cattle.
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