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Cooper, J. J. (1998). Comparative learning theory and its application in the training of horses. Equine Vet J Suppl, (27), 39–43.
Abstract: Training can best be explained as a process that occurs through stimulus-response-reinforcement chains, whereby animals are conditioned to associate cues in their environment, with specific behavioural responses and their rewarding consequences. Research into learning in horses has concentrated on their powers of discrimination and on primary positive reinforcement schedules, where the correct response is paired with a desirable consequence such as food. In contrast, a number of other learning processes that are used in training have been widely studied in other species, but have received little scientific investigation in the horse. These include: negative reinforcement, where performance of the correct response is followed by removal of, or decrease in, intensity of a unpleasant stimulus; punishment, where an incorrect response is paired with an undesirable consequence, but without consistent prior warning; secondary conditioning, where a natural primary reinforcer such as food is closely associated with an arbitrary secondary reinforcer such as vocal praise; and variable or partial conditioning, where once the correct response has been learnt, reinforcement is presented according to an intermittent schedule to increase resistance to extinction outside of training.
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Mills, D. S. (1998). Applying learning theory to the management of the horse: the difference between getting it right and getting it wrong. Equine Vet J Suppl, (27), 44–48.
Abstract: Horses constantly modify their behaviour as a result of experience. This involves the creation of an association between events or stimuli. The influence of people on the modification and generation of certain behaviour patterns extends beyond the intentional training of the horse. The impact of any action depends on how it is perceived by the horse, rather than the motive of the handler. Negative and positive reinforcement increase the probability of specific behaviours recurring i.e. strengthen the association between events, whereas punishment reduces the probable recurrence of a behaviour without providing specific information about the desired alternative. In this paper the term 'punishers' is used to refer to the physical aids, such as a whip or crop, which may be used to bring about the process of punishment. However, if their application ceases when a specific behaviour occurs they may negatively reinforce that action. Intended 'punishers' may also be rewarding (e.g. for attention seeking behaviour). Therefore, contingency factors (which define the relationship between stimuli, such as the level of reinforcement), contiguity factors (which describe the proximity of events in space or time) and choice of reinforcing stimuli are critical in determining the rate of learning. The many problems associated with the application of punishment in practice lead to confusion by both horse and handler and, possibly, abuse of the former. Most behaviour problems relate to handling and management of the horse and can be avoided or treated with a proper analysis of the factors influencing the behaviour.
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Harman, A. M., Moore, S., Hoskins, R., & Keller, P. (1999). Horse vision and an explanation for the visual behaviour originally explained by the 'ramp retina'. Equine Vet J, 31(5), 384–390.
Abstract: Here we provide confirmation that the 'ramp retina' of the horse, once thought to result in head rotating visual behaviour, does not exist. We found a 9% variation in axial length of the eye between the streak region and the dorsal periphery. However, the difference was in the opposite direction to that proposed for the 'ramp retina'. Furthermore, acuity in the narrow, intense visual streak in the inferior retina is 16.5 cycles per degree compared with 2.7 cycles per degree in the periphery. Therefore, it is improbable that the horse rotates its head to focus onto the peripheral retina. Rather, the horse rotates the nose up high to observe distant objects because binocular overlap is oriented down the nose, with a blind area directly in front of the forehead.
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Yokoyama, S., & Radlwimmer, F. B. (1999). The molecular genetics of red and green color vision in mammals. Genetics, 153(2), 919–932.
Abstract: To elucidate the molecular mechanisms of red-green color vision in mammals, we have cloned and sequenced the red and green opsin cDNAs of cat (Felis catus), horse (Equus caballus), gray squirrel (Sciurus carolinensis), white-tailed deer (Odocoileus virginianus), and guinea pig (Cavia porcellus). These opsins were expressed in COS1 cells and reconstituted with 11-cis-retinal. The purified visual pigments of the cat, horse, squirrel, deer, and guinea pig have lambdamax values at 553, 545, 532, 531, and 516 nm, respectively, which are precise to within +/-1 nm. We also regenerated the “true” red pigment of goldfish (Carassius auratus), which has a lambdamax value at 559 +/- 4 nm. Multiple linear regression analyses show that S180A, H197Y, Y277F, T285A, and A308S shift the lambdamax values of the red and green pigments in mammals toward blue by 7, 28, 7, 15, and 16 nm, respectively, and the reverse amino acid changes toward red by the same extents. The additive effects of these amino acid changes fully explain the red-green color vision in a wide range of mammalian species, goldfish, American chameleon (Anolis carolinensis), and pigeon (Columba livia).
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McDonnell, S. M., Freeman, D. A., Cymbaluk, N. F., Schott, H. C. 2nd, Hinchcliff, K., & Kyle, B. (1999). Behavior of stabled horses provided continuous or intermittent access to drinking water. Am J Vet Res, 60(11), 1451–1456.
Abstract: OBJECTIVE: To compare quantitative measures and clinical assessments of behavior as an indication of psychologic well-being of stabled horses provided drinking water continuously or via 1 of 3 intermittent delivery systems. ANIMALS: 22 Quarter Horse (QH) or QH-crossbred mares and 17 Belgian or Belgian-crossbred mares (study 1) and 24 QH or QH-crossbred mares and 18 Belgian or Belgian-crossbred mares (study 2). PROCEDURE: Stabled horses were provided water continuously or via 1 of 3 intermittent water delivery systems in 2 study periods during a 2-year period. Continuous 24-hour videotaped samples were used to compare quantitative measures and clinical assessments of behavior among groups provided water by the various water delivery systems. RESULTS: All horses had clinically normal behavior. Significant differences in well being were not detected among groups provided water by the various delivery systems. CONCLUSIONS AND CLINICAL RELEVANCE: Various continuous and intermittent water delivery systems can provide adequately for the psychologic well-being of stabled horses.
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Pell, S. M., & McGreevy, P. D. (1999). Prevalence of stereotypic and other problem behaviours in thoroughbred horses. Aust Vet J, 77(10), 678–679.
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Feist, J. D., & McCullough, D. R. (1975). Reproduction in feral horses. J Reprod Fertil Suppl, (23), 13–18.
Abstract: A behavioural study of feral horses was conducted on the Pryor Mountain Wild Horse Range in the western United States. All 270 horses on the Range were identified individually. The sex ratio was nearly balanced. Foal to adult female ratio was 43-2:100. Morality was concentrated among foals and old horses. Horses were organized as forty-four harem groups each with a dominant stallion, one to two immature stallions, one to three immature mares, one to three adult mares and their yearling and foal offspring, and 23 bachelor groups of one to eight stallions. Harem groups were quite stable year-round because of dominance and leadership by the stallions and group fidelity by mares and their offsring. Most changes occurred during the breeding season and involved immature females. Defeat of dominant stallions was infrequent. Immature males were tolerated because of their submissive behaviour. Bachelor stallion groups were inherently unstable. Mares came into heat after foaling in May/June, and were mated by harem stallions only.
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Belonje, P. C., & van Niekerk, C. H. (1975). A review of the influence of nutrition upon the oestrous cycle and early pregnancy in the mare. J Reprod Fertil Suppl, (23), 167–169.
Abstract: Attention is drawn to the beneficial effect of improved nutrition during winter and early spring on the ovarian activity of mares. Furthermore, the necessity of an adequate plane of nutrition during early pregnancy to prevent embryonic resorption is stressed.
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Abbruzzetti, S., Crema, E., Masino, L., Vecli, A., Viappiani, C., Small, J. R., et al. (2000). Fast events in protein folding: structural volume changes accompanying the early events in the N-->I transition of apomyoglobin induced by ultrafast pH jump. Biophys J, 78(1), 405–415.
Abstract: Ultrafast, laser-induced pH jump with time-resolved photoacoustic detection has been used to investigate the early protonation steps leading to the formation of the compact acid intermediate (I) of apomyoglobin (ApoMb). When ApoMb is in its native state (N) at pH 7.0, rapid acidification induced by a laser pulse leads to two parallel protonation processes. One reaction can be attributed to the binding of protons to the imidazole rings of His24 and His119. Reaction with imidazole leads to an unusually large contraction of -82 +/- 3 ml/mol, an enthalpy change of 8 +/- 1 kcal/mol, and an apparent bimolecular rate constant of (0.77 +/- 0.03) x 10(10) M(-1) s(-1). Our experiments evidence a rate-limiting step for this process at high ApoMb concentrations, characterized by a value of (0. 60 +/- 0.07) x 10(6) s(-1). The second protonation reaction at pH 7. 0 can be attributed to neutralization of carboxylate groups and is accompanied by an apparent expansion of 3.4 +/- 0.2 ml/mol, occurring with an apparent bimolecular rate constant of (1.25 +/- 0.02) x 10(11) M(-1) s(-1), and a reaction enthalpy of about 2 kcal/mol. The activation energy for the processes associated with the protonation of His24 and His119 is 16.2 +/- 0.9 kcal/mol, whereas that for the neutralization of carboxylates is 9.2 +/- 0.9 kcal/mol. At pH 4.5 ApoMb is in a partially unfolded state (I) and rapid acidification experiments evidence only the process assigned to carboxylate protonation. The unusually large contraction and the high energetic barrier observed at pH 7.0 for the protonation of the His residues suggests that the formation of the compact acid intermediate involves a rate-limiting step after protonation.
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Clayton, H. M., Lanovaz, J. L., Schamhardt, H. C., & van Wessum, R. (1999). The effects of a rider's mass on ground reaction forces and fetlock kinematics at the trot. Equine Vet J Suppl, 30, 218–221.
Abstract: Ground reaction force (GRF) measurements are often normalised to body mass to facilitate inter-individual comparisons. The objective of this study was to explore the effect of a rider on the GRFs and fetlock joint kinematics of trotting horses. The subjects were 5 dressage-trained horses and 3 experienced dressage riders. Ground reaction force measurements and sagittal view videotapes were recorded as the horses trotted at the same velocity in hand (3.49 +/- 0.52 m/s) and with a rider (3.49 +/- 0.46 m/s). Data were time-normalised to stance duration. Ground reaction force measurements were expressed in absolute terms and normalised to the system mass (horse or horse plus rider). All the horses showed changes in the same direction when comparing the ridden condition with the in-hand condition. There was an increase in the absolute peak vertical GRFs of the fore- and hindlimbs with a rider. However, the mass-normalised peak vertical GRFs were lower for the ridden condition, with the peak occurring later in the forelimbs and earlier in the hindlimbs compared with the inhand condition. Maximal fetlock angle and its time of occurrence were similar for the 2 conditions, but the fore fetlock joint was more extended during the later part of the stance phase in ridden horses. The presence of a rider appeared to affect the GRFs and fetlock joint kinematics differently in the fore- and hindlimbs, and the ridden horse did not seem to be equivalent to a proportionately larger horse. This should be considered when normalising for body mass in studies comparing horses in hand and ridden horses.
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