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Hagen, S. J., & Eaton, W. A. (2000). Two-state expansion and collapse of a polypeptide. J Mol Biol, 301(4), 1019–1027.
Abstract: The initial phase of folding for many proteins is presumed to be the collapse of the polypeptide chain from expanded to compact, but still denatured, conformations. Theory and simulations suggest that this collapse may be a two-state transition, characterized by barrier-crossing kinetics, while the collapse of homopolymers is continuous and multi-phasic. We have used a laser temperature-jump with fluorescence spectroscopy to measure the complete time-course of the collapse of denatured cytochrome c with nanosecond time resolution. We find the process to be exponential in time and thermally activated, with an apparent activation energy approximately 9 k(B)T (after correction for solvent viscosity). These results indicate that polypeptide collapse is kinetically a two-state transition. Because of the observed free energy barrier, the time scale of polypeptide collapse is dramatically slower than is predicted by Langevin models for homopolymer collapse.
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Dunbar, R. I. M. (1998). The social brain hypothesis. Evol. Anthropol., 6(5), 178–190.
Abstract: Conventional wisdom over the past 160 years in the cognitive and neurosciences has assumed that brains evolved to process factual information about the world. Most attention has therefore been focused on such features as pattern recognition, color vision, and speech perception. By extension, it was assumed that brains evolved to deal with essentially ecological problem-solving tasks. © 1998 Wiley-Liss, Inc.
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Tyler, S. J. (1972). The behaviour and social organisation of the new Forest ponies. Anim. Behav. Monogr., 5(2), 85–196.
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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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Klingel H,. (1977). Observations on Social Organization and Behaviour of African and Asiatic Wild Asses (Equus africanus and E. hemionus). Z. Tierpsychol., 44, 323–331.
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van Heel, M. C. V., Kroekenstoel, A. M., van Dierendonck, M. C., van Weeren, P. R., & Back, W. (2006). Uneven feet in a foal may develop as a consequence of lateral grazing behaviour induced by conformational traits. Equine. Vet. J., 38(7), 646–651.
Abstract: REASONS FOR PERFORMING STUDY: Conformational traits are important in breeding, since they may be indicative for performance ability and susceptibility to injuries. OBJECTIVES: To study whether certain desired conformational traits of foals are related to lateralised behaviour while foraging and to the development of uneven feet. METHODS: Twenty-four Warmblood foals, born and raised at the same location, were studied for a year. Foraging behaviour was observed by means of weekly 10 min scan-sampling for 8 h. A preference test (PT) was developed to serve as a standardised tool to determine laterality. The foals were evaluated at age 3, 15, 27 and 55 weeks. The PT and distal limb conformation were used to study the relation between overall body conformation, laterality and the development of uneven feet. Pressure measurements were used to determine the loading patterns under the feet. RESULTS: About 50% of the foals developed a significant preference to protract the same limb systematically while grazing, which resulted in uneven feet and subsequently uneven loading patterns. Foals with relatively long limbs and small heads were predisposed to develop laterality and, consequently unevenness. CONCLUSIONS: Conformational traits may stimulate the development of laterality and therefore indirectly cause uneven feet.
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Rhodin, M., Johnston, C., Holm, K. R., Wennerstrand, J., & Drevemo, S. (2005). The influence of head and neck position on kinematics of the back in riding horses at the walk and trot. Equine Vet J, 37(1), 7–11.
Abstract: REASONS FOR PERFORMING STUDY: A common opinion among riders and in the literature is that the positioning of the head and neck influences the back of the horse, but this has not yet been measured objectively. OBJECTIVES: To evaluate the effect of head and neck position on the kinematics of the back in riding horses. METHODS: Eight Warmblood riding horses in regular work were studied on a treadmill at walk and trot with the head and neck in 3 different predetermined positions achieved by side reins attached to the bit and to an anticast roller. The 3-dimensional movement of the thoracolumbar spine was measured from the position of skin-fixed markers recorded by infrared videocameras. RESULTS: Head and neck position influenced the movements of the back, especially at the walk. When the head was fixed in a high position at the walk, the flexion-extension movement and lateral bending of the lumbar back, as well as the axial rotation, were significantly reduced when compared to movements with the head free or in a low position. At walk, head and neck position also significantly influenced stride length, which was shortest with the head in a high position. At trot, the stride length was independent of head position. CONCLUSIONS: Restricting and restraining the position and movement of the head and neck alters the movement of the back and stride characteristics. With the head and neck in a high position stride length and flexion and extension of the caudal back were significantly reduced. POTENTIAL RELEVANCE: Use of side reins in training and rehabilitation programmes should be used with an understanding of the possible effects on the horse's back.
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Gomez Alvarez, C. B., Rhodin, M., Bobber, M. F., Meyer, H., Weishaupt, M. A., Johnston, C., et al. (2006). The effect of head and neck position on the thoracolumbar kinematics in the unridden horse. Equine Vet J Suppl, (36), 445–451.
Abstract: REASONS FOR PERFORMING STUDY: In many equestrian activities a specific position of head and/or neck is required that is dissimilar to the natural position. There is controversy about the effects of these positions on locomotion pattern, but few quantitative data are available. OBJECTIVES: To quantify the effects of 5 different head and neck positions on thoracolumbar kinematics of the horse. METHODS: Kinematics of 7 high level dressage horses were measured walking and trotting on an instrumented treadmill with the head and neck in the following positions: HNP2 = neck raised, bridge of the nose in front of the vertical; HNP3 = as HNP2 with bridge of the nose behind the vertical; HNP4 = head and neck lowered, nose behind the vertical; HNP5 = head and neck in extreme high position; HNP6 = head and neck forward and downward. HNP1 was a speed-matched control (head and neck unrestrained). RESULTS: The head and neck positions affected only the flexion-extension motion. The positions in which the neck was extended (HNP2, 3, 5) increased extension in the anterior thoracic region, but increased flexion in the posterior thoracic and lumbar region. For HNP4 the pattern was the opposite. Positions 2, 3 and 5 reduced the flexion-extension range of motion (ROM) while HNP4 increased it. HNP5 was the only position that negatively affected intravertebral pattern symmetry and reduced hindlimb protraction. The stride length was significantly reduced at walk in positions 2, 3, 4 and 5. CONCLUSIONS: There is a significant influence of head/neck position on back kinematics. Elevated head and neck induce extension in the thoracic region and flexion in the lumbar region; besides reducing the sagittal range of motion. Lowered head and neck produces the opposite. A very high position of the head and neck seems to disturb normal kinematics. POTENTIAL RELEVANCE: This study provides quantitative data on the effect of head/neck positions on thoracolumbar motion and may help in discussions on the ethical acceptability of some training methods.
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Weishaupt, M. A., Wiestner, T., von Peinen, K., Waldern, N., Roepstorff, L., van Weeren, R., et al. (2006). Effect of head and neck position on vertical ground reaction forces and interlimb coordination in the dressage horse ridden at walk and trot on a treadmill. Equine Vet J Suppl, (36), 387–392.
Abstract: REASONS FOR PERFORMING STUDY: Little is known in quantitative terms about the influence of different head-neck positions (HNPs) on the loading pattern of the locomotor apparatus. Therefore it is difficult to predict whether a specific riding technique is beneficial for the horse or if it may increase the risk for injury. OBJECTIVE: To improve the understanding of forelimb-hindlimb balance and its underlying temporal changes in relation to different head and neck positions. METHODS: Vertical ground reaction force and time parameters of each limb were measured in 7 high level dressage horses while being ridden at walk and trot on an instrumented treadmill in 6 predetermined HNPs: HNP1 – free, unrestrained with loose reins; HNP2 – neck raised, bridge of the nose in front of the vertical; HNP3 – neck raised, bridge of the nose behind the vertical; HNP4 – neck lowered and flexed, bridge of the nose considerably behind the vertical; HNP5 – neck extremely elevated and bridge of the nose considerably in front of the vertical; HNP6 – neck and head extended forward and downward. Positions were judged by a qualified dressage judge. HNPs were assessed by comparing the data to a velocity-matched reference HNP (HNP2). Differences were tested using paired t test or Wilcoxon signed rank test (P<0.05). RESULTS: At the walk, stride duration and overreach distance increased in HNP1, but decreased in HNP3 and HNP5. Stride impulse was shifted to the forehand in HNP1 and HNP6, but shifted to the hindquarters in HNP5. At the trot, stride duration increased in HNP4 and HNP5. Overreach distance was shorter in HNP4. Stride impulse shifted to the hindquarters in HNP5. In HNP1 peak forces decreased in the forelimbs; in HNP5 peak forces increased in fore- and hindlimbs. CONCLUSIONS: HNP5 had the biggest impact on limb timing and load distribution and behaved inversely to HNP1 and HNP6. Shortening of forelimb stance duration in HNP5 increased peak forces although the percentage of stride impulse carried by the forelimbs decreased. POTENTIAL RELEVANCE: An extremely high HNP affects functionality much more than an extremely low neck.
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Krueger, K. (Ed.). (2008). Proceedings of the International Equine Science Meeting 2008. Wald: Xenophon Verlag.
Abstract: Target group: Biologists, Psychologists, Veterinarians and Professionals
Meeting target: Because the last international meeting on Equine Science took place a couple years ago, there is an urgent need for equine scientists to exchange scientific knowledge, coordinate research provide knowledge for practical application, and discus research results among themselves and with professionals who work with horses. Additionally, dialog concerning the coordination of the study “Equitation Science” in Europe is urgently needed. Coordination and cooperation shall arise from the meeting, enrich the research, and advance the application of scientific knowledge for the horses` welfare.
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