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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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Hall, C., Rigg, V., Truswell, M., & Owen, H. (2012). Picture recognition of con-specifics and facial expression in the horse (Equus caballus). In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: The management of the domestic horse often requires them to be kept in isolation from con-specifics. Installing a picture of a horse (generally head and neck view) with a view to providing surrogate companionship has been shown to reduce the negative impact of this isolation. This study aimed firstly to compare the spontaneous response of horses (N=10) to a 2-D image of a horse’s face (FP) with their response to a comparable abstract 2-D image (AP). Secondly, the spontaneous response of horses (N=20) to a 2-D image of a horse’s face with the ears forward (PFP positive) was compared with the response to a 2-D image of a horse’s face with the ears back (NFP negative). The posters were A1 sized and displayed in the horse’s own stable. In study 1, one poster was displayed for 5 minutes and the horse’s behaviour video-recorded. This was removed and the second poster was displayed for 5 minutes and the behaviour video-recorded. FP was displayed first for 5 of the horses and AP displayed first for the other 5. The video footage was observed and the behaviour of the horses and number of times they touched the poster recorded. For the purpose of identifying the area of the poster that was touched by the horse it was divided into 4 equal quarters (TL, TR, BL, BR). In FP the nose of the horse in the 2-D image was located in BL, eyes and ears in TL, chest and lower neck in BR and upper neck in TR. In AP each area contained similar but unique abstract patterns of comparable colour to FP. Differences in behaviour were found according to which poster was displayed. FP was touched significantly more than AP (p=0.001) and was looked at more often (p=0.008). With FP the horses spent significantly longer with their ears forward (p=0.008) and licking and chewing (p=0.016). When the number of touches per poster area was compared (FP and AP) a significant difference was found in the number of times that BL (nose) and BR (chest/lower neck) were touched (p=0.011). Both areas were touched more frequently on FP, with BL being touched the most. In study 2 the same experimental protocol was used to compare responses to positive (PFP) or negative (NFP) 2-D images of a horse’s face (same horse in both PFP and NFP). Again, differences in behaviour were found in response to the two posters. PFP was touched significantly more than NFP (p=0.002) and on both posters the area BL (nose) was touched more frequently than the other areas (PFP: p=0.02, NFP: p=0.01). More ears back behaviour (p<0.001) and more ear locked on behaviour (p=0.008) was shown with NFP. The results of these studies indicate that horses can recognize 2-D images as con-specifics as well as responding to differences in facial expression. There is now the potential for further investigation into the importance of other visual cues in recognition and social interaction as well as the application of findings to enhance equine welfare.
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Hall, C. A., & Cassaday, H. J. (2006). An investigation into the effect of floor colour on the behaviour of the horse. Appl. Anim. Behav. Sci., 99(3-4), 301–314.
Abstract: Adverse reactions of the domestic horse to environmental stimuli can be problematic in training and management. Hesitation and alarm reactions to visual features of the ground can occur in both ridden work and when handling horses. To assess the effect of one visual feature (colour) on the behaviour of the domestic horse, the reactions of 16 riding horses to 8 different coloured mats were recorded. The effect of stimulus position on these reactions was assessed by presenting them in two different positions, either on the ground (where the horses had to walk over them) or against a wall (where the horses walked past them). Each colour/position combination was presented twice in order to assess the effect of previous experience. An alleyway was constructed to allow the horses to be tested unconstrained and freely walking throughout. The time taken to traverse the alleyway and the observed reaction to the colour was recorded. Significant differences in both measures were found in relation to the position of the colour and whether the subject had previous experience of that colour/position combination. The initial presentation of the colours on the ground produced the highest percentage of adverse reactions. Certain colours encountered for the first time on the ground (yellow, white, black and blue) were found to cause a greater number of adverse reactions than others (green, red, brown and grey) and an increase in time taken to traverse the alleyway. Although a significant difference in relation to colour was found in the behaviour observed during the second presentation on the ground, no difference was found in relation to the time taken to traverse the alleyway. No significant effect of colour was found when the coloured stimuli were presented against the wall. These findings have important implications for situations where the colour of flooring could be controlled in order to minimise adverse behavioural reactions, in particular during initial training.
Keywords: Horse; Colour aversion; Flooring; Vision
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Hall, C. A., Cassaday, H. J., & Derrington, A. M. (2003). The effect of stimulus height on visual discrimination in horses. J. Anim Sci., 81(7), 1715–1720.
Abstract: This study investigated the effect of stimulus height on the ability of horses to learn a simple visual discrimination task. Eight horses were trained to perform a two-choice, black/white discrimination with stimuli presented at one of two heights: ground level or at a height of 70 cm from the ground. The height at which the stimuli were presented was alternated from one session to the next. All trials within a single session were presented at the same height. The criterion for learning was four consecutive sessions of 70% correct responses. Performance was found to be better when stimuli were presented at ground level with respect to the number of trials taken to reach the criterion (P < 0.05), percentage of correct first choices (P < 0.01), and repeated errors made (P < 0.01). Thus, training horses to carry out tasks of visual discrimination could be enhanced by placing the stimuli on the ground. In addition, the results of the present study suggest that the visual appearance of ground surfaces is an important factor in both horse management and training.
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Hall, C. A., Cassaday, H. J., Vincent, C. J., & Derrington, A. M. (2006). Cone excitation ratios correlate with color discrimination performance in the horse (Equus caballus). J Comp Psychol, 120(4), 438–448.
Abstract: Six horses (Equus caballus) were trained to discriminate color from grays in a counterbalanced sequence in which lightness cues were irrelevant. Subsequently, the pretrained colors were presented in a different sequence. Two sets of novel colors paired with novel grays were also tested. Performance was just as good in these transfer tests. Once the horse had learned to select the chromatic from the achromatic stimulus, regardless of the specific color, they were immediately able to apply this rule to novel stimuli. In terms of the underlying visual mechanisms, the present study showed for the first time that the spectral sensitivity of horse cone photopigments, measured as cone excitation ratios, was correlated with color discrimination performance, measured as accuracy, repeated errors, and latency of approach.
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Hall, R. A., Broom, A. K., Smith, D. W., & Mackenzie, J. S. (2002). The ecology and epidemiology of Kunjin virus. Curr Top Microbiol Immunol, 267, 253–269.
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Hampson, B. A., Morton, J. M., Mills, P. C., Trotter, M. G., Lamb, D. W., & Pollitt, C. C. (2010). Monitoring distances travelled by horses using GPS tracking collars. Aust. Vet. J., 88(5), 176–181.
Abstract: Objective The aims of this work were to (1) develop a low-cost equine movement tracking collar based on readily available components, (2) conduct preliminary studies assessing the effects of both paddock size and internal fence design on the movements of domestic horses, with and without foals at foot, and (3) describe distances moved by mares and their foals. Additional monitoring of free-ranging feral horses was conducted to allow preliminary comparisons with the movement of confined domestic horses. Procedures A lightweight global positioning system (GPS) data logger modified from a personal/vehicle tracker and mounted on a collar was used to monitor the movement of domestic horses in a range of paddock sizes and internal fence designs for 6.5-day periods. Results In the paddocks used (0.8-16 ha), groups of domestic horses exhibited a logarithmic response in mean daily distance travelled as a function of increasing paddock size, tending asymptotically towards approximately 7.5 km/day. The distance moved by newborn foals was similar to their dams, with total distance travelled also dependent on paddock size. Without altering available paddock area, paddock design, with the exception of a spiral design, did not significantly affect mean daily distance travelled. Feral horses (17.9 km/day) travelled substantially greater mean daily distances than domestic horses (7.2 km/day in 16-ha paddock), even when allowing for larger paddock size. Conclusions Horses kept in stables or small yards and paddocks are quite sedentary in comparison with their feral relatives. For a given paddock area, most designs did not significantly affect mean daily distance travelled.
Keywords: behaviour; feral horses; foals; global positioning system (GPS); horses; paddock design
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Hanggi, E. B. (2003). Discrimination learning based on relative size concepts in horses (Equus caballus). Appl. Anim. Behav. Sci., 83(3), 201–213.
Abstract: This study explored whether or not horses (Equus caballus) could respond to stimuli using a concept based on relative size. In Experiment 1, after learning to respond to the larger of the two stimuli for six sets of two-dimensional (2D) training exemplars, one horse was tested for size transposition that used novel larger and smaller stimuli as well as three-dimensional (3D) objects (5 two-dimensional sets and 5 three-dimensional sets with large, medium, small, and tiny sizes). The horse correctly chose (significantly above chance) the larger of two stimuli regardless of novelty or dimension or combination. In Experiment 2, two additional horses were tested using a subset of the stimuli from Experiment 1. One horse was required to select the larger stimulus--as in Experiment 1--and the other the smaller stimulus. After learning the task, both horses responded correctly to new stimuli and showed size transposition. These results suggest that at least some horses are capable of solving problems based on relative size concepts. Moreover, they are able to generalize across situations that vary from flat, black shapes to objects of different materials and colors including balls, flower pots, and PVC connectors. These findings support earlier research that showed that horses could categorize certain stimuli, and provide new evidence that they are capable of using some form of concept for problem solving. Understanding that horses have more advanced learning abilities than was previously believed should help improve training methods and management.
Keywords: Horse; Concept; Size transposition; Generalization; Learning; Training
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Hanggi, E. B., & Ingersoll, J. F. (2009). Long-term memory for categories and concepts in horses (Equus caballus). Anim. Cogn., 13(3), 451–462.
Abstract: Three horses (Equus caballus) with a history of performing cognitive tasks including discrimination learning, categorization, and concept use were tested to evaluate their long-term memory (LTM) in three experiments. In addition, use of LCD multi-displays for stimulus presentation was incorporated into cognition testing protocol for the first time with horses. Experiment 1 tested LTM for discrimination learning that originally occurred 6 years earlier. Five sets of stimuli were used and the two horses tested showed no decrement in performance on four of the sets; however, both horses did score below chance on one set. Experiment 2 examined long-term categorization recall 10 years after horses had demonstrated the ability to make stimulus selections based on shared characteristics within a given category. The horse tested for LTM after the decade-long interval immediately and consistently applied the previously learned categorization rule to not only familiar but also novel sets of stimuli. Experiment 3 tested another horse for LTM for a relative size concept. This horse had originally demonstrated concept rule use in order to select stimuli based on their relative size to one another. More than 7 years later and without further training, this horse reliably applied the previously established size concept to both familiar and novel sets of stimuli. These findings are the first reports of long-term categorical and conceptual memory in horses and are consistent with observations of domestic and wild horses, which indicate that behavioral and ecological events may be remembered for long periods of time. These studies also demonstrate the adaptive nature of horses with regard to their ability to generalize over several different testing conditions.
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Hanggi, E. B., & Ingersoll, J. F. (2009). Stimulus discrimination by horses under scotopic conditions. Behav. Process., 82(1), 45–50.
Abstract: Scotopic vision in horses (Equus caballus) was investigated using behavioral measurements for the first time. Four horses were tested for the ability to make simple visual discriminations of geometric figures (circles and triangles) under various brightness levels within an enclosed building. Measurements of brightness ranging from 10.37 to 24.12 magnitudes per square arcsecond (mag/arcsec2; in candelas per square meter--7.70 to 2.43E-05 cd/m2) were taken using a Sky Quality Meter. These values approximated outdoor conditions ranging from twilight in open country to a dark moonless night in dense forest. The horses were able to solve the discrimination problems in all brightness settings up to 23.77 mag/arcsec2 (3.35E-05 cd/m2). Moreover, they easily navigated their way around obstacles located within the testing area in extremely dim light (>23.50 mag/arcsec2; 4.30E-05 cd/m2), which were in conditions too dark for the human experimenters to see. These findings support physiological data that reveal a rod-dominated visual system as well as observations of equine activity at night.
Keywords: Discrimination learning; Equine; Horse; Night vision; Scotopic vision
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