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Hanggi, E. B., Ingersoll, J. F., & Waggoner, T. L. (2007). Color vision in horses (Equus caballus): deficiencies identified using a pseudoisochromatic plate test. J. Comp. Psychol., 121(1), 65–72.
Abstract: In the past, equine color vision was tested with stimuli composed either of painted cards or photographic slides or through physiological testing using electroretinogram flicker photometry. Some studies produced similar results, but others did not, demonstrating that there was not yet a definitive answer regarding color vision in horses (Equus caballus). In this study, a pseudoisochromatic plate test--which is highly effective in testing color vision both in small children and in adult humans--was used for the first time on a nonhuman animal. Stimuli consisted of different colored dotted circles set against backgrounds of varying dots. The coloration of the circles corresponded to the visual capabilities of different types of color deficiencies (anomalous trichromacy and dichromacy). Four horses were tested on a 2-choice discrimination task. All horses successfully reached criterion for gray circles and demonstration circles. None of the horses were able to discriminate the protan-deutan plate or the individual protan or deutan plates. However, all were able to discriminate the tritan plate. The results suggest that horses are dichromats with color vision capabilities similar to those of humans with red-green color deficiencies.
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Mills, D. S. (2007). Comments about the importance of behaviour to equine clinicians. Equine Vet J, 39(1), 95.
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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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Kobayashi, K., Jackowiak, H., Frackowiak, H., Yoshimura, K., Kumakura, M., & Kobayashi, K. (2005). Comparative morphological study on the tongue and lingual papillae of horses (Perissodactyla) and selected ruminantia (Artiodactyla). Ital J Anat Embryol, 110(2 Suppl 1), 55–63.
Abstract: A common characteristic of horses, Rocky Mountain goats, and cattle is that they all have a well developed lingual prominence on the dorsal surface of the posterior area of the tongue. Foliate papillae were found in the horse studied but not in the goat or in cattle. The horse filiform papillae had a long and slender external form with a thin and slender CTC, while in the goat and cattle the external form consisted of a large thick main process and the CTC consisted of a bundle of numerous rod-shaped protrusions. The special papilla found on the lingual prominence resembled larger filiform-like papillae in the horses; however, in the goat and cattle it was a very thick and large tongue like papillae. The horses had two large vallate papillae, while the goat and cattle had 15 or more vallate papillae at the posterior area of the lingual prominence. This suggests that the fine structure of horse tongues may display a more primitive pattern than that present in goats and cattle.
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Lemasson, J. J., Fontenille, D., Lochouarn, L., Dia, I., Simard, F., Ba, K., et al. (1997). Comparison of behavior and vector efficiency of Anopheles gambiae and An. arabiensis (Diptera:Culicidae) in Barkedji, a Sahelian area of Senegal. J Med Entomol, 34(4), 396–403.
Abstract: The ecology, population dynamics, and malaria vector efficiency of Anopheles gambiae and An. arabiensis were studied for 2 yr in a Sahelian village of Senegal. Anophelines were captured at human bait and resting indoors by pyrethrum spray. Mosquitoes belonging to the An. gambiae complex were identified by polymerase chain reaction. Of 26,973 females, An. arabiensis represented 79% of the mosquitoes captured and remained in the study area longer than An. gambiae after the rains terminated. There were no differences in nocturnal biting cycles or endophagous rates between An. gambiae and An. arabiensis. Based on an enzyme-linked immunosorbent assay test of bloodmeals, the anthropophilic rate of these 2 vectors were both approximately 60%, when comparisons were made during the same period. Overall, 18% of the resting females had patent mixed bloodmeals, mainly human-bovine. The parity rates of An. gambiae and An. arabiensis varied temporally. Despite similar behavior, the Plasmodium falciparum circumsporozoite protein (CSP) rates were different between An. gambiae (4.1%) and An. arabiensis (1.3%). P. malariae and P. ovale only represented 4% of the total Plasmodium identified in mosquitoes. Transmission was seasonal, occurring mainly during 4 mo. The CSP entomological inoculation rates were 128 bites per human per year for the 1st yr and 100 for the 2nd yr. Because of the combination of a high human biting rate and a low CSP rate, An. arabiensis accounted for 63% of transmission. Possible origin of differences in CSP rate between An. gambiae and An. arabiensis is discussed in relation to the parity rate, blood feeding frequency, and the hypothesis of genetic factors.
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Spadavecchia, C., Arendt-Nielsen, L., Andersen, O. K., Spadavecchia, L., Doherr, M., & Schatzmann, U. (2003). Comparison of nociceptive withdrawal reflexes and recruitment curves between the forelimbs and hind limbs in conscious horses. Am J Vet Res, 64(6), 700–707.
Abstract: OBJECTIVE: To compare nociceptive withdrawal reflexes (NWRs) evoked from the distal aspect of the left forelimb and hind limb in conscious standing horses and to investigate NWR recruitment for graded electrical stimulation intensities. ANIMALS: 20 adult horses. PROCEDURE: Surface electromyographic (EMG) activity evoked by transcutaneous electrical stimulation of the digital palmar (or plantar) nerve was recorded from the common digital extensor and cranial tibial muscles. Stimuli consisted of 25-millisecond train-of-5 constant current pulses. Current intensity was gradually increased until NWR threshold intensity was reached. The EMG signal was analyzed for quantification of the NWR. Behavioral responses accompanying the reflex were scored (scale, 0 to 5). The NWR recruitment curves were determined at 0.9, 1.1, 1.2, and 1.3 times the NWR threshold intensity. RESULTS: The NWR threshold was significantly higher for the hind limb (median value, 6.6 mA; range, 3 to 10 mA) than the forelimb (median, 3 mA; range, 1.7 to 5.5 mA). The NWR of the hind limb had a significantly longer latency (median, 122.8 milliseconds; range, 106 to 172 milliseconds), compared with the forelimb (median, 98 milliseconds; range, 86 to 137 milliseconds), and it was associated with significantly stronger behavioral reactions. Gradual increase of NWR amplitude was evident at increasing stimulation intensities and supported by the behavioral observations. CONCLUSIONS AND CLINICAL RELEVANCE: We documented NWRs evoked from the forelimb and hind limb and their recruitment with stimuli of increasing intensity in horses. These results provide a basis for use of NWRs in studies on nociceptive modulation in horses.
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Clayton, H. M. (1995). Comparison of the stride kinematics of the collected, medium, and extended walks in horses. Am J Vet Res, 56(7), 849–852.
Abstract: Six horses, highly trained for dressage competition, were used to study the stride kinematics of the walk, and to compare the kinematics of the collected, medium, and extended walks. Horses were filmed in a sagittal plane at a rate of 150 frames/s; temporal, linear, and angular data were extracted from the films. Results of ANOVA and Duncan's multiple range test indicated that the speed of the collected walk (1.37 m/s) was significantly (P < 0.01) slower than that of the medium (1.73 m/s) and extended (1.82 m/s) walks, values for which were not significantly different from each other. The increase in speed was associated with a significant increase in stride length, from 157 cm in the collected walk to 193 cm in the extended walk. This was a result of an increase in the over-tracking distance, whereas there was no significant difference in the distance between lateral placements of the limbs. Stride duration decreased (P < 0.01) from the collected walk (1,159 ms) to the extended walk (1,064 ms). Angles of the metacarpal and metatarsal segments, measured on the palmar/ plantar aspect, were higher at impact and lower at lift off in the collected than in the extended walk (P < 0.01). This indicated greater range of angular motion of this segment during the stance phase in the extended walk. Only 1 of the 6 horses had a regular 4-beat rhythm of the footfalls, with equal time elapsing between the lateral and diagonal footfalls.
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Clayton, H. M. (1994). Comparison of the stride kinematics of the collected, working, medium and extended trot in horses. Equine Vet J, 26(3), 230–234.
Abstract: Highly-trained dressage horses were studied to test the hypothesis that stride length is altered independently of stride duration in the transitions between the collected, working, medium and extended trot. Six well-trained dressage horses were filmed at a frame rate of 150 frames/s performing the collected, working, medium and extended trots in a sand arena. Temporal, linear and angular data were extracted from the films, with 4 strides being analysed for each horse and gait type. There were no significant asymmetries between the left and rights limbs or diagonals when data from the whole group were pooled, but 3 horses showed asymmetries in one or more variables (P < 0.01). Analysis of variance and post-hoc tests indicated that the speed increased significantly (P < 0.01) from the collected (3.20 m/s) to the working (3.61 m/s) to the medium (4.47 m/s) to the extended (4.93 m/s) trot. The increases in speed were associated with a significant increase in stride length from 250 cm in the collected trot, to 273 cm in the working trot, 326 cm in the medium trot and 355 cm in the extended trot (P < 0.01). The lengthening of the stride was a result of increases between each gait type in the over-reach distance, whereas the diagonal distance was significantly longer in the extended than the collected trot only (P < 0.01). The stride duration tended to decrease as speed increased, and the difference became significant between the collected and extended trots (P < 0.01).
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Burns, T. E., & Clayton, H. M. (1997). Comparison of the temporal kinematics of the canter pirouette and collected canter. Equine Vet J Suppl, (23), 58–61.
Abstract: The objectives were to compare the temporal characteristics of canter pirouette strides with collected canter strides in elite dressage horses, and to determine whether the stride kinematics of the canter pirouettes fulfilled the requirements specified in the Federation Equestre Internationale Rules for Dressage Events. Eleven horses were videotaped (60 fields/s) during the individual medal competition at the 1992 Olympic Games. Temporal variables were extracted from the videotapes using standard methods. Two strides were analysed on each of the left and right leads and these were pooled to give mean values for the collected canter and the pirouettes. The pirouettes were completed in 4-9 strides, (mean of 6.4). In the collected canter strides, mean duration of the suspension was 0.013 s. There was no suspension in any of the pirouette strides, instead the stance phases of the leading forelimb and trailing hindlimb overlapped by a mean of 0.163 s. In 9 horses the trailing forelimb contacted the ground before the diagonal leading hindlimb in the collected canter, whereas in the pirouettes the leading hindlimb always made contact before the trailing forelimb (mean dissociation 0.164 s), giving the strides a distinct 4 beat rhythm. Due to increases in advanced placement between the diagonal limb pair and between the 2 forelimbs, the stride duration was longer in the pirouette (0.879 s) than the collected canter (0.629 s). It is concluded that the canter pirouette strides did not maintain the rhythm and timing of the the collected canter strides in any of the 11 horses.
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Galloux, P., & Barrey, E. (1997). Components of the total kinetic moment in jumping horses. Equine Vet J Suppl, (23), 41–44.
Abstract: Thirty horses were filmed with a panning camera operating at 50 frames/s as they jumped over a 1.20 x 1.20 m fence. The markers of 9 joints on the horse and 7 joints on the rider were tracked in 2D with the TrackEye system. The centre of gravity and moment of inertia of each segment were calculated using a geometric algorithm and a cylindric model, respectively. The kinetic moment of each part of the horse was calculated after filtering, and resampling of data. This method showed the relative contribution of each body segment to the body overall rotation during the take-off, jump and landing phases. It was found that the trunk, hindlimbs and head-neck had the greatest influence. The coordination between the motion of the body segments allowed the horse to control its angular speed of rotation over the fence. This remained nearly constant during the airborne phase (120 +/- 5 degrees/s). During the airborne phase, the kinetic moment was constant because its value was equal to the moment of the external forces (722 +/- 125 kg x m2/s).
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