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Smith, S., & Goldman, L. (1999). Color discrimination in horses. Appl. Anim. Behav. Sci., 62(1), 13–25.
Abstract: Four Arabian horses and one Thoroughbred were presented with a series of two-choice color vs. gray discrimination problems. Testing was done in a stall containing a wall with two translucent panels that were illuminated from behind by light projected through color or gray filters to provide the discriminative stimuli. Horses first learned to push one of the panels in order to receive the food reward behind the positive stimulus in an achromatic light-dark discrimination task, and were then tested on their ability to discriminate between gray and four individual colors: red (617 nm), yellow (581 nm), green (538 nm), and blue (470 nm). The criterion for learning was set at 85% correct responses, and final testing for all color vs. gray discriminations involved grays of varying intensities, making brightness an irrelevant cue. Three subjects were tested with all four colors. Two of those subjects successfully reached the criterion for learning on all four color vs. gray discriminations, while the third reached criterion with red and blue, but performed at chance levels for yellow and green. A fourth horse was only tested with green and yellow, and a fifth only with blue, and both of those horses successfully reached criterion on the discriminations they attempted. With the exception of the one subject's poor performance with yellow and green, there was no significant difference between horses on any of the discrimination tasks, and no significant difference in their performance with different colors. The results suggest that horses have color vision that is at least dichromatic, although partial color-blindness may occur in some individuals.
Keywords: Horses; Vision; Color; Discrimination; Behavior
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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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Baumgartner, M., Boisson, T., Erhard, M. H., & Zeitler-Feicht, M. H. (2020). Common Feeding Practices Pose A Risk to the Welfare of Horses When Kept on Non-Edible Bedding. Animals, 10, 441.
Abstract: During the evolution of the horse, an extended period of feed intake, spread over the entire 24-h period, determined the horses� behaviour and physiology. Horses will not interrupt their feed intake for more than 4 h, if they have a choice. The aim of the present study was to investigate in what way restrictive feeding practices (non ad libitum) affect the horses� natural feed intake behaviour. We observed the feed intake behaviour of 104 horses on edible (n = 30) and non-edible bedding (n = 74) on ten different farms. We assessed the duration of the forced nocturnal feed intake interruption of horses housed on shavings when no additional roughage was available. Furthermore, we comparatively examined the feed intake behaviour of horses housed on edible versus non-edible bedding. The daily restrictive feeding of roughage (2 times a day: n = 8; 3 times a day: n = 2), as it is common in individual housing systems, resulted in a nocturnal feed intake interruption of more than 4 hours for the majority (74.32%, 55/74) of the horses on shavings (8:50 ± 1:25 h, median: 8:45 h, minimum: 6:45 h, maximum: 13:23 h). In comparison to horses on straw, horses on shavings paused their feed intake less frequently and at a later latency. Furthermore, they spent less time on consuming the evening meal than horses on straw. Our results of the comparison of the feed-intake behaviour of horses on edible and non-edible bedding show that the horses� ethological feeding needs are not satisfied on non-edible bedding. If the horses accelerate their feed intake (also defined as �rebound effect�), this might indicate that the horses� welfare is compromised. We conclude that in addition to the body condition score, the longest duration of feed intake interruption (usually in the night) is an important welfare indicator of horses that have limited access to roughage.
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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.
Keywords: Animals; Artiodactyla/*anatomy & histology/physiology; Cattle; Connective Tissue/physiology/ultrastructure; Feeding Behavior/physiology; Goats/anatomy & histology/physiology; Horses/anatomy & histology/physiology; Mastication/physiology; Microscopy, Electron, Scanning; Perissodactyla/*anatomy & histology/physiology; Tongue/physiology/*ultrastructure
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Peeters, M., Sulon, J., Beckers, J. - F., Ledoux, D., & Vandenheede, M. (2011). Comparison between blood serum and salivary cortisol concentrations in horses using an adrenocorticotropic hormone challenge. Equine Veterinary Journal, 43(4), 487–493.
Abstract: Reasons for performing study: In horses, serum cortisol concentration is considered to provide an indirect measurement of stress. However, it includes both free and bound fractions. The sampling method is also invasive and often stressful. This is not the case for salivary cortisol, which is collected using a more welfare-friendly method and represents a part of the free cortisol fraction, which is the biologically active form. Objectives: To compare salivary and serum cortisol assays in horses, in a wide range of concentrations, using an adrenocorticotropic hormone (ACTH) stimulation test, in order to validate salivary cortisol for stress assessment in horse. Methods: In 5 horses, blood samples were drawn using an i.v. catheter. Saliva samples were taken using swabs. Cortisol was assayed by radioimmunoassay. All data were treated with a regression method, which pools and analyses data from multiple subjects for linear analysis. Results: Mean ± s.d. cortisol concentrations measured at rest were 188.81 ± 51.46 nmol/l in serum and 1.19 ± 0.54 nmol/l in saliva. They started increasing immediately after ACTH injection and peaks were reached after 96 ± 16.7 min in serum (356.98 ± 55.29 nmol/l) and after 124 ± 8.9 min in saliva (21.79 ± 7.74 nmol/l, P<0.05). Discharge percentages were also different (225% in serum and 2150% in saliva, P<0.05). Correlation between serum and salivary cortisol concentrations showed an adjusted r2= 0.80 (P<0.001). The strong link between serum and salivary cortisol concentrations was also estimated by a regression analysis. Conclusions: The reliability of both RIAs and regression found between serum and salivary cortisol concentrations permits the validation of saliva-sampling as a noninvasive technique for cortisol level assessment in horses.
Keywords: horse; cortisol; ACTH challenge; saliva; stress
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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.
Keywords: Animals; Anopheles/*parasitology; *Behavior, Animal; Cattle; Desert Climate; Horses/parasitology; Humans; Insect Vectors/*parasitology; Longitudinal Studies; Malaria/*transmission; Malaria, Falciparum/transmission; Periodicity; Plasmodium malariae/isolation & purification; Protozoan Proteins/analysis; Rain; Seasons; Senegal; Sheep/parasitology; Species Specificity
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Bonin, S. J., Clayton, H. M., Lanovaz, J. L., & Johnston, T. (2007). Comparison of mandibular motion in horses chewing hay and pellets. Equine Vet. J., 39(3), 258–262.
Abstract: Summary Reasons for performing study: Previous studies have suggested that temporomandibular joint (TMJ) kinematics depend on the type of food being masticated, but accurate measurements of TMJ motion in horses chewing different feeds have not been published. Hypothesis: The temporomandibular joint has a larger range of motion when horses chew hay compared to pellets. Methods: An optical motion capture system was used to track skin markers on the skull and mandible of 7 horses as they chewed hay and pellets. A virtual marker was created on the midline between the mandibles at the level of the 4th premolar teeth to represent the overall motion of the mandible relative to the skull during the chewing cycle. Results: Frequency of the chewing cycles was lower for hay than for pellets. Excursions of the virtual mandibular marker were significantly larger in all 3 directions when chewing hay compared to pellets. The mean velocity of the virtual mandibular marker during the chewing cycle was the same when chewing the 2 feeds. Conclusions: The range of mediolateral displacement of the mandible was sufficient to give full occlusal contact of the upper and lower dental arcades when chewing hay but not when chewing pellets. Potential relevance: These findings support the suggestion that horses receiving a diet high in concentrate feeds may require more frequent dental prophylactic examinations and treatments to avoid the development of dental irregularities associated with smaller mandibular excursions during chewing.
Keywords: horse; temporomandibular joint; mastication; kinematics
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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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