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Bobbert, M. F., & Santamaria, S. (2005). Contribution of the forelimbs and hindlimbs of the horse to mechanical energy changes in jumping. J Exp Biol, 208(2), 249–260.
Abstract: The purpose of the present study was to gain more insight into the contribution of the forelimbs and hindlimbs of the horse to energy changes during the push-off for a jump. For this purpose, we collected kinematic data at 240 Hz from 23 5-year-old Warmbloods (average mass: 595 kg) performing free jumps over a 1.15 m high fence. From these data, we calculated the changes in mechanical energy and the changes in limb length and joint angles. The force carried by the forelimbs and the amount of energy stored was estimated from the distance between elbow and hoof, assuming that this part of the leg behaved as a linear spring. During the forelimb push, the total energy first decreased by 3.2 J kg(-1) and then increased again by 4.2 J kg(-1) to the end of the forelimb push. At the end of the forelimb push, the kinetic energy due to horizontal velocity of the centre of mass was 1.6 J kg(-1) less than at the start, while the effective energy (energy contributing to jump height) was 2.3 J kg(-1) greater. It was investigated to what extent these changes could involve passive spring-like behaviour of the forelimbs. The amount of energy stored and re-utilized in the distal tendons during the forelimb push was estimated to be on average 0.4 J kg(-1) in the trailing forelimb and 0.23 J kg(-1) in the leading forelimb. This means that a considerable amount of energy was first dissipated and subsequently regenerated by muscles, with triceps brachii probably being the most important contributor. During the hindlimb push, the muscles of the leg were primarily producing energy. The total increase in energy was 2.5 J kg(-1) and the peak power output amounted to 71 W kg(-1).
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Powers, P., & Harrison, A. (2002). Effects of the rider on the linear kinematics of jumping horses. Sports Biomech, 1(2), 135–146.
Abstract: This study examined the effects of the rider on the linear projectile kinematics of show-jumping horses. SVHS video recordings (50 Hz) of eight horses jumping a vertical fence 1 m high were used for the study. Horses jumped the fence under two conditions: loose (no rider or tack) and ridden. Recordings were digitised using Peak Motus. After digitising the sequences, each rider's digitised data were removed from the ridden horse data so that three conditions were examined: loose, ridden (including the rider's data) and riderless (rider's data removed). Repeated measures ANOVA revealed significant differences between ridden and loose conditions for CG height at take-off (p < 0.001), CG distance to the fence at take-off (p = 0.001), maximum CG during the suspension phase (p < 0.001), CG position over the centre of the fence (p < 0.001), CG height at landing (p < 0.001), and vertical velocity at take-off (p < 0.001). The results indicated that the rider's effect on jumping horses was primarily due to behavioural changes in the horses motion (resulting from the rider's instruction), rather than inertial effects (due to the positioning of the rider on the horse). These findings have implications for the coaching of riders and horses.
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Ratzlaff, M. H., Wilson, P. D., Hyde, M. L., Balch, O. K., & Grant, B. D. (1993). Relationship between locomotor forces, hoof position and joint motion during the support phase of the stride of galloping horses. Acta Anat (Basel), 146(2-3), 200–204.
Abstract: Three methods were used simultaneously to determine the relationships between the vertical forces exerted on the hooves and the positions of the limbs and hooves at the times of peak vertical forces from 2 horses galloping on a track straightaway. Vertical forces were recorded from an instrumented shoe, fetlock joint motion was measured with an electrogoniometer and the angles of the carpus, fetlock and hoof were determined from slow-motion films. At hoof contact, the mean angles of the carpus and fetlock were 181-182 degrees and 199-206 degrees, respectively. Peak vertical forces on the heel occurred at or near maximum extension of the carpal and fetlock joints. Peak forces on the toe occurred during flexion of the fetlock joint and at mean hoof angles of 28-31 degrees from the horizontal. The mean angles of the hoof from the horizontal at the time of heel contact were 6-7 degrees. Hoof lift occurred at mean carpal angles of 173-174 degrees and mean fetlock angles of 199-200 degrees.
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Visser, E. K., van Reenen, C. G., Schilder, M. B. H., Barneveld, A., & Blokhuis, H. J. (2003). Learning performances in young horses using two different learning tests. Appl. Anim. Behav. Sci., 80, 311–326.
Abstract: To achieve optimal performance in equine sports as well as in leisure not only the physical abilities of the horse should be considered, but also the horse's personality. Besides temperamental aspects, like emotionality, or the horse's reactivity towards humans in handling situations, the learning ability of the horse is another relevant personality trait. To study whether differences in learning performance are consistent over time and whether individual learning performance differs between learning tests or is affected by emotionality, 39 young horses (Dutch Warmblood) were tested repeatedly in two learning tests. An aversive stimulus (AS) was used in one learning test (the avoidance learning test) and a reward was used in the other learning test (the reward learning test). During both learning tests behaviour as well as heart rate were measured. Each test was executed four times, twice when horses were 1 year of age, and twice when they were 2 years of age. Half of the horses received additional physical training from 6 months onwards. In both tests horses could be classified as either performers, i.e. completing the daily session, or as non-performers, i.e. returning to the home environment without having completed the daily session. There were some indications that emotionality might have caused non-performing behaviour, but these indications are not convincing enough to exclude other causes. Furthermore, there seem to be no simple relationships between measures of heart rate, behavioural responses putatively related to emotionality and learning performance. Horses revealed consistent individual learning performances within years in both tests, and in the avoidance learning test also between years. There was no significant correlation between learning performances in the avoidance learning test and the learning performances in the reward learning test. It is concluded that individual learning abilities are consistent over a short time interval for an avoidance learning test and a reward learning test and over a longer time for the avoidance learning test. Furthermore, results indicate that some horses perform better when they have to learn to avoid an aversive stimulus while others perform better when they are rewarded after a correct response. It is suggested that these differences may be relevant to design optimal individual training programmes and methods.
Keywords: Horse; Personality; Learning performance; Consistency; Emotionality
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Krause Hoare, Hemelrijk, & Rubenstein. (2000). Leadership in fish shoals. Fish Fish, 1, 82–89.
Abstract: Leadership is not an inherent quality of animal groups that show directional locomotion. However, there are other factors that may be responsible for the occurrence of leadership in fish shoals, such as individual differences in nutritional state between group members. It appears that front fish have a strong influence on directional shoal movements and that individuals that occupy such positions are often characterised by larger body lengths and lower nutritional state. Potential interactions between the two factors and their importance for positioning within shoals need further attention. Initiation of directional movement in stationary shoals and position preferences in mobile shoals need to be addressed separately because they are potentially subject to different constraints. Individuals that initiate a swimming direction may not necessarily be capable of the sustained high swimming performance required to keep the front position or have the motivation to do so, for that matter. More empirical and theoretical work is necessary to look at the factors controlling positioning behaviour within shoals, as well as overall shoal shape and structure. Tracking of marked individuals whose positioning behaviour is monitored over extended time periods of hours or days would be useful. There is an indication that shoal positions are rotated by individuals according to their nutritional needs, with hungry fish occupying front positions only for as long as necessary to regain their nutritional balance. This suggests that shoal members effectively take turns at being leaders. There is a need for three-dimensional recordings of shoaling behaviour using high-speed video systems that allow a detailed analysis of information transfer in shoals of different size. The relationship between leadership and shoal size might provide an interesting field for future research. Most studies to date have been restricted to shoals of small and medium size and more information on larger shoals would be useful.
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Wilson, A. M., McGuigan, M. P., Su, A., & van Den Bogert, A. J. (2001). Horses damp the spring in their step. Nature, 414(6866), 895–899.
Abstract: The muscular work of galloping in horses is halved by storing and returning elastic strain energy in spring-like muscle-tendon units.These make the legs act like a child's pogo stick that is tuned to stretch and recoil at 2.5 strides per second. This mechanism is optimized by unique musculoskeletal adaptations: the digital flexor muscles have extremely short fibres and significant passive properties, whereas the tendons are very long and span several joints. Length change occurs by a stretching of the spring-like digital flexor tendons rather than through energetically expensive length changes in the muscle. Despite being apparently redundant for such a mechanism, the muscle fibres in the digital flexors are well developed. Here we show that the mechanical arrangement of the elastic leg permits it to vibrate at a higher frequency of 30-40 Hz that could cause fatigue damage to tendon and bone. Furthermore, we show that the digital flexor muscles have minimal ability to contribute to or regulate significantly the 2.5-Hz cycle of movement, but are ideally arranged to damp these high-frequency oscillations in the limb.
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Moehlman, P. D. (1998). Behavioral patterns and communication in feral asses (Equus africanus). Appl. Anim. Behav. Sci., 60(2-3), 125–169.
Abstract: The behavior of feral populations of the African wild ass (Equus africanus) were studied in the Northern Panamint Range of Death Valley National Monument for 20 months from 1970 to 1973 [Moehlman, P.D., 1974. Behavior and ecology of feral asses (Equus asinus). PhD dissertation, University of Wisconsin, Madison, 251 pp.; Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Natl. Geogr. Soc. Res. Reports, 1970: 405-411]. Maintenance behavior is described and behavior sequences that were used in social interactions are quantified by sex and age class. Agonistic, sexual, and greeting behavior patterns are described and analyzed in conjunction with the responses they elicited. Mutual grooming mainly occurred between adult males, and between females and their offspring. Five types of vocalizations were distinguished: brays, grunts, growls, snorts, and whuffles. A second population was studied for 1 month on Ossabaw Island, GA (Moehlman, 1979). This population had more permanent social groups and had a higher rate of mutual grooming and foal social play.
Keywords: Equids; Feral asses; Behavior patterns; Facial expressions; Postures; Locomotion
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Fiset, S., Landry, F., & Ouellette, M. (2006). Egocentric search for disappearing objects in domestic dogs: evidence for a geometric hypothesis of direction. Anim. Cogn., 9(1), 1–12.
Abstract: In several species, the ability to locate a disappearing object is an adaptive component of predatory and social behaviour. In domestic dogs, spatial memory for hidden objects is primarily based on an egocentric frame of reference. We investigated the geometric components of egocentric spatial information used by domestic dogs to locate an object they saw move and disappear. In experiment 1, the distance and the direction between the position of the animal and the hiding location were put in conflict. Results showed that the dogs primarily used the directional information between their own spatial coordinates and the target position. In experiment 2, the accuracy of the dogs in finding a hidden object by using directional information was estimated by manipulating the angular deviation between adjacent hiding locations and the position of the animal. Four angular deviations were tested: 5, 7.5, 10 and 15 degrees . Results showed that the performance of the dogs decreased as a function of the angular deviations but it clearly remained well above chance, revealing that the representation of the dogs for direction is precise. In the discussion, we examine how and why domestic dogs determine the direction in which they saw an object disappear.
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Iversen, I. H., & Matsuzawa, T. (2003). Development of interception of moving targets by chimpanzees (Pan troglodytes) in an automated task. Anim. Cogn., 6(3), 169–183.
Abstract: The experiments investigated how two adult captive chimpanzees learned to navigate in an automated interception task. They had to capture a visual target that moved predictably on a touch monitor. The aim of the study was to determine the learning stages that led to an efficient strategy of intercepting the target. The chimpanzees had prior training in moving a finger on a touch monitor and were exposed to the interception task without any explicit training. With a finger the subject could move a small “ball” at any speed on the screen toward a visual target that moved at a fixed speed either back and forth in a linear path or around the edge of the screen in a rectangular pattern. Initial ball and target locations varied from trial to trial. The subjects received a small fruit reinforcement when they hit the target with the ball. The speed of target movement was increased across training stages up to 38 cm/s. Learning progressed from merely chasing the target to intercepting the target by moving the ball to a point on the screen that coincided with arrival of the target at that point. Performance improvement consisted of reduction in redundancy of the movement path and reduction in the time to target interception. Analysis of the finger's movement path showed that the subjects anticipated the target's movement even before it began to move. Thus, the subjects learned to use the target's initial resting location at trial onset as a predictive signal for where the target would later be when it began moving. During probe trials, where the target unpredictably remained stationary throughout the trial, the subjects first moved the ball in anticipation of expected target movement and then corrected the movement to steer the ball to the resting target. Anticipatory ball movement in probe trials with novel ball and target locations (tested for one subject) showed generalized interception beyond the trained ball and target locations. The experiments illustrate in a laboratory setting the development of a highly complex and adaptive motor performance that resembles navigational skills seen in natural settings where predators intercept the path of moving prey.
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Goto, K., Lea, S. E. G., & Dittrich, W. H. (2002). Discrimination of intentional and random motion paths by pigeons. Anim. Cogn., 5(3), 119–127.
Abstract: Twelve pigeons ( Columba livia) were trained on a go/no-go schedule to discriminate between two kinds of movement patterns of dots, which to human observers appear to be “intentional” and “non-intentional” movements. In experiment 1, the intentional motion stimulus contained one dot (a “wolf”) that moved systematically towards another dot as though stalking it, and three distractors (“sheep”). The non-intentional motion stimulus consisted of four distractors but no stalker. Birds showed some improvement of discrimination as the sessions progressed, but high levels of discrimination were not reached. In experiment 2, the same birds were tested with different stimuli. The same parameters were used but the number of intentionally moving dots in the intentional motion stimulus was altered, so that three wolves stalked one sheep. Despite the enhanced difference of movement patterns, the birds did not show any further improvement in discrimination. However, birds for which the non-intentional stimulus was associated with reward showed a decline in discrimination. These results indicated that pigeons can discriminate between stimuli that do and do not contain an element that human observer see as moving intentionally. However, as no feature-positive effect was found in experiment 1, it is assumed that pigeons did not perceive or discriminate these stimuli on the basis that the intentional stimuli contained a feature that the non-intentional stimuli lacked, though the convergence seen in experiment 2 may have been an effective feature for the pigeons. Pigeons seem to be able to recognise some form of multiple simultaneously goal-directed motions, compared to random motions, as a distinctive feature, but do not seem to use simple “intentional” motion paths of two geometrical figures, embedded in random motions, as a feature whose presence or absence differentiates motion displays.
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