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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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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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Dutto, D. J., Hoyt, D. F., Clayton, H. M., Cogger, E. A., & Wickler, S. J. (2004). Moments and power generated by the horse (Equus caballus) hind limb during jumping. J Exp Biol, 207(Pt 4), 667–674.
Abstract: The ability to jump over an obstacle depends upon the generation of work across the joints of the propelling limb(s). The total work generated by one hind limb of a horse and the contribution to the total work by four joints of the hind limb were determined for a jump. It was hypothesized that the hip and ankle joints would have extensor moments performing positive work, while the knee would have a flexor moment and perform negative work during the jump. Ground reaction forces and sagittal plane kinematics were simultaneously recorded during each jumping trial. Joint moment, power and work were determined for the metatarsophalangeal (MP), tarsal (ankle), tibiofemoral (knee) and coxofemoral (hip) joints. The hip, knee and ankle all flexed and then extended and the MP extended and then flexed during ground contact. Consistent with our hypothesis, large extensor moments were observed at the hip and ankle joints and large flexor moments at the knee and MP joints throughout ground contact of the hind limb. Peak moments tended to occur earlier in stance in the proximal joints but peak power generation of the hind limb joints occurred at similar times except for the MP joint, with the hip and ankle peaking first followed by the MP joint. During the first portion of ground contact (approximately 40%), the net result of the joint powers was the absorption of power. During the remainder of the contact period, the hind limb generated power. This pattern of power absorption followed by power generation paralleled the power profiles of the hip, ankle and MP joints. The total work performed by one hind limb was 0.71 J kg(-1). Surprisingly, the knee produced 85% of the work (0.60 J kg(-1)) done by the hind limb, and the positive work performed by the knee occurred during the first 40% of the take-off. There is little net work generated by the other three joints over the entire take-off. Velocity of the tuber coxae (a landmark on the pelvis of the animal) was negative (downward) during the first 40% of stance, which perhaps reflects the negative work performed to decrease the potential energy during the first 40% of contact. During the final 60% of contact, the hip, ankle and MP joints generate positive work, which is reflected in the increase of the animal's potential energy.
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Witte, T. H., Knill, K., & Wilson, A. M. (2004). Determination of peak vertical ground reaction force from duty factor in the horse (Equus caballus). J Exp Biol, 207(Pt 21), 3639–3648.
Abstract: Measurement of peak vertical ground reaction force (GRFz) from multiple limbs simultaneously during high-speed, over-ground locomotion would enhance our understanding of the locomotor mechanics of cursorial animals. Here, we evaluate the accuracy of predicting peak GRFz from duty factor (the proportion of the stride for which the limb is in contact with the ground). Foot-mounted uniaxial accelerometers, combined with UHF FM telemetry, are shown to be practical and accurate for the field measurement of stride timing variables, including duty factor. Direct comparison with the force plate produces a mean error of 2.3 ms and 3.5 ms for the timing of foot on and foot off, respectively, across all gaits. Predictions of peak GRFz from duty factor show mean errors (with positive values indicating an overestimate) of 0.8+/-0.04 N kg(-1) (13%; N=42; mean +/- S.E.M.) at walk, -0.3+/-0.06 N kg(-1) (3%; N=75) at trot, -2.3+/-0.27 N kg(-1) (16%; N=18) for the non-lead limb at canter and +2.1+/-0.7 N kg(-1) (19%; N=9) for the lead limb at canter. The substantial over- and underestimate seen at canter, in the lead and non-lead limbs, respectively, is attributed to the different functions performed by the two limbs in the asymmetrical gaits. The difference in load experienced by the lead and non-lead limbs decreased with increasing speed.
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Etienne, A. S., Maurer, R., & Seguinot, V. (1996). Path integration in mammals and its interaction with visual landmarks. J Exp Biol, 199(Pt 1), 201–209.
Abstract: During locomotion, mammals update their position with respect to a fixed point of reference, such as their point of departure, by processing inertial cues, proprioceptive feedback and stored motor commands generated during locomotion. This so-called path integration system (dead reckoning) allows the animal to return to its home, or to a familiar feeding place, even when external cues are absent or novel. However, without the use of external cues, the path integration process leads to rapid accumulation of errors involving both the direction and distance of the goal. Therefore, even nocturnal species such as hamsters and mice rely more on previously learned visual references than on the path integration system when the two types of information are in conflict. Recent studies investigate the extent to which path integration and familiar visual cues cooperate to optimize the navigational performance.
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Buchner, H. H. F., Obermuller, S., & Scheidl, M. (2000). Body Centre of Mass Movement in the Sound Horse. The Veterinary Journal, 160(3), 225–234.
Abstract: The body centre of mass (BCM) is a key factor in the analysis of equine locomotion, as its position and movement determines the distribution and magnitude of loads on the limbs. In this study, the three-dimensional (3D) movement of the BCM in walking and trotting horses was assessed using a kinematic, segmental method. Thirty markers representing 20 body segments were recorded in 12 sound horses while standing, walking and trotting on a treadmill using a high-speed video system. Based on segmental inertial data, 3D positions of the segmental centres of mass as well as the total BCM were calculated. The position within the trunk during square standing and the movements of the BCM were determined for the three planes. The position of the BCM in the standing horse is presented relative to external reference points. At the trot, vertical displacement amplitude of the BCM amounted to 53 (6) mm as mean (sd), which was 27% smaller than external trunk movement. Medio-lateral displacement amplitude of the BCM was 19 (4) mm, 34% less than trunk amplitude. Sagittal forward-backward oscillations of the BCM independent from general forward movement were 13 (3) mm, being 24% less than trunk movements. At the walk, vertical, medio-lateral and sagittal BCM movements were smaller than trunk movements by 43, 65 and 65% respectively. The results show reduced and efficient BCM movements compared to the trunk and form a basis for the assessment of various clinical conditions such as lameness, the influence of a rider and various dressage performances.
Keywords: Horse; centre of mass; kinematics; segment model; locomotion.
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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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Argue, C. K., & Clayton, H. M. (1993). A preliminary study of transitions between the walk and trot in dressage horses. Acta Anat (Basel), 146(2-3), 179–182.
Abstract: The object of this study was to determine the limb support sequence during the transitions from walk to trot and from trot to walk in dressage horses under saddle and to test the null hypothesis that the limb support sequence during the transitions is not related to the level of training. Sixteen dressage horses training at novice to FEI Grand Prix level were videotaped performing an average of 9 transitions each from walk to trot and from trot to walk. The 30-Hz videotapes were viewed in slow motion, and based on the limb support sequence the transitions were categorized into two types. In type 1 transitions there were no intermediate steps between the walk and trot sequences. Type 2 transitions were characterized by intermediate steps, including a single support phase. The Kendall rank-order correlation coefficient showed that a higher level of training was positively associated with an increased percentage of type 1 transitions for both walk-to-trot transitions (p < or = 0.05) and trot-to-walk transitions (p < or = 0.01). No significant preference for initiating or completing the trot on the left or right diagonal was found using the binomial test for individual horses and the Wilcoxon signed-ranks test for the group.
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Wolff, A., & Hausberger, M. (1994). Behaviour of foals before weaning may have some genetic basis. Ethology, 96(1), 1–10.
Abstract: In this preliminary study on foal behaviour, 13 French saddlebred foals (2-3 mo old) and their dams were observed on pasture. The most important findings are the interindividual quantitative differences in foal behaviour patterns as well as in the amount of mainly foal-initiated time spent at given distances from their mares. Interindividual differences seem in part due to a sire effect
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Pearce, G. P., May-Davis, S., & Greaves, D. (2005). Femoral asymmetry in the Thoroughbred racehorse. Aust Vet J, 83(6), 367–370.
Abstract: OBJECTIVE: To investigate the occurrence of geometrical asymmetries in the macro-architecture of left and right femurs from Thoroughbred racehorses previously used in competitive training and racing in New South Wales, Australia. METHODS: Detailed postmortem measurements were made of 37 characteristics of left and right femurs from eleven Thoroughbred racehorses euthanased for reasons unrelated to the study. Measurements focused on articulating surfaces and sites of attachment of muscles and ligaments known to be associated with hindlimb locomotion. RESULTS: Five measurements were significantly larger in left compared to right femurs (P < 0.05). The regions showing significant differences between left and right limbs were proximal cranial and overhead medio-lateral widths, greater trochanter depth, depth of the fovea in the femoral head and distal inter-epicondylar width. CONCLUSION: The left-right differences in femoral morphology were associated with sites of muscle and ligament attachment known to be involved with hindlimb function in negotiating turns. These differences may be the result of selection pressure for racing performance on curved race tracks and/or adaptations related to asymmetrical loading of the outside hindlimb associated with repeated negotiation of turns on such tracks.
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