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Author |
Wilson, A.M.; McGuigan, M.P.; Su, A.; van Den Bogert, A.J. |
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Title |
Horses damp the spring in their step |
Type |
Journal Article |
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Year |
2001 |
Publication |
Nature |
Abbreviated Journal |
Nature |
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Volume  |
414 |
Issue |
6866 |
Pages |
895-899 |
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Keywords |
Animals; Biomechanics; Elasticity; Forelimb; Gait; Horses/anatomy & histology/*physiology; Leg Bones/*physiology; Locomotion; Models, Biological; Muscle Fibers/physiology; Muscle, Skeletal/anatomy & histology/*physiology; Tendons/anatomy & histology/*physiology; Vibration |
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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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Department of Veterinary Basic Sciences, The Royal Veterinary College, Hatfield, Herts AL9 7TA, UK. awilson@rvc.ac.uk |
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0028-0836 |
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PMID:11780059 |
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no |
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Equine Behaviour @ team @ |
Serial |
2300 |
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Author |
Real, L.A. |
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Title |
Animal choice behavior and the evolution of cognitive architecture |
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Journal Article |
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Year |
1991 |
Publication |
Science (New York, N.Y.) |
Abbreviated Journal |
Science |
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Volume |
253 |
Issue |
5023 |
Pages |
980-986 |
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Keywords |
Animals; Bees/genetics/*physiology; Biomechanics; *Choice Behavior; *Cognition; *Evolution; Mathematics; Models, Genetic; Probability |
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Abstract |
Animals process sensory information according to specific computational rules and, subsequently, form representations of their environments that form the basis for decisions and choices. The specific computational rules used by organisms will often be evolutionarily adaptive by generating higher probabilities of survival, reproduction, and resource acquisition. Experiments with enclosed colonies of bumblebees constrained to foraging on artificial flowers suggest that the bumblebee's cognitive architecture is designed to efficiently exploit floral resources from spatially structured environments given limits on memory and the neuronal processing of information. A non-linear relationship between the biomechanics of nectar extraction and rates of net energetic gain by individual bees may account for sensitivities to both the arithmetic mean and variance in reward distributions in flowers. Heuristic rules that lead to efficient resource exploitation may also lead to subjective misperception of likelihoods. Subjective probability formation may then be viewed as a problem in pattern recognition subject to specific sampling schemes and memory constraints. |
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Department of Biology, University of North Carolina, Chapel Hill 27599-3280 |
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0036-8075 |
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PMID:1887231 |
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Equine Behaviour @ team @ |
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2846 |
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Author |
Skedros, J.G.; Dayton, M.R.; Sybrowsky, C.L.; Bloebaum, R.D.; Bachus, K.N. |
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Title |
The influence of collagen fiber orientation and other histocompositional characteristics on the mechanical properties of equine cortical bone |
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Journal Article |
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Year |
2006 |
Publication |
The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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Volume |
209 |
Issue |
Pt 15 |
Pages |
3025-3042 |
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Keywords |
Animals; Biomechanics; Bone and Bones/*physiology; Collagen/*physiology; Forelimb; Horses/*physiology |
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Abstract |
This study examined relative influences of predominant collagen fiber orientation (CFO), mineralization (% ash), and other microstructural characteristics on the mechanical properties of equine cortical bone. Using strain-mode-specific (S-M-S) testing (compression testing of bone habitually loaded in compression; tension testing of bone habitually loaded in tension), the relative mechanical importance of CFO and other material characteristics were examined in equine third metacarpals (MC3s). This model was chosen since it had a consistent non-uniform strain distribution estimated by finite element analysis (FEA) near mid-diaphysis of a thoroughbred horse, net tension in the dorsal/lateral cortices and net compression in the palmar/medial cortices. Bone specimens from regions habitually loaded in tension or compression were: (1) tested to failure in both axial compression and tension in order to contrast S-M-S vs non-S-M-S behavior, and (2) analyzed for CFO, % ash, porosity, fractional area of secondary osteonal bone, osteon cross-sectional area, and population densities of secondary osteons and osteocyte lacunae. Multivariate multiple regression analyses revealed that in S-M-S compression testing, CFO most strongly influenced total energy (pre-yield elastic energy plus post-yield plastic energy); in S-M-S tension testing CFO most strongly influenced post-yield energy and total energy. CFO was less important in explaining S-M-S elastic modulus, and yield and ultimate stress. Therefore, in S-M-S loading CFO appears to be important in influencing energy absorption, whereas the other characteristics have a more dominant influence in elastic modulus, pre-yield behavior and strength. These data generally support the hypothesis that differentially affecting S-M-S energy absorption may be an important consequence of regional histocompositional heterogeneity in the equine MC3. Data inconsistent with the hypothesis, including the lack of highly longitudinal collagen in the dorsal-lateral ;tension' region, paradoxical histologic organization in some locations, and lack of significantly improved S-M-S properties in some locations, might reflect the absence of a similar habitual strain distribution in all bones. An alternative strain distribution based on in vivo strain measurements, without FEA, on non-Thoroughbreds showing net compression along the dorsal-palmar axis might be more characteristic of the habitual loading of some of the bones that we examined. In turn, some inconsistencies might also reflect the complex torsion/bending loading regime that the MC3 sustains when the animal undergoes a variety of gaits and activities, which may be representative of only a portion of our animals, again reflecting the possibility that not all of the bones examined had similar habitual loading histories. |
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Utah Bone and Joint Center, 5323 S. Woodrow Street #202, Salt Lake City, UT 84107, USA. jskedros@utahboneandjoint.com |
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0022-0949 |
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PMID:16857886 |
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Call Number |
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Serial |
1868 |
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Author |
Bobbert, M.F.; Santamaria, S. |
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Title |
Contribution of the forelimbs and hindlimbs of the horse to mechanical energy changes in jumping |
Type |
Journal Article |
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Year |
2005 |
Publication |
The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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Volume |
208 |
Issue |
2 |
Pages |
249-260 |
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Keywords |
Animals; Biomechanics; Forelimb/*physiology; Hindlimb/*physiology; Horses/*physiology; Locomotion/*physiology; Muscle, Skeletal/*physiology; Time Factors |
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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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Address |
Institute for Fundamental and Clinical Human Movement Sciences, Vrije Universiteit, van der Boechorstraat 9, NL-1081 BT Amsterdam, The Netherlands. MFBobbert@fbw.vu.nl |
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0022-0949 |
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Notes |
PMID:15634844 |
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Call Number |
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Serial |
1895 |
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Author |
Dutto, D.J.; Hoyt, D.F.; Clayton, H.M.; Cogger, E.A.; Wickler, S.J. |
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Title |
Moments and power generated by the horse (Equus caballus) hind limb during jumping |
Type |
Journal Article |
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Year |
2004 |
Publication |
The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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Volume |
207 |
Issue |
Pt 4 |
Pages |
667-674 |
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Keywords |
Animals; Biomechanics; Hindlimb/*physiology; Horses/*physiology; Locomotion/*physiology |
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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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Department of Kinesiology and Health Promotion, California State Polytechnic University, Pomona, CA 91768, USA. ddutto@csupomona.edu |
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0022-0949 |
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Notes |
PMID:14718509 |
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no |
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Call Number |
Equine Behaviour @ team @ |
Serial |
3654 |
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Author |
Witte, T.H.; Knill, K.; Wilson, A.M. |
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Title |
Determination of peak vertical ground reaction force from duty factor in the horse (Equus caballus) |
Type |
Journal Article |
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Year |
2004 |
Publication |
The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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Volume |
207 |
Issue |
Pt 21 |
Pages |
3639-3648 |
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Keywords |
*Acceleration; Animals; Biomechanics; Forelimb/physiology; *Gait; Hindlimb/physiology; Horses/*physiology; Locomotion/*physiology; Telemetry; Time Factors |
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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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Structure and Motion Lab, The Royal Veterinary College, Hawkshead Lane, Hatfield, Hertfordshire, AL9 7TA, UK |
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0022-0949 |
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PMID:15371472 |
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no |
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Call Number |
Equine Behaviour @ team @ |
Serial |
3658 |
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Author |
McGuigan, M.P.; Wilson, A.M. |
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Title |
The effect of gait and digital flexor muscle activation on limb compliance in the forelimb of the horse Equus caballus |
Type |
Journal Article |
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Year |
2003 |
Publication |
The Journal of Experimental Biology |
Abbreviated Journal |
J Exp Biol |
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Volume |
206 |
Issue |
Pt 8 |
Pages |
1325-1336 |
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Keywords |
Animals; Biomechanics; Forelimb/anatomy & histology/*physiology; Gait/*physiology; Horses/anatomy & histology/*physiology; Muscle Contraction/*physiology; Running |
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Abstract |
A horse's legs are compressed during the stance phase, storing and then returning elastic strain energy in spring-like muscle-tendon units. The arrangement of the muscle-tendon units around the lever-like joints means that as the leg shortens the muscle-tendon units are stretched. The forelimb anatomy means that the leg can be conceptually divided into two springs: the proximal spring, from the scapula to the elbow, and the distal spring, from the elbow to the foot. In this paper we report the results of a series of experiments testing the hypothesis that there is minimal scope for muscle contraction in either spring to adjust limb compliance. Firstly, we demonstrate that the distal, passive leg spring changes length by 127 mm (range 106-128 mm) at gallop and the proximal spring by 12 mm (9-15 mm). Secondly, we demonstrate that there is a linear relationship between limb force and metacarpo-phalangeal (MCP) joint angle that is minimally influenced by digital flexor muscle activation in vitro or as a function of gait in vivo. Finally, we determined the relationship between MCP joint angle and vertical ground-reaction force at trot and then predicted the forelimb peak vertical ground-reaction force during a 12 m s(-1) gallop on a treadmill. These were 12.79 N kg(-1) body mass (BM) (range 12.07-13.73 N kg(-1) BM) for the lead forelimb and 15.23 N kg(-1) BM (13.51-17.10 N kg(-1) BM) for the non-lead forelimb. |
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Structure and Motion Laboratory, Veterinary Basic Sciences, The Royal Veterinary College, Hawkshead Lane, North Mymms, Hatfield, Hertfordshire AL9 7TA, UK. m.p.mcguigan@leeds.ac.uk |
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0022-0949 |
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PMID:12624168 |
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Call Number |
Equine Behaviour @ team @ |
Serial |
3655 |
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Author |
Sloet van Oldruitenborgh-Oosterbaan, M.M.; Blok, M.B.; Begeman, L.; Kamphuis, M.C.D.; Lameris, M.C.; Spierenburg, A.J.; Lashley, M.J.J.O. |
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Title |
Workload and stress in horses: comparison in horses ridden deep and round ('rollkur') with a draw rein and horses ridden in a natural frame with only light rein contact |
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Journal Article |
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Year |
2006 |
Publication |
Tijdschrift Voor Diergeneeskunde |
Abbreviated Journal |
Tijdschr Diergeneeskd |
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Volume |
131 |
Issue |
5 |
Pages |
152-157 |
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Keywords |
Animal Husbandry/methods; Animals; Biomechanics; Blood Glucose/analysis; Female; Heart Rate/physiology; Hematocrit/veterinary; Horses/blood/*physiology; Hydrocortisone/blood; Lactic Acid/blood; Physical Conditioning, Animal/adverse effects/*physiology; Stress, Physiological/blood/etiology/veterinary |
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'Rollkur' or 'overbending' is the low and deep riding of a dressage horse during training or warming up. Lately, this technique has been criticized, and not necessarily objectively, on welfare grounds. To be able to evaluate these criticisms, more needs to be known about the workload and stress of horses being ridden 'rollkur'. The aim of the present study was to compare the workload of eight riding-school horses when being ridden deep and round with a draw rein ('rollkur') and when being ridden in a natural frame with only light rein contact ('free'). Workload (as measured by heart rate and blood lactate concentration) was slightly higher when horses were ridden 'rollkur' than when they were ridden 'free'. There were no differences in packed cell volume, or glucose and cortisol concentrations. No signs of uneasiness or stress could be determined when the horses were ridden 'rollkur'. Subjectively, all horses improved their way of moving during 'rollkur' and were more responsive to their rider. |
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Department of Equine Sciences, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 12, 3584 CM Utrecht, The Netherlands. m.sloet@vet.uu.nl |
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0040-7453 |
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PMID:16532786 |
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no |
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Call Number |
Equine Behaviour @ team @ |
Serial |
5638 |
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Author |
Santamaria, S.; Bobbert, M.F.; Back, W.; Barneveld, A.; van Weeren, P.R. |
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Title |
Effect of early training on the jumping technique of horses |
Type |
Journal Article |
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Year |
2005 |
Publication |
American Journal of Veterinary Research |
Abbreviated Journal |
Am J Vet Res |
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Volume |
66 |
Issue |
3 |
Pages |
418-424 |
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Keywords |
Age Factors; Analysis of Variance; Animals; Biomechanics; Forelimb/physiology; Gait/*physiology; Hindlimb/physiology; Horses/growth & development/*physiology; Locomotion/*physiology; Models, Biological; Physical Conditioning, Animal/*methods |
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Abstract |
OBJECTIVE: To investigate the effects of early training for jumping by comparing the jumping technique of horses that had received early training with that of horses raised conventionally. ANIMALS: 40 Dutch Warmblood horses. PROCEDURE: The horses were analyzed kinematically during free jumping at 6 months of age. Subsequently, they were allocated into a control group that was raised conventionally and an experimental group that received 30 months of early training starting at 6 months of age. At 4 years of age, after a period of rest in pasture and a short period of training with a rider, both groups were analyzed kinematically during free jumping. Subsequently, both groups started a 1-year intensive training for jumping, and at 5 years of age, they were again analyzed kinematically during free jumping. In addition, the horses competed in a puissance competition to test maximal performance. RESULTS: Whereas there were no differences in jumping technique between experimental and control horses at 6 months of age, at 4 years, the experimental horses jumped in a more effective manner than the control horses; they raised their center of gravity less yet cleared more fences successfully than the control horses. However, at 5 years of age, these differences were not detected. Furthermore, the experimental horses did not perform better than the control horses in the puissance competition. CONCLUSIONS AND CLINICAL RELEVANCE: Specific training for jumping of horses at an early age is unnecessary because the effects on jumping technique and jumping capacity are not permanent. |
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Department of Equine Sciences, Faculty of Veterinary Medicine, Utrecht University, NL-3584 CM Utrecht, The Netherlands |
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English |
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0002-9645 |
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Notes |
PMID:15822585 |
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Call Number |
Equine Behaviour @ team @ |
Serial |
4037 |
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Author |
Santamaria, S.; Bobbert, M.E.; Back, W.; Barneveld, A.; van Weeren, P.R. |
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Title |
Variation in free jumping technique within and among horses with little experience in show jumping |
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Journal Article |
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Year |
2004 |
Publication |
American Journal of Veterinary Research |
Abbreviated Journal |
Am J Vet Res |
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Volume |
65 |
Issue |
7 |
Pages |
938-944 |
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Keywords |
*Acceleration; Analysis of Variance; Animals; Biomechanics; Forelimb/physiology; Gait/*physiology; Hindlimb/physiology; Horses/*physiology; Locomotion/*physiology; Models, Biological; Video Recording |
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Abstract |
OBJECTIVE: To quantify variation in the jumping technique within and among young horses with little jumping experience, establish relationships between kinetic and kinematic variables, and identify a limited set of variables characteristic for detecting differences in jumping performance among horses. ANIMALS: Fifteen 4-year-old Dutch Warmblood horses. PROCEDURE: The horses were raised under standardized conditions and trained in accordance with a fixed protocol for a short period. Subsequently, horses were analyzed kinematically during free jumping over a fence with a height of 1.05 m. RESULTS: Within-horse variation in all variables that quantified jumping technique was smaller than variation among horses. However, some horses had less variation than others. Height of the center of gravity (CG) at the apex of the jump ranged from 1.80 to 2.01 m among horses; this variation could be explained by the variation in vertical velocity of the CG at takeoff (r, 0.78). Horses that had higher vertical velocity at takeoff left the ground and landed again farther from the fence, had shorter push-off phases for the forelimbs and hind limbs, and generated greater vertical acceleration of the CG primarily during the hind limb push-off. However, all horses cleared the fence successfully, independent of jumping technique. CONCLUSIONS AND CLINICAL RELEVANCE: Each horse had its own jumping technique. Differences among techniques were characterized by variations in the vertical velocity of the CG at takeoff. It must be determined whether jumping performance later in life can be predicted from observing free jumps of young horses. |
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Address |
Department of Equine Sciences, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 12, NL-3584 CM Utrecht, The Netherlands |
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English |
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ISSN |
0002-9645 |
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Notes |
PMID:15281652 |
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no |
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Call Number |
Equine Behaviour @ team @ |
Serial |
3772 |
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