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Spadavecchia, C., Arendt-Nielsen, L., Spadavecchia, L., Mosing, M., Auer, U., & van den Hoven, R. (2007). Effects of butorphanol on the withdrawal reflex using threshold, suprathreshold and repeated subthreshold electrical stimuli in conscious horses. Vet Anaesth Analg, 34(1), 48–58.
Abstract: OBJECTIVE: To assess the effects of a single intravenous dose of butorphanol (0.1 mg kg(-1)) on the nociceptive withdrawal reflex (NWR) using threshold, suprathreshold and repeated subthreshold electrical stimuli in conscious horses. STUDY DESIGN: 'Unblinded', prospective experimental study. ANIMALS: Ten adult horses, five geldings and five mares, mean body mass 517 kg (range 487-569 kg). METHODS: The NWR was elicited using single transcutaneous electrical stimulation of the palmar digital nerve. Repeated stimulations were applied to evoke temporal summation. Surface electromyography was performed to record and quantify the responses of the common digital extensor muscle to stimulation and behavioural reactions were scored. Before butorphanol administration and at fixed time points up to 2 hours after injection, baseline threshold intensities for NWR and temporal summation were defined and single suprathreshold stimulations applied. Friedman repeated-measures analysis of variance on ranks and Wilcoxon signed-rank test were used with the Student-Newman-Keul's method applied post-hoc. The level of significance (alpha) was set at 0.05. RESULTS: Butorphanol did not modify either the thresholds for NWR and temporal summation or the reaction scores, but the difference between suprathreshold and threshold reflex amplitudes was reduced when single stimulation was applied. Upon repeated stimulation after butorphanol administration, a significant decrease in the relative amplitude was calculated for both the 30-80 and the 80-200 millisecond intervals after each stimulus, and for the whole post-stimulation interval in the right thoracic limb. In the left thoracic limb a decrease in the relative amplitude was found only in the 30-80 millisecond epoch. CONCLUSION: Butorphanol at 0.1 mg kg(-1) has no direct action on spinal Adelta nociceptive activity but may have some supraspinal effects that reduce the gain of the nociceptive system. CLINICAL RELEVANCE: Butorphanol has minimal effect on sharp immediate Adelta-mediated pain but may alter spinal processing and decrease the delayed sensations of pain.
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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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Kraus-Hansen, A. E., Fackelman, G. E., Becker, C., Williams, R. M., & Pipers, F. S. (1992). Preliminary studies on the vascular anatomy of the equine superficial digital flexor tendon. Equine Vet J, 24(1), 46–51.
Abstract: The vascular and microvascular anatomy of normal equine superficial digital flexor tendons was studied by dissection of vinyl-perfused specimens and by microangiography on high detail film. The presence of an extensive intratendinous vascular latticework was confirmed, and a 'nutrient artery' described closely associated with the accessory ligament of the superficial digital flexor tendon (proximal check ligament). Circumferential stripping of the paratenon from the tendon to eliminate afferent vessels was performed bilaterally in three horses and unilaterally in a fourth, followed by a treadmill training regimen. No resulting intratendinous lesions could be documented on gross post mortem and histological examination at three, 10, or 35 days post operatively. There was mild paratendinous proliferation in all instances. In one horse, four intratendinous ligatures were placed within the medial and lateral borders of the contralateral tendon to isolate further from its blood supply a 10 cm segment. Gross lesions at 35 days post operatively included a marked paratendinous response involving the entire 10 cm segment, and a darkened, soft focus within the core of the tendon. Histopathology and electron microscopy demonstrated focal degeneration. It was concluded that the blood supply of the normal equine superficial digital flexor tendon is primarily intratendinous, rather than paratendinous as previously thought. The lesions in one horse similar to those in naturally occurring tendinitis supported a vascular aetiology of the disease, and set the groundwork for studies aimed at the development of a clinically relevant tendinitis model.
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Colahan, P., Lindsey, E., & Nunier, C. (1993). Determination of the center of pressure of the hoofs of the forelimbs of horses standing on a flat level surface. Acta Anat (Basel), 146(2-3), 175–178.
Abstract: The pressure exerted on a flat level surface by recently trimmed, unshod hoofs of the front limbs of 23 sound, adult horses was measured using pressure-sensitive film and a specially built cassette. The horses were tranquilized and stood with one foot on the 2.9-cm-thick cassette and the other on a block of equal height. The hoofs were observed for motion during the measurement, and the developed film was examined for improper alignment of the film or slipping of the hoof. The center of pressure was located using the method of weighted proportions of Barrey. This static measurement system with a long measurement time and the number of measurements reduced the influence of variables inherent in the horses' behavior and the measuring system. The calculated point was recorded as falling medial to, lateral to or on a line bisecting the central sulcus of the frog. In the dorsal to palmar orientation the point was classified with reference to a line drawn halfway between the most dorsal and the most palmar mark on the film. Forty-six percent of the calculated centers of pressure were located in the medial heel area. Binomial analysis for large samples indicates that this was a significant variation from a random distribution. Seventy-six percent of the centers were located in or on the borders of the medial heel.
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McGreevy, P. D., & Rogers, L. J. (2005). Motor and sensory laterality in thoroughbred horses. Appl. Anim. Behav. Sci., 92(4), 337–352.
Abstract: We investigated lateralisation in horses because it is likely to be important in training and athletic performance. Thoroughbred horses (n = 106) were observed every 60 s for 2 h, when they were at pasture, and the position of the forelimbs in relation to one another was recorded. There was a population bias skewed to standing with the left forelimb advanced over the right (i.e. directional lateralisation). Using the first 50 observations, the distribution of preferences was 43 significantly left, 10 significantly right with 53 being non-significant (i.e. ambidextextrous). The strength of motor bias increased with age, suggesting maturation or an influence of training. The horses were also presented with an olfactory stimulus (stallion faeces) to score the tendency to use one nostril rather than the other. A significant preference to use the right nostril first was shown in horses under 4 years of age (n = 61) but not in older horses. Of the 157 horses tested for nostril bias, 76 had been assessed for motor bias and so were used for analysis of the relationship between laterality in the two modalities. There was no significant relationship between direction of foreleg motor bias and first nostril used, total number of inhalations or laterality index of nostril use. The absence of a correlation between laterality of nostril use and motor bias indicates that lateralisation of the equine brain occurs on at least two levels of neural organisation--sensory and motor--a finding that is consistent with other examples of lateralisation in species that have been examined in more detail.
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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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McGuigan, M. P., & Wilson, A. M. (2003). The effect of gait and digital flexor muscle activation on limb compliance in the forelimb of the horse Equus caballus. J Exp Biol, 206(Pt 8), 1325–1336.
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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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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Bystrom, A., Roepstorff, L., & Johnston, C. (2006). Effects of draw reins on limb kinematics. Equine Vet J Suppl, (36), 452–456.
Abstract: REASONS FOR PERFORMING STUDY: No data exist on the GRF-kinematics relation due to changes caused by equestrian interventions. HYPOTHESIS: Through the judicious use of draw reins the rider can influence the kinematics of the horse to meet stated goals of dressage training. Relating the results to previously published kinetic data of the same experiment implies a possible relationship between kinetics and kinematics. METHODS: The kinematics of 8 sound Swedish Warmblood horses were measured whilst the horses were being ridden with and without draw reins. Three conditions were evaluated: 1) draw reins only (DR), 2) combination of draw reins and normal reins (NR+DR) and 3) normal reins only (NR). RESULTS: Head and neck angles were significantly decreased by the draw rein but 4-5 times more so for DR when with NR+DR. The forelimb position at hoof lift-off was significantly more caudal with DR. In the hind limb the hip joint extended more quickly and the hock joint flexed more with NR+DR than with NR. Compared to DR the hip joint angular pattern was not significantly different, but the pelvis was more horizontal. CONCLUSION: Riding with a draw rein can have significant influence on the kinematics of the horse. Some of the observed changes can be coupled to changes in kinetics. The hock joint angle seems to be a fairly reliable indicator of load on the hind limb and the angle of femur appears important for hind limb propulsion, when considered in conjunction with the orientation of the pelvis. POTENTIAL RELEVANCE: These findings are important for riders and trainers, as kinematic changes are what trainers observe. It is thereby important to ascertain which kinematic changes are consistently coupled to changes in kinetics in order for trainers to be able to judge correctly the success of intended goals. Further studies are warranted to validate and confirm suggested relationships between kinetics and kinematics.
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Weishaupt, M. A., Wiestner, T., von Peinen, K., Waldern, N., Roepstorff, L., van Weeren, R., et al. (2006). Effect of head and neck position on vertical ground reaction forces and interlimb coordination in the dressage horse ridden at walk and trot on a treadmill. Equine Vet J Suppl, (36), 387–392.
Abstract: REASONS FOR PERFORMING STUDY: Little is known in quantitative terms about the influence of different head-neck positions (HNPs) on the loading pattern of the locomotor apparatus. Therefore it is difficult to predict whether a specific riding technique is beneficial for the horse or if it may increase the risk for injury. OBJECTIVE: To improve the understanding of forelimb-hindlimb balance and its underlying temporal changes in relation to different head and neck positions. METHODS: Vertical ground reaction force and time parameters of each limb were measured in 7 high level dressage horses while being ridden at walk and trot on an instrumented treadmill in 6 predetermined HNPs: HNP1 – free, unrestrained with loose reins; HNP2 – neck raised, bridge of the nose in front of the vertical; HNP3 – neck raised, bridge of the nose behind the vertical; HNP4 – neck lowered and flexed, bridge of the nose considerably behind the vertical; HNP5 – neck extremely elevated and bridge of the nose considerably in front of the vertical; HNP6 – neck and head extended forward and downward. Positions were judged by a qualified dressage judge. HNPs were assessed by comparing the data to a velocity-matched reference HNP (HNP2). Differences were tested using paired t test or Wilcoxon signed rank test (P<0.05). RESULTS: At the walk, stride duration and overreach distance increased in HNP1, but decreased in HNP3 and HNP5. Stride impulse was shifted to the forehand in HNP1 and HNP6, but shifted to the hindquarters in HNP5. At the trot, stride duration increased in HNP4 and HNP5. Overreach distance was shorter in HNP4. Stride impulse shifted to the hindquarters in HNP5. In HNP1 peak forces decreased in the forelimbs; in HNP5 peak forces increased in fore- and hindlimbs. CONCLUSIONS: HNP5 had the biggest impact on limb timing and load distribution and behaved inversely to HNP1 and HNP6. Shortening of forelimb stance duration in HNP5 increased peak forces although the percentage of stride impulse carried by the forelimbs decreased. POTENTIAL RELEVANCE: An extremely high HNP affects functionality much more than an extremely low neck.
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