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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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Rizzolatti, G., Fogassi, L., & Gallese, V. (2006). Mirrors of the mind. Sci Am, 295(5), 54–61.
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Rilling, M. E., & Neiworth, J. J. (1991). How animals use images. Sci Prog, 75(298 Pt 3-4), 439–452.
Abstract: Animal cognition is a field within experimental psychology in which cognitive processes formerly studied exclusively with people have been demonstrated in animals. Evidence for imagery in the pigeon emerges from the experiments described here. The pigeon's task was to discriminate, by pecking the appropriate choice key, between a clock hand presented on a video screen that rotated clockwise with constant velocity from a clock hand that violated constant velocity. Imagery was defined by trials on which the line rotated from 12.00 o'clock to 3.00 o'clock, then disappeared during a delay, and reappeared at a final stop location beyond 3.00 o'clock. After acquisition of a discrimination with final stop locations at 3.00 o'clock and 6.00 o'clock, the evidence for imagery was the accurate responding of the pigeons to novel locations at 4.00 o'clock and 7.00 o'clock. Pigeons display evidence of imagery by transforming a representation of movement that includes a series of intermediate steps which accurately represent the location of a moving stimulus after it disappears.
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McCall, C. A., Hall, S., McElhenney, W. H., & Cummins, K. A. (2006). Evaluation and comparison of four methods of ranking horses based on reactivity. Appl. Anim. Behav. Sci., 96(1-2), 115–127.
Abstract: Four methods of ranking horses on reactivity were evaluated and compared: isolation from conspecifics, presentation of a static novel stimulus, traversing a novel stimulus in a runway (isolation, novel stimulus and runway tests, respectively) and assigning subjective emotionality scores. In all tests, horses' heart rates were recorded and behaviour was videotaped. To be considered a valid test of reactivity, at least one heart rate and one behavioural measurement in the test had to change significantly between treatments (tranquilizer administation versus sham tranquilizer administration), and behavioural measures had to be displayed in at least 75% of the trials. Forty horses performed each of the three tests daily on three different days in a switchback design. Horses were assigned randomly to a daily test sequence, which was maintained throughout the study. In the runway test, no significant difference in heart rate values in tranquilized and non-tranquilized horses was found, and no behavioural attribute was displayed in more than 52% of the trials; therefore it was rejected as a valid test of reactivity. Both isolation and novel stimulus tests produced valid measurements. Mean heart rate was the most precise physiological measure for these tests, and walking and defecation frequency were the most precise behavioural measures for novel stimulus and isolation tests, respectively. Mean heart rates on the novel stimulus and isolation tests were correlated (rs = 0.79, P < 0.01) indicating that these tests produced similar rankings based on physiological responses. However, behavioural measures ranked horses differently (rs = 0.27, P < 0.10) on the tests. Rank correlations between mean heart rates and behavioural measures were higher in the novel stimulus (rs = 0.66, P < 0.01) than the isolation test (rs = 0.55, P < 0.01), indicating that the novel stimulus test ranked horses based on either physiological or behavioural responses more similarly than did the isolation test. Therefore, the novel stimulus test was considered the more accurate evaluation of reactivity. Subjective emotionality scores were correlated moderately with mean heart rates (rs > 0.33, P < 0.01) from the novel stimulus and isolation tests and with walking scores (rs = 0.47, P < 0.01) from the novel stimulus test. Assignment of subjective emotionality scores was not as accurate as the novel stimulus or isolation tests in ranking horses for reactivity. Using physiological data alone, combining physiological and behavioural measurements or using more than one behavioural measurement in reactivity tests may reflect the reactivity of the horse better than a single behavioural measurement.
Keywords: Horse; Reactivity tests; Heart rate; Emotionality; Temperament
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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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Wennerstrand, J., Johnston, C., Roethlisberger-Holm, K., Erichsen, C., Eksell, P., & Drevemo, S. (2004). Kinematic evaluation of the back in the sport horse with back pain. Equine Vet J, 36(8), 707–711.
Abstract: REASONS FOR PERFORMING STUDY: Earlier studies have developed a clinical tool to evaluate objectively the function of the equine back. The ability to differentiate horses with back pain from asymptomatic, fully functioning horses using kinematic measures from this tool has not been evaluated. OBJECTIVES: To compare the kinematics of the back at walk and trot in riding horses with back dysfunction to the same parameters in asymptomatic sport horses. METHODS: The kinematics of the back in 12 horses with impaired performance and back pain were studied at walk and trot on a treadmill. Data were captured for 10 sees at 240 Hz. Range of movement (ROM) and intravertebral pattern symmetry of movement for flexion and extension (FE), lateral bending (LB) and axial rotation (AR) were derived from angular motion pattern data and the results compared to an earlier established database on asymptomatic riding horses. RESULTS: At walk, horses with back dysfunction had a ROM smaller for dorsoventral FE in the caudal thoracic region (T13 = 7.50 degrees, T17 = 7.71 degrees; P<0.05), greater for LB at T13 (8.13 degrees; P<0.001) and smaller for AR of the pelvis (10.97 degrees; P<0.05) compared to asymptomatic horses (FE-T13 = 8.28 degrees, FE-T17 = 8.49 degrees, LB-T13 = 6.34 degrees, AR-pelvis = 12.77 degrees). At trot, dysfunctional horses had a smaller (P<0.05) ROM for FE at the thoracic lumbar junction (T17 = 2.46 degrees, L1 = 2.60 degrees) compared to asymptomatic horses (FE-T17 = 3.07 degrees, FE-L1 = 3.12 degrees). CONCLUSIONS: The objective measurement technique can detect differences between back kinematics in riding horses with signs of back dysfunction and asymptomatic horses. The clinical manifestation of back pain results in diminished flexion/extension movement at or near the thoracic lumbar junction. However, before applying the method more extensively in practice it is necessary to evaluate it further, including measurements of patients whose diagnoses can be confirmed and long-term follow-ups of back patients after treatment. POTENTIAL RELEVANCE: Since the objective measurement technique can detect small movement differences in back kinematics, it should help to clinically describe and, importantly, objectively detect horses with back pain and dysfunction.
Keywords: Animals; Back/*physiology; Back Pain/diagnosis/physiopathology/*veterinary; Biomechanics; Exercise Test/veterinary; Gait/*physiology; Horse Diseases/diagnosis/*physiopathology; Horses/anatomy & histology/*physiology; Locomotion/physiology; Lumbar Vertebrae/physiology; Range of Motion, Articular; Stress, Mechanical; Thoracic Vertebrae/physiology; Weight-Bearing
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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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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.
Keywords: Animals; Biomechanics; Exercise Test/instrumentation/methods/*veterinary; Forelimb/physiology; Gait; Head/physiology; Hindlimb/physiology; Horses/*physiology; Locomotion/*physiology; Male; Neck/physiology; Physical Conditioning, Animal/methods/*physiology; Posture; Statistics, Nonparametric; Walking/*physiology
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Faber, M. J., van Weeren, P. R., Schepers, M., & Barneveld, A. (2003). Long-term follow-up of manipulative treatment in a horse with back problems. J Vet Med A Physiol Pathol Clin Med, 50(5), 241–245.
Abstract: In order to objectively quantify the effect of manipulation on back-related locomotion anomalies in the horse, a recently developed kinematic measuring technique for the objective quantification of thoracolumbar motion in the horse was applied in a dressage horse that was suffering from a back problem. In this horse, clinically, a right-convex bending (scoliosis) from the 10th thoracic vertebra to the second lumbar vertebra was diagnosed. As a result, there was a marked asymmetric movement of the thoracolumbar spine. Functionally, there was severe loss of performance. Thoracolumbar motion was measured in terms of ventrodorsal flexion, lateral flexion, and axial rotation using an automated gait analysis system. Measurements were repeated before and 2 days after treatment, before the second treatment 3 weeks later, and at 4 weeks and 8 months after the second treatment to assess long-term effect. At the same time, performance of the horse was assessed subjectively by the trainer as well. Symmetry of movement improved dramatically after the first treatment. After this, there was a slight decrease in symmetry, but 8 months after the last treatment the symmetry indexes for the various joints were still considerably better than during the first (pre-treatment) measuring session. Subjectively, the trainer did not notice improvement until after measurement session 4. Between sessions 4 and 5 (at 4 weeks and 8 months after the second treatment) there was a change of trainer. The new trainer did not report any back problem, and succeeded in bringing the horse back to its former level in competition. It is concluded that manipulation had a measurable influence on the kinematics of the thoracolumbar spine. However, it is recognized that an improvement in symmetry of motion is not equivalent to clinical improvement and that other measures, such as changes in management, may be more decisive.
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Holmstrom, M., & Drevemo, S. (1997). Effects of trot quality and collection on the angular velocity in the hindlimbs of riding horses. Equine Vet J Suppl, (23), 62–65.
Abstract: The angular velocities of the hindlimb angles of 14 horses, including 6 Grand Prix dressage horses, 4 horses judged as good at the trot and 4 horses judged as poor, were analysed. The horse material was the same as previously used by Holmstrom (1994) in studies on conformation and trotting gaits in the Swedish Warmblood riding horse. Four consecutive strides of each horse and the corresponding pace were analysed and mean velocity curves (Xh) for each angle were calculated. Before calculation the data were filtered forwards and backwards with a Butterworth third order filter with a cut off frequency of 60 Hz. During the last 60% of the stance phase there were differences between the horses judged as good and poor at the trot in all the analysed hindlimb angles except the femur inclination. The angular velocity in the hock joint, pelvis inclination and hindlimb pendulation was larger in the good horses. The angular velocity of the hindlimb pendulation decreased with collection in the Grand Prix horses. During parts of the stance phase, there was also a gradual decrease in the femur angular velocity from trot at hand to piaffe. In the hock joint, there was no difference in angular velocity between trot at hand and passage during the last 30%. The higher compression of the hock angle and pelvic angle to the horizontal plane probably reflects a higher compression of the whole hindlimb. It probably contributes to the greater springiness in the movements of good young horses and Grand Prix dressage horses. The results from the present study confirmed the importance of storing elastic strain energy for the quality of the dressage horse gaits.
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