[Show abstract][Hide abstract] ABSTRACT: Results: When pressure on the sole of the hindlimb was present, horses were judged as lame (grade 2 on the AAEP scale 1–5) by an experienced clinician. No significant unloading of this limb was found in the group of horses (unloading was observed in 4 animals, but was not detectable in the other 2), but statistically significant effects on back kinematics were detected. The overall flexion-extension (FE) range of motion (ROM) of the vertebral column was increased at walk, especially in the thoracic segments. Axial rotation (AR) ROM of the pelvis was also increased. At trot, the FE ROM was decreased only in the segment L3-L5-S3. During the stance phase of the lame limb, the segment T6-T10-T13 was more flexed and the neck was lowered at both gaits; the thoracolumbar segments were more extended at walk and trot. There were no significant changes in the stride length or protraction-retraction angles in any of the limbs.
[Show abstract][Hide abstract] ABSTRACT: This paper presents a simulation study that was conducted to investigate whether the stereotyped motion pattern observed in human sub-maximal jumping can be interpreted from the perspective of energy expenditure. Human sub-maximal vertical countermovement jumps were compared to jumps simulated with a forward dynamic musculo-skeletal model. This model consisted of four interconnected rigid segments, actuated by six Hill-type muscle actuators. The only independent input of the model was the stimulation of muscles as a function of time. This input was optimized using an objective function, in which targeting a specific sub-maximal height value was combined with minimizing the amount of muscle work produced. The characteristic changes in motion pattern observed in humans jumping to different target heights were reproduced by the model. As the target height was lowered, two major changes occurred in the motion pattern. First, the countermovement amplitude was reduced; this helped to save energy because of reduced dissipation and regeneration of energy in the contractile elements. Second, the contribution of rotation of the heavy proximal segments of the lower limbs to the vertical velocity of the centre of gravity at take-off was less; this helped to save energy because of reduced ineffective rotational energies at take-off. The simulations also revealed that, with the observed movement adaptations, muscle work was reduced through improved relative use of the muscle's elastic properties in sub-maximal jumping. According to the results of the simulations, the stereotyped motion pattern observed in sub-maximal jumping is consistent with the idea that in sub-maximal jumping, subjects are trying to achieve the targeted jump height with minimal energy expenditure.
Full-text · Article · Mar 2008 · Journal of Electromyography and Kinesiology
[Show abstract][Hide abstract] ABSTRACT: In the literature, it has been hotly debated whether the brain uses internal models or equilibrium point (EP) control to generate arm movements. EP control involves specification of EP trajectories, time series of arm configurations in which internal forces and external forces are in equilibrium; if the arm is not in a specified EP, it is driven toward this EP by muscle forces arising due to central drive, reflexes, and muscle mechanics. EP control has been refuted by researchers claiming that EP trajectories underlying movements of subjects were complex. These researchers used an approach that involves applying force perturbations during movements of subjects and fitting a mass-spring-damper model to the kinematic responses, and then reconstructing the EP trajectory using the estimated stiffness, damping, and measured kinematics. In this study, we examined the validity of this approach using an EP-controlled musculoskeletal model of the arm. We used the latter model to simulate unperturbed and perturbed maximally fast movements and optimized the parameter values of a mass-spring-damper model to make it reproduce as best as possible the kinematic responses. It was shown that estimated stiffness not only depended on the "true" stiffness of the musculoskeletal model but on all of its dynamical parameters. Furthermore it was shown that reconstructed EP trajectories were in agreement with those presented in the literature but did not resemble the simple EP trajectories that had been used to generate the movement of the model. It was concluded that the refutation of EP control on the basis of results obtained with mass-spring-damper models was unjust.
No preview · Article · Oct 2007 · Journal of Neurophysiology
[Show abstract][Hide abstract] ABSTRACT: The purpose of this study was to determine whether individual limb forces could be calculated accurately from kinematics of trotting and walking horses. We collected kinematic data and measured vertical ground reaction forces on the individual limbs of seven Warmblood dressage horses, trotting at 3.4 m s(-1) and walking at 1.6 m s(-1) on a treadmill. First, using a segmental model, we calculated from kinematics the total ground reaction force vector and its moment arm relative to each of the hoofs. Second, for phases in which the body was supported by only two limbs, we calculated the individual reaction forces on these limbs. Third, we assumed that the distal limbs operated as linear springs, and determined their force-length relationships using calculated individual limb forces at trot. Finally, we calculated individual limb force-time histories from distal limb lengths. A good correspondence was obtained between calculated and measured individual limb forces. At trot, the average peak vertical reaction force on the forelimb was calculated to be 11.5+/-0.9 N kg(-1) and measured to be 11.7+/-0.9 N kg(-1), and for the hindlimb these values were 9.8+/-0.7 N kg(-1) and 10.0+/-0.6 N kg(-1), respectively. At walk, the average peak vertical reaction force on the forelimb was calculated to be 6.9+/-0.5 N kg(-1) and measured to be 7.1+/-0.3 N kg(-1), and for the hindlimb these values were 4.8+/-0.5 N kg(-1) and 4.7+/-0.3 N kg(-1), respectively. It was concluded that the proposed method of calculating individual limb reaction forces is sufficiently accurate to detect changes in loading reported in the literature for mild to moderate lameness at trot.
[Show abstract][Hide abstract] ABSTRACT: Lameness has often been suggested to result in altered movement of the back, but there are no detailed studies describing such a relationship in quantitative terms.
To quantify the effect of induced subtle forelimb lameness on thoracolumbar kinematics in the horse.
Kinematics of 6 riding horses was measured at walk and at trot on a treadmill before and after the induction of reversible forelimb lameness grade 2 (AAEP scale 1-5). Ground reaction forces (GRF) for individual limbs were calculated from kinematics.
The horses significantly unloaded the painful limb by 11.5% at trot, while unloading at walk was not significant. The overall flexion-extension range of back motion decreased on average by 0.2 degrees at walk and increased by 3.3 degrees at trot (P<0.05). Changes in angular motion patterns of vertebral joints were noted only at trot, with an increase in flexion of 0.9 degrees at T10 (i.e. angle between T6, T10 and T13) during the stance phase of the sound diagonal and an increase in extension of the thoracolumbar area during stance of the lame diagonal (0.7degrees at T13, 0.8 degres at T17, 0.5 degres at L1, 0.4 degrees at L3 and 0.3 degrees at L5) (P<0.05). Lameness further caused a lateral bending of the cranial thoracic vertebral column towards the lame side (1.3 degrees at T10 and 0.9 degrees at T13) (P<0.05) during stance of the lame diagonal.
Both range of motion and vertebral angular motion patterns are affected by subtle forelimb lameness. At walk, the effect is minimal, at trot the horses increased the vertebral range of motion and changed the pattern of thoracolumbar motion in the sagittal and horizontal planes, presumably in an attempt to move the centre of gravity away from the lame side and reduce the force on the affected limb.
Subtle forelimb lameness affects thoracolumbar kinematics. Future studies should aim at elucidating whether the altered movement patterns lead to back and/or neck dysfunction in the case of chronic lameness.
Full-text · Article · Jun 2007 · Equine Veterinary Journal
[Show abstract][Hide abstract] ABSTRACT: According to the equilibrium point theory, the control of posture and movement involves the setting of equilibrium joint positions (EP) and the independent modulation of stiffness. One model of EP control, the alpha-model, posits that stable EPs and stiffness are set open-loop, i.e. without the aid of feedback. The purpose of the present study was to explore for the elbow joint the range over which stable EPs can be set open-loop and to investigate the effect of co-contraction on intrinsic low-frequency elbow joint stiffness (K (ilf)). For this purpose, a model of the upper and lower arm was constructed, equipped with Hill-type muscles. At a constant neural input, the isometric force of the contractile element of the muscles depended on both the myofilamentary overlap and the effect of sarcomere length on the sensitivity of myofilaments to [Ca2+] (LDCS). The musculoskeletal model, for which the parameters were chosen carefully on the basis of physiological literature, captured the salient isometric properties of the muscles spanning the elbow joint. It was found that stable open-loop EPs could be achieved over the whole range of motion of the elbow joint and that K (ilf), which ranged from 18 to 42 N m.rad(-1), could be independently controlled. In the model, LDCS contributed substantially to K (ilf) (up to 25 N m.rad(-1)) and caused K (ilf) to peak at a sub-maximal level of co-contraction.
[Show abstract][Hide abstract] ABSTRACT: Vertical jumping was used to assess muscle mechanical output in bonobos and comparisons were drawn to human jumping. Jump height, defined as the vertical displacement of the body centre of mass during the airborne phase, was determined for three bonobos of varying age and sex. All bonobos reached jump heights above 0.7 m, which greatly exceeds typical human maximal performance (0.3-0.4m). Jumps by one male bonobo (34 kg) and one human male (61.5 kg) were analysed using an inverse dynamics approach. Despite the difference in size, the mechanical output delivered by the bonobo and the human jumper during the push-off was similar: about 450 J, with a peak power output close to 3000 W. In the bonobo, most of the mechanical output was generated at the hips. To account for the mechanical output, the muscles actuating the bonobo's hips (directly and indirectly) must deliver muscle-mass-specific power and work output of 615 Wkg-1 and 92 Jkg-1, respectively. This was twice the output expected on the basis of muscle mass specific work and power in other jumping animals but seems physiologically possible. We suggest that the difference is due to a higher specific force (force per unit of cross-sectional area) in the bonobo.
Full-text · Article · Oct 2006 · Proceedings of the Royal Society B: Biological Sciences
[Show abstract][Hide abstract] ABSTRACT: We hypothesized that the initial rate (first 40 ms) of unilateral knee extensor torque development during a maximally fast isometric contraction would depend on the subjects' ability for fast neural activation and that it would predict bilateral jumping performance.
Nine males (21.8 +/- 0.9 yr, means +/- SD) performed unilateral fast isometric knee extensions (120 degrees knee angle) without countermovement on a dynamometer and bilateral squat jumps (SJ) and countermovement jumps (CMJ) starting from 90 and 120 degrees knee angles (full extension = 180 degrees ). The dynamometer contractions started either from full relaxation or from an isometric pre-tension (15% maximal isometric torque, Tmax). Torque time integral for the first 40 ms after torque onset (TTI-40, normalized to Tmax) and averaged normalized rectified knee extensor EMG for 40 ms before fast torque onset (EMG-40) were used to quantify initial torque rise and voluntary muscle activation.
TTI-40 without pre-tension (range: 0.02-0.19% Tmax per second) was significantly lower than TTI-40 with pre-tension, and both were significantly (r = 0.81 and 0.80) related to EMG-40. During jumping, similar significant positive relations were found between jump height and knee extensor EMG during the first 100 ms of the rise in ground reaction force. There also were significant positive linear relations between dynamometer TTI-40 and jump height (r = 0.75 (SJ 90), 0.84 (SJ 120), 0.76 (CMJ 90), and 0.86 (CMJ 120)) but not between dynamometer Tmax and jump height (-0.16 < r < 0.02).
One-legged TTI-40 to a large extent explained the variation in jump height. The ability to produce a high efferent neural drive before muscle contraction seemed to dominate performance in both the simple single-joint isometric task and the complex multijoint dynamic task.
Full-text · Article · Oct 2006 · Medicine & Science in Sports & Exercise
[Show abstract][Hide abstract] ABSTRACT: There are several ways to quantify jumping performance, a common definition being the height gained by the body's centre of mass (CM) in the airborne phase. Under this definition, jump height is determined by take-off velocity. According to the existing literature on jumping and scaling, take-off velocity, and hence jumping performance is independent of size because the energy that differently sized geometrically scaled jumpers can generate with their muscles is proportional to their mass. In this article it is shown, based on a simple energy balance, that it is incorrect to presume that jump height does not depend on size. Contrary to common belief, size as such has does have an effect on take-off velocity, putting small jumpers at a mechanical advantage, as is shown analytically. To quantify the effect of size on take-off velocity, a generic jumper model was scaled geometrically and evaluated numerically. While a 70-kg jumper took off at 2.65 m/s and raised its CM by 0.36 m after take-off, a perfectly geometrically similar jumper of 0.7 g reached a take-off velocity of 3.46 m/s and raised its CM by 0.61 m. The reason for the better performance of small jumpers is their higher efficacy in transforming the energy generated by the actuators into energy due to vertical velocity of the CM. Considering the ecological and evolutionary relevance of different definitions of jump height, size-dependent efficacy might explain why habitual jumping is especially prominent among small animals such as insects.
[Show abstract][Hide abstract] ABSTRACT: Several types of equilibrium point (EP) controllers have been proposed for the control of posture and movement. EP controllers are appealing from a computational perspective because they do not require solving the "inverse dynamic problem" (i.e., computation of the torques required to move a system along a desired trajectory). It has been argued that EP controllers are not capable of controlling fast single-joint movements. To refute this statement, several extensions have been proposed, although these have been tested using models in which only the tendon compliance, force-length-velocity relation, and mechanical interaction between tendon and contractile element were not adequately represented. In the present study, fast elbow-joint movements were measured and an attempt was made to reproduce these using a realistic musculoskeletal model of the human arm. Three types of EP controllers were evaluated: an open-loop alpha-controller, a closed-loop lambda-controller, and a hybrid open- and closed-loop controller. For each controller we considered a continuous version and a version in which the control signals were sent out intermittently. Only the intermittent hybrid EP controller was capable of generating movements that were as fast as those of the subjects. As a result of the nonlinear muscle properties, the hybrid EP controller requires a more detailed representation of static muscle properties than generally assumed in the context of EP control. In sum, this study shows that fast single-joint movements can be realized without explicitly solving the inverse dynamics problem, but in a less straightforward manner than implied by proponents of conventional EP controllers.
No preview · Article · Jun 2006 · Journal of Neurophysiology
[Show abstract][Hide abstract] ABSTRACT: Evaluation of free jumping at sub-maximal heights is common practice within selection procedures for young breeding stallions. Early training might cause an unjustified bias. To investigate this, data from a 5-year longitudinal study on 30 horses were used. Half of these horses (experimental group) had received early training between 6months and 4years, the other half (control group) had not. Between 4 and 5years, all horses had received standard training under saddle. At the age of 5years, the horses were tested in a puissance competition and the 7 best and 6 worst jumpers were used for the present study. Kinematic variables that were different at ages 5years and 6months, and had been shown to be predictive for performance in earlier studies, were analysed at the age of 4years. It showed that early training had effaced the differences between potentially good and less good show jumpers in 3 of 3 predictive variables and had introduced a (false, because not related to performance) difference between the trained and untrained horses in one of them and a nearly significant trend in another.Early training may to a certain extent obscure differences in talent among individuals at the age at which selection events occur. Experienced judges may be able to account for this, but studbooks and judges should be aware of this possible pitfall.
Full-text · Article · Jun 2006 · Livestock Science
[Show abstract][Hide abstract] ABSTRACT: This study investigated whether changes in lower limb muscle activity occurred in anticipation of a possible perturbation in 11 young (mean age 27 years) and 11 older (mean age 68 years) adults. Altered muscle activity could affect tripping responses and consequently the ecological validity of experimental results of studies on tripping. It was hypothesized that anticipatory muscle activity would be present immediately after a trip, and decrease after several subsequent unperturbed (forewarned) walking trials. Electromyograms of lower limb muscles were measured in 3 conditions: during normal walking, during forewarned walking immediately after a trip, and during forewarned walking several trials after a trip had occurred. Small but statistically significant differences in averaged muscle activity over a stride were found among conditions. Young adults showed slightly increased activity immediately after tripping (co-contraction) in hamstrings, quadriceps and tibialis anterior muscles. This increased activity diminished after several unperturbed trials, although it did not return to the baseline activity levels during normal walking. In older adults, an increased muscle activity among conditions was only discerned in tibialis anterior and soleus muscles. This suggested that older adults prefer to avoid contact with the obstacle over joint stiffening. Yet, for both age-groups, the increases in muscle activity were very small when compared to tripping responses reported in the literature. Therefore, anticipatory effects are not expected to jeopardize the validity of experiments in which subjects are perturbed more than once.
Full-text · Article · May 2006 · Journal of Electromyography and Kinesiology
[Show abstract][Hide abstract] ABSTRACT: In the literature, it has been reported that the mechanical output per leg is less in two-leg jumps than in one-leg jumps. This so-called bilateral deficit has been attributed to a reduced neural drive to muscles in two-leg jumps. The purpose of the present study was to investigate the possible contribution of nonneural factors to the bilateral deficit in jumping. We collected kinematics, ground reaction forces, and electromyograms of eight human subjects performing two-leg and one-leg (right leg) squat jumps and calculated mechanical output per leg. We also used a model of the human musculoskeletal system to simulate two-leg and one-leg jumps, starting from the initial position observed in the subjects. The model had muscle stimulation as input, which was optimized using jump height as performance criterion. The model did not incorporate a reduced maximal neural drive in the two-leg jump. Both in the subjects and in the model, the work of the right leg was more than 20% less in the two-leg jump than in the one-leg jump. Peak electromyogram levels in the two-leg jump were reduced on average by 5%, but the reduction was only statistically significant in m. rectus femoris. In the model, approximately 75% of the bilateral deficit in work per leg was explained by higher shortening velocities in the two-leg jump, and the remainder was explained by lower active state of muscles. It was concluded that the bilateral deficit in jumping is primarily caused by the force-velocity relationship rather than by a reduction of neural drive.
Preview · Article · Mar 2006 · Journal of Applied Physiology
[Show abstract][Hide abstract] ABSTRACT: The purpose of this study was to evaluate a Hill-based mathematical model of muscle energetics and to disclose inconsistencies in existing experimental data. For this purpose, we simulated iso-velocity contractions of mouse fast twitch EDL and slow twitch SOL fibers, and we compared the outcome to experimental results. The experimental results were extracted from two studies published in the literature, which were based on the same methodology but yielded different outcomes (B96 and B93). In the model, energy cost was modeled as the sum of heat and work. Parameters used to model heat rate were entirely independent of the experimental data-sets. Parameters describing the mechanical behavior were derived from both experimental studies. The model was found to accurately predict the muscle energetics and mechanical efficiency of data-set B96. The model could not, however, replicate the energetics and efficiency of SOL and EDL that were found in data-set B93. The model overestimated the shortening heat rate of EDL but, surprisingly, also the mechanical work rate for both muscles. This was surprising since mechanical characteristics of the model were derived directly from the experimental data. It was demonstrated that the inconsistencies in data-set B93 must have been due to some unexplained confounding artifact. It was concluded that the presented model of muscle energetics is valid for iso-velocity contractions of mammalian muscle since it accurately predicts experimental results of an independent data-set (B96). In addition, the model appeared to be helpful in revealing inconsistencies in a second data-set (B93).
Full-text · Article · Feb 2006 · Journal of Biomechanics
[Show abstract][Hide abstract] ABSTRACT: The purpose of this study was to quantify performance characteristics of good jumping horses, and to determine whether these were already detectable at foal age. Kinematic data were collected of horses performing free jumps over a 0.60 m high fence at six months of age and of these same horses jumping with a rider over a 1.15 m high fence at five years of age. At five years of age the horses were divided into three groups on the basis of a puissance competition: a group of seven best jumpers that made no errors and in the end cleared a 1.50 m high fence, a group of nine worst jumpers that were unable to clear a 1.40 m high fence, and an intermediate group of 13 horses. Longitudinal kinematic data was available for all seven best jumpers and for six of the nine worst jumpers. Average values of variables for the best jumpers were compared with those of the worst jumpers for the jumps over 1.15 m. In the group of best jumpers, the forelimbs were shorter at forelimb clearance due to increased elbow flexion, and the hind limbs were further retroflexed at hind limb clearance. The same superior technique in clearing fences with the limbs was also found in this group at six months of age. Nevertheless, for individual horses it turned out to be too far-fetched to predict adult jumping capacity on the basis of kinematic variables collected during submaximal jumps at foal age.
Full-text · Article · Oct 2005 · The Veterinary Journal
[Show abstract][Hide abstract] ABSTRACT: It is well documented that muscle fibers become more sensitive for [Ca2+] with increasing sarcomere length. In mechanical terms this length-dependent [Ca2+] sensitivity (LDCS) adds to the stiffness of muscle fibers, because muscle force, normalized for the force-length relationship at maximal stimulation, increases with contractile element (CE) length. Although LDCS is well-documented in the physiological literature, it is ignored in most motor control studies. The aim of the present study was to investigate the importance of LDCS as a contributor to the stiffness of a muscle. Comparison of experimental data with predictions derived from the model of activation dynamics proposed by Hatze (Myocybernetic Control Models of Skeletal Muscle, University of South Africa, Pretoria, 1981, pp. 31-42) indicated that this model captures the main characteristics of LDCS well. It was shown that LDCS accounts for the experimentally observed shifts in optimum length at sub-maximal stimulation levels. Furthermore, it was shown that in conditions with low-to-medium muscle stimulation, the contribution of LDCS to the total amount of stiffness provided by the muscle is substantial. It was concluded that LDCS is an important muscle property and should be taken into account in studies concerning motor control.
Preview · Article · Oct 2005 · Journal of Biomechanics