Intrinsic Muscle Stiffness Research Articles

Evidence for symmetrically reduced foot mechanics and energetics in patients after trimalleolar fracture repair: A cross-sectional study

BackgroundTrimalleolar fractures osteosynthesis is associated with a suboptimal outcome. It is hypothesized that patients with trimalleolar fractures face different ankle joint kinetics at mid- and long-term causing compensations at the distal foot joints. Research questionDo patients with a history of a trimalleolar fracture demonstrate different foot joint mechanics and energetics (1) between their affected side and their matched controls? (2) between their unaffected side and their matched controls? (3) between their affected side and their unaffected side? MethodsFifteen patients who sustained a trimalleolar fracture and underwent osteosynthesis for both the lateral, medial and posterior malleolus were compared to a asymptomatic control group which was matched for sex, age and walking speed. Three-dimensional gait analysis was used to quantify kinetic parameters in the Ankle, Chopart, Lisfranc and first metatarsophalangeal joint through a multi-segment kinetic foot model. Statistical analysis was performed using a Univariate Analysis of Covariance and/or a paired t-test. ResultsThe peak internal ankle moment was significantly lower in patients when compared to the control group (p < 0.001). Mean peak power generation and total positive work were significantly lower for the Chopart joint when comparing the patients to the control group (p < 0.001). These results were observed for both the affected and unaffected side of the patients compared to the control group, showing symmetrical changes in the patient group. SignificanceDespite adequate radiographic quality of reduction and the fact that all patients were treated according to a fixed postoperative protocol, this study indicates that patients with a history of a trimalleolar fracture demonstrate reduced foot joint kinetics. It is hypothesized that these findings originate from extrinsic and intrinsic foot muscle strength, stiffness and pain. Future research is needed to validate this hypothesis.

Constraints on the deformation of the vibrissa within the follicle

Nearly all mammals have a vibrissal system specialized for tactile sensation, composed of whiskers growing from sensor-rich follicles in the skin. When a whisker deflects against an object, it deforms within the follicle and exerts forces on the mechanoreceptors inside. In addition, during active whisking behavior, muscle contractions around the follicle and increases in blood pressure in the ring sinus will affect the whisker deformation profile. To date, however, it is not yet possible to experimentally measure how the whisker deforms in an intact follicle or its effects on different groups of mechanoreceptors. The present study develops a novel model to predict vibrissal deformation within the follicle sinus complex. The model is based on experimental results from a previous ex vivo study on whisker deformation within the follicle, and on a new histological analysis of follicle tissue. It is then used to simulate whisker deformation within the follicle during passive touch and active whisking. Results suggest that the most likely whisker deformation profile is “S-shaped,” crossing the midline of the follicle right below the ring sinus. Simulations of active whisking indicate that an increase in overall muscle stiffness, an increase in the ratio between deep and superficial intrinsic muscle stiffness, and an increase in sinus blood pressure will all enhance tactile sensitivity. Finally, we discuss how the deformation profiles might map to the responses of primary afferents of each mechanoreceptor type. The mechanical model presented in this study is an important first step in simulating mechanical interactions within whisker follicles.

Molecular Mechanisms of Muscle Tone Impairment under Conditions of Real and Simulated Space Flight.

Kozlovskaya et al. [1] and Grigoriev et al. [2] showed that enormous loss of muscle stiffness (atonia) develops in humans under true (space flight) and simulated microgravity conditions as early as after the first days of exposure. This phenomenon is attributed to the inactivation of slow motor units and called reflectory atonia. However, a lot of evidence indicating that even isolated muscle or a single fiber possesses substantial stiffness was published at the end of the 20th century. This intrinsic stiffness is determined by the active component, i.e. the ability to form actin-myosin cross-bridges during muscle stretch and contraction, as well as by cytoskeletal and extracellular matrix proteins, capable of resisting muscle stretch. The main facts on intrinsic muscle stiffness under conditions of gravitational unloading are considered in this review. The data obtained in studies of humans under dry immersion and rodent hindlimb suspension is analyzed. The results and hypotheses regarding reduced probability of cross-bridge formation in an atrophying muscle due to increased interfilament spacing are described. The evidence of cytoskeletal protein (titin, nebulin, etc.) degradation during gravitational unloading is also discussed. The possible mechanisms underlying structural changes in skeletal muscle collagen and its role in reducing intrinsic muscle stiffness are presented. The molecular mechanisms of changes in intrinsic stiffness during space flight and simulated microgravity are reviewed.

Contractile properties of superficial skeletal muscle affect postural control in healthy young adults: A test of the rambling and trembling hypothesis.

The rambling and trembling analysis separates the center of pressure (COP) fluctuations into two components: rambling (supraspinal contribution) and trembling (muscle stiffness / reflexive properties contribution). We examined whether the trembling component is correlated to the contractile properties (muscle stiffness and contraction time) of lower limb superficial skeletal muscles to experimentally test the rambling and trembling hypothesis. We hypothesized that muscle stiffness and contraction time, would be: (a) more correlated with; and (b) have a greater impact on the trembling component compared to the rambling component. Thirty-two healthy young adults were recruited for the study and tensiomyography was used to assess mechanical muscle responses to a single electrical stimulus to calculate muscle stiffness and contraction time based on radial muscle belly displacement measurements of lower limb muscles unilaterally. Moreover, upright postural control was assessed using a force plate to record ground reaction forces and moments and calculate the COP fluctuations during two 30 seconds trials. From the COP fluctuations, rambling and trembling time series were extracted, and all fluctuation time series were described using a number of different time-domain and frequency-domain parameters in both the anterior-posterior and medial-lateral directions. Our results demonstrated that both muscle stiffness and contraction time were moderately correlated with time-domain and frequency-domain parameters of the trembling component, as compared with those of the rambling component which was not as well correlated. Moreover, they also predicted the trembling component better. Overall, these results imply that postural control during quiet stance is, in part, related to intrinsic muscle stiffness in the lower extremities. Moreover, we showed that the rambling and trembling hypothesis is effective in separating postural sway fluctuations during upright posture to extract the contributions of muscle stiffness / reflexive properties (trembling), and likely the supraspinal contribution (rambling).

Manipulation of visual information affects control strategy during a visuomotor tracking task

Proper understanding of motor control requires insight into the extent and manner in which task performance and control strategy are influenced by various aspects of visual information. We therefore systematically manipulated the visual presentation (i.e., scaling factor and optical flow density) of a visuomotor tracking task without changing the task itself, and investigated the effect on performance, effort, motor control strategy (i.e., anticipatory or corrective steering) and underlying neuromechanical parameters (i.e., intrinsic muscle stiffness and damping, and proprioceptive and visual feedback). Twenty healthy participants controlled the left-right position of a virtual car (by means of wrist rotations in a haptic robot) to track a slightly curved virtual road (presented on a 60” LED screen), while small torque perturbations were applied to the wrist (1.25–20Hz multisine) for quantification of the neuromechanical parameters. This visuomotor tracking task was performed in conditions with low/medium/high scaling factor and low/high optical flow density. Task performance was high in all conditions (tracking accuracy 96.6%–100%); a higher scaling factor was associated with slightly better performance. As expected, participants did adapt their control strategy and the use of proprioceptive and visual feedback in response to changes in the visual presentation. These findings indicate that effects of visual representation on motor behavior should be taken into consideration in designing, interpreting and comparing experiments on motor control in health and disease. In future studies, these insights might be exploited to assess the sensory-motor adaptability in various clinical conditions.

Effects of plyometric training on passive stiffness of gastrocnemii muscles and Achilles tendon

Plyometric training is commonly used to improve athletic performance; however, it is unclear how each component of the muscle-tendon complex (MTC) is affected by this intervention. The effects of 14weeks of plyometric training on the passive stiffness of the gastrocnemii muscles and Achilles tendon was determined simultaneously to assess possible local adaptations of elastic properties. The passive force-length relationship of the gastrocnemii MTC and elongation of the gastrocnemii muscles were determined using ultrasonography during passive cyclic stretching in 19 subjects divided into trained (n=9) and control (n=10) groups. An upward trend in stiffness of the gastrocnemii MTC (P=0.09) and a significant increase in the intrinsic gastrocnemii muscle stiffness were found (P<0.05). In contrast, no significant change in gastrocnemii tendon stiffness, or in muscle and tendon geometry, was determined (P>0.05). Considering the lack of change in gastrocnemii muscle geometry, the change in the gastrocnemii muscle stiffness may be mainly due to a change in the intrinsic mechanical properties of the muscular tissues.

Quantification of the effects of an alpha-2 adrenergic agonist on reflex properties in spinal cord injury using a system identification technique

BackgroundDespite numerous investigations, the impact of tizanidine, an anti-spastic medication, on changes in reflex and muscle mechanical properties in spasticity remains unclear. This study was designed to help us understand the mechanisms of action of tizanidine on spasticity in spinal cord injured subjects with incomplete injury, by quantifying the effects of a single dose of tizanidine on ankle muscle intrinsic and reflex components.MethodsA series of perturbations was applied to the spastic ankle joint of twenty-one spinal cord injured subjects, and the resulting torques were recorded. A parallel-cascade system identification method was used to separate intrinsic and reflex torques, and to identify the contribution of these components to dynamic ankle stiffness at different ankle positions, while subjects remained relaxed.ResultsFollowing administration of a single oral dose of Tizanidine, stretch evoked joint torque at the ankle decreased significantly (p < 0.001) The peak-torque was reduced between 15% and 60% among the spinal cord injured subjects, and the average reduction was 25%. Using systems identification techniques, we found that this reduced torque could be attributed largely to a reduced reflex response, without measurable change in the muscle contribution. Reflex stiffness decreased significantly across a range of joint angles (p < 0.001) after using tizanidine. In contrast, there were no significant changes in intrinsic muscle stiffness after the administration of tizanidine.ConclusionsOur findings demonstrate that tizanidine acts to reduce reflex mechanical responses substantially, without inducing comparable changes in intrinsic muscle properties in individuals with spinal cord injury. Thus, the pre-post difference in joint mechanical properties can be attributed to reflex changes alone. From a practical standpoint, use of a single "test" dose of Tizanidine may help clinicians decide whether the drug can helpful in controlling symptoms in particular subjects.

POSTURAL RESPONSES EVOKED BY PLATFORM PERTUBATIONS ARE DOMINATED BY CONTINUOUS FEEDBACK

This paper reviews different approaches for explaining body sway while quiet standing that directly address the instability of the human inverted pendulum. We argue that both stiffness control [Winter, D. A., Patla, A. E., Riedtyk, S., & Ishac, M. (2001). Ankle muscle stiffness in the control of balance during quiet standing. Journal of Neurophysiology, 85, 2630–2633] and continuous feedback control by means of a PID (Proportional, Integral, Derivative) mechanism [Peterka, R. J. (2000). Postural control model interpretation of stabilogram diffusion analysis. Biological Cybernetics, 83, 335–343.] can guarantee asymptotic stability of controlled posture at the expense of unrealistic assumptions: the level of intrinsic muscle stiffness in the former case, and the level of background noise in the latter, which also determines an unrealistic level of jerkiness in the sway. We show that the decomposition of the control action into a slow and a fast component (rambling and trembling, respectively, as proposed by [Zatsiorsky, V. M., & Duarte, M. (1999). Instant equilibrium point and its migration in standing tasks: Rambling and trembling components of the stabilogram. Motor Control, 4, 185–200; Zatsiorsky, V. M., & Duarte, M. (2000). Rambling and trembling in quiet standing. Motor Control, 4, 185–200.]) is useful but must be modified in order to take into account that rambling is not a stable equilibrium trajectory. We address the possibility of a form of stability weaker than asymptotic stability in light of the intermittent stabilization mechanism outlined by [Loram, I. D., & Lakie, M. (2002a). Human balancing of an inverted pendulum: position control by small, ballistic-like, throw and catch movements. Journal of Physiology, 540, 1111–1124.], and propose an indicator of intermittent stabilization that is related to the phase portrait of the human inverted pendulum. This indicator provides a further argument against the plausibility of PID-like control mechanisms. Finally, we draw attention to the sliding mode control theory that provides a useful theoretical framework for formulating realistic intermittent, stabilization models.

Role of reflex dynamics in spinal stability: Intrinsic muscle stiffness alone is insufficient for stability

Spinal stability is related to both the intrinsic stiffness of active muscle as well as neuromuscular reflex response. However, existing analyses of spinal stability ignore the role of the reflex response, focusing solely on the intrinsic muscle stiffness associated with voluntary activation patterns in the torso musculature. The goal of this study was to empirically characterize the role of reflex components of spinal stability during voluntary trunk extension exertions. Pseudorandom position perturbations of the torso and associated driving forces were recorded in 11 healthy adults. Nonlinear systems-identification analyses of the measured data provided an estimate of total systems dynamics that explained 81% of the movement variability. Proportional intrinsic response was less than zero in more than 60% of the trials, e.g. mean value of P INT during the 20% maximum voluntary exertion trunk extension exertions −415±354 N/m. The negative value indicated that the intrinsic muscle stiffness was not sufficient to stabilize the spine without reflex response. Reflexes accounted for 42% of the total stabilizing trunk stiffness. Both intrinsic and reflex components of stiffness increased significantly with trunk extension effort. Results reveal that reflex dynamics are a necessary component in the stabilizing control of spinal stability.

Body sway during quiet standing: Is it the residual chattering of an intermittent stabilization process?

This paper reviews different approaches for explaining body sway while quiet standing that directly address the instability of the human inverted pendulum. We argue that both stiffness control [Winter, D. A., Patla, A. E., Riedtyk, S., & Ishac, M. (2001). Ankle muscle stiffness in the control of balance during quiet standing. Journal of Neurophysiology, 85, 2630–2633] and continuous feedback control by means of a PID (Proportional, Integral, Derivative) mechanism [Peterka, R. J. (2000). Postural control model interpretation of stabilogram diffusion analysis. Biological Cybernetics, 83, 335–343.] can guarantee asymptotic stability of controlled posture at the expense of unrealistic assumptions: the level of intrinsic muscle stiffness in the former case, and the level of background noise in the latter, which also determines an unrealistic level of jerkiness in the sway. We show that the decomposition of the control action into a slow and a fast component (rambling and trembling, respectively, as proposed by [Zatsiorsky, V. M., & Duarte, M. (1999). Instant equilibrium point and its migration in standing tasks: Rambling and trembling components of the stabilogram. Motor Control, 4, 185–200; Zatsiorsky, V. M., & Duarte, M. (2000). Rambling and trembling in quiet standing. Motor Control, 4, 185–200.]) is useful but must be modified in order to take into account that rambling is not a stable equilibrium trajectory. We address the possibility of a form of stability weaker than asymptotic stability in light of the intermittent stabilization mechanism outlined by [Loram, I. D., & Lakie, M. (2002a). Human balancing of an inverted pendulum: position control by small, ballistic-like, throw and catch movements. Journal of Physiology, 540, 1111–1124.], and propose an indicator of intermittent stabilization that is related to the phase portrait of the human inverted pendulum. This indicator provides a further argument against the plausibility of PID-like control mechanisms. Finally, we draw attention to the sliding mode control theory that provides a useful theoretical framework for formulating realistic intermittent, stabilization models.

2.16 Dynamic electromyographic activities of lower extremity muscles in patients with osteoarthritis of the knees

This paper reviews different approaches for explaining body sway while quiet standing that directly address the instability of the human inverted pendulum. We argue that both stiffness control [Winter, D. A., Patla, A. E., Riedtyk, S., & Ishac, M. (2001). Ankle muscle stiffness in the control of balance during quiet standing. Journal of Neurophysiology, 85, 2630–2633] and continuous feedback control by means of a PID (Proportional, Integral, Derivative) mechanism [Peterka, R. J. (2000). Postural control model interpretation of stabilogram diffusion analysis. Biological Cybernetics, 83, 335–343.] can guarantee asymptotic stability of controlled posture at the expense of unrealistic assumptions: the level of intrinsic muscle stiffness in the former case, and the level of background noise in the latter, which also determines an unrealistic level of jerkiness in the sway. We show that the decomposition of the control action into a slow and a fast component (rambling and trembling, respectively, as proposed by [Zatsiorsky, V. M., & Duarte, M. (1999). Instant equilibrium point and its migration in standing tasks: Rambling and trembling components of the stabilogram. Motor Control, 4, 185–200; Zatsiorsky, V. M., & Duarte, M. (2000). Rambling and trembling in quiet standing. Motor Control, 4, 185–200.]) is useful but must be modified in order to take into account that rambling is not a stable equilibrium trajectory. We address the possibility of a form of stability weaker than asymptotic stability in light of the intermittent stabilization mechanism outlined by [Loram, I. D., & Lakie, M. (2002a). Human balancing of an inverted pendulum: position control by small, ballistic-like, throw and catch movements. Journal of Physiology, 540, 1111–1124.], and propose an indicator of intermittent stabilization that is related to the phase portrait of the human inverted pendulum. This indicator provides a further argument against the plausibility of PID-like control mechanisms. Finally, we draw attention to the sliding mode control theory that provides a useful theoretical framework for formulating realistic intermittent, stabilization models.

Active stiffness of the ankle in response to inertial and elastic loads

Effective stiffness of the musculoskeletal system was examined as a function of the characteristics of an external load. Thirteen healthy subjects provided active contraction of the ankle plantarflexion musculature in a neutral ankle posture to support an external load. Musculoskeletal stiffness was computed from kinetic data recorded in response to dorsiflexion/plantarflexion perturbations. Ankle dynamics were recorded while supporting external loads of 19 and 38 kg with and without antagonistic co-contraction. External loads were applied using pure gravitational mass. In separate trials external loads were applied from stretch of steel springs in parallel with the plantarflexion musculature that also provided added parallel stiffness to the system. Adding external stiffness of 4.9 and 8.1 kN/m surprisingly failed to significantly change the stiffness of the ankle-plus-spring system. This suggests contributions from intrinsic muscle stiffness and reflex stiffness declined in response to added external stiffness. This could not be explained by load magnitudes, ankle postures, or co-activation as these were similar between the inertial and elastic loading conditions. However, non-linear parametric analyses suggest mean intrinsic stiffness of 35.5 kN/m and reflex gain of 11.6 kN/m with a constant reflex delay of 70 ms accurately described the empirical results. The phase response between the mechanical dynamics of the musculoskeletal system and delayed neuromotor feedback combine to provide robust control of system behavior.

Bifurcation and stability analysis in musculoskeletal systems: a study in human stance.

Reflexes are important in the control of such daily activities as standing and walking. The goal of this study is to establish how reflexive feedback of muscle length, velocity, and force can lead to stable equilibria (i.e., posture) and limit cycles (e.g., ankle clonus and gait). The influence of stretch reflexes on the behavior and stability of musculoskeletal systems was examined using a model of human stance. We computed branches of fold and Hopf bifurcations by numerical bifurcation analysis of the model. These fold and Hopf branches divide the parameter space, constructed by the reflexive feedback gains, into regions of different behavior: unstable posture, stable posture, and stable limit cycles. These limit cycles correspond to a neural deficiency, termed ankle clonus. We also linked bifurcation analysis to known biomechanical concepts by linearizing the model: the fold branch corresponds to zero ankle stiffness and defines the minimal muscle length feedback necessary for stable posture; the Hopf branch is related to unstable reflex loops. Crossing the Hopf branch can lead to the above-mentioned stable limit cycles. The Hopf branch reduces with increasing time delays, making the subject's posture more susceptible to unstable reflex loops. This might be one of the reasons why elderly people, or those with injuries to the central nervous system, often have trouble with standing and other posture tasks. The influence of cocontraction and force feedback on the behavior of the posture model was also investigated. An increase in cocontraction leads to an increase in ankle stiffness (i.e., intrinsic muscle stiffness) and a decrease in the effective reflex loop gain. On the one hand, positive force feedback increases the ankle stiffness (i.e., intrinsic and reflexive muscle stiffness); on the other hand it makes the posture more susceptible to unstable reflex loops. For negative force feedback, the opposite is true. Finally, we calculated areas of reflex gains for perturbed stance and quiet stance in healthy subjects by fitting the model to data from the literature. The overlap of these areas of reflex gains could indicate that stretch reflexes are the major control mechanisms in both quiet and perturbed stance. In conclusion, this study has successfully combined bifurcation analysis with the more common biomechanical concepts and tools to determine the influence of reflexes on the stability and quality of stance. In the future, we will develop this line of research to look at rhythmic tasks, such as walking.

Elbow impedance during goal-directed movements.

The mechanical properties and reflex actions of muscles crossing the elbow joint were examined during a 60-deg voluntary elbow extension movement. Brief unexpected torque pulses of identical magnitude and time-course (20-Nm extension switching to 20-Nm flexion within 30 ms) were introduced at various points of a movement in randomly selected trials. Single pulses were injected in different trials, some before movement onset and some either during early, mid, late or ending stages of the movement. Changes in movement trajectory induced by a torque pulse were determined over the first 50 ms by a nearest-neighbor prediction algorithm, and then a modified K-B-I (stiffness-damping-inertia) model was fit to the responses. The stiffness and damping coefficients estimated during voluntary movements were compared to values recorded during trials in which subjects were instructed to strongly co-contract while maintaining a static posture. This latter protocol was designed to help determine the maximum impedance a subject could generate. We determined that co-contraction increased joint stiffness greatly, well beyond that recorded under control conditions. In contrast, the stiffness magnitudes were quite small during routine voluntary movements, or when the subjects relaxed their limb. Furthermore, the damping coefficients were always significant and increased measurably at the end of movement. Reflex activity, as measured by EMG responses in biceps and triceps brachii, showed highly variable responses at latencies of 160 ms or greater. These reflexes tended to activate both elbow flexors and extensors simultaneously. These findings suggest that very low intrinsic muscle stiffness values recorded during point-to-point motion render an equilibrium point or impedance control approach implausible as a means to regulate movement trajectories. In particular, muscle that is shortening against inertial loads seems to exhibit much smaller stiffness than similarly active isometric muscle, although some degree of damping is always present and does not simply co-vary with stiffness. Although the limb muscles can be co-contracted statically or during movement with an observable increase in stiffness and even task performance, this control strategy is rarely utilized, presumably due to the greater energetic cost.

The effect of eccentric exercise on intrinsic and reflex stiffness in the human hand

Objective. The purpose of this work was to determine the effect of strenuous eccentric exercise on joint stiffness and to separate joint stiffness into components due to intrinsic muscle mechanics and delayed reflex muscle activation. Design. Subjects performed 100 maximal eccentric contractions, using the first dorsal interosseus muscle to abduct the index finger while undergoing a 20° displacement of the metacarpophalangeal joint. Joint stiffness was measured 24 h later during 15% and 65% maximal voluntary contraction and during electrical muscle stimulation at 15% of maximal voluntary contraction torque. Background. Joint stiffness can be varied by changing voluntary muscle activation and thereby serves an important role in joint stabilization. Eccentric exercise has been shown to result in muscle fiber injury, reducing maximal muscle force. However, it is not known whether intrinsic muscle stiffness or reflex stiffness is also affected. Methods. Displacements of 3° amplitude were used to estimate joint stiffness about the neutral angle of the index finger. The difference between measurements made during voluntary muscle activation and electrical muscle stimulation was used to obtain reflex stiffness. Results. There was no change in the passive joint stiffness nor was there any change in either the intrinsic or reflex stiffness at 15% maximal voluntary contraction. However, intrinsic stiffness for the electrically stimulated muscle was higher post-exercise than pre-exercise, while active joint stiffness at 65% maximal voluntary contraction (comprising intrinsic and reflex stiffness) was lower. Conclusion. The observed mechanical changes are compatible with the hypothesis that type II muscle fibers are more susceptible to injury than type I muscle fibers, which have higher intrinsic stiffness. Relevance Muscle stiffness is important for maintaining mechanical stability of a joint. The effects of eccentric exercise on muscle stiffness are likely analogous to the effects of muscle injury, making this a good model for biomechanical changes associated with muscle injury.

Impedance characteristics of a neuromusculoskeletal model of the human arm I. Posture control.

The mechanical impedance of neuromusculoskeletal models of the human arm is studied in this paper. The model analysis provides a better understanding of the contributions of possible intrinsic and reflexive components of arm impedance, makes clear the limitations of second-order mass-viscosity-stiffness models and reveals possible task effects on the impedance. The musculoskeletal model describes planar movements of the upper arm and forearm, which are moved by six lumped muscles with nonlinear dynamics. The motor control system is represented by a neural network which combines feedforward and feedback control. It is optimized for the control of movements or for posture control in the presence of external forces. The achieved impedance characteristics depend on the conditions during the learning process. In particular, the impedance is adapted in a suitable way to the frequency content and direction of external forces acting on the hand during an isometric task. The impedance characteristics of a model, which is optimized for movement control, are similar to experimental data in the literature. The achieved stiffness is, to a large extent, reflexively determined whereas the approximated viscosity is primarily due to intrinsic attributes. It is argued that usually applied Hill-type muscle models do not properly represent intrinsic muscle stiffness.

Assessing muscle stiffness from quiet stance in Parkinson's disease.

In previous studies, we developed a postural stiffness measure that is extracted from foot center-of-pressure (COP) trajectories from quietly standing individuals and is based on an analytical mechanical model of posture control. Here we apply this measure to patients with Parkinson's disease (PD). We correlated the postural stiffness measure with different clinical rating scales, obtained from patients. Kendall's rank correlation was highly significant between the stiffness measure and rigidity, bradykinesia, posture impairment, gait, and leg agility, respectively, as rated by the Unified Parkinson's Disease Rating Scale. These results provide further evidence that a higher intrinsic muscle stiffness may contribute to the aforementioned clinically defined symptoms. From a clinical standpoint, this work indicates that the proposed postural stiffness measure may be useful as an assessment tool for the evaluation of PD patients subsequent to pharmacological and surgical treatment.

Gain of the triceps surae stretch reflex in decerebrate and spinal cats during postural and locomotor activities.

1. The triceps surae (TS) stretch reflex was measured in decerebrate cats during crossed extensor stimulation and after spinalization during rhythmic locomotor activity induced by clonidine and manual perineal stimulation. The TS force in response to sinusoidal stretch was measured at a given contraction level before and after deafferentation, and the 'reflex force' was computed by subtracting these two responses. Reflex 'gain' was computed as the ratio of the reflex and deafferented force responses (a unitless estimate of the open loop feedback gain). 2. Prior to locomotion the spontaneous muscle activity was low (less than 15% of maximum), but the reflex gain was relatively high (close to 1.0 with a 5 Hz stretch). When locomotion commenced the reflex gain was markedly lowered when measured at the same contraction level as before locomotion (25% of the gain prior to locomotion). At higher contraction levels the reflex gain was not significantly increased. The reflex force and EMG responses to stretch increased with the contraction level, but their effect on the total reflex gain was cancelled by an associated increase in the intrinsic muscle stiffness. 3. In the decerebrate cat, during weak tonic contractions (spontaneous), the reflex gain was high and comparable with the gain in the resting spinal cat. However, with increased tonic contractions produced by crossed extensor stimulation the reflex gain dropped. At higher contraction levels the gain was not significantly different from the gain during spinal locomotion. 4. When the frequency of stretch was increased from 3 to 20 Hz, EMG responses to stretch increased, but the reflex force decreased, since a more fused contraction developed with the more frequent reflex activations. Overall, the reflex gain decreased with frequency in both spinal and decerebrate cats. The phase lag of the reflex force, relative to the intrinsic muscle force, increased with increasing frequency, due to reflex delays, with a 180 deg lag occurring between 12 and 18 Hz (tremor frequencies). The mean gain was significantly lower and the phase lag was significantly greater during locomotion than during tonic crossed extensor contractions, suggesting different reflex mechanisms. 5. In conclusion, during locomotion in spinal cats afferent feedback from low frequency ankle movements, similar to those occurring during the normal step cycle, reflexly produces a small but significant fraction of the extensor force (about a quarter of the stretch-related force modulation). This fraction is remarkably constant at the different contraction levels of the step cycle. Afferent feedback during higher frequency movement is less effective, minimizing the chance of instability and tremor. In contrast during tonic contractions afferent feedback produces half of the total muscle force during perturbations, clearly contributing to the maintenance of posture.

Catching a ball: contributions of intrinsic muscle stiffness, reflexes, and higher order responses

In three sets of experiments in nine normal subjects and a patient with a percutaneous wrist-stabilizing splint, we quantified the open-loop gain (OLG) of the stretch reflex acting about the elbow. The subjects exerted a steady mean flexing force and were instructed not to intervene (i.e., not to resist actively) when force or displacement perturbations were imposed on the forearm. The method was either to reconstruct transmission around the entire loop in a two-part experiment, or to use the attenuation of external perturbations in normal and electrically stimulated muscle to compute gain. Across all experiments, the mean magnitude of stretch reflex OLG was close to unity in the frequency range 1-2 Hz, and declined at higher frequencies, as required to ensure stability, given that the phase lag approached 180 degrees at 5 Hz. Inherent muscle stiffness was approximately equal to reflex stiffness. In functional terms, an OLG of 1 means that the yield caused by a force perturbation is approximately halved by reflex action (prevailing inherent muscle stiffness is doubled). Automatic scaling of reflex transmission at Ia/alpha-motoneuronal synapses ensures that the OLG remains close to unity as inherent stiffness increases. Trials in the patient with the wrist fixator gave similar results, indicating that the reflexes were proprioceptive ly mediated. In a fourth experiment in which the task was to catch a heavy ball, we compared the efficacy of inherent muscle stiffness and reflexes alone, with the subject's intentional reactions, which included predictive and voluntary components of response. The latter were far more effective in maintaining the position of the hand after the ball was caught than inherent and reflex stiffnesses alone. We conclude that stability requirements limit the extent to which stretch reflexes can augment inherent muscle stiffness. When inherent muscle stiffness is low, such as in our ball-catching task, the reflex stiffness is also low, and predictive and pre-programmed reactions predominate in load compensation, thus shifting the emphasis from automatic servo or equilibrium-point behaviour to higher order control.

Stretch reflex responses in the human elbow joint during a voluntary movement.

1. The responsiveness of the stretch reflex is modulated during human voluntary limb movements. The influence of this modulation on the limb mechanical properties (stiffness) was investigated. 2. Subjects were taught to replicate accurately a rapid (4.0 rad s-1) targeted elbow flexion movement of 1 rad. From the onset of 12% of the trials a sinusoidal position disturbance (0.05 rad) was superimposed on the normal (trained) movement trajectory. The net joint torque (muscle torque) resisting these stretches was computed from measurements of applied torque, acceleration and limb inertia. Electromyographic (EMG) responses in the triceps brachii (TB), brachialis (Br) and biceps brachii (BB) were monitored. 3. The EMG responses to sinusoidal stretches applied early in the movement were less than those responses to perturbations applied when the arm neared the target (especially in the antagonist muscle TB). These EMG responses caused fluctuations in the resistance to the perturbation (stiffness), as described below. 4. When the perturbation frequency was low (< 4 Hz) the resistance of the elbow muscles to the stretch increased as the arm approached the target (48% increase). In contrast, when the stretch frequency was 7 Hz the resistance decreased by 63%. This decrease can be explained by the increased reflex response, since at 7 Hz the reflex response is probably timed so that it assists, rather than resists, the stretching as a result of loop delays. This reflex timing was confirmed by observing that, after abruptly stopping the sinusoidal stretch, the reflex response persisted for 100 ms and was indeed in a direction that would have reduced the resistance, had the perturbation continued. 5. The time course of the net muscle stiffness was estimated for frequencies ranging from 4 to 8 Hz and for each 40 ms interval a Nyquist plot was constructed, forming a C-shaped curve as frequency was varied. The size of this curve gave a measure of the stiffness resulting from reflex activity. When the arm neared the target this reflexive stiffness reached a maximum, and was probably comparable in size to the intrinsic (non-reflexive) muscle stiffness. Also, in four of the five subjects the viscous component of stiffness at 7 Hz dropped significantly below zero when the arm neared the target, again indicating that at this frequency the reflex was large and acted inappropriately.(ABSTRACT TRUNCATED AT 400 WORDS)