Wrist splint effects on muscle activity and force during a handgrip task

Wrist splints are commonly prescribed to limit wrist motion and provide support at night and during inactive periods but are often used in the workplace. In theory, splinting the wrist should reduce wrist extensor muscle activity by stabilizing the joint and reducing the need for co-contraction to maintain posture. Ten healthy volunteers underwent a series of 24 10-s gripping trials with surface electromyography on 6 forearm muscles. Trials were randomized between splinted and nonsplinted conditions with three wrist postures (30 degrees flexion, neutral, and 30 degrees extension) and four grip efforts. Custom-made Plexiglas splints were taped to the dorsum of the hand and wrist. It was found that when simply holding the dynamometer, use of a splint led to a small (<1% MVE) but significant reduction in activity for all flexor muscles and extensor carpi radialis (all activity <4% maximum). At maximal grip, extensor muscle activity was significantly increased with the splints by 7.9-23.9% MVE. These data indicate that splinting at low-to-moderate grip forces may act to support the wrist against external loading, but appears counterproductive when exerting maximal forces. Wrist bracing should be limited to periods of no to light activity and avoided during tasks that require heavy efforts.     

Co-contraction of the pronator teres and extensor carpi radialis during wrist extension movements in humans.

In order to elucidate the functional significance of excitatory spinal reflex arcs (facilitation) between musculus (M.) pronator teres (PT) and M. extensor carpi radialis (ECR, longus: ECRL, brevis: ECRB) in humans, activities of the muscles were studied with electromyography (EMG) and electrical neuromuscular stimulation (ENS). In EMG study, activities of PT, ECRL, ECRB, and M. flexor carpi radialis during repetitive static (isometric) wrist extension and a series of a dynamic motion of wrist flexion/extension in the prone, semiprone, and supine positions of the forearm were recorded in 12 healthy human subjects. In the prone, semiprone, and supine positions, PT and ECR showed parallel activities during the static extension in all, eight, and eight subjects, respectively, and at the extension phase during the dynamic motion in all, eight and five subjects, respectively. These findings suggest that co-contraction of PT and ECR occurs during wrist extension movements at least with the prone forearm. The facilitation must be active during the co-contraction. In ENS study, ENS to PT was examined in 11 out of the 12 and that to ECRL was in the 12 subjects. Before ENS, the forearm was in the prone, semiprone, and supine positions. In all the subjects, ENS to PT induced a motion of forearm pronation to the maximum pronation. ENS to ECRL induced motions of wrist extension to the maximum extension and abduction (radial flexion) to 5-20 degrees of abduction regardless of the positions of the forearm. Moreover, it induced 30-80 degrees supination of the forearm from the prone position. Consequently, combined ENS to PT and ECRL resulted in motions of the extension and abduction while keeping the maximum pronation. These findings suggest that the co-contraction of PT and ECR during wrist extension movements occurs to prevent supinating the forearm. Forearm supination from the prone position should be added to one of the actions of ECRL.     

Posture and hand load alter muscular response to sudden elbow perturbations

Joint stiffness and stability are reliant on coordinated muscle activity which may differ depending on initial posture and loading during sudden perturbations. This study investigated the effects of arm posture and hand load on muscle activity during perturbations of the arm. Fifteen male participants experienced perturbations to the wrist causing elbow extension using a combination of three body postures (standing, supine, sitting) and three hand load conditions (no, solid, and fluid loads), with known and unknown timing. Surface EMG was collected from eight muscles of the right upper extremity. The response to sudden loading was examined using muscle activities pre (baseline) and post (reflex) perturbation. During the baseline period, known perturbation timing resulted in greater muscular activity than for unknown timing, while the opposite was found for the reflex period. During the reflex period with fluid load, biceps brachii and brachioradialis demonstrated increased activity of 2.4% and 4.0% of maximum respectively, from supine to standing. During the reflex period, the fluid load resulted in forearm co-contraction 23% and 47% greater than the solid and no load conditions. Body orientation and hand loading influenced muscular response to elbow perturbations. Muscle co-contraction at the elbow during known timing suggests a contribution to elbow joint stability that may reduce injury risk caused by sudden elbow loading.     

Crosstalk in surface electromyography of the proximal forearm during gripping tasks.

Electromyographic (EMG) crosstalk was systematically analyzed to evaluate the magnitude of common signal present between electrode pairs around the forearm. Surface EMG was recorded and analyzed from seven electrode pairs placed circumferentially around the proximal forearm in six healthy individuals. The cross-correlation function was used to determine the amount of common signal, which was found to decrease as the distance between electrode pairs increased, but was not significantly altered by forearm posture (pronation, neutral, supination). Overall, approximately 40% common signal was detected between adjacent electrode pairs (3 cm apart), dropping to about 10% at 6 cm spacing and 2.5% at 9 cm. The magnitude of common signal approached 50% between adjacent electrode pairs over the extensor muscles, while over 60% was observed between neighbouring sites on the flexor aspect of the forearm. Although flexor and extensor EMG amplitude was similar, less than 2% common signal was present between flexor and extensor electrode pairs during both pinch and grasp tasks. Maximum grip force production was not affected by forearm rotation for pinch, but reduced 18% from neutral (mid-prone) to pronation during grasp (p=0.01). In spite of differences in grip force, mean muscle activity did not vary between the three forearm postures during maximum pinch or grasp trials. While this study improved our knowledge of crosstalk and electrode spacing issues, further examination of forearm EMG is required to improve understanding of muscle loading, EMG properties and motor control during gripping tasks.     

A new technique for the selective recording of extensor carpi radialis longus and brevis EMG.

Tennis Elbow or Lateral Epicondylalgia is manifested by pain over the region of the lateral epicondyle of the humerus, related to use of the wrist extensor muscles. Extensor carpi radialis longus (ECRL) and brevis (ECRB) have been implicated in the dysfunction associated with Lateral Epicondylalgia. For muscles in the human forearm, particularly those in close proximity, selective recordings are nearly impossible without the use of fine wire, indwelling electrodes. These can be inserted in precise locations and have small recording areas. Standard electromyography texts indicate, however, that the activity of ECRL and ECRB cannot be distinguished, even with intramuscular electrodes. We present a new technique for determining the most appropriate sites at which to insert intramuscular electrodes for selective recordings of ECRB and ECRL. The location of ECRB and ECRL was measured on 10 cadaver specimens, 5 right arms and 5 left arms. The distance from the muscle origin to (1) insertion, (2) largest portion of the muscle belly, (3) most proximal fibres and (4) most distal fibres were measured and expressed relative to forearm length. The mean distance and 95% confidence interval was calculated for each of the four measures. These data indicated a significant separation of the belly of each muscle along the length of the forearm. These relative distances were used to mark electrode insertion points on three volunteers. Fine wire electrodes were used to record the electromyogram in three participants. Each participant was required to perform isometric contractions to produce (1) wrist extension torque, (2) radial deviation torque, (3) elbow flexion torque and (4) finger extension. The electromyographic recordings show clear differentiation of ECRB and ECRL with the relative activation patterns reflecting the underlying anatomical organisation of the two muscles. This technique provides an important objective method that can be used in conjunction with manual muscle testing to provide a means of ensuring accurate intramuscular electromyographic recording from these two muscles.     

Muscle activity during maximal isometric forearm rotation using a power grip

This study aimed to provide quantitative activation data for muscles of the forearm during pronation and supination while using a power grip. Electromyographic data was collected from 15 forearm muscles in 11 subjects while they performed maximal isometric pronating and supinating efforts in nine positions of forearm rotation. Biceps brachii was the only muscle with substantial activation in only one effort direction. It was significantly more active when supinating (µ = 52.1%, SD = 17.5%) than pronating (µ = 5.1%, SD = 4.8%, p < .001). All other muscles showed considerable muscle activity during both pronation and supination. Brachioradialis, flexor carpi radialis, palmaris longus, pronator quadratus and pronator teres were significantly more active when pronating the forearm. Abductor pollicis longus and biceps brachii were significantly more active when supinating. This data highlights the importance of including muscles additional to the primary forearm rotators in a biomechanical analysis of forearm rotation. Doing so will further our understanding of forearm function and lead to the improved treatment of forearm fractures, trauma-induced muscle dysfunction and joint replacements.     

Ia presynaptic inhibition in human wrist extensor muscles: effects of motor task and cutaneous afferent activity.

The task-dependence of the presynaptic inhibition of the muscle spindle primary afferents in human forearm muscles was studied, focusing in particular on the modulation associated with the co-contraction of antagonist muscles and the activation of cutaneous afferents. The changes known to affect the motoneuron proprioceptive assistance during antagonist muscle co-activation in human leg and arm muscles were compared. The evidence available so far that these changes might reflect changes in the presynaptic inhibition of the muscle spindle afferent is briefly reviewed. The possible reasons for changes in presynaptic inhibition during the antagonist muscle co-contraction are discussed. Some new experiments on the wrist extensor muscles are briefly described. The results showed that the changes in the Ia presynaptic inhibition occurring during the co-contraction of the wrist flexor and extensor muscles while the hand cutaneous receptors were being activated (the subject's hand was clenched around a manipulandum) could be mimicked by contracting the wrist extensor muscles alone while applying extraneous stimulation to the hand cutaneous receptors. It is concluded that besides the possible contribution of inputs generated by the co-contraction of antagonist muscles and by supraspinal pathways, cutaneous inputs may play a major role in modulating the proprioceptive assistance during manipulatory movements.     

Fatigue effect on cross-talk in mechanomyography signals of extensor and flexor forearm muscles during maximal voluntary isometric contractions

Objective:This paper presents the analyses of the fatigue effect on the cross-talk in mechanomyography (MMG) signals of extensor and flexor forearm muscles during pre- and post-fatigue maximum voluntary isometric contraction (MVIC).Methods:Twenty male participants performed repetitive submaximal (60% MVIC) grip muscle contractions to induce muscle fatigue and the results were analyzed during the pre- and post-fatigue MVIC. MMG signals were recorded on the extensor digitorum (ED), extensor carpi radialis longus (ECRL), flexor digitorum superficialis (FDS) and flexor carpi radialis (FCR) muscles. The cross-correlation coefficient was used to quantify the cross-talk values in forearm muscle pairs (MP1, MP2, MP3, MP4, MP5 and MP6). In addition, the MMG RMS and MMG MPF were calculated to determine force production and muscle fatigue level, respectively.Results:The fatigue effect significantly increased the cross-talk values in forearm muscle pairs except for MP2 and MP6. While the MMG RMS and MMG MPF significantly decreased (p<0.05) based on the examination of the mean differences from pre- and post-fatigue MVIC.Conclusion:The presented results can be used as a reference for further investigation of cross-talk on the fatigue assessment of extensor and flexor muscles’ mechanic.     

Individual muscle force parameters and fiber operating ranges for elbow flexion-extension and forearm pronation-supination

We have quantified individual muscle force and moment contributions to net joint moments and estimated the operating ranges of the individual muscle fibers over the full range of motion for elbow flexion/extension and forearm pronation/supination. A three dimensional computer graphics model was developed in order to estimate individual muscle contributions in each degree of freedom over the full range of motion generated by 17 muscles crossing the elbow and forearm. Optimal fiber length, tendon slack length, and muscle specific tension values were adjusted within the literature range from cadaver studies such that the net isometric joint moments of the model approximated experimental joint moments within one standard deviation. Analysis of the model revealed that the muscles operate on varying portions of the ascending limb, plateau region, and descending limb of the force-length curve. This model can be used to further understand isometric force and moment contributions of individual muscles to net joint moments of the arm and forearm and can serve as a comprehensive reference for the forces and moments generated by 17 major muscles crossing the elbow and wrist.     

Trunk muscle contributions of to L4–5 joint rotational stiffness following sudden trunk lateral bend perturbations

The purpose of this research was to investigate the contributions of individual muscles to joint rotational stiffness and total joint rotational stiffness about the lumbar spine’s L_4–5 joint prior to, and following, sudden dynamic lateral perturbations to the trunk. Kinematic and surface EMG data were collected while subjects maintained a kneeling posture on a robotic platform, while restrained so that motions caused by the perturbation were transferred to the pelvis, causing motion of the trunk and head. The robotic platform caused sudden inertial trunk lateral perturbations to the right or left, with or without timing and direction knowledge. An EMG-driven model of the lumbar spine was used to calculate the muscle forces and contributions to joint rotational stiffness during the perturbations. Data showed 95% and 106% increases in total joint rotational stiffness_, about the lateral bend and axial twist axes, when subjects had knowledge of the timing of the perturbation. Also, the contralateral muscles exhibited a significantly larger total joint rotational stiffness about the lateral bend axis, and earlier surface EMG responses, than the ipsilateral muscles. The results indicate that, when the timing of the perturbation was unknown, subjects relied more on delayed muscle forces following the perturbation to stiffen the L_4–5 joint.     

Evaluating protocols for normalizing forearm electromyograms during power grip

Many studies use a reference task of an isometric maximum voluntary power grip task in a mid-pronated forearm posture to normalize their forearm electromyographic (EMG) signal amplitude. Currently there are no recommended protocols to do this. In order to provide guidance on the topic, we examined the EMG amplitude of six forearm muscles (three flexors and three extensors) during twenty different maximal voluntary efforts that included various gripping postures, force and moment exertions and compared them to a frequently used normalization task of exerting a maximum grip force, termed the reference task. 16 participants (8 male and 8 female, aged 18–26) were recruited for this study. Overall, maximal muscle activity was produced during the resisted moment tasks. When contrasted with the reference task, the resisted moment tasks produced EMG activity that was up to 2.8 times higher (p < 0.05). Although there was no one task that produced greater EMG values than the reference task for all forearm muscles, the resisted flexor and extensor moment tasks produced similar, if not higher EMG activity than the reference task for the three flexors and three extensor muscles, respectively. This suggests that researchers wishing to normalize forearm EMG activity during power gripping prehensile tasks should use resisted flexor and extensor moment tasks to obtain better estimates of the forearm muscles’ maximum electrical activation magnitudes.     

Contribution of muscle short-range stiffness to initial changes in joint kinetics and kinematics during perturbations to standing balance: A simulation study

Simulating realistic musculoskeletal dynamics is critical to understanding neural control of muscle activity evoked in sensorimotor feedback responses that have inherent neural transmission delays. Thus, the initial mechanical response of muscles to perturbations in the absence of any change in muscle activity determines which corrective neural responses are required to stabilize body posture. Muscle short-range stiffness, a history-dependent property of muscle that causes a rapid and transient rise in muscle force upon stretch, likely affects musculoskeletal dynamics in the initial mechanical response to perturbations. Here we identified the contributions of short-range stiffness to joint torques and angles in the initial mechanical response to support surface translations using dynamic simulation. We developed a dynamic model of muscle short-range stiffness to augment a Hill-type muscle model. Our simulations show that short-range stiffness can provide stability against external perturbations during the neuromechanical response delay. Assuming constant muscle activation during the initial mechanical response, including muscle short-range stiffness was necessary to account for the rapid rise in experimental sagittal plane knee and hip joint torques that occurs simultaneously with very small changes in joint angles and reduced root mean square errors between simulated and experimental torques by 56% and 47%, respectively. Moreover, forward simulations lacking short-range stiffness produced unreasonably large joint angle changes during the initial response. Using muscle models accounting for short-range stiffness along with other aspects of history-dependent muscle dynamics may be important to advance our ability to simulate inherently unstable human movements based on principles of neural control and biomechanics.     

Active trunk stiffness increases with co-contraction.

Trunk dynamics, including stiffness, mass and damping were quantified during trunk extension exertions with and without voluntary recruitment of antagonistic co-contraction. The objective of this study was to empirically evaluate the influence of co-activation on trunk stiffness. Muscle activity associated with voluntary co-contraction has been shown to increase joint stiffness in the ankle and elbow. Although biomechanical models assume co-active recruitment causes increase trunk stiffness it has never been empirically demonstrated. Small trunk displacements invoked by pseudorandom force disturbances during trunk extension exertions were recorded from 17 subjects at two co-contraction conditions (minimal and maximal voluntary co-contraction recruitment). EMG data were recorded from eight trunk muscles as a baseline measure of co-activation. Increased EMG activity confirms that muscle recruitment patterns were different between the two co-contraction conditions. Trunk stiffness was determined from analyses of impulse response functions (IRFs) of trunk dynamics wherein the kinematics were represented as a second-order behavior. Trunk stiffness increased 37.8% (p < 0.004) from minimal to maximal co-activation. Results support the assumption used in published models of spine biomechanics that recruitment of trunk muscle co-contraction increases trunk stiffness thereby supporting conclusions from those models that co-contraction may contribute to spinal stability.     

Targeted gripping reduces shoulder muscle activity and variability

The purpose of this study was to determine if the effect of visually targeted gripping on shoulder muscle activity was maintained with repeated exposures. Eleven healthy males had eight shoulder muscles monitored via surface electromyography while maintaining shoulder elevation at 90° in the scapular plane with and without a 30% grip force. Three non-gripping trials were followed by 15 gripping trials and another 3 non-gripping control trials. Gripping significantly decreased the activity of the anterior deltoid, trapezius, and latissimus dorsi over the exposure of 15 trials. Gripping also reduced variability in all muscles' activity. The changes in shoulder muscle activity are likely in response to forces being transferred through multi-articular muscles spanning from the forearm to the shoulder. Targeted gripping during shoulder elevation resulted in small but significant decreases in muscle activity and reduced variability which supports previous evidence for increased risk of upper extremity disorders in occupational settings.     

Effects of posture,movement and hand load on shoulder muscle activity

The influence of external factors such as arm posture, hand loading and dynamic exertion on shoulder muscle activity is needed to provide insight into the relationship between internal and external loading of the shoulder joint. Surface electromyography was collected from 8 upper extremity muscles on 16 participants who performed isometric and dynamic shoulder exertions in three shoulder planes (flexion, mid-abduction and abduction) covering four shoulder elevation angles (30°, 60°, 90° and 120°). Shoulder exertions were performed under three hand load conditions: no load, holding a 0.5 kg load and 30% grip. It was found that adding a 0.5 kg load to the hand increased shoulder muscle activity by 4% maximum voluntary excitation (MVE), across all postures and velocities. Performing a simultaneous shoulder exertion and hand grip led to posture specific redistribution of shoulder muscle activity that was consistent for both isometric and dynamic exertions. When gripping, anterior and middle deltoid activity decreased by 2% MVE, while posterior deltoid, infraspinatus and trapezius activity increased by 2% MVE and biceps brachii activity increased by 6% MVE. Increased biceps brachii activity with gripping may be an initiating factor for the changes in shoulder muscle activity. The finding that hand gripping altered muscle activation, and thus the internal loading, of the shoulder may play an important role in shoulder injury development and rehabilitation.     

Strength and fatigability of selected muscles in upper limb: assessing muscle imbalance relevant to tennis elbow.

PURPOSE: The aetiology of tennis elbow has remained uncertain for more than a century. To examine muscle imbalance as a possible pathophysiological factor requires a reliable method of assessment. This paper describes the development of such a method and its performance in healthy subjects. We propose a combination of surface and fine-wire EMG of shoulder and forearm muscles and wrist strength measurements as a reliable tool for assessing muscle imbalance relevant to the pathophysiology of tennis elbow. METHODS: Six healthy volunteers participated. EMG data were acquired at 50% maximal voluntary isometric contraction from five forearm muscles during grip and three shoulder muscles during external rotation and abduction, and analysed using normalized median frequency slope as a fatigue index. Wrist extension/flexion strength was measured using a purpose-built dynamometer. RESULTS: Significant negative slope of median frequency was found for all muscles, with good reproducibility, and no significant difference in slope between the different muscles of the shoulder and the wrist. (Amplitude slope showed high variability and was therefore unsuitable for this purpose.) Wrist flexion was 27+/-8% stronger than extension (mean+/-SEM, p=0.006). CONCLUSION: This is a reliable method for measuring muscle fatigue in forearm and shoulder. EMG and wrist strength studies together can be used for assessing and identifying the muscle balance in the wrist-forearm-shoulder chain.     

Changes in joint stability with muscle contraction measured from transmission of mechanical vibration

A non-invasive in vivo technique was developed to evaluate changes in wrist joint stability properties induced by increased co-activation of the forearm muscles in a gripping task. Mechanical vibration at 45, 50 and 55 Hz was applied to the radial head in ten healthy volunteers. Vibrations of the styloid process of the radius and the distal end of the metacarpal bone of the index finger were measured with triaxial accelerometers. Joint stability properties were quantified by the transfer function gain between accelerations on either side of the wrist-joint. Gain was calculated with the muscles at rest and at five force levels ranging from 5% to 25% of maximum grip force (%MF). During contraction the gain was significantly greater than in control trial (0%MF) for all contractions levels at 45 and 50 Hz and a trend for 15%MF and higher at 55 Hz. Group means of contraction force and gain were significantly correlated at 45 (R(2)=0.98) and 50 Hz (R(2)=0.72), but not at 55 Hz (R(2)=0.10). In conclusion, vibration transmission gain may provide a method to evaluate changes in joint stability properties.     

Intra-session and inter-day reliability of forearm surface EMG during varying hand grip forces

Surface electromyography (EMG) is widely used to evaluate forearm muscle function and predict hand grip forces; however, there is a lack of literature on its intra-session and inter-day reliability. The aim of this study was to determine reliability of surface EMG of finger and wrist flexor muscles across varying grip forces. Surface EMG was measured from six forearm flexor muscles of 23 healthy adults. Eleven of these subjects undertook inter-day test–retest. Six repetitions of five randomized isometric grip forces between 0% and 80% of maximum force (MVC) were recorded and normalized to MVC. Intra- and inter-day reliability were calculated through the intraclass correlation coefficient (ICC) and standard error of measurement (SEM).Normalized EMG produced excellent intra-session ICC of 0.90 when repeated measurements were averaged. Intra-session SEM was low at low grip forces, however, corresponding normalized SEM was high (23–45%) due to the small magnitude of EMG signals. This may limit the ability to evaluate finer forearm muscle function and hand grip forces in daily tasks. Combining EMG of functionally related muscles improved intra-session SEM, improving within-subject reliability without taking multiple measurements. Removing and replacing electrodes inter-day produced poor ICC (ICC < 0.50) but did not substantially affect SEM.     

Uncovering patterns of forearm muscle activity using multi-channel mechanomyography

A coordinated activation of distal forearm muscles allows the hand and fingers to be shaped during movement and grasp. However, little is known about how the muscle activation patterns are reflected in multi-channel mechanomyogram (MMG) signals. The purpose of this study is to determine if multi-site MMG signals exhibit distinctive patterns of forearm muscle activity. MMG signals were recorded from forearm muscle sites of nine able-bodied participants during hand movement. By using 14 features selected by a genetic algorithm and classified by a linear discriminant analysis classifier (LDA), we show that MMG patterns are specific and consistent enough to identify 7 ± 1 hand movements with an accuracy of 90 ± 4%. MMG-based movement recognition required a minimum of three recording sites. Further, by classifying five classes of contraction patterns with 98 ± 3% accuracy from MMG signals recorded from the residual limb of an amputee participant, we demonstrate that MMG shows pattern-specificity even in the absence of typical musculature. Multi-site monitoring of the RMS of MMG signals is suggested as a method of estimating the relative contributions of muscles to motor tasks. The patterns in MMG facilitate our understanding of the mechanical activity of muscles during movement.     

Muscle material properties in passive and active stroke-impaired muscle

Stroke survivors routinely experience long-term motor and sensory impairments. In parallel with neurological changes, material properties of muscles in the impaired limbs, such as muscle stiffness, may also change progressively. However, these stiffness measures are routinely derived from individual joint stiffness, representing whole muscle groups. Here, we use shear wave (SW) ultrasound elastography to measure SW velocity, as a surrogate measure of stiffness, to quantify material properties in individual muscles. Accordingly, the purpose of this study was to compare muscle material properties of the bicep brachii in stroke survivors and in age-matched control subjects by measuring SW velocity at rest and different voluntary activation levels. Our main findings show that at rest, the SW velocity was on average 41% greater in the paretic muscle compared the contralateral non-paretic muscle. The mean passive SW velocity across all subjects were 2.34 ± 0.41 m/s for the non-paretic side, 3.30 ± 1.20 m/s for the paretic side, and 2.24 ± 0.18 for controls. SW velocity was significantly different in muscles of the paretic and non-paretic side (p < 0.001), but not between muscles of the non-paretic and controls (p = 0.47). As voluntary activation increased, SW velocity increased non-linearly, with an average power fit of r² = 0.83 ± 0.09 for the non-paretic side, r² = 0.61 ± 0.24 for the paretic side, and r² = 0.24 ± 0.15 for the healthy age-matched controls. In active muscle (10, 25, 50, 75, 100% maximum voluntary contraction), there was no significant difference in SW velocity between the non-paretic, paretic, and control muscles.These findings suggest that stroke-impaired muscles have potentially altered muscle material properties, specifically stiffness, and that passive and active stiffness may contribute differently to total muscle stiffness.